Bulletin | British Museum (Natural History ) Entomology Series _ VOLUME 59 NUMBER 2 27 SEPTEMBER 1990 The Bulletin of the British Museum (Natural History ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. The Entomology Series is produced under the editorship of the Keeper of Entomology: Dr L. A. Mound Publications Manager (Entomology): Mr W. G. Tremewan Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. A volume contains about 288 pages, made up of two numbers: published Spring and Autumn. Subscriptions may be placed for one or more of the series on an Annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Sales Department, Natural History Museum Publications, British Museum (Natural History), Cromwell Road, London SW7 5BD Telephone: 071-938-9386 Telex: 929437 NH PUBS G Fax: 071-938-8709 World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1990 ISBN 0 565 06040 6 Entomology Series ISSN 0524-6431 Vol 59, No. 2, pp. 117 — 286 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 27 September 1990 Typeset by Computer Typesetting Services, Redhill, Surrey Printed in Great Britain by Henry Ling Ltd, Dorchester, Dorset Bull. Br. Mus. nat. Hist. (Ent.) 59(2):117—286 Issued 27 September 1990 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae) S. J. BROOKS & P. C. BARNARD Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD CONTENTS MALEK OCU CEL OM oy-7- 00 pa tavs ay at =. ay stacs Secvaraie ec ofa isve apes se avavnvtce tebe AVI nage, Well eel eeran tray alas eyo. tG17/ FAISLOLICAl ME VLC Ww reqcatisrstaee ie aero eietnnn: aes araereranctch eee seen ery ACRE Saletan ees 118 Discussionioftaxonomic characters). 2-1 «.-- ee. o-Riee ele nila stale sivee ie «cis te ee 119 Materials ANGE MOUS «cuclscisttiteneieicusts sa.c! ssetarnee cee eee tks ay cel iver ckietsy lesa aes 123 A summary of the generic classification of the Chrysopidae ....................--. 125 he tribal/classification/ofthe family ios... . « cchinyete ayers sieyerspoteter ele» wlaletever siecle Gieinye an 126 Key to the subfamilies of the Chrysopidae.................. 00000. e ee eee eee eee 132 Subfamily Apochrysinae Handlirsch ................ 000 cece cece eee eee eee ee 132 Ney tothe seneraiof Apochrysinde saa. ..... <1 elisa slersl> «le eieetel aie slo sreceieiens eretarels 134 Subfamily Chrysopinae Schneider iss sacs = «15:08 aleve = cn cle = ysllaieloe ei sianad ocleee- Heels 149 Key to the genera and subgenera of the Chrysopinae...................-..22-4. 149 irbeAnkylopteryeinilINAVaStiaer sel ae + wile si Se ewe aati aletel niepeleloa’s saatiaae 155 inibesBelonoptery oiniiNavas erasers -.< 3s sreris « ee ele lauert wea dtu chetoeiaie Anti. Sua ah: 163 Minibe Chrysopini Scuneldenesae ae <--- «; «ics aratatesietevere © cyusselaeye Se avatars te gel Ae 184 MinibedbeucochrysimiA Gams iacrees ayslo's «APs ale re aie tis loose ava recauevarereutt are Siepwiaes @ wf 243 Subfamily Nothochrysinae NAVas (ice. eieciscciececiche carne sin wisieiejeue mpere)e epeuere ete 252 Key to the genera of the Nothochrysinae................. 0000s cece eee eee eee 254 Checklistofextant species of Chrysopidae 2.062062 e cee eee eels nent eens nae 264 PNCKTO WIE CP EIMEMES ee cio Fasten See foy-paie = ene ereterduspeeieno eeee tiene ceon's pac tinea oe 280 PRE LETEM CES iene rerscg dearest NF Ae Need fee 6y 2 Fapeasielere ms eereiss ceed qs Ober o's ¢ sie eee 8 5 281 INGLES cance APR Ske aad, Crete Page Odeo ect ce UCR Meat EPP Ceca NCE or OR agen retin iar 286 SYNOPSIS. The systematic relationships of the world genera of green lacewings, family Chrysopidae, are discussed. Each of the 75 genera and 11 subgenera currently recognised is redescribed, and a cladistic analysis of the main groups of genera is made. The resultant higher classification recognises the three subfamilies Apochrysinae, Chrysopinae and Nothochrysinae, with the Chrysopinae subdivided into four tribes, the Ankylopterygini, Belonopterygini, Chrysopini and Leucochrysini. Keys are provided to the subfamilies and to the genera within each subfamily and a check list of the 1200 currently recognised species and subspecies is given. Three new genera are described and two are given replacement names because of homonymy; there are 18 new generic synonyms and ten genera have new status. In addition, 233 new generic combinations of species are given. INTRODUCTION The Chrysopidae is one of the largest and economically most important families of the Neuroptera. The family includes over 1200 cur- rently recognized species and subspecies that are divided between 86 genera and subgenera. The larvae of all species and the adults of a few genera are predaceous and most feed on aphids, coccids and other soft-bodied insects which they en- counter on foliage. For this reason some species have been reared and successfully used for the biological control of agricultural pests (New, 1975). The classification of the family is confused and many of the species and genera, especially those described by L. Navas during the first 30 years of this century, are difficult to interpret from the original descriptions. Many of these genera were based on single species (frequently on single 118 specimens) and have not been examined or redescribed by modern systematists. The present study is the first to attempt a generic revision of the entire Chrysopidae on a world-wide basis. However, the two smallest subfamilies, the Apochrysinae and Nothochrysinae which include no more than 3% of the des- cribed species between them, were revised by Kimmins (1952a) and Adams (1967) respec- tively. There have also been several faunistic accounts (Zimmerman, 1957: Hawaii; Adams, 1959: Micronesia; Kuwayama, 1962: Japan; Tjeder, 1966: South Africa; H6élzel, 1967: Asia; Aspock, Aspock & H6lzel, 1980; Europe; Holzel, 1980: Arabia; New, 1980: Australia), although the majority of these were concerned with the identification of species and did not critically review or examine the natural relationships of the genera. The need for a world revision of the family was recognized over 50 years ago by Killington (1937). This need is just as apparent now and has been discussed by New (1984). During the present study representatives of nearly all the described chrysopid genera with examples of almost 50% of the described species (in addition to many un- described species) have been critically examined. As a result of this work 75 genera and 11 sub- genera are now recognized as valid and these have been redescribed and figured. Three new genera are described and two have been given replace- ment names as a result of homonymy. There are 18 new generic synonyms and ten genera have new status. In addition, 233 new generic combinations of species are listed. Although this review will not be the last word on the classification of this large and important group, it should provide a framework for future taxonomic work on the family. The main prob- lems yet to be resolved would seem to be the exact relationships of all the genera within each tribe and subfamily. HISTORICAL REVIEW Suprageneric classification The Chrysopidae were first recognized as a supra- generic taxon by Schneider (1851) who proposed Chrysopina as a subgroup of the Hemerobiidae. Two genera were included, Chrysopa Leach which comprised 53 species, and Apochrysa Schneider of which only one species was known. Handlirsch (1908) regarded Apochrysa and re- S. J. BROOKS & P. C. BARNARD lated genera as sufficiently distinct from the rest of the Chrysopidae to merit a family of their own. This view was supported by Esben-Petersen (1918) who based his reasons on venational characters, but Tillyard (1926) regarded the group as a subfamily of the Chrysopidae. The Apochrysinae were subsequently revised by Kimmins (19525), who also based his study on the wing venation. He concluded that the group were highly derived chrysopids and should be treated as a subfamily of the Chrysopidae. Most later workers have followed Kimmins but some (e.g. Adams, 1978a) have argued that the group should have no more than tribal status, being related to the Neotropical chrysopine genus Leucochrysa McLachlan. Navas (1910a; 1913a; 1914a) divided the Chrysopidae into several tribes many of which were distinguished by relatively trivial venational characters and have been ignored by subsequent authors. However, one of these tribes, the Nothochrysini, has been shown to be valid and was revised by Adams (1967). He established the Nothochrysinae as a subfamily of the Chrysopidae and demonstrated that it was probably the most primitive group of extant chrysopids. Another tribe proposed by Navas (1910a) was the Ankylopterygini and this was one of three tribes into which H6lzel (1970) divided the Old World Chrysopinae. The Chrysopinae is the largest of the three subfamilies of Chrysopidae, comprising more than 97% of the described species, and yet H6lzel’s was the first critical attempt to assess the tribal classification of the subfamily. H6lzel included the majority of genera in the Chrysopini but also recognized the Ankylopterygini, which was further characterized by Brooks (1983), and the Italochrysini, which was also redescribed by Brooks (1984). Adams (1978a) listed two additional New World chryso- pine tribes, the Belonopterygini which was origin- ally erected by Navas (1913a) and a new tribe, the Leucochrysini. The tribes were not described but the constituent genera were given. Later, Adams & Penny (1986) synonymized the Italochrysini with the Belonopterygini. The Chrysopini and Leucochrysini have, until now, never been fully described. Generic classification Until about twenty years ago most of the generic classification of the Chrysopidae had relied on characters of the wing venation. This resulted in the erection of many genera containing few species with unusual venational characteristics but THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 119 left nearly 1000 species with more conservative venation in the unwieldy and phylogenetically meaningless genus Chrysopa Leach. However, additional characters were available, particularly those of the genitalia. The morphology of neurop- teran genitalia was elucidated by Stitz (1909a; 1909b) and their possible significance in chrysopid taxonomy was recognized by Smith (1932) and Killington (1937), but it was not until Tjeder’s (1966) important work on the South African lacewings that their true value in chrysopid systematics was realized. Tjeder noted that there is a great variety in the male genitalic components of Chrysopa s.1. Furthermore, he was able to demonstrate that the presence or absence of certain structures in the male genitalia united groups of allied species which he considered as subgenera of Chrysopa. However, he was unable to discern many signifi- cant external characters or differences in the female genitalia which would also distinguish most of these subgenera. This implied that the male genitalic characters had limited application, since there would be difficulty in assigning females which were not associated with males to the correct subgenus. Some subsequent authors (New, 1980; Barnard, 1984) have argued that the separation of taxa based almost entirely on the composition of the male genitalia can be simplistic and that these characters do not necessarily delimit natural groups. However, Principi (1977) demonstrated that useful external and female genitalic characters do exist which help to define these genera and Séméria (1977) has observed biological differences. Hdlzel (1970) and Aspéck et al. (1980) have followed Tjeder’s lead and raised many of his subgenera to generic rank. Most of the recent work on the genera of the New World fauna has been carried out by Adams (1975; 1977; 1982a; 1982b). He has also shown the usefulness of the male genitalia for delimiting genera and has linked many external and female genitalic characters with them. More than any other contemporary chrysopid worker, Adams has tried to demonstrate phylogenetic relation- ships between the genera. This has necessitated a close scrutiny of the characters used to define genera in order to assess whether or not they are derived. In addition to using adult morphological characters to investigate chrysopid taxonomy, some authors (Tauber, 1974; 1975; Tsukaguchi, 1978; Gepp, 1983) have examined the larvae while others (Séméria, 1983) have looked at the karyo- types. Although these methods can help to elucidate particular problems when used in con- junction with morphological evidence, only a few Palaearctic and North American chrysopid species have so far been examined using these techniques. It is therefore difficult to draw any general conclusions from these data until biologi- cal information from species occupying a wider zoogeographical range is available. DISCUSSION OF TAXONOMIC CHARACTERS Almost all the chrysopid genera that were des- cribed before Tjeder’s (1966) work were based on wing venational characters. Since 1966 authors have adopted a more integrated approach, using as many characters as possible and placing an emphasis on male genitalic characters. However, almost invariably authors have not taken a phylo- genetic view when describing genera and usually have not attempted to assess whether the charac- ters used to define a taxon are apomorphic or plesiomorphic. This has led to the erection of many polyphyletic genera based on convergent characters and also to the unnecessary splitting of taxa when undue weight is placed on a particular derived character, leaving sister groups poorly defined without unique apomorphies. In this study we have thoroughly examined as many specimens as possible in order to assess the maximum number of characters from all parts of the insect. Like previous authors, we have found that the male genitalia provide the most useful and phylogenetically meaningful characters. However, there are many external and female genitalic characters of great significance and we have succeeded in finding external (non-genitalic) autapomorphies for most genera. This has en- abled the construction of a key to the genera with which most chrysopid specimens can be correctly assigned without recourse to dissecting the genitalia. Wings (Fig. 1) Barnard (1984) has discussed the taxonomically useful imaginal chrysopid characters and advised caution when interpreting the venation. During this study it has become apparent that many genera which have been distinguished on vena- tional characters are invalid when other charac- ters are taken into consideration. The most frequent errors have arisen when undue emphasis has been placed on the number of rows of gradate crossveins. It seems likely that multiple series of gradates have arisen, or been lost, independently 120 S. J. BROOKS & P. C. BARNARD Fig. 1 Chrysopa perla, right fore wing. bsx = basal subcostal crossvein, C = costa, c; and c, = Ist and 2nd cubital cells, Cu, = 2nd cubital vein, dec = distal cubital cell, i.g. = inner gradate crossveins, im = intramedian cell, M = media, m, and m, = Ist and 2nd median cells, o.g. = outer gradate crossveins, pmx = posterior median crossveins, Psc = pseudocubitus, Psm = pseudomedia, R = radius, Rs = radial sector, rx = radial crossveins, Sc = subcosta, St = pterostigma, 1A —3A = 1st—3rd anal veins. on many occasions in the Chrysopidae and that the number of gradate series has little significance at the generic level. Indeed, the number of gradate rows sometimes varies even between individuals of the same species (Tsukaguchi, 1985). The shape of the intramedian cell in the fore wing has also been used on several occasions to distinguish certain genera. Again, this has proved to have little significance when other characters are taken into account and it is apparent that the shape of cell im, and sometimes even its presence or absence, can vary within genera or species. However, a long median fork is probably the plesiomorphic condition and a short fork is apomorphic within the family. Therefore, a broad, quadrangular intramedian cell is plesio- morphic and a narrow, short, ovate im is apomor- phic. The absence of im is the most derived condition. However, there are many wing venational characters which are helpful in determining phylo- genies. In the Nothochrysinae the absence of a tympanal organ is plesiomorphic. The tympanal organ is small, spherical and bulging in the Chrysopinae and is in its most derived condition in the Apochrysinae where it is elongate but not bulging. The basal subcostal crossvein is posi- tioned relatively distally in the Nothochrysinae. This is probably the primitive condition in the family Chrysopidae; in the more advanced sub- families it occupies a more basal position. In the Chrysopinae the subcostal crossvein is situated closer to the base of the fore wing than in the Nothochrysinae, but within the subfamily Chrysopinae it is most distal in the Belonopterygini and most basal in the Ankylopterygini. The most derived condition occurs in the Apochrysinae in which the crossvein is completely lacking. The relative lengths of cells c, and cy are also sig- nificant. In most Chrysopidae c, is shorter than c> and this is seen at its most extreme in the Nothochrysinae where c, may be less than half the length of c>. In the Apochrysinae and most of the Chrysopinae cells c, and cy are about equal in length but in most of the belonopterygine genera cell c,; is considerably longer than cz and this probably represents the most derived condition. The plesiomorphic condition for wing setation which occurs widely throughout the Chrysopidae is for short, inclined setae. However, the apo- morphic condition occurs in the Apochrysinae, Ankylopterygini and a few Chrysopini and Leucochrysini in which the setae are long and erect. Description of plesiomorphic chrysopid wing Fore wing unmarked; length : breadth 2.5-3.0: 1; costal setae short, inclined; costal area narrow; Sc long; Sc and R widely separated; basal Sc crossvein > 0.75 mm; im broad, quadrangular; radial crossveins straight; Rs sinuate; gradates in two parallel series; inner and outer gradates equal in number; basal inner gradate meeting Psm; inner gradates not extending basally; veins not crassate in 6; c, shorter than cj; c> narrow; dcc open at margin. Head The head also provides many useful characters and the shape of the mandibles and apical segment THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 121 of the palps are of considerable significance when considering generic relationships. Plesiomorphic- ally the mandibles are broad, asymmetrical with a basal tooth on the left mandible, and this con- dition occurs in all three subfamilies. However, in some Chrysopini genera there is a basal tooth on both mandibles and this would appear to be an apomorphic condition. The most derived con- dition occurs in some species of Chrysopodes Navas and the Ankylopterygini in which the mandibles are narrow, scythe-like and lack basal teeth. The morphology of the apical segment of the palps varies considerably in the Chrysopidae but is consistent within certain groups. The plesiomorphic condition probably occurs in the Nothochrysinae and the Belonopterygini in which the palps are broad and rounded apically. In the Apochrysinae and Chrysopini the apical segment is compressed dorsoventrally and the apex is asymmetrical since it tapers more strongly on the inner margin than outer. The palps of the Apochrysinae are more truncate than those of the Chrysopinae. The most derived condition occurs in the Ankylopterygini in which the apical seg- ment is rounded but narrows abruptly subapically and is considerably elongated. This condition is approached in some Chrysopini genera such as Eremochrysa Banks. The relative size of the eyes and width of the head can also suggest possible relationships. The Nothochrysinae have the smallest eyes amongst the Chrysopidae and this apparently represents the apomorphic condition since it rarely occurs in the rest of the family. Small eyes also occur in the Chrysopinae but they are restricted to a few genera such as Chrysopa Leach and Chrysopiella Banks. The largest eyes occur in the Apochrysinae. Similarly, the relative width of the flagellar segments and the number of rings of setae can provide useful clues. The plesiomorphic condition for the width of the flagellar segments in the Chrysopidae is probably 2-3 times as long as broad. In the Belonopterygini the trend has been for the segments to become broader and the most derived condition occurs in Belonopteryx Gerstaecker in which the segments are twice as broad as long. In other taxa the segments are elongated and in Ankylopteryx Brauer they are over 4 times as long as broad. It seems likely that there has been a reduction in the number of setal rings on the flagellar segments during the evolu- tion of the Chrysopidae. In the Nothochrysinae there are six rings, with five in the Apochrysinae, but this has reduced to four in the Chrysopinae. In some Belonopterygini the apical ring is pre- sent but sparse, suggesting the most derived condition. Description of plesiomorphic chrysopid head Head with black markings; palps broad, short, rounded apically; galea broad with large apical papilla; mandibles broad, asymmetrical with basal tooth on left mandible; labrum emarginate; vertex flattened; toruli small; eyes large; scape about as long as broad; pedicel hardly constricted medially; flagellar segments 2-3 times as long as broad; flagellar setae in six rings; antenna shorter than fore wing. Thorax There are few useful characters on the thorax and legs although the degree of dilation at the base of the claw is sometimes important. A simple claw probably represents the plesiomorphic condition, but the basal dilation appears to have arisen in the Chrysopidae on many separate occasions. A broad pronotum is probably plesiomorphic in the family, and occurs in the Nothochrysinae and Belonopterygini. The most derived condition occurs in the Apochrysinae and certain Chrysopini such as Plesiochrysa Adams and Ceratochrysa Tjeder in which the pronotum is elongate. Abdomen The morphology of the apical abdominal seg- ments and genitalia provide the most numerous characters of phylogenetic significance. The shape of the ectoprocts, the degree of fusion between them and tergite 9, and the presence or absence of a dorsal suture between the ectoprocts in both males and females have proved to be very useful in establishing generic relationships. Of similar importance is whether or not sternites 8 and 9 are fused in males, fusion between the sternites prob- ably being the derived condition. Description of plesiomorphic chrysopid abdomen Abdominal setae short, sparse; microtholi absent; trichobothria few; ectoprocts with slight dorso- apical invagination; ectoprocts short and rounded apically; ectoprocts fused dorsally; ectoprocts entirely separate from tergite 9; sternites 8 and 9 separate; sternite 9 short, broad. Male genitalia (Figs 2, 3) The morphology and complement of the male genitalia are of fundamental significance in assess- ing the higher classification of the Chrysopidae. The plesiomorphic male genitalia simply consist of a broadly arcuate gonarcus with entoprocessus and a short, hooked arcessus surrounded by a small gonosaccus with a few short gonosetae; this S. J. BROOKS & P. C. BARNARD sg Figs 2-6 2, 3, diagrammatic representation of apex of abdomen and genitalia of male chrysopid; (2) lateral view, (3) caudal view. ap = apodemes, arc = arcessus, cc = callus cerci, ent = entoprocessus, gc = gonarcus, ger = gonocristae, gps = gonapsis, gsc = gonosaccus, gst = gonosetae, hp = hypandrium internum, mt = microtholi, mu = median plate, pm = parameres, ti = tignum, 7s = 7th sternite, 7t — 8t = 7th — 8th abdominal tergites, 8+9s = 8th + 9th fused sternites, 9t+ep = 9th tergite plus ectoproct. 4, typical female chrysopid spermatheca, lateral view; 5, 6, apical abdominal segments of typical female chrysopid; (5) ventral view, (6) lateral view. cc = callus cerci, d = duct, lg = lateral gonapophyses, pg = praegenitale, sg = subgenitale, v = vela, vi = ventral impression, 7s = 7th sternite, 7t and 8t = 7th and 8th abdominal tergites, 9t+ep = 9th abdominal tergite plus ectoproct. (After Barnard, 1984.) occurs largely in the Nothochrysinae and the Apochrysinae. In the Chrysopinae the genitalia are more complex. In the most primitive tribe, the Belonopterygini, the gonarcus is transverse and hardly arcuate, often with a pair of lateral horns (gonocornua), and the entoprocessus have been lost. The arcessus is short and is flanked apically by membranous lateral lobes. In some of the more derived genera a pair of parameres is present and the gonocornua are absent. The male genitalia of the Leucochrysini are similar to those of the Belonopterygini, although parameres are never present, and the shared apomorphies suggest that the two tribes may be closely related. In the Chrysopini, the gonarcus is acutely arched and in many genera entoprocessus are present. These structures can be distinguished from gonocornua because they articulate freely with the gonarcus whereas gonocornua are fused with the gonarcus. Where the entoprocessus are absent in the Chrysopini they have probably been secondarily lost. Many Chrysopinae possess a tignum and/or a gonapsis and this may be plesiomorphic within the tribe Chrysopini. This seems to be the most plausible explanation; otherwise one would have to postulate the repeated independent THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 123 development of these structures in genera which are otherwise clearly related to genera lacking a tignum or gonapsis. This seems to be highly unlikely especially considering the close similarity in the shape of the tignum throughout the tribe. Parameres do not occur in the Chrysopini, but the gonapsis, which may have evolved more than once in the family (Adams, 1982a), can resemble Parameres in some genera such as Chrysocerca Weele and Himalochrysa H6lzel; however, para- meres consist of a paired structure articulated basally whereas the gonapsis is a single entity. In some Chrysopini the arcessus is detached from the gonarcus and is termed the pseudopenis; this is the apomorphic condition. Similarly, a large and strongly setose gonosaccus is the derived con- dition for the family. Gonocristae have arisen in several genera of Chrysopini. They are situated on a membranous flap at the apex of sternite 8+9 and probably represent modified microsetae. Female genitalia (Figs 4-6) The female genitalia offer fewer characters than the male genitalia. The basic pattern repeated throughout the family is a bilobed subgenitale and a pillbox-shaped spermatheca with long, coiled duct, conical ventral impression and curved vela. However, many of the Belonopterygini have a lobate praegenitale situated basad of the sub- genitale. This may be derived from a basal elongation of the subgenitale which has become detached; an intermediate condition is probably present in Nacarina Navas and is perhaps sug- gested in Leucochrysa McLachlan in which many species have a ventrally curving basal extension of the subgenitale. The ventral impression is very broad and hardly tapers apically in the Notho- chrysinae. In the Chrysopinae it gradually tapers but is exceptionally deep in the Belonopterygini. In the Apochrysinae the ventral impression is deep and abruptly tapers apically. These character states probably represent a progression from the plesiomorphic nothochrysine condition to the most derived in the Apochrysinae. Description of plesiomorphic female genitalia in the Chrysopidae Apex of sternite 7 straight; gonopophyses narrow; praegenitale absent; subgenitale bilobed apically, not extended basally; spermatheca broad, pillbox- shaped; ventral impression broad, not tapering apically; vela and duct moderately long. Biological characters By examining the gut contents of dissected specimens we have been able to ascertain that most adult chrysopids do not feed on insects, unlike the Hemerobiidae. The gut contents of most species are amorphous and these are prob- ably nectar feeders, representing the plesio- morphic condition within the family. Several of the Nothochrysinae feed on pollen and the grains are easily recognizable in the guts. In some genera, especially Chrysopa and Atlantochrysa, the guts are crammed with insect remains. This probably represents the most derived condition and the species have secondarily returned to a carnivorous diet. Stridulation has arisen on at least three separate occasions in the Chrysopinae (Brooks, 1987) and courtship displays, which have led to morpho- logical modifications, have appeared in Meleoma (Tauber, 1969) and Mallada (Duelli & Johnson, 1981). The larval morphology offers several useful characters. Larvae in all three chrysopid sub- families carry debris and, although this is un- doubtedly a derived character for the Chrysopidae, probably represents the plesiomorphic condition within the family. The most plesiomorphic con- dition occurs in the Nothochrysinae in which a few large particles of debris are carried, the abdomen is hardly humped, and the thoracic tubercles are only slightly enlarged and bear a few short setae. The most apomorphic of the debris-carriers occur in genera such as Chrysotropia Navas and Ceraeochrysa Adams in which the larva is almost entirely covered by the debris packet, the abdomen is strongly humped, and the thoracic tubercles are greatly elongated and bear numer- ous, long, hooked setae. In some Chrysopini the larvae do not carry debris and are very active. This is probably apomorphic and the larvae have secondarily lost the debris-carrying habit. In these genera the abdomen is not humped and the thoracic tubercles are reduced in size, the most extreme state being present in Nineta in which the tubercles are virtually absent. In the Belonopterygini it is likely that the larvae of some genera, like /talochrysa, live parasitically in ant colonies and this also represents an apomorphic modification. MATERIALS AND METHODS This work has been based primarily on the extensive chrysopid collections held at the British Museum (Natural History) (abbreviated to BMNH hereafter) although additional specimens have been kindly loaned from many institutions around 124 the world, and visits were made by one of us (SJB) to the Muséum National d’Histoire Naturelle, Paris; the Musée Royal de l'Afrique Centrale, Tervuren; and the Institut Royal des Sciences Naturelles de Belgique, Brussels. The initial stages of the project involved a complete exam- ination of all the specimens held in the BMNH. First, the material was identified and sorted into appropriate genera and species which involved dissecting the male genitalia of large numbers of specimens. After initial sorting, the material was re-examined genus by genus. At least one specimen from each zoogeographical region in which the genus occurred was thoroughly studied and notes were made of morphological characters from all over the body. In this way a complete picture of the generic characters was built up. In addition, reliably identified examples of the type species of almost all the available generic names were examined. In many cases it was possible to examine the type species of each genus. We have examined nearly 50% of the described species of Chrysopidae together with many undescribed species. Some of the characters used in this study were assessed by making comparative measurements and are explained below. All measurements were taken with a calibrated graticule. Head width : eye width The head width was taken as the distance between the eyes across the front of the vertex. The eye width was taken as a mean of the distance between the inner and outer edge of both eyes from the dorsal side (Fig. 16). Flagellar segment width : length A flagellar segment from the middle of the antenna was measured because it was found that the segments were broadest at the base of the antenna and narrower towards the apex. Wing length : breadth The wing length was measured from the base to the extreme apex. The breadth was measured across the broadest part of the wing, perpendicu- lar to the costa. Basal subcostal crossvein (Fig. 1) This was taken as the distance in millimetres along the radius from the crossvein between m, and m, to the first subcostal crossvein. Cy : C2 (Fig. 1) The lengths of c, and cp were measured along the posterior margin of the cells. Genitalia The terminology for the genitalia largely follows Tjeder (1966) although it has been necessary to S. J. BROOKS & P. C. BARNARD modify this slightly. To avoid confusion we have redefined the terms used in the present work below. Males (Figs 2, 3). ARCESsUS. Attached medially to the gonarcus, usually long, narrow and tapering to a hook. ENTOPROCESSUS (plural). Short, paired structures attached laterally to the gonarcus but articulat- ing with the gonarcus and not fused like the gonocornua. Gonapsis. Single plate-like structure closely associated with sternite 8+9. Gonarcus. A large arcuate structure situated medially. GONOCORNUA. A pair of medio-lateral horns fused to the gonarcus. GONOCRISTAE. Patch of coarse, enlarged micro- setae on membrane at apex of sternite 8+9. GOnosaAccus. Membranous sac surrounding the gonarcus complex, often bearing long gonosetae. MEDIAN PLATE [= mediuncus of Tjeder]. Like the arcessus this structure is also attached medially to the gonarcus but lies above the arcessus. It is usually transverse and bears horns. PARAMERES. Like the gonapsis these are also associated with sternite 8+9 but are formed from paired cylindrical structures weakly fused and articulating basally. PSEUDOPENIS. A long, narrow, arcuate structure, lying ventrad of the gonarcus, loosely associated with the gonosaccus. TIGNUM. A transverse, often arcuate, structure situated dorsad of the gonarcus. The acumen is the small median projection. Females (Figs 4-6). CRUMENA. Small pocket situated medially and near base of subgenitale. PRAEGENITALE. Ventral lobed structure situated subapically of sternite 7. Not fused with subgenitale. SPERMATHECA. Large, heavily sclerotized, doughnut-shaped structure situated within 7th abdominal segment. SPERMATHECAL DUCT. Long, narrow, often highly coiled tube emanating posteriorly from sperma- theca. Towards apex duct is fringed with minute glandular ducts. SUBGENITALE. Bilobed structure, sometimes with short median projection, positioned at apex of long membranous tube posteriorly to sternite 7. VELA. Tubular structure extending dorsally from the centre of the spermatheca. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) VENTRAL IMPRESSION. Ventral median invagina- tion in spermatheca. A SUMMARY OF THE GENERIC CLASSIFICATION OF THE CHRYSOPIDAE A full synonymic check list of the world Chrys- opidae is given at the end of this paper, but in order to facilitate an understanding of the classi- fication of the family there follows a simple list of the valid subfamilies, tribes, genera and sub- genera. Names within in each group are in alpha- betical order. Subfamily APOCHRYSINAE Handlirsch Anapochrysa Kimmins Apochrysa Schneider Claverina Navas Domenechus Navas Joguina Navas Lainius Navas Loyola Navas Nacaura Navas Nobilinus Navas Nothancyla Navas Oligochrysa Esben-Petersen Synthochrysa Needham Subfamily CHRYSOPINAE Schneider Tribe ANKYLOPTERYGINI Navas Ankylopteryx Brauer Subgenus Ankylopteryx Subgenus Sencera Navas Parankylopteryx Tjeder Retipenna Brooks Semachrysa Brooks Signochrysa gen. n. Tribe BELONOPTERYGINI Navas Abachrysa Banks Belonopteryx Gerstaecker Calochrysa Banks Chrysacanthia Lacroix Chrysaloysia Navas Dysochrysa Tjeder Evanochrysa gen. n. Italochrysa Principi Nacarina Navas Nesochrysa Navas Nodochrysa Banks Oyochrysa Brooks Stigmachrysa Navas Turnerochrysa Kimmins Tribe CHRYSOPINI Schneider Anomalochrysa McLachlan Apertochrysa Tjeder Atlantochrysa Holzel Austrochrysa Esben-Petersen Borniochrysa nom. n. Brinckochrysa Tjeder Ceraeochrysa Adams Ceratochrysa Tjeder Chrysemosa nom. n. Chrysocerca Weele Chrysopa Leach Chrysoperla Steinmann Chrysopidia Navas Subgenus Anachrysa Holzel Subgenus Chrysopidia Subgenus Chrysotropia Navas Chrysopodes Navas Subgenus Chrysopodes Subgenus Neosuarius Adams & Penny Cunctochrysa Holzel Eremochrysa Banks Subgenus Chrysopiella Banks Subgenus Eremochrysa Glenochrysa Esben-Petersen Himalochrysa Holzel Kostka Navas Mallada Navas Meleoma Fitch Nineta Navas Parachrysopiella gen. n. Peyerimhoffina Lacroix Plesiochrysa Adams Rexa Navas Suarius Navas Tumeochrysa Needham Ungla Navas Yumachrysa Banks Tribe LEUCOCHRYSINI Adams Berchmansus Navas Cacarulla Navas Gonzaga Navas Leucochrysa McLachlan Subgenus Leucochrysa Subgenus Nodita Navas Neula Navas Nuvol Navas Vieira Navas 125 126 Subfamily NOTHOCHRYSINAE Navas Dictyochrysa Esben-Petersen Hypochrysa Hagen Kimochrysa Tjeder Nothochrysa McLachlan Pamochrysa Tjeder Pimachrysa Adams Triplochrysa Kimmins THE TRIBAL CLASSIFICATION OF THE FAMILY As described in the Historical Review (p. 118) most recent authors have divided the family Chrysopidae into three subfamilies, the Apoch- rysinae, Chrysopinae and Nothochrysinae, with a varying number of ill-defined tribes within the Chrysopinae. In order to attempt a clarification of the relationships between these various groups we studied the distribution of 120 characters among as many genera and subgenera as possible: only Neula Navas and Nuvol Navas were omitted from the analysis owing to lack of suitable material. The list of characters is given below, with the pre- sumed apomorphic (+) and plesiomorphic (—) states indicated. The character polarity is based on outgroup comparisons with other Neuropteran families, principally the closely _ related Hemerobiidae. 1 Palps truncate (+); tapered (—) 2 Palps elongate (+); rounded (—) 3 Labrum straight (+); indented (—) 4 Mandibles narrow (+); broad (—) 5 Both mandibles toothed (+); only left mandible toothed (—) 6 Galea narrow (+); broad, swollen apically (—) 7 Vertex raised (+); flat or domed (—) 8 Toruli small (+); large (—) 9 Head width : eye width large, >2.8:1 (+); small ts) 10 Scape elongate (+); squared (—) 11 Flagellar segments narrow (+); broad, length : breadth < 2: 1 (—) 12 Flagellar segments with more than 4 rings of setae (+); 4 rings (—) 13 Antenna longer than fore wing (+); shorter or same length (—) 14 Head with red markings (+); black markings or unmarked (—) 15 Prothorax broad (+); narrow or elongate (—) 16 Thorax marked with red or black (+); unmarked (—) S. J. BROOKS & P. C. BARNARD 17 Prothoracic setae long (+); short (—) 18 Tarsal claws dilated (+); simple (—) 19 Setae on legs long (+); short (—) 20 Legs marked (+); unmarked (—) 21 Fore wing narrow, length : breadth >3.0 (+); broad (—) 22 Costal setae long (+); short (—) 23 Subcosta short (+); long (—) 24 Pterostigma marked (+); unmarked (—) 25 Subcosta and radius close (+); widely separated Ge) 26 Basal subcostal crossvein proximal (+); distal, >1.4 (-) 27 Fore wing gradate crossveins in 3 series (+); 2 series (—) 28 Hind wings gradates in 3 series (+); 2 series (—) 29 Gradates divergent (+); parallel (—) 30 Gradates unequal in number (+); equal (—) 31 Basal inner gradate not meeting Psm (+); meeting Psm (—) 32 Cell im ovate (+); rectangular (—) 33 Rs straight (+); sinuate (—) 34 Veins crassate in male (+); not crassate (—) 35 Basal inner gradate series extended basally (+); not basally extended (—) 36 Basal costal area narrow (+); broad (—) 37 Costal cells broad, >0.4 (+); narrow (—) 38 Cell c,; longer than cy (+); shorter or same length (—) 39 Cu, forked (+); unforked (—) 40 1A forked (+); unforked (—) 41 Cell c, broad (+); narrow (—) 42 Fore wing extensively marked (+); unmarked 43 Basal costal spot present (+); absent (—) 44 Radial crossveins sinuate (+); straight (—) 45 Cell dec closed (+); open (—) 46 Basal post-marginal crossvein forked (+); unforked (—) 47 Hind wing narrow (+); broad (—) 48 Male abdominal setae long (+); short (—) 49 Abdominal setae dense (+); sparse (—) 50 Microtholi present (+); absent (—) 51 Callus cerci rounded (+); ovate (—) 52 Trichobothria numerous (+); few (—) 53 Ectoproct dorsal invagination deep (+); shallow (—) 54 Ectoprocts extended apically (+); rounded (—) 55 Ectoprocts fused dorsally (+); not fused or with suture (—) 56 Sternites 8 and 9 fused (+); not fused (—) 57 Sternites 8 and 9 long (+); short (—) 58 Male ectoproct and tergite fused (+); not fused GC [59 & 60: characters not used in final analysis] 61 Tignum present (+); absent (—) 62 Gonapsis present (+); absent (—) THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 127 63 Median plate present (+); absent (—) 64 Entoprocessus present (+); absent (—) 65 Parameres present (+); absent (—) 66 Gonarcus transverse (+); arcuate (—) 67 Gonarcus with gonocornua (+); gonocornua absent (—) 68 Pseudopenis present (+); absent (—) 69 Additional ventral process present (+); absent 70 Arcessus long and narrow (+); broad and short (—) 71 Apical hook of arcessus strong (+); weak (—) 72 Gonosaccus long (+); short (—) 73 Gonosetae present (+); absent (—) 74 Gonosetae in lateral clumps (+); evenly dis- persed (—) 75 Gonosetae numerous (+); sparse (—) 76 Gonocristae present (+); absent (—) 77 Apex of sternite 7 in female indented (+); straight (—) 78 Praegenitale present (+); absent (—) 79 Crumena present (+); absent (—) 80 Subgenitale trilobed (+); bilobed (—) 81 Spermatheca narrow (+); broad (—) 82 Ventral impression of spermatheca deep (+); shallow (—) 83 Vela long (+); short (—) 84 Spermathecal duct long (+); short (—) 85 Adult insectivorous (+); not insectivorous (—) 86 Stridulatory structure present (+); absent (—) 87 Cell mz short (+); long (—) 88 Sternites enlarged (+); small (—) 89 Subgenitale extended basally (+); short (—) 90 Papilla on galea large (+); small (—) 91 Basal Sc crossvein present (+); absent (—) 92 Cell im present (+); absent (—) 93 Thorax with median yellow stripe (+); stripe absent (—) 94 Cell im narrow (+); broad (—) 95 Arcessus with dorsal striations (+); striations absent (—) 96 Tympanal organ present (+); absent (—) 97 Tympanal organ swollen (+); narrow (—) 98 Psc and Psm close (+); widely separated (—) 99 Psm meeting outer gradates (+); meeting inner gradates (—) 100 Rs arises close to wing base (+); arises distally =) 101 Ectoproct and tergite fused in female (+); not fused (—) 102 Arcessus trifurcate apically (+); simple (—) 103 Ectproct not hinged, no basal extension (+); hinged (—) 104 Gonosetae long (+); short (—) 105 Arcessus with dorsal microsetae (+); no microsetae (—) 106 Male with additional short setae on sclerites (+); setae absent (—) 107 Larvae naked (+); larvae debri-carriers (—) 108 Ventral apodeme projecting dorsally (+); not projecting (—) 109 Tergite 9 broad (+); narrow (—) 110 Male ectoprocts lobed (+); simple (—) 111 Flagellar segments with 3 (or reduced 4) rings of setae (+); more than 3 rings (—) 112 Costal crossveins sinuous at base (+); straight a 113 Prothoracic setae absent (+); present (—) 114 Cell im short (+); long (—) 115 Spermatheca with lateral striations (+); no striations (—) 116 Eighth abdominal spiracle on membrane (+); spiracle on tergite (—) 117 Ventral impression of spermatheca narrow (+); broad (—) 118 Psm upturned at outer gradates (+); down- turned (—) 119 Jugal lobe reduced (+); well developed (—) 120 Larva associated with ants (+); not ant- associated (—) The distribution of these character states throughout the 80 genera and subgenera exam- ined is given in Table 1. It should be noted that where a name represents both a genus and its nominate subgenus (e.g. Ankylopteryx) then it is the subgenus which is referred to: this was to see whether the presumed relationships of subgenera were confirmed by the analysis. The data were analysed using G. S. Farris’s phylogenetic program ’Hennig-86’. The full results are discussed in detail in a separate paper by I. J. Kitching & S. J. Brooks (in prep.), but the resultant relationships of the subfamilies and tribes are shown as a cladogram in Fig. 7. The cladogram confirms that the three subfamilies Apochrysinae, Chrysopinae and Nothochrysinae are monophyletic; the Nothochrysinae is appar- ently the most plesiomorphic group, which cor- responds to many authors’ description of them as the most primitive subfamily. Within the Chrysopinae the tribe Belonopterygini is the most primitive, but the relationships of the other three tribes are impossible to determine with any certainty, largely because it appears that the tribe Chrysopini is not monophyletic. However, our aim is to provide a practical framework for the classification of the family: the final classification simply recognises the three subfamilies, with four tribes within the Chrysopinae, and we are con- fident that this basic classification will serve the needs of taxonomists working on this group for some time to come. 128 Abachrysa Anachrysa Ankylopteryx Anapochrysa Anomalochrysa Apertochrysa Apochrysa Atlantochrysa Austrochrysa Belonopteryx Berchmansus Borniochrysa Brinckochrysa Cacarulla Calochrysa Ceraeochrysa Ceratochrysa Chrysacanthia Chrysaloysia Chrysemosa Chrysocerca Chrysopa Chrysoperla Chrysopidia Chrysopiella Chrysopodes Chrysotropia Claverina Cunctochrysa Dictyochrysa Domenechus Dysochrysa Eremochrysa Evanochrysa Glenochrysa Gonzaga Himalochrysa Hypochrysa Italochrysa Joguina Kimochrysa Kostka Lainius Leucochrysa Loyola Mallada Meleoma Nacarina Nacaura Neosuarius Nesochrysa Nineta Nobilinus Nodita Nodochrysa Nothancyla Nothochrysa Oligochrysa Oyochrysa Pamochrysa Parachrysopiella Parankylopteryx Peyerimhoffina Pimachrysa Plesiochrysa Retipenna Rexa Semachrysa Sencera Signochrysa Stigmachrysa Suarius Synthochrysa Triplochrysa Turnerochrysa Tumeochrysa Ungla Vieira Yumachrysa S. J. BROOKS & P. C. BARNARD DHNMTNOF DNS KNMTOOF DONO —AMTNOFDHNOK—NMTNOFONOK— NM t+—---+-—-+4+-+77? 2am ttt +--+ 4—4+--- $+ ——t—-4+ 2444 ----- -- ++=+772 2?--222?--+----- thee ta tt tt 2-2 ttt — $4 4—---- +4+----24+-+27 —--2724+4+4+4+-4+----- Fata t ttm tt tat tan tt ttt Ht ttt $2? ----- 24+4--4+-4+--4+----4+----— + -- - — — $$ 2 $$ Ht ttt 2? 2-272 44--277 2 -4+4444—--4+-4+-4+--4+-- 4+ --- 4 -- -t+4---- = +—-+----++-+77 2-27 244-—4---- + 24-7 4+----4+----- +4—--+4+--+------— ++—-+—-—-+4+4+4+-4+77 Table 1 The distribution of characters among the 80 genera and subgenera examined in the cladistic analysis, showing presence (+) or absence (—) ofa character, or state unknown (?) due to lack of information. See text for list of characters 1-120. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 129 Re eh IO ESO Ole ICSF ICT S00 OIG USP LESOO ON oe CUI tN KOIS COONS Se en oe eS = - ceree ce nN GOOSCOOCOR REA ARRRE NO DD ODODDOOR AARAAARDAAS Seer reer errr eereee Abachrysa —-+4+4+4+----- 4-22 -44--—4+44+4--4+27-24+4---4+4-4+-4+----- 2-— 2222277272942 Anachrysa ol a ce a a ata 244—4—44—+—-—- + 72--7-777227222742 Anieghae el fa ?------+4+------ +4 —4—-4+4—4+—4+—-—+—-—-27-277797727272777427 Anap ochr SSSES Qa tt tt 4—t 2 2-77 772772777974+2 ttt —— 4+ 4-4-4 —--— 2 --— $- 9 2727777777 4- Anone liar a pes dae +—+4—-—4+—--4¢-F4+44—--—-4+444+4+4+444-4-4+------- 222729729977 4— ochrysa 242-2 2-44 —-- 777272277277 -- 27-79 -—-- 2-4 - 4+ 4+4---- 2=-227727992997974+27 Pei eackryea mtn ttt ta tt tt +4+--4+44-4------- DD PND 21D = Austrochrysa ‘pa at Ge. Ea eee 222227292742 Belonopteryx ee tie eg a 2 A ARDS SS a EY Gad i ea NTE Tae Tae ee eae Berchmansus Borniochrysa Brinckochrysa eee Calochr Ceraeoc rysa Ceratochr Chrysacan ares ---+4+4+----+-+---+4+-4+- LY Pe area ok aa) 4D —+-+----- 2—4——— 99999742 Chrysaloysia ENR UA Uh iy Dah pi Nat 222 —4-9 hrysemosa SS eae mee fi ++++------ ++-+-+4+- 5 eee sane 27222243 Chrysocerca Chrysopa Chrysoperla Chrysopidia ? Chrysopiella Pa tht $4—F— Ft —$——— $-F + 2 -- $-- 279977 Chrysopodes Te ae alee DN a a Chrysotropia oo -=— tanta tata ta ta ttn tt ttt tat ita tenn Claverina | =. 222222227222222222222222-2-22-—-224—+44-2722229222-- 22277 Cunctochrysa 7 Sine See ee ee SE 2222224- Dictyochrysa aan te—t—-t+--+t4---- t+ ---- = +2 +4--+-2--------- ee 292229—— DomenechuS = -=---------- +4+------- $a $$ 0 a 4 $$ 2 --- +? +4444? Dysochrysa hf een en Dh pa amb aa a — 0 ee a fa a pe) em Pap a —— 997999949 Eremochrysa +4+—------- $+—-+4+—------- +------ +o tt —t+—+4—-4-—--—— t— ae) 29 Pwanochrysa © 9 -—-—-=—— FH —-—4+444—---4t-4+-4+4+4—--4+-- 2 44---4¢4-4-4----- 2-4---4+4+772774+2? Glenochrysa -+++----- +—t$4—4+4---— 4 -- + -- --— $- 4 4-4 - $4 - 4-4 4-4 -- 2 ------- 2272242 Gonzaga matt t—4+-4+--- ta—tttt 2 ——4—4+44—-—-4+4+4+—-4+-4+4+-4+-— 2? --4+---- 2777747 Himalochrysa 2 4+--4+—-- t+ 4+4+4+------ —---+----- $24+444-44—-4-=—4+-—--- 2? ——------ 222942 Hypochrysa ---+--++----- 22?-2---++------ +2 44—----?-4----?--+4+------- 299 9 —— Italochrysa aS NG Sc re eg ee Se ey ner ee Bey iy i]t ING SERE AE sta 1 1g ad SN eh 27 ---2-4+-4+44+7-2--+7------ 2222742 Kimochrysa ---+--------- PD se ee re SER ea eee 2999-9 Kostka SSS SSS +--+-+----- $$ t$4——4—4+ 2 $4 ——— + 4-4 -— + 444-42 ------- tee Og ike Pee 8) een = 2222799272 —— P= P= 9 tt $$ 2-7 2-4 2 -— = 2222942 Leucochrysa Soy See B= PSE eae oe oh A ee eer 29792 4= re ola GL aa di it a Sa a ia Bd $$ 2-2 = 72H FF4=— 72772797 - 9 -- = 77277942 fede Gea 2 Ta --277272?------ ++4+4+4+44+—-4+-4------------ +72274— hoteone 24+ —4+—----+4+—--4+-— Eire el ey) PSG ENR lat rece —+-2?--4+--+4+------- 292724+— Nacarina matte tH $$ +e4——t+—— 2 +44 —— $+ — 4-4 $2 7720 Naeadteas . ————— eS $— == $a a aa a 444 -- $--- -- P+ttt+— Neosuarius oo <--------- +--+---2?----+4++4+------ +442 —4+4—-4+-4+-2-4-------- = 222242 Nesochrysa 4 $$ = $= 2 0 4 $$ at at ta 9 = 999947 Nineta manta tt tt +-++----- +44+—-4-44-4+—---4+4-44--4----77774- Nobilinus ---<---------- 2?------ See Se ae a So 22299242 NoG@utda +> Soe ba nlm aoe Ae eg oe Se SO) ee +$22274— Nodochrysa an $a $a a Pa tt tt tt tt ttt 2 7 4 - - - - 799747 Nothancyla ---+----- +=—--2?2?---=---=-- PySoy sade) SS aotearoa SaaS 2224+? Nothochrysa ---+-------- +4------- Ae SS hao ee oe ee eS RaQ Oligochrysa —2?---2-+-4+4+4+--+744+2------ 2-+4+24? Oyochrysa +4+4+---+4+-4+-4--2--2-+---2--+4+274+7 Pamochrysa ++++----? Parachrysopiella ? foe SS Be ear Paranky peer yx -------------- 24+4+—4—4+4—-4+=-+—-—+——7? eee fina -+-+-----++-++4+------+-+4------ Le ee at Aa Pimachr pS ey Plesioc faa —+-+4+44+-----+-------- DPS ae ee et Retipenna ?------------ Dtt—+—t4+—4—-4--4 4-24 +7 Rexa = ff mn +4+-+------ tt t$$4———— + 2+ 444+—4+4+—-4+—-4+4—--- = 2 -- $+ - 7 + Semachrysa 2---4+-4+=------- +4+—-4+244-4+-4+--4+--2-------- +4247 Sencera 24+--2?-+4-4+-+--+--------- tenn Ot SuPnOChEySa 0 Saat ee +4 —4—44— 444 —4 $= — 9 — Stismachzysa Suarius Synthochrysa Triplochrysa Turnerochrysa Tumeochrysa Ungla Vieira Yumachrysa Table 1 (contd) 130 S. J. BROOKS & P. C. BARNARD Chrysopini Ankylopterygini Leucochrysini , Belonopterygini APOCHRYSINAE NOTHOCHRYSINAE CHRYSOPINAE Fig. 7 Cladogram of subfamilies and tribes of Chrysopidae. See text for explanation. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 131 10 Figs 8-10 Base of fore wing of three chrysopid genera showing variation in shape of intramedian cell (im). Pe yi. f. Figs 11-15 11, chrysopid claw with basal dilation; 12, claw without basal dilation; 13-15, apical segment of maxillary palps showing variation between the tribes of Chrysopinae. 2 &D Phy 21 19 Figs 16-21 16, chrysopid head showing head width : eye width measurements; 17, 18, variation in morphology of chrysopid mandibles; 19, prothoracic markings in Austrochrysa; 20, position of stridulatory structure in certain genera; 21, apex of 2 7th sternite in Brinckochrysa, ventral view. 132 amen Fig. 22 Apex of d abdomen of Chrysemosa, dorsal view, showing suture between ectoprocts. Key to the Subfamilies of Chrysopidae 1 In fore wing Psm continuous with inner row of gradates (Fig. 545); tympanal organ absent; jugal lobe well developed (Fig. 527) ... NOTHOCHRYSINAE — In fore wing Psm meets outer gradates (Fig. 1); tympanal organ present (but sometimes small); jugal lobe reduced or absent (Fig. 1)............. 2 2 In fore wing basal subcostal crossvein present; cell im present (Fig. 1); flagellar setae in four (or less) TUNGS | svccnin cteiesempsgeeeera ea onan ais CHRYSOPINAE — In fore wing basal subcostal crossvein absent; cell im absent (Fig. 27); flagellar setae in five PUD OS) 5 aries thefeieiy agence eos eeu APOCHRYSINAE Subfamily APOCHRYSINAE Handlirsch Apochrysidae Handlirsch, 1908: genus: Apochrysa Schneider. Apochrysinae Tillyard, 1926: 318. Apochrysini Adams, 1978a: 211. 1251. Type DISTRIBUTION. Afrotropical, eastern Palaearctic, Oriental, Australian, Neotropical. DiAGcnosis. Adult. Large lacewings, fore wing 18-34 mm. Head width : eye width = 1.6-2.3: 1, head relatively narrow; palps broad, short, trun- cate (Fig. 83); toruli small; vertex flattened; scape with long, coarse frontal setae; pedicel deeply constricted medially; antenna longer than fore wing; flagellar segments narrow, 3 times as long as broad; setae arranged in five rings. Pronotum narrow. Legs long; claws with basal dilation. Fore wing broad (length : breadth = 2.1-2.6: 1), often densely reticulated; marked with black pustulate spots in discal area; costal area broad; costal setae long, erect; Sc short; basal Sc crossvein absent; im absent; tympanal organ narrow, elongate; Rs straight, arising near to wing base; c, shorter than C2; Psm and Psc very close, upturned apically; posterior margin broad; jugal lobe small, 1A forked or unforked. Hind wing longer and much narrower than fore wing (length : breadth = S. J. BROOKS & P. C. BARNARD 2.9-3.3 : 1). Abdomen long, elongate, swollen apically; trichobothria 32-63; d: sternite 8+9 usually fused, short, broad; ectoprocts rounded basally, not hinged; microtholi usually absent; apodemes short, weakly sclerotized; ¢ : sternite 7 straight apically; gonapophyses narrow, bearing coarse setae. GENITALIA 6. Weakly sclerotized; tignum and gonapsis, median plate absent; entoprocessus absent; gonarcus narrow, arcuate; arcessus short, triangular with apical hook; pseudopenis, gono- cristae and spinellae absent; gonosetae usually absent or very short. GENITALIA @. Praegenitale absent; subgenitale bilobed; spermatheca long, very broad with lateral striations; ventral impression deep, often abruptly tapered apically; duct long narrow; vela short, straight. REMARKS. At present the Apochrysinae includes 12 genera. From the Afrotropics are Anapochrysa Kimmins and Apochrysa Schneider, from the eastern Palaearctic is Nacaura Navas, from the Oriental region and Australia are Oligochrysa Esben-Petersen, Nobilinus Navas, Nothancyla Navas, Synthochrysa Needham and Joguina Navas, and from the Neotropics are Loyola Navas, Claverina Navas, Domenechus Navas and Lainius Navas. Adams (1967) suggested that the Apochrysinae may be specialized derivatives of the Leucochrysini and should have no more than tribal status (Adams, 1978a). He suggested that the zig-zag of MP>, which forms the intramedian cell in Gonzaga, could easily have become straightened in the apochrysines. However, this does not seem to be a particularly convincing argument since MP) is zig-zagged in all chrysopids and so does not necessarily indicate a close relationship between the Apochrysinae and Leucochrysini. Indeed the Apochrysinae lack several apomorphic characters that are present in the Chrysopinae which suggests that the Apochrysinae may have arisen before the advent of the Chrysopinae. In the Apochrysinae the setae on the flagellar segments are arranged in five concentric rings, 1A is often unforked and the male genitalia are simple. There are several synapomorphies exhibited in the Apochrysinae which suggest that the group is monophyletic and distinct from the rest of the Chrysopinae, fully justifying its retention as a subfamily. These include the deeply constricted pedicel; the elong- ate tympanal organ; lack of basal subcostal cross- vein; the close proximity of Psm and Psc; the long, narrow hind wing; the lateral striations on the spermatheca. Within the subfamily the male genitalia are very THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 133 A SN Se NOTE Cy iw, CY Figs 23-26 23, 24, base of fore wing in (23) Plesiochrysa, (24) Chrysopa; 25, 26, front of head of (25) Semachrysa, (26) Plesiochrysa. conservative and offer few generic characters. However, the wing venation is diverse and it is on these characters that the genera are based. The most primitive genera are distributed in the Old World. In the Australian and African genera the wing venation is open but becomes more reticu- lated in the Oriental genera. The Neotropical genera appear to be the most derived and have densely reticulated venation. The genera of the Apochrysinae are apparently closely related to each other since they all share a large number of characters, both externally and in the genitalia. They seem to form a much more cohesive group than any other suprageneric taxon in the family. Although the genitalia differ in detail between most of the genera, it is really only in Apochrysa that the male genitalia differ signifi- cantly from the usual Apochrysinae pattern. It is therefore in details of the wing venation that the genera can best be distinguished. To an extent this is unsatisfactory since it has been shown that genera in the rest of the family can only con- vincingly be distinguished by the male genitalia and that venational characters alone are not sufficient to delimit the genera. One could there- fore argue that many of these monophyletic genera of Apochrysinae should be synonymized, but we have retained them in this work in the 134 absence of strong evidence either way: when more is known of the biology and larvae of this group such decisions may become easier to make. Key to the Genera of Apochrysinae This key is based on that of Kimmins (1952b) with the addition of Nothancyla Navas. 1 Fore wing with quadrangular cell im (Fig. 70) NOTHANCYLA Navas — Hore wing lacking celle. ee l e e 2 2 Only twoseries of gradatesin both wings ........ 3 — More than two series of gradates, at least in fore wing, or basal half of wing with dense reticulation of CROSSVGINS = eas oneness onsaka sand eee es 5) 3 Basal half of fore wing with dense irregular reticu- lation\(Bigs59) ic seek eee: NACAURA Navas — Basal half of both wings with no more than three rows of cellsbetween RandPsm............... 4 4 Costal area of fore wing almost equally broad from near base to apex; inner gradate series sinuous, abruptly bent twice at about middle of wing (Fig. dis) Vee Cea OLIGOCHRYSA Esben-Petersen — Costal area of fore wing tapering towards apex; inner gradate series bent forwards at about two- thirds from base of wing (Fig. 27) ANAPOCHRYSA Kimmins 5 Radial area of fore wing consisting of single row of cells (sometimes with an occasional divided cell) (BigtGD) Pace cee cen oreo eon meets meee 6 — Radial area of fore wing with at least the basal half consisting of two or more rows of cells (Fig. 35) . . .8 6 Radial area of fore wing with crossveins almost to apex; venation dense (Fig. 65) NOBILINUS Navas — Radial area of fore wing with distinct space free of crossveins behind pterostigma................. 7 7 Outer gradates in both wings more or less parallel to wing margin (Fig. 82) SYNTHOCHRYSA Needham — Outer gradates in both wings bent inwards towards costal margin near apex of wing (Fig. 32) APOCHRYSA Schneider 8 Costal area of fore wing with virtually all cells woverhnreeral( ETS SS) caooocacgruccdwagerssucces 9 — Costal area of fore wing consisting of two or more rows of cells, or densely reticulated (Fig. 44) ....11 9 Fore wing with three to seven elevated pustules, often arranged in an arc from midway along Psm to apex of wing (Fig. 55) .......... LOYOLA Navas — Fore wing with not more than two elevated pustules visa aves Sera ane me eee a aaa Se ae 10 S. J. BROOKS & P. C. BARNARD 10 Fore wing with 1A forked, 2A and 3A simple (Fig. 35); hind wing with small elevated pustule CLA VERINA Navas — Forewing with 1A and 2A forked, 3A simple; no elevated pustule in hind wing (Fig. 39) DOMENECHUS Navas 11 Fore wing with Rs straight, distinct; an accessory row of crossveins present in marginal area in apical half of both wings (Fig. 44) ...... JOGUINA Navas — Fore wing with Rs irregular, not distinct from general reticulation of veins; no accessory row of crossveins in marginal area in apical half of either Wing(Bigs4 9) oie. siecle LAINIUS Navas Genus ANAPOCHRYSA Kimmins Anapochrysa Kimmins, 1952b: 932. Type species: Anapochrysa africana Kimmins, by monotypy. DISTRIBUTION. East Africa, Madagascar. Two species have been described in the genus. DIAGNosIs. Adults. Large lacewings, fore wing (Fig. 27) 18-20 mm. Head marked with red spot between base of scape and eye; palps truncate, brown stripe on scape; labrum indented; man- dibles broad, asymmetrical with basal tooth on left mandible; galea narrow with median constric- tion but widely expanded apically; apical papilla very small; vertex flattened; head width : eye width = 1.8-1.9 : 1; scape elongate; pedicel very narrow medially; antenna longer than fore wing; flagellar segments 3 times as long as broad; setae arranged in five rings. Pronotum marked with red lateral stripe; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.6 : 1), pointed apically; membrane with several pustules which are marked with dark shading; costal setae long, erect; stigma short; Sc short; Sc and R quite close; basal Sc crossvein absent; im absent; tympanal organ narrow, elongate; Rs straight, arising near base of wing; gradates in two parallel series; inner gradates greatly extended basally, not meeting Psm; veins not crassate in 6; costal area broad; c, shorter than c); Psm and Pcu very close, con- verging apically, upturned apically; posterior margin broad, marginal crossveins forked towards base of wing; 1A not forked. Abdomen elongate (Figs 28, 29), swollen apically; unmarked; setae long, sparse; callus cerci rounded; trichbothria 32-34; ectoprocts slightly invaginate apico-dorsally, fused dorsally, fused with tergite 9; atria small; 3: microtholi present; sternite 8+9 fused, short; apodemes weakly sclerotized; @: sternite 7 straight apically; gonapophyses long, narrow. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 135 28 Figs 27-31 Anapochrysa africana. 27, fore wing (from Kimmins); 28, apex of d abdomen; 29, apex of 2 abdomen; 30, ¢ genitalia, dorsal; 31, 2 spermatheca, lateral. GENITALIA 6 (Fig. 30). Tignum, gonapsis and median plate absent; entoprocessus absent; gonarcus narrow, arcuate; arcessus short, tri- angular with short apical hook; pseudopenis, gonosaccus, gonosetae, gonocristae and spinellae absent. GENITALIA @ (Fig. 31). Praegenitale absent; subgenitale bilobed; spermatheca broad, long with annulations; ventral impression deep, broad, abruptly tapered apically; vela short, straight, tapered; duct very long, narrow, highly coiled. REMARKS. Anapochrysa possesses few apomor- phic apochrysine characters and the lack of multi- plicity in the gradates is notable. However, the posterior marginal crossveins are forked at the base of the fore wing and the costa narrows towards the wing apex, and these are probably apomorphic characters. BIoLoGy. Unknown. Genus APOCHRYSA Schneider Apochrysa Schneider, 1851: 38. Type species: Hemerobius leptalea Rambur, by monotypy. DISTRIBUTION. Southern Africa. The genus is monotypic. DiaGnosis. Adult. Large lacewings, fore wing (Fig. 32) 21-22 mm. Head marked with red stripe e Were SY between eyes and vertex, scape with red lateral stripe; palps tapered; labrum indented; clypeus with long setae; vertex flattened; head width : eye width = 2.2-2.3 : 1; scape elongate with long setae; pedicel constricted medially; antenna as long as fore wing; flagellar segments 3 times as long as broad; setae arranged in five rings. Pronotum elongate; marked with black baso- lateral spot; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae very long, pale; claws with slight basal dilation. Fore wing broad (length : breadth = 2.52.6: 1), pointed apically; with pustules shaded black; costal setae long, erect; Sc short; Sc and R widely separated; basal Sc crossvein absent; im absent; tympanal organ narrow, elongate; Rs straight, arises near wing base; gradates in four irregular series; inner gradates greatly extended basally; veins slightly crassate in d; costal area broad; c; shorter than cz; Psm and Pcu very close, parallel, upturned apically; posterior margin broad; 1A not forked. Hind wing much narrower than fore wing (length : breadth = 3.2 : 1). Abdomen (Fig. 33) elongate, swollen apically; marked with dorsal red/brown spot on tergite 1; setae short and dense on tergites, long and dense on sternites; callus cerci rounded; trichobothria 62; ectoprocts with slight dorso-apical invagination, fused dorsally, with deep broad suture between ectoproct and tergite 9; ectoprocts positioned ventrally; atria 136 S.J. BROOKS & P. C. BARNARD 72S Figs 32-34 Apochrysa leptalea. 32, fore wing (from Kimmins); 33, apex of d abdomen, lateral; 34, ¢ genitalia, ventral. small; ¢: microtholi absent; sternite 8+9 fused, short broad, indented apically; apodemes weakly sclerotized, very short. GENITALIA 6 (Fig. 34). Tignum, gonapsis, median plate absent; gonosaccus short; gonosetae few, very short in lateral clump; gonocristae and spinellae absent. GENITALIA 2. Unknown. REMARKS. The wing venation of Apochrysa has two apomorphies which help to distinguish the genus. The gradates are arranged in three series and the posterior marginal crossveins are forked towards the base of the wing. The male genitalia are particularly significant, however. In the rest of the subfamily the male genitalia have the same basic pattern with an arcuate gonarcus and tri- angular arcessus. In Apochrysa the gonarcus or arcessus are totally different. Indeed, the genitalia are so distinct that homologous structures are difficult to interpret. Therefore it would appear that although externally Apochrysa resembles the rest of the subfamily, it is in fact quite distantly related to the other genera. BIoLoGy. Unknown. There were no insect re- mains in the guts of any of the adults examined during this study. Genus CLA VERINA Navas Claverina Navas, 1913a: 164. Type species: Apochrysa beata Walker, by monotypy. DISTRIBUTION. Brazil (Amazonia). The genus is monotypic. D1AGnosis. Adult. Large lacewings, fore wing (Fig. 35) 24.5 mm; ground colour green. Head marked with broad red stripe in front of vertex; palps tapered apically; vertex flat, steeply raised anteriorly; head width : eye width = 1.6: 1; toruli small; scape slightly elongate; antenna about 1.5 times length of fore wing; flagellar segments about 3 times as long as broad; setae arranged in five rings. Pronotum unmarked; dorsal setae long, pale; meso— and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing marked with pustules, veins in pustules shaded dark brown; wing broad (length : breadth = 2.2 : 1); rounded apically; costal setae long, erect; stigma very short, marked with dark brown spot; Sc short; Sc and R widely separated; basal Sc crossvein absent; im absent; Rs sinuate; radial cells doubled; gradates in 5—6 poorly defined rows; inner gradates greatly extended basally, not meeting Psm; costal area broad; c; THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 137 TM 37 nates Ds we osceet GER ey ANN AN Figs 35-38 Claverina beata. 35, fore wing (from Kimmins); 36, apex of 2 abdomen, lateral; 37, 2 subgenitale, ventral; 38, spermatheca, lateral. shorter than c; c, narrow, squared apically; posterior margin very broad; 1A forked. Hind wing 24.5 mm, much narrower than fore wing (length : breadth = 3.1 : 1). Abdomen (Fig. 36) long, unmarked; setae long, dense but sparser on sternites than tergites; ectoprocts with slight dorso-apical invagination, fused dorsally with fine suture between ectoprocts and tergite 9; tricho- bothria 24; sternite 7 straight apically. GENITALIA 3d. Unknown. GENITALIA 2 (Figs 37, 38). Praegenitale absent; spermatheca large rounded, lateral striations absent; vela prominent; duct very long, coiled; ventral impression deep, tapering abruptly apically. REMARKS. Claverina can be distinguished from other apochrysine genera by the following apo- morphies: the gradates are arranged in 5-6 poorly defined rows, the radial area is subdivided and 1A is forked. BIoLoGy. Unknown. The gut contents of adults examined during this study did not include insect remains. Genus DOMENECHUS Navas Domenechus Navas, 1913b: 298. Type species: Domenechus sigillatus Navas, by monotypy. DISTRIBUTION. Brazil, central America. Only two species of this genus have been described. Dracnosis. Adult. Large lacewings, fore wing (Fig. 39) 28-29 mm; ground colour green. Head with red stripe on frons and front of vertex; palps tapered apically; labrum invaginate; vertex flat; head width : eye width = 1.4-1.5 : 1; antenna considerably longer than fore wing; flagellar segments about 3 times as long as broad; setae 138 41 S.J. BROOKS & P. C. BARNARD Figs 39-43. Domenechus mirifica. 39, fore wing (from Kimmins); 40, apex of ¢ abdomen, lateral; 41, apex of ? abdomen, lateral; 42, d genitalia, dorsal; 43, 2 spermatheca, lateral. arranged in five rings. Pronotum unmarked; slightly elongate; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with short basal dilation. Fore wing marked with two black pustules in discal area; costal area narrow at base; costal setae long, slightly inclined; stigma marked with brown spot; Sc short; Sc and R widely separated; radial area subdivided; gradates arranged in six rows; gradates greatly extended basally; c, and cz sub- equal in length; cubital cells very short; 1A and2A forked; posterior marginal area subdivided by crossveins. Abdomen (Figs 40, 41) unmarked; setae long, quite dense; callus cerci rounded; trichobothria 36-58; ectoprocts with deep, broad dorso-apical invagination, fused dorsally; 3d: additional short setae present on all sclerites; microtholi absent; suture present between ster- nites 8 and 9; deep suture present between ectoproct and tergite 9; 2: short suture between ectoprocts and tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 42). Tignum, gonapsis, median plate and entoprocessus absent; gonarcus narrow, broadly arcuate; arcessus short, triangu- lar, tapering to short apical hook; pseudopenis absent; gonosaccus short; gonosetae few, very short, positioned in small lateral clump at base of arcessus; gonocristae absent. GENITALIA @ (Fig. 43). Praegenitale absent; subgenitale bilobed apically; spermatheca broad with lateral striations; ventral impression narrow, deep; vela short; duct long, narrow, highly coiled. REMARKS. Species of Domenechus can be recog- nized by the numerous rows of gradates, the subdivided radial area, 1A and 2A forked and the partially divided posterior marginal area in the fore wing. BioLoGy. Unknown. Insect remains were not present in the guts of any of the adults examined during this study. Genus JOGUINA Navas Joguina Navas, 1912: 98. Type species: Apochrysa nicobarica Brauer, by original designation. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 139 ie ES ry WWD SRS — ee ee ee 47 ———— = h Mh Figs 44-48 Joguina nicobarica. 44, fore wing (from Kimmins); 45, apex of d abdomen, lateral; 46, apex of @ abdomen, lateral; 47, d genitalia, dorsal; 48, 2 spermatheca, lateral. DISTRIBUTION. Assam, Burma, Malaysia, Borneo. The genus includes three described species and there is a further undescribed species present in the BMNH collections. DIAGNosIs. Adult. Large lacewings, fore wing (Fig. 44) 26-27 mm. Head marked with brown stripe on frons, vertex scape red; palps broad, truncate; toruli small; vertex flat with anterior fringe of long setae; head width : eye width = 1.7 : 1; scape broad with long setae; pedicel constricted medially; antenna 1.5 times length of fore wing; flagellar segments about 3 times as long as broad; setae arranged in five rings. Pronotum unmarked; elongate; dorsal setae long, pale; mesonotum marked with broad red anterior stripe; metanotum unmarked. Fore wing broad (length : breadth = 2.1—2.3 : 1), rounded apically; marked with dark shading on pustules; costal setae long, erect; stigma unmarked; Sc short; Sc and R widely separated; basal Sc crossvein absent; numerous Sc crossveins in apical half of wing; im absent; tympanal organ very narrow, elongate; Rs straight, arising close to wing base; wings highly reticulated, gradates indistinct; veins slightly crassate in 6; costal area broad; c, and c) indistinct; Psm and Pcu very close, parallel, curving apically; posterior margin broad; 1A deeply forked; 2A forked. Hind wing much narrower than fore wing (length : breadth = 3.1: 1). Abdomen (Figs 45, 46) elongate, swollen apically; marked with red/brown dorsal band at 140 S. J. BROOKS & P. C. BARNARD , IAA 52 Figs 49-54 Lainius constellatus. 49, fore wing (from Kimmins); 50, apex of d abdomen, lateral; 51, apex of ¢ abdomen, lateral; 52, d genitalia, dorsal; 53, 2 spermatheca, lateral; 54, 2 subgenitale, ventral. apex of each segment; setae long, sparse; ecto- procts quite deeply invaginated dorso-apically, not fused dorsally; d: additional short, dense setae present on sternites; callus cerci ovate; trichobothria 52; ectoprocts not fused with tergite 9; atria small; microtholi absent; sternite 8+9 fused but with very short dorsal suture, short, broad; apodemes absent; @: sternite 7 straight apically; callus cerci rounded; trichobothria 38. GENITALIA 6 (Fig. 47). Genitalia weakly sclerot- ized; tignum, gonapsis, median plate and ento- processus absent; gonarcus barely arcuate, very narrow; arcessus short, broadly triangular with apical hook; pseudopenis, gonosaccus absent; gonosetae very short, situated ventro-laterally of arcessus; gonocristae and spinellae absent; hypandrium small. GENITALIA 2 (Fig. 48). Praegenitale absent; subgenitale bilobed; spermatheca broad with lateral striations; ventral impression deep, narrow; vela short; duct long, narrow, highly coiled. REMARKS. Joguina is apparently the most highly evolved of the Old World apochrysine genera. There are multiple gradates series, the radial area is subdivided, the costal area is subdivided, 1A and 2A are forked, and the posterior and apical marginal areas are subdivided. BIoLoGy. Unknown. Insect remains were not included in any of the adult guts examined during this study. Genus LAINIUS Navas Lainius Navas, 1913b: 300. Type species: Lainius constellatus Navas, by original designation. DISTRIBUTION. Central and South America. The genus includes two described species. DIAGNOSIS. Adult. Large lacewings, fore wing (Fig. 49) 21-22 mm. Head marked with red stripe on vertex, scape, brown stripe on pedicel and basal flagellomeres; palps truncate; labrum slightly indented; vertex flat, steeply rising anteriorly; head width : eye width = 2.0—2.1 : 1; toruli small; THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 141 pedicel constricted medially; antenna slightly longer than fore wing; flagellar segments 3 times as long as broad; setae arranged in five rings. Pronotum marked with red baso-lateral spot; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.0—2.2 : 1), rounded apically; pustules marked with dark shading; costal setae long, erect; stigma marked with brown spot; Sc short; Sc and R quite close; basal Sc crossvein absent, five Sc crossveins present in apical half of wing; im absent; tympanal organ narrow, elongate; Rs sinuate, arising basally; wing highly reticulated particularly in basal half, gradates indistinct; veins slightly crassate in d ; costal area broad; c;, c> indistinct; Psm and Pcu very close together, upturned apically; posterior margin broad; 1A forked. Hind wing narrower than fore wing (length : breadth = 3.3 : 1). Abdomen (Figs 50, 51) swollen apically; marked with red dorsal spot at apex of tergite; d setae short, dense with long, sparse setae interspersed; callus cerci rounded; trichobothria 37; ectoprocts slightly invaginated dorso-apically, not fused dorsally, suture between ectoprocts and tergite 9; , narrow; 1A forked; 2A unforked. Hind wing narrow (length : breadth = 3.1 : 1); with two distinct gradate series; radial area undivided; posterior apical margin marked with faint black suffusion. Abdomen (Figs 60, 61) unmarked; elongate; setae long sparse; callus rounded; trichobothria 42-45; ectoprocts with slight dorso-apical invagination, fused dorsally; 3d: additional short, dense setae present on all sclerites; microtholi absent; ectoproct separated from tergite 9 by deep suture; sternite 8+9 fused, short, broad; 2: sternite 7 straight apically. GENITALIA 6 (Fig. 62). Tignum, gonapsis and median plate absent; gonarcus narrow almost straight, hardly arcuate; arcessus weakly sclero- tized, triangular, bearing a few short lateral gonosetae; entoprocessus absent; gonosaccus very short. GENITALIA @ (Figs 63, 64). Praegenitale absent; subgenitale bilobed; spermatheca large, slightly THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 143 62 Figs 59-64 Nacaura matsumurai. 59, fore wing (from Kimmins); 60, apex of d abdomen, lateral; 61, apex of ? abdomen, lateral; 62, d genitalia, dorsal; 63, 2 spermatheca, lateral; 64, 2 subgenitale, ventral. flattened, bearing several lateral grooves; duct very long, highly coiled; vela short; ventral impression deep with irregular outline. Larva. Abdomen fusiform, weakly humped; debris-carrier (Tsukaguchi, in litt.). REMARKS. Nacaura can be distinguished from | other apochrysine genera by the elongate scape, the dense reticulation of the basal half of the fore wing (although the hind wing has no additional crossveins), and the forking of the first anal vein. A single undescribed species represented in the BMNH collection by three males reared as a predator of the mealybug Rastrococcus invadens from Bangalore, India may belong to this genus. It | has the dense reticulation in the basal half of the fore wing, but has three distinct rows of gradates | in the apical half, and hence would not key out to Nacaura. BIOLOGY. Insect remains were not included in the gut contents of any adults examined during this study. According to Tsukaguchi (in litt.), Nacaura matsumurae Okamoto usually occurs in laurel forests and passes the winter in the adult stage. Genus NOBILINUS Navas Nobilinus Navas, 1913b: 295. Type species: Nobilinus insignitus Navas, by original designation. DISTRIBUTION. Oriental. The genus includes six described species and subspecies with a further two undescribed species represented in the BMNH collections. Most of the species are Indonesian. DIAGNOsISsS. Adult. Large lacewings, fore wing (Fig. 65) 23-31 mm; ground colour pale green. 144 Figs 65-69 GS MAIS S. J. BROOKS & P. C. BARNARD 68 Nobilinus. 65, N. albardae insignitus, fore wing (from Kimmins); 66-69, N. phantoma; (66) apex of 3 abdomen, lateral; (67) apex of 2 abdomen, lateral; (68) 3d genitalia, ventral; (69) 2 spermatheca, lateral. Head marked with red transverse stripes on frons and vertex; palps truncate apically; labrum invagi- nate; vertex flattened; head width : eye width = 1.82.0 : 1; antenna considerably longer than fore wing; flagellar segments 3 times as long as broad; setae in five rings. Pronotum elongate; unmarked; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.1—2.3 : 1); marked with dark brown spots in discal area; costal area narrow at base; costal setae long, erect; stigma unmarked; Sc short; numerous Sc crossveins present in apical half of wing; Sc and R widely separated; gradates in three divergent series; inner gradates greatly extended basally; basal inner gradate not meeting Psm; crossveins not crassate in 6; c, slightly shorter than c>, cubital cells broad; 1A and 2A forked. Abdomen (Figs 66, 67) unmarked; setae long, sparse; callus cerci rounded; trichobothria 50; ectoprocts with slight dorso-apical invagi- nation, fused dorsally, fused with tergite 9; d: all sclerites with dense covering of additional short setae; microtholi absent; sternites 8+9 separated by short suture, broad; @: sternite 7 straight apically. GENITALIA 6 (Fig. 68). Tignum, gonapsis, median plate and entoprocessus absent; gonarcus hardly arcuate; arcessus triangular with weak apical hook bearing a few short setae laterally; pseudopenis absent; gonosaccus short; gonocristae absent. GENITALIA @ (Fig. 69). Praegenitale absent; subgenitale bilobed apically; spermatheca broad with lateral striations; ventral impression deep; vela short; duct long, narrow, sinuous. REMARKS. Nobilinus has four apomorphies which help distinguish it from other apochrysine genera: the gradates are arranged in three series, the radial area below the pterostigma is completely filled with crossveins, and both 1A and 2A are forked. ert THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 145 Figs 70-75 Nothancyla verreauxi. 70, fore wing (from Kimmins); 71, apex of d abdomen, lateral; 72, apex of 2 abdomen, lateral; 73, d genitalia, lateral; 74, 2 spermatheca, lateral; 75, 2 subgenitale, ventral. BIoLoGy. Unknown. The gut contents of adults examined during this study did not include insect | remains. | Genus NOTHANCYLA Navas | Nothancyla Navas, 1910a: 51. Type species: Nothancyla verreauxi Navas, by monotypy _ DIsTRIBUTION. Southern and western Australia, | Tasmania. The single known species has a well-defined Bassian distribution. DIAGNosIs. Adult. Large lacewings, fore wing (Fig. 70) d 13-15 mm, 2 17-20 mm; ground colour pale yellowish green. Head unmarked or with red stripe on front of vertex; palps tapered apically; labrum slightly indented; mandibles broad, symmetrical with large basal tooth on each mandible; vertex slightly raised; head width : eye width = 2.3-2.8 : 1; antenna shorter than fore wing; flagellar segments 3 times as long as broad; 146 setae arranged in five rings; scape and pedicel bearing coarse setae. Pronotum unmarked; dorsal setae long, pale, very coarse; mesonotum with red spot above wing base or unmarked; metanotum unmarked. Legs unmarked; setae short, dark; claws without basal dilation. Fore wing broad (length : breadth = 2.5-3.0 : 1); unmarked or red at base; costal area broad at base; costal setae long, erect; stigma unmarked; Sc quite short; some costal crossveins forked; Sc and R widely separated; basal Sc crossvein absent; im short, quadrangular; Rs slightly sinuate; gradates in two parallel series; basal inner gradate not meeting Psm, extending basally; veins not crassate in 3; c about same length as cz or slightly longer; dcc closed at margin, short. Hind wing broad (length : breadth = 2.9-3.0 : 1). Abdomen (Figs 71, 72) unmarked; setae short, sparse; ectoproct fused with tergite 9, fused dorsally; 3: callus cerci ovate; trichobothria 27; ectoprocts greatly elong- ated apically, grooved ventrally; tergites small, sternites broad; sternites 8 and 9 not fused; sternite 9 short, squared apically; microtholi absent; microtrichia very small; @: callus cerci rounded; trichobothria 30; ectoprocts with slight apical invagination; microtrichia present; sternite 7 straight apically; gonapophyses laterales small. GENITALIA 6 (Fig. 73). Tignum, gonapsis and median plate absent; entoprocessus broad, quite long; arcessus narrow, dorsally grooved; pseudo- penis absent; gonarcus narrow, long; gonosaccus, gonosetae, spinellae and gonocristae absent. GENITALIA @ (Figs 74, 75). Praegenitale absent; subgenitale bilobed apically, very broad with basal extension; spermatheca very broad; ventral impression very small; duct very long, sinuous; vela very short. REMARKS. Nothancyla can be readily distin- guished from other chrysopid genera by the quad- rangular intramedian cell and broad basal costal margin in the fore wing, and the coarse setae on the scape, pedicel, pronotum and mesonotum. The greatly extended male ectoprocts and the large basal lobe on the female subgenitale are also distinctive. When New (1980) redescribed the genus he mentioned the presence of a tignum, but we have been unable to find this structure in any of the males we examined. The phylogenetic position of Nothancyla is difficult to ascertain. The possession of five rows of flagellar setae and the absence of a basal subcostal crossvein are apomorphic for the Apochrysinae. However, Nothancyla does not fit comfortably in this subfamily since it lacks the other apochrysine venational apomorphies, such as Psm and Psc in close proximity or a short Sc. S.J. BROOKS & P. C. BARNARD Nothancyla also lacks apochrysine apomorphies in the male and female genitalia, such as the absence of entoprocessus or the presence of lateral striations on the spermatheca, although the large spermatheca is reminiscent of the Apochrysinae. Although the genus sits rather uneasily in this subfamily, it cannot be placed in any other group on existing evidence and is therefore tentatively placed in the Apochrysinae until further information is available to clarify its relationships. BioLoGy. Unknown. There were no insect re- mains in the guts of any of the adults examined during this study. Genus OLIGOCHRYSA Esben-Petersen Oligochrysa Esben-Petersen, 1914: 639. Type species: Oligochrysa gracilis Esben-Petersen, by original designation. DISTRIBUTION. Australia (Queensland, New South Wales), Norfolk Island. The genus is monotypic. DraGnosis. Adult. Large lacewings, fore wing (Fig. 76) 17-23 mm; ground colour pale greyish green. Head unmarked; palps short, broad, taper- ing apically; labrum indented; vertex flattened; head width : eye width = 2.1-2.3 : 1; scape elongate, marked with red apical annulation; antenna considerably longer than fore wing; flagellar segments about 3 times as long as broad; setae arranged in five rings. Pronotum marked with red lateral stripe; dorsal setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.3-2.5 : 1); marked with black spot in middle of inner gradates and on radial crossvein below stigma, outer gradates black; costal setae long, erect; stigma unmarked; Sc short; Sc and R quite close; gradates in two divergent series; inner gradates greatly extended basally; basal inner gradate not meeting Psm; veins not crassate in 6; costal area broad at base; c; about half the length of c,; 1A not forked. Hind wing narrow (length : breadth = 3.1-3.4 : 1). Abdomen (Figs 77, 78) unmarked; setae long, sparse; callus cerci rounded; tricho- bothria 32-37; ectoprocts slightly invaginated dorso-apically, fused dorsally, fused with tergite 9; 6: additional short setae present on sternites 5-9 and ectoprocts; microtholi absent; sternites 8+9 partially separated by narrow suture; 9: sternite 7 straight apically. GENITALIA <6 (Fig. 79). Tignum, gonapsis, median plate and entoprocessus absent; gonarcus THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 147 arcuate; arcessus short, triangular, tapering to short apical hook; pseudopenis absent; gono- | saccus short; few short setae present latero- ventrally on arcessus; gonocristae absent. GENITALIA 2 (Figs 80, 81). Praegenitale absent; subgenitale bilobed apically; spermatheca long, broad; ventral impression deep, broad; vela short; duct long, narrow, highly coiled. REMARKS. We have been unable to find any autapomorphic characters for Oligochrysa and it may be the most primitive genus of the subfamily. It may be most closely related to Synthochrysa, which is distributed in the western Pacific and Indonesia, and from which it can be distinguished by the lack of a third series of gradate crossveins in the fore wing. BIOLOGY. Unknown. $1 Figs 76-81 Oligochrysa lutea. 76, fore wing (from Kimmins); 77, apex of d abdomen, lateral; 78, apex of ? abdomen, lateral; 79, 3d genitalia, dorsal; 80, 2 spermatheca, lateral; 81, 2 subgenitale, ventral. Genus SYNTHOCHRYSA Needham Synthochrysa Needham, 1909: 202. Type species: Hemerobius_ stigma Girard, by _ original designation. DISTRIBUTION. Western Pacific and Indonesia. Four species have been described in the genus. DraGnosis. Adult. Large lacewings, fore wing (Fig. 82) 20-24 mm; ground colour pale green. Head unmarked; palps truncate (Fig. 83); man- dibles broad, asymmetrical with basal tooth on left mandible (Fig. 84); labrum indented; vertex slightly raised; head width : eye width = 1.62.2 : 1; antenna longer than fore wing; flagel- lar segments 3 times as long as broad; setae arranged in five rings. Pronotum marked with red lateral stripe; dorsal setae long, pale; meso- and 148 i oe S . J. BROOKS & P. C. BARNARD 83 88 84 Figs 82-88 Synthochrysa. 82, 85-88, S. montrouzieri; 83, 84, S. salomonis. 82, fore wing (from Kimmins); 83, apical segment of maxillary palp, dorsal; 84, mandibles, dorsal; 85, apex of d abdomen, lateral; 86, apex of @ abdomen, lateral; 87, d genitalia, dorsal; 88, 2 spermatheca, lateral. metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.2-2.5 : 1); marked with black spot on central inner gradates; costal area narrow at base; costal setae long, erect; stigma unmarked; Sc short; Sc and R widely separated; gradates in three divergent series; inner gradates greatly extended basally; basal inner gradate not meeting Psm; veins not crassate in 6; c; shorter than c); 1A not forked. Abdomen (Figs 85, 86) unmarked; setae long, sparse; callus cerci rounded; trichobothria 35-63; ectoprocts with slight dorso-apical invagination, fused dor- sally; d: additional short setae on sternites 5-9, tergites 7-9 and ectoprocts; microtholi absent; ectoprocts fused with tergite 9; sternite 8+-9 fused, short; 2: sternite 7 straight apically with long subapical median horn. GENITALIA 6 (Fig. 87). Tignum, gonapsis, median plate and entoprocessus absent; gonar- cus arcuate; arcessus triangular with apical hook and lateral lobes; pseudopenis absent; gonosaccus short; gonosetae and gonocristae absent. GENITALIA @ (Fig. 88). Praegenitale absent; subgenitale bilobed with median projection; spermatheca long, broad with lateral annulations; ventral impression deep, broad, abruptly narrow- ing apically; vela short; duct long, narrow, sinuous. REMARKS. Synthochrysa is one of the least derived genera in the Apochrysinae and the wing venation closely resembles Oligochrysa Esben-Petersen. Synthochrysa may be distinguished from other genera by the presence of three series of gradate crossveins, the median projection at the apex of sternite 7 in females and the lateral lobes at the apex of the arcessus in males. BIoLoGy. Unknown. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 149 Subfamily CHRYSOPINAE Schneider Chrysopina Schneider, 1851: 35. Type genus: Chrysopa Leach. Chrysopinae Esben-Petersen, 1918: 27. DISTRIBUTION. World-wide. D1AGnosis. Adult. Small to large lacewings, fore wing 9-31 mm; head width : eye width = 1.64.9 : 1; pedicel slightly constricted medially; setae on flagellar segments arranged in four or less rings; basal Sc crossvein in fore wing positioned relatively basad (—0.08 to +0.96 mm); im quad- rangular or ovate; basal Rs crossvein meets im in apical half; tympanal organ swollen, spherical; c, longer or shorter than cz; Psm and Psc widely separated; Psm long, fusing with outer gradates; gradates arranged in 04 rows; jugal lobe small. Genitalia variable. REMARKS. This is by far the largest of the three subfamilies of Chrysopidae, containing 56 of the 75 genera currently recognised in the family. It is also a very diverse group, which has led several authors (e.g. Adams, 1967) to suggest that it may be a convenient but paraphyletic assemblage. However, our studies show that it seems to be a monophyletic group, recognised by the autapo- morphy of possessing only four rings of setae on the flagellar segments. Key to the genera and subgenera of the Chrysopinae Three genera, Chrysaloysia Navas (p. 170), Nuvol Navas (p. 251) and Neula Navas (p. 251), have been omitted from this key because we have not been able to examine suitable specimens. The key ignores the tribal classification of the Chrysopinae and goes straight to generic and subgeneric level: this is because we believe that keys should be practical and easy to use, rather than conforming to the phylogeny of the group concerned. A key to tribes would necessitate an examination of male genitalia and mouthparts of every specimen, whereas the key presented here uses non-genitalic ‘external’ characters as far as possible. Some of the more complex genera key out in more than one place. The captions of Figs 1-6 explain the termin- ology of many of the characters used in the key. 1 Sc very short; 1-2 crossveins below pterostigma (RG a aoe el a ane Oa 2 — Sc long; at least 34 crossveins below pterostigma (VETS) eee 5. cron 0 Ottis Bcc Hee See eee 4 2 Cell im absent in fore wing (Fig. 95) ANK YLOPTERYX (subgenus SENCERA Navas) 10 11 Cell im present in fore wing (Fig. 89)............ 3 Costa at base of fore wing unmarked; apex of tarsus marked black; d: entoprocessus long, usually fused apically; gonosaccus simple; pseudopenis present; arcessus absent (Fig. 92) ANKYLOPTER YX Brauer Costa at base of fore wing marked black; apex of tarsus unmarked; d: entoprocessus short, not fused apically; gonosaccus paired; pseudopenis absent; arcessus present (Fig. 104) PARANKYLOPTER YX Tjeder Gradates absent from fore and hind wing (or only one gradate crossvein present) (Fig. 215) TURNEROCHRYSA Kimmins At least one complete series of gradate crossveins present in either fore orhind wing .............. 5 Only one series of gradate crossveins present in hind wing, inner gradates absent from all wings . . .6 Two or more series of gradate crossveins in hind WEDD ere na cree mane ist adsts etc cra cis Pereieiearse eveuniele’ it 3 Old World species; cell im absent from fore wing CHRYSOPA Leach [part: minuta McLachlan only] New World species; cell im present in fore wing. . .7 One series of gradate crossveins in fore wing ..... 8 Two series of gradate crossveins in fore wing (Fig. 363) EREMOCHRYSA Banks (subgenus EREMOCHRYSA) Claws dilated (Fig. 11); pedicel elongate; <6: ectoprocts extended apically, bearing strong, apical teeth (Fig. 426); gonapsis short, broad, bifurcate (Fig. 428); 2: subgenitale extended basally (Fig. ABN) wie io atierivgst PARACHRYSOPIELLA gen. n. Claws undilated (Fig. 12); pedicel about as long as broad; 6: ectoprocts rounded, not extended apically with long, fine setae (Fig. 356); gonapsis long, narrow (Fig. 359); ¢: subgenitale not extended basally but projects ventrally (Fig. 360) EREMOCHRYSA Banks (subgenus CHRYSOPIELLA Banks) Three or more regular series of gradate crossveins INnikoLe wine (Bigs 220)... anatase cet eialtone tote a, 10 Two series of gradate crossveins in fore wing, although occasional crossveins may be doubled (Big 3322). 26 seis cccconne. oktecioe eeaigmpgpatee 21 Two series of gradate crossveins in hind wing... .11 More than two regular series of gradate crossveins ral ovine Wh6Te coe, ORR Gaeta g Atco Oaks cc cient 13 Neotropical species; antenna equal to or longer than fore wing; fore wing with dark spots or shading LiF St SAR ENON ic SENECA ORT eee Re Pe 12 Oriental species; antenna shorter than fore wing; fore wing unmarked (Fig. 253) BORNIOCHRYSA nom. n. [part] 150 15 16 17 18 19 20 21 Antenna 1.5 times length of fore wing; cell im quadrangular; fore wing with numerous dark spots (Hig:486) age cele... CACARULLA Navas Antenna about as long as fore wing; cell im ovate; fore wing with pale brown shading around cross- VEINS) saga tc See CHR YSOPODES Navas [part] Cell im quadrangular (Fig.8)................. 14 GCellimovate (Fic 210) eee erate eer 15 Claws with basal dilation (Fig. 11); scape elongate (Fig. 16). AUSTROCHRYSA Esben-Petersen [part] Claws without basal dilation (Fig. 12); scape about as broadaslong ............. REXA Navas [part] Scape grossly enlarged; costa convex in hind wing TUMEOCHRYSA Needham Scape as broad as long or slightly elongate; costa Straight bindiwinei esse sea ee ee 16 Eyes small (head width : eye width = 2.5 : 1) (Fig. 16); 3: ectoprocts flattened caudally; apex of sternite 8+9 hooked dorsally (Fig. 223); species confined to Hawaiian Islands ANOMALOCHRYSA McLachlan Eyes larger (head width : eye width < 2.5: 1); d: ectoprocts not flattened; sternite 8+9 straight; Orientallspecies#-t: See ee 2s eee eres = 17 Costal setae long, erect or slightly inclined towards fore wing apex; d: sternite 8+9 elongate, extend- ing beyond apex of ectoprocts (Fig. 317)........ 20 Costal setae short, strongly inclined towards wing apexes Go) SLenMILe 6-1-9 SHORE ene eee re 18 Gradates arranged in four series; radial crossveins sinuate in apical half of fore wing; d: gonosaccus bearing teeth; arcessus with paired ventro-apical hooks (Fig. 384) HIMALOCHRYSA Haizel [part] Gradates arranged in three series; radial crossveins straight; d: gonosaccus untoothed; arcessus with- out paired apical hooks...................... 19 Small species, fore wing < 15 mm; d: sternites 8+9 fused; gonapsis, tignum and arcessus present; pseudopenis absent... .. MALLADA Navas [part] Large species, fore wing > 16 mm; d: sternites 8+ 9 not fused; gonapsis, tignum and arcessus absent; pseudopenis present PLESIOCHRYSA Adams [part] Radial crossveins in fore wing straight (Fig. 308); 3 with tignum (Fig. 312)...... ANACHRYSA Hoizel Radial crossveins in fore wing sinuous (Fig. 316); d without tignum .......... CHRYSOPIDIA Navas Cell im absent in fore wing (Fig. 133) NESOCHRYSA Navas [Madagascar species] and BELONOPTERYX Gerstaecker [New World species] 24 Pes) 26 27 28 30 S.J. BROOKS & P. C. BARNARD Celli presentinitore witty: 0 2472 Cell im quadrangular (Fig. 8)................. 23 Cellim ovate (Fig. 10) or triangular (Fig.9)..... 34 Pterostigma of fore wing marked with dark spot Pterostigma of fore wing unmarked Pronotum narrow; flagellar segments at least 3 times as long as broad in mid-antenna; palps narrow, asymmetrical apically, flattened apically in lateral view (Fig. 14); vertex raised; New World species Pronotum broad; flagellar segments at most 2 times as long as broad in mid-antenna; palps broad, symmetrical apically, cylindrical not flattened apically in lateral view (Fig. 13); vertex flat; Old World species:.. 2 sys: 3:8 6c Soe eee eee 26 Fore wing marked, often extensively, with black or dark brown spots (Fig. 492) . ... GONZAGA Navas Fore wing usually unmarked but if markings are present these are restricted to dark shading adjacent to crossveins (Fig. 498) LEUCOCHRYSA McLachlan (subgenus LEUCOCHRYSA) Inner gradate crossveins not extended basally; cz narrow (Fig. 201); d: gonarcus arcuate (Fig. 205); 2: praegenitale absent; subgenitale supported on elongate sclerotized plate (Fig. 203) OYOCHRYSA Brooks Inner gradate crossveins extended basally; c, broad (Fig. 208); 3d: gonarcus transverse (Fig. 211); 2: praegenitale present (Fig. 210); subgenitale not supported on sclerotized plate ................ 27 Antenna shorter than or same length as fore wing; flagellar segments in mid-antenna less than twice as long as broad; c, 2.0-2.5 times longer than c); 3d: parameres short, paired (Fig. 211); 2: small median projection at apex of subgenitale (Fig. 7118) ree eed} 5: STIGMACHRYSA Navas Antenna longer than fore wing; flagellar segments in mid-antenna twice as long as broad; c, at most 1.5 times longer than c,; d: parameres long, 2-3 pairs (Fig. 190); 2: without median projection at apex of subgenitale (Fig. 194) ..... NESOCHRYSA Navas Flagellar segments twice as broad as long; c; shorter than cz; pronotum narrow ................... 29 Flagellar segments about as broad as long; c, at least 1.5 times longer than cz; pronotum broad . .32 Claws without basal tooth (Fig. 12); radial cross- VelnSStralp hte rani ee fee ate REXA Navas Claws with basal tooth (Fig. 11); radial crossveins SINUOUS 8.24 Sis Jee Ete PA eee 30 Old World species; pronotum elongate, marked with black medio-lateral spot (Fig. 19); antennae longer than fore wing; mandibles (Fig. 18) and palps (Fig. 14) broad)... 0.02 -2.-..-05---- eee 31 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 31 New World species; pronotum almost square, marked with yellow median stripe; antennae at most as long as fore wing; mandibles (Fig. 17) and palps (Fig. 15) narrow CHR YSOPODES Navas (subgenus CHRYSOPODES) Fore wing marked with a few large spots in discal area; inner gradate series irregular and widely divergent from outer gradates; outer gradates green (ICSE Sie lacie Gade Alelstf sere KOSTKA Navas Fore wing without large spots; outer gradates black; inner and outer gradates parallel (Fig. 246) AUSTROCHRYSA Navas New World species; claws without basal tooth (Fig. 12) oo chye tO eee NACARINA Navas Old World species; claws with basal dilation (Fig. LIT sc ete 3 Mec ROS RSI sr eee 33 ‘ Cell im short (Fig. 162); ¢ parameres absent; @ praegenitale present (Fig. 166) EVANOCHRYSA gen. n. Cell im long (Fig. 169); d parameres present; 2 praegenitale absent..... ITALOCHRYSA Principi Basal radial crossvein, which meets apex of cell im, leaves radius before origin of Rs (Fig. 480) BERCHMANSUS Navas No radial crossvein basal to originofRs ........ 35 Cellicymuchilongerthantcs:i....---+ee8sen--- 36 Cell c, shorter (rarely slightly longer) thanc, ... .39 Claws without distinct basal dilation (Fig. 12) . . .37 Claws with strongly toothed basal dilation (Fig. Ih 0 cence RSIS AP eS ie ar 38 Vein Cu, in fore wing forked (Fig. 135) CALOCHRYSA Banks Vein Cu, in fore wing unforked (Fig. 154) DYSOCHRYSA Tjeder Both wings and pterostigma extensively marked with black spots (Fig. 147); scape slightly elongate; 3d: parameres long, tapering apically (Fig. 151); ?: sternite 7 without subapical spines (Fig. 149) CHRYSACANTHIA Lacroix Crossveins shaded in basal half of fore wing, pterostigma unmarked (Fig. 195); scape broad; d: parameres short, swollen apically (Fig. 198); 2: sternite 7 with short subapical spines (Fig. Si) aaa reeaaeee NODOCHRYSA Banks Female (N.B. this part of the key should be used with caution as it is not always possible to identify unassociated females)....................... 74 Gonapsis present (Figs 2, 3) but sometimes very small and indistinct in Glenochrysa (Fig. 377), most specimens of which have extensively marked wings (Bi 953 73) ie ates rato le cen eB ans agen is 41 41 42 43 45 46 47 48 49 Gonapsis'absent) 22% feisty an eh cles signee: 's 50 Pseudopenis present as additional hooked structure belowarcessus\(Figs408)0sn soe ele ae ss 42 Pseudopenis absent, only arcessus present...... 43 Fore wing narrow (length : breadth = 3.0: 1) and unmarked; scapes elongate and widely separated at base; New World species, often with head ornamentation or stridulatory structure on second Stemmite (Fir. 20). soon vane - MELEOMA Fitch Fore wing broad (length : breadth < 3.0 : 1) and marked with small black spot at base of costa; scapes as long as broad and close together at base; species endemic to Canary Is and Madeira; head without ornamentation; stridulatory structure ADSM ects sucesso: ATLANTOCHRYSA Holzel TSA PTESOM Ee res as) ops alto MALLADA Navas WDIPMIMMAOSERL BN ante Sai teks ches. keene = 44 Gonapsis large, elongate (Fig. 270); median plate PLESENLMHMAe ee er boMincibae lencidled cad et. 6)s 45 Gonapsis small, short; median plate absent ..... 46 Fore wing narrow (length : breadth > 2.9 : 1); anal veins not crassate; radial crossveins straight; median plate with dorsal horns (Fig. 268) CERAEOCHRYSA Adams Fore wing broad (length : breadth < 2.8 : 1); anal veins crassate; radial crossveins sinuate; median plate without dorsal horns CHR YSOPODES Navas Fore wing marked, often extensively, with black and pale brown shading; Sc short; basal branch of Cu, recurrent (Fig. 373) GLENOCHRYSA Esben-Petersen Fore wing unmarked or markings restricted to small black spots on cell dec and pterostigma; Sc long; basal branch of Cu, only slightly curved ........ 47 Fore wing usually (but not invariably) marked; gonapsis narrow, broadly arcuate; ectoprocts separated by dorsal suture (Fig. 22)............ 48 Fore wing unmarked; gonapsis V- or T-shaped; ectoproctsitused dorsally 7-5. 2-0-2 -s50---- =o 49 Fore wing usually marked with red basal suffusion; ectoprocts very long and narrow (Fig. 289); gonap- sis broad, arcuate with apical teeth (Fig. 291); antenna considerably longer than fore wing CHR YSOCERCA Weele Fore wing usually marked with black spot on dcc and pterostigma; ectoprocts short and broad (Fig. 281); gonapsis narrow, simple (Fig. 284); antenna short, at most only slightly longer than fore wing CHRYSEMOSA nom. n. Ectoprocts narrow, pointed dorso-apically (Fig. 433); gonapsis V-shaped (Fig. 435); inner gradates more numerous than outer; wings very narrow (length : breadth > 3.5 : 1) (Fig. 432) PEYERIMHOFFINA Lacroix 152 — Ectoprocts broad (Fig. 231); gonapsis small, T-shaped (Fig. 234); outer gradates more numerous than inner; wings broad (length : breadth < 3.5: 1) (Big 230) is oy cca APERTOCHRYSA Tjeder 50) Wignumipresent(Fies3)) eae eae eee eee 51 — ei hignumiabsent\s. sagen whee ee ese 53 : : ; : 60 51 First Rs crossvein usually meets Psm distal to cell im (Fig. 301); arcessus present; pseudopenis absent (Fig. 304); sternites 8+-9 fused; apex of sternite 8+9 with distinct lip (Fig. 302) CHRYSOPERLA Steinmann - — First Rs crossvein meets cell im subapically; arcessus absent; pseudopenis present; sternites 8 and 9 not fused; apex of sternite 9 without lip . . . .52 52 Head marked with black spot on gena and between antennae; radial crossveins straight; tignum small, OVate DAI cena. CHRYSOPA Leach [part] 61 — Head and pronotum marked with orange or red spots (Fig. 26); radial crossveins sinuous or oblique; tignum large, transverse (Fig. 446) PLESIOCHRYSA Adams [part] 62 53 Stridulatory structure situated laterally on abdomi- nal segment 2 (Fig. 20) = BRINCKOCHRYSA Tjeder — Stridulatory structure absent ................. 54 63 54 Gonarcus short, broad, transverse (Fig. 523); arcessus broad (pseudopenis absent) .......... 55 — Gonarcus long, narrow, arcuate; arcessus or pseudopenis narrow, tapering apically ......... 57 55 Palps broad, rounded apically (Fig. 13); pronotum very broad; parameres present (Fig. 129) ABACHRYSA Banks 64 — Palps narrow, tapered apically (Fig. 14); pronotum Narrow: pParamMeresidWscit sae ae eee - 56 56 Fore wing (except pterostigma) unmarked or witha few small black spots; gradates parallel; basal costal crossveins straight; radial crossveins parallel (Fig. 512) LEUCOCHRYSA (subgenus NODITA Navas ) 65 — Fore wing marked extensively with large black spots; inner gradates widely divergent from outer; basal costal crossveins sinuate; radial crossveins 66 below pterostigma divergent (Fig. 520) VIEIRA Navas 57 Sternites 8 and 9 not fused (Fig. 274)........... 58 — Stemnitesé--9 fusedi(Higs2) me aereeeecnaen eee 61 58 Entoprocessus very long and narrow, protruding beyond apex of abdomen (Fig. 276); at least twice as many outer gradates as inner gradates in all wings (Bi8827/3)) ae aeeeras CERATOCHRYSA Tjeder 67 — Entoprocessus short; less than twice as many outer gradates as inner gradates.................... 59 59 Pseudopenis present, arcessus absent; ectoprocts S. J. BROOKS & P. C. BARNARD short (Fig. 2); inner gradates not extended basally, basal inner gradate meeting Psm (Fig. 1) ....... 60 Pseudopenis absent, arcessus present; ectoprocts elongated apically (Fig. 417); inner gradates greatly extended basally, basal inner gradate not meeting Psmi(Big, 414) p20 ccna x. oe oes NINETA Navas Head marked (often extensively) with black spots; entoprocessus with dorsal horn (Fig. 297); horns absent from gonarcus; eyes small (head width : eye width= 2.6): 1) oe CHRYSOPA Leach Head usually marked with large red or yellow spots (Fig. 26); entoprocessus without horn; gonarcus with paired median horns (Fig. 444); eyes large (head width : eye width < 2.6: 1) PLESIOCHRYSA Adams [part] Ectoprocts narrow and projecting apically, bearing tuft of long, coarse setae at apex (Fig. 254) BORNIOCHRYSA nom. n. Ectoprocts rounded apically, not extended, tuft of setae absent 7-03 cients. See eee 62 Radial crossveins oblique or sinuate (Fig. 380); scape usually elongate (Fig. 16) ............... 63 Radial crossveins straight (Fig. 107); scape about as long:as Dioad jg. se ce eee 67 Median radial crossveins sinuate or oblique, radial crossveins below pterostigma straight (Fig. 380); stermite 8-9 short (Higs38)) heeeee sete eee ae 64 Median radial crossveins straight, radial crossveins below pterostigma sinuate or oblique (Fig. 322); sternite 8+9 elongate (Fig. 325)............... 66 Old World species; dorsal apodeme short, not extending beyond apex of abdomen; sternite 8+9 without ventral swelling ..................... 65 New World species; dorsal apodeme long, extend- ing beyond apex of abdomen; small mid-ventral swelling on sternite 8+9 (Fig. 333) CHR YSOPODES Navas [part] Gonocristae present (Fig. 247); gonosaccus without apicalitecthier sapere AUSTROCHRYSA Navas Gonocristae absent; gonosaccus with apical teeth (Riss384)\ ace eee HIMALOCH8HRYSA Hoizel Costal setae in fore wing long, erect; arcessus narrow, tapering apically (Fig. 327); head un- marked or with red spots; wings unmarked (Fig. B22) a sys Naege Mioangs) = Cee CHR YSOTROPIA Navas Costal setae short, inclined towards wing apex; arcessus with apico-ventral forked process (Fig. 122); head with black markings; wings with exten- sive black shading (Fig. 119) SIGNOCHRYSA gen. n. Mandibles narrow, scythe-like (Fig. 17); palps narrow, cylindrical, elongate apically (Fig. 15) . .68 Mandibles broad (Fig. 18); palps broad, flattened laterally, abruptly tapering apically (Fig. 14) ... .70 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 68 69 70 ph 72 73 74 75 76 New World species; arcessus weakly sclerotized, triangular with apical hook, lateral rods and dorsal microsetae (Fig. 335) CHR YSOPODES Navas [part] Old World species; arcessus strongly sclerotized, linear, tapering apically without strong apical hook; dorsal microsetae and lateral rods absent ....... 69 Antenna shorter than fore wing; frons unmarked; apodeme of sternite 8+9 with prominent apical tooth (Fig. 108); genitalia with median plate and membranous sac dorsal of arcessus present (Fig. 110); gonosetae numerous .. RETIPENNA Brooks Antenna longer than fore wing; frons marked with row of 2-3 black spots below antennae (Fig. 25); apodeme of sternite 8+-9 short, not projecting (Fig. 114); genitalia with median plate and dorsal membranous sac absent; gonosaccus with 4 (rarely 2) long, lateral gonosetae (Fig. 116) SEMACHRYSA Brooks Basal inner gradate not meeting Psm (Fig. 230) . .71 Basal inner gradate meeting Psm(Fig.1)....... 72 Old world species; ectoprocts separated dorsally by deep groove or narrow suture (Fig. 22); fore wing often marked, particularly cell dec (Fig. 456) (fore wing marked in all species with fore wing > 9 mm); arcessus trifurcate apically (Fig. 459); ¢ ectoprocts extended and hinged basally .... SUARIUS Navas New World species; ectoprocts fused dorsally, dorsal groove absent; fore wing unmarked (Fig. 468); arcessus tapering to single point (Fig. 471); 3d ectoprocts rounded, not hinged, basally (Fig. AOD eles Sos cas yee UNGLA Navas Arcessus axe-head shaped when viewed laterally (EG: C574) eee eee CUNCTOCHRYSA Holzel Arcessus narrow, tapering apically to short hook Gonocristae present (Fig. 475); entoprocessus present with dorsal hooks; arcessus entirely sclerot- ized, large (Fig. 477) ..... YUMACHRYSA Banks Gonocristae absent; entoprocessus absent; arcessus small, membranous with lateral sclerotized rods (ig 346)... 2 +: NEOSUARIUS Adams & Penny Praegenitale present (Fig. 6); pronotum very broad TS seereyh i aiS Stays Atkosagaasle Pee aE be oe 75 Praegenitale absent; pronotum narrow ........ 76 New World species; pronotum marked with two transverse rows of black spots; fore wing < 20 mm; claws without basal dilation (Fig. 12) ABACHRYSA Banks Old World species; pronotum marked with broad red/brown longitudinal stripe; fore wing > 20 mm; claws basally dilated (Fig. 11) NESOCHRYSA Navas Basal inner gradate not meeting Psm in fore wing (E252) Rs tot Se eae eee 77 77 78 79 80 81 82 83 84 153 Basal inner gradate meeting Psm in at least one Vay CBA ie) ae a RAEN eae Seen ae See oe 85 Sternite 7 with deep apical invagination (Fig. 21); stridulatory structure present, situated laterally on abdominal segment 2 (Fig. 20) BRINCKOCHRYSA Tjeder Sternite 7 straight or convex apically; stridulatory structure/Absents yoga Ne hase 78 Radial crossveins sinuous or oblique (Fig. 119) . .79 Radial crossveins straight (Fig.1) ............. 81 Large species (fore wing = 16 mm); fore wing unmarked; mandibles broad, symmetrical with basal tooth on both mandibles (Fig. 416); palps broad, tapering abruptly apically (Fig. 14) ...... 80 Smaller species (fore wing < 13 mm); fore wing marked with black spots; mandibles narrow, scythe-like, symmetrical without basal tooth (Fig. 17); palps narrow, elongate apically (Fig. 15) SIGNOCHRYSA gen. n. Scape elongate; inner gradates extended basally; head broad (head width : eye width > 2.7 : 1) LOG PERO RE. OS A NINETA Navas Scape about as broad as long; inner gradates not extended basally; head narrow (head width : eye width <2.6: 1)... PLESIOCHRYSA Adams [part] Sc short; basal branch of Cu, recurrent; wing veins usually with extensive black and pale brown shading; wing membrane with pustules (Fig. 373) GLENOCHRYSA Esben-Petersen Sc long; basal branch of Cu, not recurrent; wing unmarked or marked with small spots only; wing Membrane NOE DUSTMNALCC nc. anne osetia ae 2 82 Ectoprocts not fused dorsally or with deep dorsal groove (Fig. 22); basal lobe of subgenitale elongate or with V-shaped indentation; fore wing usually marked with black spots, particularly on cell dcc; longitudinal veins often unforked apically ...... 83 Ectoprocts fused dorsally; subgenitale straight or tapered basally; fore wing usually unmarked; longi- tudinal veins forked apically; ................. 84 Fore wing marked with black spots on dcc and pterostigma (Fig. 280); claws with basal dilation (Fig. 11); basal lobe of subgenitale with V-shaped indentation (Fig. 286); ectoprocts separated by dorsalsuture? =... .0.-- CHRYSEMOSA nom. n. Fore wing unmarked or marked with numerous small black spots throughout wing (Fig. 456); claws undilated (Fig. 12); basal lobe of subgenitale elongate (Fig. 460); ectoprocts separated by deep domsalisroovemn arise se SUARIUS Navas Subgenitale tapering basally (Figs 236, 472) APERTOCHRYSA Tjeder [Old World species] and UNGLA Navas [New World species] Subgenitale straight basally (Fig. 400) MALLADA Navas 87 88 89 90 91 92 93 94 Radial crossveins oblique or sinuate (Fig. 245) . .86 Radial crossveins straight (Fig. 1) Costal crossveins sinuate; fore wing extensively marked, with spot on stigma; inner gradates widely divergent from outer gradates (Fig. 520) VIEIRA Navas Costal crossveins straight; stigma unmarked; fore wing unmarked or with small spots; inner and outer pradatessubparalleli 3.5 eee 87 Median radial crossveins sinuate, radial crossveins below pterostigma straight (Fig. 245) Median radial crossveins straight, radial crossveins below pterostigma oblique (Fig. 322) Fore wing marked with black spot at base of costa; head marked with broad black X-shaped marking between antennae; species endemic to Canary Is and Madeira........ ATLANTOCHRYSA Holzel Fore wing with costa unmarked basally; head not marked between antennae; species not occurring in (CAM APSOHBMVECIOTED o cose cauiocesoecodueraoe 89 Pronotum marked with black medio-lateral spot (Fig. 19); callus cerci large, trichobothria widely dispersed (Fig. 247) AUSTROCHRYSA Esben-Petersen Pronotum marked with yellow median stripe; callus cerci small, trichobothria compact (Fig. 335) .. . .90 Fore wing with more than 4 outer gradates and less than 4 inner gradates and with at least twice as many outer gradates as inner gradates (Fig. 273); sub- genitale mounted on long sclerotized plate (Fig. DIB) ect csctt eect: CERATOCHRYSA Tjeder Wings with less than twice as many outer gradates as inner gradates (Fig. 1); subgenitale without sclerotizediventraliplatem-- = sees 91 New World species; radial crossveins oblique (Fig. 330); subgenitale straight basally (Fig. 338) CHR YSOPODES Navas [part] Old World species; radial crossveins sinuate (Fig. 380); subgenitale elongate basally (Fig. 386) HIMALOCH8HRYSA Hélzel Scape elongate (Fig. 16); costal setae of fore wing long, erectsier emia CHRYSOTROPIA Navas Scape as broad as long; costal setae short, inclined towards wing apex............ MALLADA Navas Sc in stigmatic region marked black; im usually triangular or broadly ovate (Fig. 512); antenna about 1.5 times as long as fore wing LEUCOCHRYSA McLachlan (subgenus NODITA Navas) Pterostigma unmarked; im ovate; antenna shorter or only a little longer than fore wing Wings usually marked, often extensively, with brown spots or shading around crossveins; palps narrow, elongate apically (Fig. 15), rounded in 95 96 97 98 99 100 101 102 103 S.J. BROOKS & P. C. BARNARD cross-section; mandibles narrow, scythe-like (Fig. LT) ich avcurectoneperaiet aoe repeuntroncie teats Ite ee ee 95 Wings unmarked; palps broad, tapering abruptly apically (Fig. 14), flattened in cross-section; man- diblesibroad|(Big 218) eee eee ee eee 97 New World species CHR YSOPODES Navas [part] OldiWorldispecies 2 5e. 2-2 = eee 96 Frons marked with row of 2-3 black spots below antennae (Fig. 25); spermatheca duct narrow (Fig. PL rete oe pee tere SEMACHRYSA Brooks Frons unmarked; spermatheca duct with broad ventral expansion (Fig. 111) RETIPENNA Brooks First Rs crossvein meets Psm distal to apical cell (Fig. 301) iy. San CHR YSOPERLA Steinmann First Rs crossvein meets cell im at apex or subapically Fore wing very narrow (length : breadth = 3.5 : 1); inner gradates more numerous than outer gradates (Big..432) accessors PEYERIMHOFFINA Lacroix Fore wing broader (length : breadth < 3.2 : 1); inner gradates fewer thanouter............... 99 Scape and/or pronotum marked with red lateral stripes; median radial crossveins black CERAEOCHRYSA Adams Not with this combination of markings ........ 100 Base of fore wing and mesoscutum marked red/ brownorblack........ CHRYSOCERCA Weele Base of fore wing unmarked................ 101 Basal fork of M (at base of im) and crossvein between m, and m, thickened; M with acute fork (Fig. 23); vertex and frons usually marked with large yellow, orange or red suffusion (Fig. 26) PLESIOCHRYSA Adams [part] Crossveins at base of wing not swollen; M with less acute fork (Fig. 24); head with black or brown markings, any red markings restricted to small Spotornarrow stripe... --------- eee 102 Scapes elongate and widely separated from each other at base or stridulatory structure present laterally on abdominal segment 2 and inner surface of hind femur (Fig. 20) MELEOMA Fitch Scapes as long as broad and close together at base; stridulatory structure absent ............... 103 Cell mz broad (width : length < 2.5 : 1); most crossveins in fore wing entirely green BORNIOCHRYSA nom. n. Cell m, narrow (width : length = 2.7 : 1); most crossveins in fore wing entirely black or black at Gachiend fits ceeaeenergeecacindtn teens ee eee 104 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 104 Spermatheca large; vela very long and coiled (Fig. BA) a ishafe nates NEOSUARIUS Adams & Penny — Spermatheca small; vela shorter, not coiled . . .105 105 Subgenitale extended basally (Fig. 478); head with extensive brown markings; pronotum with broad brown lateral stripe; antenna brown; longitudinal wing veins brown....... YUMACHRYSA Banks — Subgenitale straight basally; head and pronotal markings (if extensive) black; antenna and longi- tudinal wing veins green/yellow............. 106 106 Head markings restricted to black or red stripe on gena; fore wing narrow (length : breadth = 3.0 : 1), slightly pointed at apex; Rs sinuous (Fig. 349); subgenitale with basal crumena (Fig. B59) heeft rs lat Stes CUNCTOCHRYSA Holzel — Head usually marked with extensive black spots and/or stripes; fore wing broad (length : breadth < 3.0: 1), rounded at apex; Rs straight or slightly sinuous, especially in larger species (Fig. 294); subgenitale without crumena CHRYSOPA Leach Tribe ANKYLOPTERYGINI Navas Ancilopteryginos Navas, 1910a: 59. Type genus: Ankylopteryx Brauer. Ancylopterygini Navas, 1913b: 293. Ankylopterygini Holzel, 1970: 51. DISTRIBUTION. Palaeotropics. DIAGNOsIS. Adult. Small to medium lacewings, forewing 7-21 mm. Head short; head narrow, head width : eye width = 1.62.6 : 1; palps narrow, elongate apically (Fig. 15); galea narrow; mandibles scythe-like, symmetrical without basal teeth (Fig. 17); toruli small; vertex raised; flagel- lar segments narrow, at least 3 times as long as broad; setae arranged in four rings. Pronotum narrow, short. Legs short. Fore wing more or less marked with black spots; usually highly setose; costal area narrow or broad at base; Sc and R generally close together; basal Sc crossvein —0.08 to +0.4 mm; im usually ovate, narrow or sub- rhomboidal; gradates in two parallel series; c, shorter than c>. Hind wing usually narrow (length : breadth > 3.3 : 1). Abdomen with setae long, sparse; trichobothria less than 35; ectoprocts narrow; d: microtholi usually absent; sternite 8+9 fused. GENITALIA 6d. Tignum and gonapsis absent; median plate usually absent; entoprocessus pres- ent; long, narrow arcessus or pseudopenis present; gonarcus narrowly arcuate; gonosetae usually present; parameres absent. GENITALIA 2. Praegenitale absent; spermatheca narrow. 155 REMARKS. Ankylopterygini includes six genera and subgenera. Ankylopteryx Brauer is distri- buted throughout the Old World, Parankylopteryx Tjeder is restricted to the Afrotropics, and from the eastern Palaearctic and Oriental region are Signochrysa gen. n., Retipenna Brooks, Semachrysa Brooks and Sencera Navas. Ankylopterygini genera are characterised by narrow hind wings, highly setose wings, scythe- like mandibles and elongate apical segments of the palps. The most derived genera are Ankylopteryx, Sencera, Parankylopteryx and Semachrysa all of which have the basal costal area of the fore wing enlarged and the basal subcostal crossvein positioned more basad than any other chrysopid genus (it is often basad of the m,/m, crossvein). The presence of a pseudopenis and lack of an arcessus in Sencera and Ankylopteryx, which is the apomorphic condition in the Chrysopidae, suggests that these two genera are the most advanced of the tribe. Sencera has a further specialisation in that the intramedian cell is absent. Semachrysa is the least specialised of this subgroup because the basal costal area is only slightly expanded. In Retipenna and Signochrysa the costal area is narrow basally and the basal subcostal crossvein is positioned more distally, although it is still more basal than in most other chrysopid genera. The apomorphic ankylopterygine characters also occur in the Chrysopodes canudasi Navas group of species and initially we had considered this group to be related to the Ankylopterygini. However, after fully surveying the male genitalia of Chrysopodes it has become evident that it is a very diverse genus and also includes species such as C. divisa (Walker) and C. collaris which exhibit typically chrysopine features. It makes no sense to include two species groups which have very similar male genitalic characters in different tribes so we have decided to exclude Chrysopodes from the Ankylopterygini, thus implying that the canudasi group characters have evolved indepen- dently of the Ankylopterygini. Genus ANK YLOPTERYX Brauer Ankylopteryx Brauer, 1864: 899. Type species: Chrysopa venusta Hagen, by subsequ€nt desig- nation by Tjeder, 1966: 493. Ethiochrysa Fraser, 1952: 57. Type species: Ethiochrysa polychlora Fraser, by monotypy. Syn. n. DISTRIBUTION. Palaeotropics. There are 50 species and subspecies included in this large Palaeotropical genus. From Madagascar 156 S. J. BROOKS & P. C. BARNARD Figs 89-94 Ankylopteryx. 89, A. basalis, 90, 92, A. buttikaferi, 93, 94, A. venusta. 89, fore and hind wing (from Kimmins); 90, apex of d abdomen, lateral; 91, apex of 2 abdomen, lateral; 92, 3 genitalia, lateral; 93, 2 spermatheca, lateral; 94, 2 subgenitale, caudal. and the Afrotropics 18 species have been des- cribed, eight species are known from India and southern China, 20 from the Oriental region, with a further four Australian species. DiAGnosis. Adult. Small to medium lacewings; fore wing (Fig. 89) 9-17 mm; ground colour pale green. Head marked with black or red stripes on clypeus, gena or frons; palps elongate; labrum indented; mandibles scythe-like, symmetrical, without basal teeth; vertex slightly raised; head narrow (head width : eye width 1.9-2.6 : 1); antenna about as long as fore wing; flagellar segments 3-4 times as long as wide with four rings of setae. Pronotum narrow, sometimes marked with black lateral spot; setae long, pale, fine; mesonotum sometimes with broad black marking; metanotum unmarked. Legs often marked with black annulation on fore and mid tibia, apex of Sth tarsal segment usually black; setae long, pale; claws with basal dilation. Fore wing very broad (length : breadth = 2.1—2.5 : 1); marked with large black spots or veins with dark brown suffusion; costa usually unmarked at base; costal area at base of fore wing broad; costal setae long, erect; Sc very short; stigma often marked with black spot; Sc and R very close; basal Sc crossvein 0.08-0.12 mm; im present or absent; m, with strongly arched costal margin; Rs strongly sinuate; gradates in two slightly divergent rows, basal inner gradate meet- ing Psm; veins not crassate in d; c; shorter than C2; C7 Narrow, rounded apically. Hind wing narrow (length : breadth = 3.0-4.0 : 1). Abdomen (Figs 90, 91) unmarked; setae long, sparse; callus cerci ovate, 17-24 trichobothria; ectoprocts fused dor- sally with slight dorsal invagination; d: sternite 8+9 fused; microtholi absent; 2: sternite 7 straight apically with small setose apical tubercle. GENITALIA 6 (Fig. 92). Tignum, gonapsis and median plate absent; gonarcus long, arcuate; entoprocessus long, usually fused apically; arces- sus absent; pseudopenis narrow, tapering apically; gonosaccus long with few gonosetae; gonocristae and spinellae absent. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 157 GENITALIA 2 (Figs 93, 94). Praegenitale absent; subgenitale bilobed apically; spermatheca broad; ventral impression absent or very small; duct long, coiled; vela absent or very small. Larva. The presumed larva of the subgenus Sencera is described below. REMARKS. Ankylopteryx can be distinguished from other ankylopterygine genera by the pres- ence of a pseudopenis in the male genitalia; in all other genera in the tribe an arcessus is present rather than a pseudopenis. It is probably closely related to Parankylopteryx with which it shares several apomorphies. Both genera have a broad, setose fore wing marked with numerous black spots and females of both genera have a very small ventral impression and vela. However, Ankylopteryx can be distinguished from Paranky- lopteryx by the black tip to the tarsi, the unmarked base of the costa, and the long, apically fused entoprocessus. From Fraser’s (1952) figure of the fore wing it is apparent that Ethiochrysa is a synonym of Ankylopteryx. The type species of Ethiochrysa, E. polychlora Fraser, appears to be closely related to Ankylopteryx decorsei. Unfortunately, we were unable to locate the type of E. polychlora in the MNHN, Paris collections. BIOLOGY. There were no insect remains in the guts of any of the adults examined during this study. Subgenus ANK YLOPTERYX Brauer There are two major species groups in Ankylop- teryx s.str.: those, including A. venusta (Hagen), which have the wings marked with large black spots; and a smaller group, including A. poly- gramma Gerstaecker, in which the wing veins are suffused with dark markings but without definite spots. Ankylopteryx s.str. is apparently very closely related to subgenus Sencera, from which the generalized pattern of the male and female genitalia are indistinguishable. Species of Sencera may be separated from Ankylopteryx s.str. only by the absence of the intramedian cell in the fore wing. However, this condition is approached in some Ankylopteryx species, such as A. doleschali Brauer and A. obliqua Banks, where cell im is much reduced in size. Subgenus SENCERA Navas stat. n. Sencera Navas, 1925a: 26. Type species: Sencera scioneura Navas, by original designation and monotypy. DISTRIBUTION. Indo-Australian. The subgenus includes 4 described species and a further two undescribed species are present in the BMNH collections. The description of Sencera is the same as for the genus Ankylopteryx except in the following characters. Fore wing (Fig. 95) 10-11 mm. Head width : eye width = 1.8-1.9 : 1. Fore wing broad, length : breadth = 2.2-2.6 : 1; im absent. Abdomen (Figs 96, 97) unmarked or with black dorsal stripes. The male genitalia (Fig. 98) and female genitalia (Figs 99, 100) are as described for the genus. Larva. Description based on an unreared larva from Papua New Guinea collected by Dr N. D. Penny on croton bushes. Similar larvae taken at the same time were reared and found to be Sencera scioneura Navas. Abdomen narrow, fusi- form, humped; thoracic tubercles long and cylin- drical, bearing very long, plumose setae; transverse row of four long setae on meso- and metanotum; abdominal tubercles short and broad, with long plumose setae; dorsal setae plumose, hooked apically; dorso-lateral chalazae present; large debris packet carried. REMARKS. Sencera is very similar to Ankylopteryx s.str. and can be distinguished only by the absence of the intramedian cell in the fore wing; for this reason it can be considered as no more than a subgenus of Ankylopteryx. BIOLoGy. Adults examined lacked insect remains in the gut contents. The presumed larva of S. scioneura is described above. The pupa of an undescribed species reared from Sulawesi was covered in a large amount of debris which confirms that the larva is a debris carrier. Genus PARANKYLOPTERYX Tjeder Stat. n. Parankylopteryx Tjeder, 1966: 508 [as subgenus of Ankylopteryx Brauer]. Type species: Ankylop- teryx neavei Navas, by original designation and monotypy. DISTRIBUTION. Afrotropics and Madagascar. The genus includes 10 described species. DIAGNOsIs. Adults. Small to medium lacewings, fore wing (Fig. 101) 9-14 mm. Head marked with black stripes on gena, clypeus, labrum, frons, scape, vertex, post-ocular region; palps elongate; labrum indented; mandibles narrow, symmetrical without basal teeth; vertex flattened; head width : eye width = 1.9-2.1: 1, head narrow; antenna 1.5 158 99 Figs 95-100 Ankylopteryx (Sencera) scioneura. 95, fore wing (from Kimmins); 96, apex of d abdomen, lateral; 97, apex of 2 abdomen, lateral; 98, d genitalia, lateral; 99, 2 spermatheca, lateral; 100, 2 subgenitale, ventral. times length of fore wing; flagellar segments about 6 times as long as broad; setae arranged in four rings. Pronotum unmarked; dorsal setae very long, pale; mesonotum marked with broad black stripe; metanotum unmarked. Legs unmarked; setae very long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.3-2.4 : 1); marked with black spots, particularly at base of inner gradate series, base of costa black; costal area broad at base; costal setae long, erect; Sc short; stigma unmarked; Sc and R close; basal Sc crossvein positioned slightly basally of mj —m, crossvein (—0.04 mm); m, very short; im short, four or five-sided, occasionally ovate, costal margin angled; Rs sinuate; gradates in two parallel or divergent series; basal inner gradate meeting Psm; veins not crassate in 6; c; shorter than c); fork of 1A short; dcc open or closed at posterior margin. Hind wing narrow (length : breadth = 3.3-3.8 : 1); marked with black spots on gradates and marginal veins. Abdomen (Figs 102, 103) unmarked; setae long, sparse; callus cerci S. J. BROOKS & P. C. BARNARD ‘ - rounded; trichobothria 16-21; ectoprocts with slight apical invagination, fused dorsally, fused to tergite 9; atria large; d: sternite 8+9 fused; microtholi absent; °: sternite 7 straight apically. GENITALIA 6 (Fig. 104). Tignum, gonapsis and median plate absent; entoprocessus short, broad; parameres absent; gonarcus narrow, long; arces- sus ‘bone’-shaped; pseudopenis absent; gonosac- cus long, paired; gonosetae long, numerous; gonocristae, spinellae absent. GENITALIA 2 (Figs 105, 106). Praegenitale absent; subgenitale bilobed apically with deep invagination, extended basally; spermatheca broad; ventral impression absent; vela absent; duct short, curved. REMARKS. Species of Parankylopteryx can be distinguished from those of Ankylopteryx, which they closely resemble, by the unmarked tarsi, black base of the costa, the black spot on the basal inner gradates and the black post-ocular spot. In the male genitalia an arcessus is present and THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 159 Figs 101-106 Parankylopteryx neavei. 101, fore wing (from Kimmins); 102, apex of d abdomen; 103, apex of ? abdomen, lateral; 104, d genitalia, lateral; 105, 2 spermatheca, lateral; 106, 2 subgenitale. pseudopenis absent, and the entoprocessus are short and not fused apically with each other. In | Ankylopteryx a pseudopenis is present and | arcessus absent, and the entoprocessus are long and fused apically. Although Parankylopteryx is apparently closely related to Ankylopteryx, these | differences indicate that the genera are suf- | ficiently distantly related to regard them as separ- _ ate taxa. ' BroLocy. Unknown. Adults examined in this _ study did not have insect remains in their gut _ contents. | Genus RETIPENNA Brooks Retipenna Brooks, 1986: 36. Type species: | Chrysopa notata Navas, by original designation. | DISTRIBUTION. India, South East Asia. Eight species of Retipenna have been described. | R. notata (Navas) is the most widespread, occupy- | ing most of the generic range, the other species being apparently more localized. | Diacnosis. Adult. Large lacewings, fore wing (Fig. 107) 15-21 mm; ground colour pale green. Head unmarked or with brown markings on gena, scape, vertex; palps elongate; galea long; man- dibles narrow, symmetrical without basal teeth; labrum emarginate; head width : eye width = 2.0-2.1 : 1; vertex slightly raised; antenna shorter than fore wing; flagellar segments about 3 times as long as broad; setae arranged in four rings. Pronotum unmarked or with brown lateral spots; dorsal setae long, pale; meso- and metanotum unmarked or with brown spots. Legs with long, pale setae; claws with basal dilation. Fore wing usually broad (length : breadth = 2.7-3.0 : 1); unmarked or with dark suffusion on gradates, spot on dcc; costal area narrow at base; costal setae long, inclined; Sc and R widely separated; Sc long; stigma unmarked; basal Sc crossvein 0.2—-0.4 mm; radial crossveins straight; im small, ovate; grad- ates in two divergent series; inner basal gradate meeting Psm or basally extended and not meeting Psm; Rs sinuate; c, shorter than cy. Hind wing length : breadth = 3.0-3.4 : 1, narrow. Abdomen (Figs 108, 109) long, unmarked; setae long, sparse; callus cerci ovate; trichobothria 26-31; ectoprocts with deep apical invagination; - 146 Figs 136-146 Calochrysa extranea. 136, fore wing (from Kimmins); 137, mandibles, dorsal; 138, galea, dorsal; 139, apex of d abdomen, lateral; 140, sternite 8+9, ventral; 141, apex of 2 abdomen, lateral; 142, d gonarcus complex, dorsal; 143, d parameres, dorsal; 144, 2 subgenitale and praegenitale, ventral; 145, 2 subgenitale, caudal; 146, 2 spermatheca, lateral. REMARKS. Belonopteryx can easily be distin- guished from other Chrysopidae by the lanceolate wings, marked with two black longitudinal stripes, the concave costal margin and absence of an intra- median cell. It differs from most Belonopterygini in primitively lacking parameres but retaining entoprocessus and in having extraordinarily dense microtholi. The genus is probably most closely related to Nacarina, species of which also lack parameres, but which, like Belonopteryx, have only three rows of setae on the flagellar segments, an unstable venation with many forked and irregularly placed crossveins, and a short second medial cell. Belonopteryx is apparently unique among the Chrysopidae in having a broad but untoothed left mandible. Due to the rarity of specimens of B. arteriosa we have not made any head preparations so have been unable to examine the right mandible but it is unlikely that this possesses a tooth either. BIoLoGy. Unknown. Genus CALOCHRYSA Banks Calochrysa Banks, 1943: 100. Type-species: Chrysopa extranea Esben-Petersen, by original designation and monotypy 168 DISTRIBUTION. Australia. C. extranea Esben-Petersen is the only species included in Calochrysa and it occurs in most regions of Australia. DIAGNOSIS. Adult. Large lacewings, fore wing (Fig. 136) 14-23 mm; ground colour pale yellow- ish green. Head marked with dark stripe between antennae; palps rounded apically; labrum in- dented; mandibles broad (Fig. 137), symmetrical with basal tooth on both mandibles; galea broad (Fig. 138); vertex slightly raised; head width : eye width = 2.02.3 : 1; antenna slightly longer than fore wing; flagellar segments almost as long as wide, with setae arranged in four rings. Pronotum broad, marked with narrow black lateral stripes; setae short, dark; mesonotum marked with black stripes; metanotum unmarked. Legs unmarked; setae short, dark; claws without basal dilation. Forewing unmarked; costal area narrow at base; costal setae short, inclined; stigma unmarked; Sc moderately long; Sc and R widely separated; basal Sc crossvein 0.52—-0.72 mm; mp short; im broad, ovate; Rs sinuate; gradates in two parallel series, basal inner gradate meeting Psm; veins not crass- ate in 6; c, 1.5 times longer than c); c> slightly broadened apically; Cu, forked. Abdomen (Figs 139-141) unmarked; setae short, dense; ecto- procts with slight dorso-apical invagination; ectoprocts fused with tergite 9; ¢: callus cerci ovate, trichobothria 39; sternite 8+-9 fused, short, indented apically; microtholi absent; 2: callus cerci rounded, trichobothria 48; sternite 7 in- dented apically. GENITALIA 6 (Figs 142, 143). Tignum, gonapsis, median plate and entoprocessus absent; para- meres paired, narrow, tapering apically; gonarcus short, broad, transverse; arcessus broad with apical hook; pseudopenis absent; gonosaccus, gonosetae, spinellae and gonocristae absent. GENITALIA ¢ (Figs 144-146). Praegenitale pres- ent; subgenitale broad, bilobed apically; sperma- theca narrow; ventral impression deep; vela long; duct long, tightly coiled. Larva (first instar only). Mandibles short, curved; abdomen fusiform, not humped; thoracic tuber- cles cylindrical; abdominal tubercles undeveloped; setae rounded apically, not hooked; oviruptor with anterior process considerably enlarged. REMARKS. Calochrysa is the only genus in the Chrysopidae in which vein Cu? is forked and this is sufficient reason to continue to regard the genus as distinct. Banks (1943) included Calochrysa in the Nadivini (= Belonopterygini) and it is clearly related to /talochrysa Principi. New (1986a) suggested that the oviruptor, with its enlarged anterior process, was characteristic of the Italo- S. J. BROOKS & P. C. BARNARD chrysini (= Belonopterygini). Other apomorphies include symmetrical toothed mandibles and a bulbous lobe at the apex of the male ectoprocts. BIoLoGy. The first instar larva was described and figured by New (1986a). The larva shares many characters with the larvae of Italochrysa Principi and for this reason it is possible that those of Calochrysa are similarly specialized to have a close association with ants (Principi, 1946); indeed this may be a characteristic of the tribe. This may be the reason why New could not keep the larvae alive beyond the first instar. Genus CHRYSACANTHIA Lacroix Chrysacanthia Lacroix, 1923: 120. Type species: Chrysacanthia esbeniana Lacroix, by monotypy. Nesochrysa Fraser, 1951: 29. Type species: Nesochrysa varicella Fraser, by monotypy. [Homonym of Nesochrysa Navas, 1910a: 53; synonymized with Glenochrysa Esben-Petersen by Fraser, 1955: 134.] Syn. n. DISTRIBUTION. Madagascar, west and central Africa, southern India. The genus includes two described species and there are a further two undescribed species in the BMNH collections. DIAGNOsIS. Adult. Medium to large lacewings, fore wing (Fig. 147) 14-18 mm; ground colour brown. Head, scape almost entirely marked dark brown, with lateral pale brown stripe on vertex; palps rounded apically; labrum emarginate; vertex raised; head width : eye width = 2.02.3: 1; scape slightly elongate; antenna pale or dark brown, longer than fore wing; flagellar segments twice as long as broad; setae arranged in four rings. Pronotum broad; marked with narrow, submedian pale brown longitudinal stripe; dorsal setae long, dark; meso- and metanotum almost entirely dark brown, with pale brown suture lines. Legs marked with dark brown spots and stripes on tibiae and femora; setae long or short, pale or dark; claws with basal dilation. Fore wing marked extensively with large pale brown spots; costal area narrow at base; costal setae quite long, inclined; stigma marked dark brown; Sc and R widely separated; basal Sc crossvein 0.56— 0.64 mm; im short, broad, ovate; mz short; Rs slightly sinuate; gradates in two divergent series; basal inner gradate meeting Psm; veins not crassate in 6; c, 1.5—2.0 times longer than c); c, broad, squared apically. Hind wing marked with large pale brown spots on stigma, 2nd posterior marginal crossvein and minute spot at wing apex. Abdomen (Figs 148, 149) marked dark brown on THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 169 Se AA Wu lyy by 1) ty \ i i rl 151 Ty Ea ae SRS Ges 153 152 Figs 147-153 Chrysacanthia. 147, C. esbeniana; 148, 150, C. sp. indet.; 149, 152, 153, C. varicella. 147, fore wing; 148, apex of d abdomen, lateral; 149, apex of 2 abdomen, lateral; 150, d gonarcus complex, dorsal; 151, ¢ parameres, lateral; 152, 2 subgenitale, caudal; 153, 2 spermatheca, lateral. ectoprocts, tergites 4-8 and sternites 7-9; setae short, dense; membranal microsetae on segments 6-8 very coarse and dense but absent from tergite 8; callus cerci rounded, prominent; trichobothria 30-36; ectoprocts with deep apical invagination, not fused dorsally, fused with tergite 9; usually short; Rs sinuate; gradates in two (sometimes three) parallel, occasionally divergent, series; inner gradates extended basally; basal inner gradate usually meeting Psm; veins not crassate in 6; c, 1.2-1.7 times longer than c); venation unstable, number of crossveins and forks varies between individuals; posterior marginal veins forked. Hind wing broad (length : breadth = 2.9 : 1); anal veins sometimes forked; anal area broad. Abdomen (Figs 180, 181) sometimes marked with red dorsal stripe; setae short, dense; trichobothria 48-60; ectoprocts with moderate or deep apical invagination, fused dorsally; sternites enlarged; ectoprocts fused with tergite 9; d: sternites with microtholi; callus cerci ovate; sternite 8+9 fused, squared apically with median indentation; @: callus cerci rounded; sternite 7 straight apically or, in N. titan (Banks), deeply excised with pair of submedian lobes. GENITALIA ¢ (Fig. 182). Tignum, gonapsis, median plate and entoprocessus absent; arcessus short with apical hook; parameres absent; gonar- cus short, broad with lateral gonocornua; pseudo- penis absent; gonosaccus short; gonosetae long or short, in small lateral clump; spinellae and gonocristae absent. GENITALIA ? (Figs 183-186). Praegenitale present (in N. titan along membranous tongue is present); subgenitale bilobed apically, sometimes with median projection, base broadly expanded, some- times downcurved; spermatheca broad; ventral impression deep; vela long; duct long, sinuous. REMARKS. Nacarina can easily be distinguished from other genera by the broad, rounded apical segment of the palps, the broad flagellar seg- ments, the antennae which are positioned very close together, the very broad pronotum, the short rectangular intramedian cell and the forked posterior marginal veins. Nacarina is unusual amongst the Chrysopidae in showing marked sexual dimorphism in which males are consider- ably smaller than females. The lack of parameres in males and the unstable venation of Nacarina indicate a close relation- ship with Belonopteryx Gerstaecker. However, Nacarina also shares several characters with Evanochrysa which, intriguingly, suggests a link | between these genera. In both genera the intra- median cell is short and broad, and the forking | marginal crossveins and gradates are black. In males microtholi are present, the arcessus is short and broad with an apical hook, the gonarcus has long lateral gonocornua, gonosetae are present and parameres are absent. In females a praegeni- 177 tale close to the base of the subgenitale is present; the long membranous tube in N. titan is similar to that of Abachrysa. New (1984) treated Nacarina and Nadiva as separate taxa but there does not seem to be any justification for this. Banks (1943) erected the tribe Nadivini in which he also included Calochrysa Banks. The holotype of Mesochrysa megaptera, the type species of Mesochrysa, was deposited in Navas’s collection but we were unable to locate it; it is not listed by Monserrat (1985) and so is probably lost. However, Navas (1927a) figured the head, thorax and base of the fore wing of M. megaptera. A red stripe is shown on the scape, vertex and laterally on the pronotum, the claws are not dilated basally, cell im is broadly ovate, c> shorter than c,; and the basal Sc crossvein is relatively distal. Navas compared Mesochrysa with Nothochrysa (= Italochrysa Principi) but cell im is quadrangular in /talochrysa. He also com- pared Mesochrysa with Nodita Banks but, unlike the latter genus, M. megaptera has robust antennae which are shorter than the fore wing and the pterostigma is unmarked. These characters suggest that Mesochrysa is a belonopterygine and so is probably synonymous with Nacarina since the other New World belonopterygine genera are not marked in this way. BIOLOGY. The gut contents of adults examined during this study did not include insect remains. Weber (1942) reported that the larvae of Nacarina valida (Erichson) are myrmecophiles. Genus NESOCHRYSA Navas Nesochrysa Navas, 1910a: 53. Type species: Nesochrysa grandidieri Navas, by original designation and monotypy. Oviedus Navas, 1913b: 326. Type species: Oviedus auricollis Navas, by monotypy. Syn. n. Madachrysa Navas, 1934: 62. Type species: Madachrysa seyrigi Navas, by original designa- tion and monotypy. Syn. n. DISTRIBUTION. Madagascar, Afrotropics. The genus includes 10 described species, three of which occur in Madagascar. N. macrostigma Gerstaecker is widely distributed over central and west Africa. DIAGNosIs. Adult. Large lacewings, fore wing (Fig. 187) 16-23 mm; ground colour brown. Head with red or black stripe on labrum, scape, vertex; palps broad, pointed apically; mandibles broad, asymmetrical with basal tooth on left mandible; galea broad with small apical papilla; labrum 178 190 S. J. BROOKS & P. C. BARNARD Figs 187-194 Nesochrysa. 187, 188, 190-194, N. seyrigi; 189, N. macrostigma. 187, fore wing (from Kimmins); 188, apex of d abdomen, lateral; 189, apex of 2 abdomen, lateral; 190, d parameres, ventral; 191, d gonarcus complex, dorsal; 192, 2 spermatheca, lateral; 193, 2 praegenitale, lateral; 194, 2 subgenitale and praegenitale, ventral. deeply indented; vertex slightly raised; toruli large; head width : eye width = 2.02.8 : 1; antenna considerably longer than fore wing; flagellar segments about twice as long as broad; setae arranged in four rings. Pronotum broad; marked with broad red or black lateral band; dorsal setae long, pale; meso- and metanotum entirely red or brown. Legs unmarked or with black stripe on tibia and femur; setae long or short, dark; claws with basal dilation. Fore wing broad or narrow (length : breadth = 2.2-3.1 : 1); unmarked; costal area narrow at base; costal setae long, inclined; stigma marked red/brown; Sc and R widely separated; basal Sc crossvein 0.52—-0.88 mm; m, very short; im absent or long, quadrangular, rarely ovate; Rs slightly sinuate; gradates in two parallel or slightly divergent series; inner gradates greatly extended basally, meeting Psm; veins not crassate in d; c, up to 1.5 times longer than cz; c broad, squared apically. Abdomen (Figs 188, 189) with red/brown or black dorsal markings; setae short, dense on tergites but long, sparse on sternites; callus cerci ovate or rounded, prominent; trichobothria 29-65; ecto- procts with deep apical invagination, not fused dorsally, fused with tergite 9; atria large; 3d: sternites with microtholi; sternite 8+9 fused, short, broad, often with long lateral lobes; tergite 9 often with lateral lobe; ectoprocts often with ventral trunk-like process; 2: setae short, dense; sternite 7 straight or slightly indented apically. GENITALIA 6 (Figs 190, 191). Tignum, gonapsis, median plate and entoprocessus absent; gonarcus short, broad, transverse with long lateral gono- cornua sometimes terminating in broad swelling; arcessus very short, broad with median hook; pseudopenis, gonosaccus, gonosetae, spinellae and gonocristae absent; three pairs of long, ‘Narrow parameres fused to arcuate basal plate. GENITALIA 2 (Figs 192-194). Praegenitale pres- ent; subgenitale bilobed apically, broad; sperma- theca long, broad; ventral impression deep; vela moderate; duct short, sinuous. REMARKS. Species of Nesochrysa can be dis- tinguished from most other belonopterygine genera by their very long antennae, pointed apex of the palps, the short cell m, and the dark brown spot on the pterostigma. Males of the genus are _ characterized by the long lateral gonocornua and the three pairs of parameres. | Nesochrysa closely resembles Stigmachrysa | Navas which has similar external characteristics, / with a marked stigma and lobes on the ectoprocts. However, Stigmachrysa differs from Nesochrysa / in that males have only one pair of parameres. __ There are two species groups within Nesochrysa. _M. seyrigi and macrostigma have broad fore wings | (length : breadth < 2.5 : 1); the costal area in the | stigmatic region is broad; the stigma is marked with a large black spot; c) is very broad; males _ have trunk-like apical lobes on the ectoprocts and | sternite 8+9; the gonocornua have a broad apical | swelling; the spermatheca is broad and sternite 7 is | straight apically in the female. In contrast, the | other species have relatively narrow fore wings ‘(length : breadth > 2.5 : 1); the costal area is _ narrow; the stigma is marked with only a small | spot; C2 is only slightly broader than c,; males have 'no lobes on the ectoprocts or sternite 8+9; the | gonocornua taper apically; the spermatheca is narrow; sternite 7 is concave apically in females. When Tjeder (1973) redescribed Madachrysa he noted that it was very similar to Nesochrysa and Oviedus but hesitated to synonymize them. How- ever, the characters which separate them are trivial and the shared apomorphies of the male genitalia indicate that they are so closely related that synonymy seems to be the only reasonable option. : | | BIOLOGY. Unknown. No gut contents were present in any of the adult specimens examined during this study. i THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 179 Genus NODOCHRYSA Banks Nodochrysa Banks, 1938a: 226 [as subgenus of Chrysopa Leach]. Type species: Chrysopa necrota Banks, by monotypy. [Raised to genus by Brooks, 1984: 80.] DISTRIBUTION. Singapore, Malaysia, Brunei. Nodochrysa is monotypic. D1aGnosis. Adult. Large lacewings, fore wing (Fig. 195) 12-15 mm. Ground colour in life orange/brown (G. S. Robinson, pers. comm.). Head marked with black stripe on vertex and between antennae; palps narrow, rounded; labrum slightly indented; vertex flat; toruli large; head width : eye width = 2.0-2.2 : 1; scape broad; antenna 1.5 times longer than fore wing; flagellar segments twice as long as broad; setae arranged in four rings. Pronotum broad, periphery marked black with black median stripe; lateral setae long, pale; dorsal setae few, black, short; mesonotum with black stripe on suture; metanotum marked with black spot above wing base. Legs with brown stripe on tibia; setae long, pale; claws with basal dilation. Fore wing length : breadth = 2.6-3.0: 1; marked with brown shading on basal crossveins; costal area very narrow at base; costal setae short, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.44—0.52 mm; im broad, ovate; Rs sinuate; gradates in two parallel series; basal inner gradate meeting Psm; veins not crassate in 6; c, 1.5—2.0 times as long as c); c) slightly broadened, squared apically. Abdomen (Figs 196, 197) marked with large dark brown spots on tergites 1, 2, 5, 6 and sternites 6-9; setae short, sparse; callus cerci rounded; trichobothria 30-33; ectoprocts with slight apico-dorsal invagi- nation, fused with tergite 9; d: microtholi present on sternites 3-8; ectoprocts not fused dorsally, elongated apico-ventrally; sternite 8+9 fused, short with lateral tubercle and apical membrane bearing enlarged setal bases; 2 : sternite 7 straight apically with pair of short, subapical spines. GENITALIA o¢ (Fig. 198). Tignum, gonapsis, median plate and entoprocessus absent; para- meres short, U-shaped, slightly swollen apically with three small apical teeth; gonarcus long, narrow with lateral gonocornua; arcessus small, membranous with strong apical hook and blunt lateral projection; gonosaccus, gonosetae, gono- cristae and spinellae absent. GENITALIA ¢ (Figs 199, 200). Praegenitale present; subgenitale broad, bilobed at apex; spermatheca narrow; ventral impression deep; vela very broad, long; duct short, coiled. Larva. Unknown. =SS3 i Wey REMARKS. Species of Nodochrysa have a broad, ovate intramedian cell and very long antennae. Males of the genus are characterized by the lateral lobes on sternite 8+ 9 and the trunk-like extension of the ectoprocts. Females of Nodochrysa are readily distinguished from other genera by the short, subapical spines on sternite 7. The long antennae, narrow palps and lobes on the ectoproct and sternite 8+9 of males suggest that Nodochrysa may be related to Nesochrysa Navas. Nodochrysa was only briefly described in a key couplet (Banks, 1938a) and since then appears to have been overlooked by subsequent workers. However, the unusual male and female abdomi- nal characters indicate the validity of the genus. The short second cubital cell, broad flagellar segments and relatively distal position of the basal subcostal crossvein place the genus in the Belonopterygini and suggest only a distant S. J. BROOKS & P. C. BARNARD 198 Figs 195-200 Nodochrysa necrota. 195, fore wing; 196, apex of d abdomen, lateral; 197, apex of 2 abdomen and praegenitale, lateral; 198, d genitalia, caudal; 199, 2 spermatheca, lateral; 200, 2 subgenitale, ventral. relationship with Chrysopa Leach, therefore justifying the elevation of Nodochrysa to full generic status. BioLoGy. Unknown. The gut contents of adults examined during this study did not include insect remains. Genus OYOCHRYSA Brooks Oyochrysa Brooks, 1984: 80. Type species: Oyochrysa ancora Brooks, by _ original designation. DISTRIBUTION. West Africa. The genus is known from three species. DIAGNOsIS. Adult. Medium to large lacewings, fore wing (Fig. 201) 14-19 mm; ground colour brown. Head with extensive red or brown mark- ings; palps rounded apically; galea broad; labrum THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 181 ERA | Figs 201-207 Oyochrysa. 201, O. sanguinea, fore wing; 202-207, O. ancora; (202) apex of d abdomen, lateral; (203) | apex of 2 abdomen, lateral; (204) ¢ parameres, dorsal; (205) ¢ gonarcus complex, lateral; (206) 2 spermatheca, lateral; (207) 2 subgenitale, ventral. emarginate; mandibles broad, asymmetrical with basal tooth on left mandible; head width : eye width = 1.8-2.1 : 1; eyes large; vertex flat; toruli large; antenna slightly longer than fore wing; flagellar segments less than twice as long as broad; setae arranged in four rings. Pronotum broad, with extensive dark brown markings; meso- and metanotum marked dark brown. Legs marked with red/brown stripes on tibiae and femora; setae short, dark; claws with basal dilation. Fore wing marked in basal half with brown or red spots; costal area narrow at base; costal setae short, inclined; Sc and R widely separated; basal Sc crossvein 0.72—-0.76 mm; im quadrangular, broad; Rs sinuate; veins not crassate in 6; c; 1.5-3.0 times longer than cz; gradates in two parallel series; basal inner gradate meeting Psm; stigma marked with brown spot. Abdomen (Figs 202, 203) with prominent dark dorsal markings; setae short, dense; callus cerci rounded; trichobothria 32-49; ectoprocts with slight dorso-apical invagi- nation, not fused dorsally, fused with tergite 9; d: sternite 8+9 fused; sternites and tergites with microtholi; 2 sternite 7 with broad apico-dorsal invagination. GENITALIA 6 (Figs 204, 205). Tignum, gonapsis and median plate absent; gonarcus long, narrow; parameres broad, deeply bifid apically; ento- S. J. BROOKS & P. C. BARNARD S214 Figs 208-214 Stigmachrysa. 208-211, 213, 214, S. cladostigma; 212, S. kervillei. 208, fore wing (from Kimmins); 209, apex of ¢ abdomen, lateral; 210, apex of 2 abdomen, lateral; 211, d genitalia, dorsal; 212, 2 spermatheca, lateral; 213, 2 subgenitale and praegenitale, caudal; 214, 2 subgenitale and praegenitale, ventral. processus absent; arcessus short, with apical tooth, situated above membranous sac; pseudo- penis absent; gonosaccus short; gonosetae absent. GENITALIA @ (Figs 206, 207). Praegenitale absent; subgenitale broad, bilobed apically, con- nected to abdomen by long membranous tube supported by sclerotized apical extension of sternite 7; spermatheca large, strongly sclerot- ized; vela large; ventral impression deep; duct long, coiled. Larva. Unknown. REMARKS. Oyochrysa shares many characters with Italochrysa Principi, to which it is probably closely related, but can be readily distinguished by the long, arcuate gonarcus in the male and the long apical extension of sternite 7 in the female. In Italochrysa, and the rest of the Belonopterygini, the gonarcus is short and transverse and sternite 7 is not extended in the female. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 183 BIoLoGy. Unknown. The gut contents of adults examined during this study did not include insect remains. Genus STIGMACHRYSA Navas Stigmachrysa Navas, 1925b: 570. Type species: Stigmachrysa kervillei Navas, by original desig- nation and monotypy. DISTRIBUTION. India, Burma, Singapore, Java. Three species have been described but a fourth undescribed species is present in the Institut Royal des Sciences Naturelles de Belgique, Brussels. DIAGNOsIs. Adult. Large lacewings, fore wing (Fig. 208) 20-23 mm; ground colour brown. Head unmarked or with dark spot below anten- nae; palps rounded; labrum invaginated; vertex slightly raised; toruli large; head width : eye width = 2.02.4 : 1; antenna as long as fore wing; flagellar segments about 1.5 times as long as broad; setae arranged in four rings. Pronotum broad; unmarked; dorsal setae long, pale; meso- and metanotum unmarked or entirely black. Legs unmarked; setae long or short, pale; claws with basal dilation. Fore wing broad; unmarked or with shading in anal region; costal area narrow at base; costal setae short, inclined; Sc long; stigma marked brown; Sc and R widely separated, particularly at stigma; basal Sc crossvein 0.60— 0.72 mm; im quadrangular, broad, short; Rs slightly sinuate; gradates in two parallel or diver- gent rows; inner gradate series irregular with additional crossveins; inner gradates extended basally, meeting Psm; veins not crassate in d; c, 2.0-2.5 times longer than c,; c) broad, squared apically. Hind wing with stigma marked brown. Abdomen (Figs 209, 210) unmarked or with brown spot on sternite 7; trichobothria 41-65; ectoprocts with narrow apical invagination, not fused dorsally, fused with tergite 9; ¢: setae short, dense; sternites with microtholi; callus cerci ovate, prominent; ectoproct with ventro-apical projection; sternite 8+9 fused; ?: setae long, sparse but short on tergites 4-5; callus cerci rounded; sternite 7 broadly concave apically. GENITALIA 6 (Fig. 211). Tignum, gonapsis and median plate absent; entoprocessus absent; para- meres short, arcuate bearing short setae; gonarcus short, broad with long, tapering lateral gono- cornua; arcessus short, broad with hooked apex; pseudopenis absent; gonosaccus short; gonosetae absent; gonocristae present in small group at apex of sternite 8+9; spinellae absent. GENITALIA ¢ (Figs 212-214). Praegenitale pres- ent; subgenitale broad, bilobed apically with small median projection and basal crumena; sperma- theca short, broad; ventral impression very deep; vela enormously elongated, highly coiled; duct long, sinuate. Larva. Unknown. REMARKS. Species of Stigmachrysa are super- ficially similar to those of Nesochrysa Navas since both genera have a large dark brown spot on the pterostigma and a basally extended inner gradate series. However, they can be distinguished because males of Stigmachrysa have a single short pair of parameres, unlike the three pairs in Nesochrysa, and there is a small median pro- jection at the apex of the subgenitale in females of Stigmachrysa which is absent in Nesochrysa. Nevertheless, Stigmachrysa is probably closely related to Nesochrysa since both genera have an apical lobe on the male ectoprocts. BIioLoGy. Unknown. There were no insect re- mains in the guts of any adults examined during this study. Genus TURNEROCHRYSA Kimmins Turnerochrysa Kimmins, 1935: 576. Type species: Turnerochrysa mirifica Kimmins, by monotypy. DISTRIBUTION. Namibia. Known only from a single species. DIAGNOsIs. Adult. 2 unknown. Small lacewings, fore wing (Fig. 215) 9-10 mm. Ground colour brown. Head marked with brown stripes on vertex; palps rounded apically; labrum indented; vertex flat; toruli large; head width : eye width = 3.1: 1, eyes small; antenna about half length of fore wing; flagellar segments about 1.5 times as long as broad; setae arranged in four rows. Pronotum very broad; marked with dark spots; dorsal setae short, dark; meso- and metanotum marked with dark spots. Legs marked with brown annulations on tibia, femora and tarsi; setae short and dark; claws without basal dilation. Fore wing broad (length : breadth = 2.4 : 1); marked with dark shading around some crossveins; costal area narrow at base; costal setae short, inclined; Sc short, stigma unmarked; Sc and R widely separ- ated; basal Sc crossvein 0.56 mm.; im broad, quadrangular; Rs straight; gradates absent; veins not crassate in d; c; 1.5 times longer than c); c, broad, squared apically; posterior marginal cross- veins very long. Hind wing very broad (length : breadth = 2.4 : 1); gradates absent. Abdomen (Fig. 216) marked with sparse black spots; setae 184 Ip ler, 4 ~ es Sr SS A KA Se Tay NAS ASSIA STS S BS Rosen wwe oS RRS \ ie < aS ATER ArES S. J. BROOKS & P. C. BARNARD 2147 218 219 Figs 215-219 Turnerochrysa mirifica. 215, fore wing (from Kimmins); 216, apex of d abdomen, lateral; 217, gonarcus complex, lateral; 218, d hypandrium internum, dorsal; 219, d paramere, lateral. very short, dense (longer on ectoprocts); micro- tholi present on sternites 3-8; callus cerci ovate; trichobothria 22; ectoprocts with slight dorsal invagination, fused with tergite 9; sternite 8+9 fused, short; apodemes short, narrow, straight; atria large. GENITALIA ¢ (Figs 217-219). Tignum, gonapsis, median plate and entoprocessus absent; arcessus short, broad with strong median tooth and long, broad lateral projections; pseudopenis absent; gonarcus short, broad, transverse with short median projection; 1 pair of short parameres tapering to apical point with dorsal subapical tooth; gonosaccus very short; gonosetae absent; gonocristae and spinellae absent; hypandrium very large. Larva. Unknown. REMARKS. Turnerochrysa seems to be a highly derived genus, and may be distinguished from other Belonopterygini by the lack of gradate crossveins and the short, broad wings with a broad posterior margin. BIoLoGy. Unknown. Insect remains were not present in the adult gut contents examined. Tribe CHRYSOPINI Schneider Chrysopina Schneider, 1851: 35. Type genus: Chrysopa Leach. Crisopinos Navas, 1910a: 59. Chrysopiscini Navas, 1910a: 59. Type genus: Chrysopisca Mc Lachlan. Suarini Navas, 1914a: 73. Type genus: Suarius Navas. Chrysopini Navas, 1914a: 76. DISTRIBUTION. World-wide. DIAGNOSIS. Small to large lacewings, fore wing 9-25 mm; palps tapered apically (Fig. 14); galea broad or narrow; mandibles broad with basal tooth on left mandible (Fig. 18), sometimes tooth also present on right mandible, exceptionally mandibles scythe-like with or without basal tooth on left mandible; toruli small; head width : eye THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 185 Figs 220-229 Anomalochrysa. 220, 223-229, A. hepatica; 221, 222, A. maclachlani. 220, fore wing (from Kimmins); 221, mandibles, dorsal; 222, galea, dorsal; 223, apex of ¢ abdomen, lateral; 224, apex of 2 abdomen, lateral; 225, 3 gonarcus complex, lateral; 226, d tignum, dorsal; 227, d gonapsis, ventral; 228, ? subgenitale, caudal; 229, spermatheca, lateral. width = 1.9-4.9 : 1; flagellar segments 2-3 times as long as broad; basal Sc crossvein in fore wing 0.16-0.52 mm; im ovate, rarely quadrangular; Psm curved posteriorly at junction with outer gradate series; c, shorter than c); abdominal setae long, sparse; ¢d: gonarcus arcuate; arcessus narrow, tapering apically; pseudopenis, tignum and gonapsis sometimes present; gonosetae present; °: praegenitale absent; spermatheca narrow. REMARKS. This is not only the largest tribe in the subfamily Chrysopinae but is also the largest in the family, comprising 31 genera. The cladistic analysis suggested that the tribe is not mono- phyletic, and it seems that further study may be needed to determine the generic relationships within this group, which is here retained as a tribe for convenience. Broadly speaking the Chrysopini corresponds to early authors’ concepts of the genus Chrysopa. Nearly all the species of any economic impor- tance are to be found in the Chrysopini, and for this reason the biologies of the genera in this tribe are the best known in the family. Genus ANOMALOCHRYSA McLachlan Anomalochrysa McLachlan, 1883: 298. Type species: A. hepatica McLachlan, by subsequent designation by Zimmerman, 1957: 95. DISTRIBUTION. Hawaiian Islands. The genus includes 22 species and subspecies all of which are endemic to the Hawaiian Islands. Nine species are confined to the island of Hawaii, 186 the other major islands each have two endemic species and the remaining five are distributed more widely throughout the archipelago. DIAGNosiIs. Adult. Medium to large lacewings, fore wing (Fig. 220) 13-20 mm. In life usually coloured green but some species are bright purple or red/brown; head usually marked with black or red stripes on frons, gena, between antennae or on vertex; palps tapered; galea broad (Fig. 222); labrum straight or indented; mandibles broad, asymmetrical with tooth on left mandible (Fig. 221); vertex raised; head broad (head width : eye width = 2.5-3.1 : 1); antenna shorter than fore wing; antennae widely separated at base; flagellar segments about three times as long as wide, with four rings of setae. Pronotum unmarked or with bright yellow median stripe, with long or short dark setae; meso- and metanotum unmarked. Legs usually unmarked, with short dark setae; claws with basal dilation. Fore wing unmarked or with brown suffusion; costal area narrow at base; costal setae short, inclined; Sc and R widely separated; basal Sc crossvein 0.28—0.36 mm.; my very long; im broad, ovate; basal fork of M (at base of im) acutely angled, thickened; Rs straight or slightly sinuous; gradates in three or more parallel rows, basal inner gradate not meeting Psm; veins sometimes crassate in d; c, usually longer than or same length as c,; dcc often closed at wing margin; posterior marginal crossveins forked in basal half of fore wing. Hind wing narrow (length : breadth = 3.44.0: 1), with three or more series of gradates. Abdomen (Figs 223, 224) with setae long, sparse; 16-35 trichobothria; ectoproct and tergite 9 fused; d: setae on tergites sometimes very long, dense, projecting basally; callus cerci very small, ovate; ectoprocts narrow, flattened caudally, fused dorsally, with apical lobe bearing short, dense setae; sternite 8+9 fused, upturned apically or projecting beyond apex of ectoprocts, often with rounded mid-dorsal swell- ing; apex of abdomen ’gaping’ (ectoprocts and sternite 8+9 widely separated), tergite 7 very small; microtholi absent; ¢: callus cerci rounded, small; sternite 7 straight apically. GENITALIA 6 (Figs 225-227). Tignum straight with median projection; gonapsis broad, cross- shaped; median plate absent; gonarcus long, narrow with short lateral lobes; entoprocessus absent; arcessus long, narrow, tapering apically; pseudopenis absent; gonosaccus small, flat; gono- setae absent but numerous microsetae present. GENITALIA @ (Figs 228-229). Subgenitale bilobed apically, sometimes on sclerotized plate; praegeni- tale absent; spermatheca narrow; ventral impres- sion absent or minute; vela large; duct long. S. J. BROOKS & P. C. BARNARD Larva. Abdomen slender, fusiform, not humped. Thoracic tubercles small, spherical, bearing a few short, unhooked setae; abdominal segments 2-8 with small lateral tubercles; setae few, short, unhooked apically; debris absent. (Description based on New, 1986b.) REMARKS. Kuwayama (1962) synonymized Bornia Navas and Chrysopidia Navas with Anomalochrysa because they all possessed more than two series of gradate crossveins. However, it is now apparent that this character has arisen many times in the Chrysopidae and does not necessarily indicate a close relationship between the genera in which it occurs. Anomalochrysa can be distinguished from most genera of Chrysopidae by the presence of at least three series of gradates in the fore and hind wing. The genus is endemic to the Hawaiian Islands and is the only one to have multiple gradate series there. The apex of the male abdomen is also distinctive since it is considerably flattened caudally, with tergite 7 greatly reduced; in addition there is a membranous lobe at the apex of the ectoprocts and sternite 8+9 is hooked apically. Although species of Anomalochrysa possess many derived characters and so are very distinc- tive, the genus seems to be most closely related to Mallada Navas, and in particular to the boninensis-group of species. Like Mallada but unlike most other chrysopine genera, males of Anomalochrysa possess both a tignum and gonapsis. The gonapsis is broad with a narrow, but deep, apical invagination and a short, narrow, median transverse bar. Morphologically, this is very similar to the gonapsis of M. boninensis (Okamoto) but dissimilar to the gonapsis of any other chrysopid and it is unlikely that this form of gonapsis would have evolved independently more than once in the family. Another character linking Anomalochrysa with the boninensis-group is the mid-lateral lobe on sternite 8+9 (Fig. 396). M. boninensis (Okamoto) and M. basalis (Walker) are both widely distributed throughout the Oriental and Pacific regions; indeed M. basalis currently occurs on the Hawaiian Islands, and it is possible that the species of Anomalochrysa origin- ally arose from a population of M. basalis which became isolated on the Hawaiian Islands. BIOLOGY. Zimmermann (1957) stated that all the species are confined to the forests. Eggs are not stalked and are laid singly or in small groups. Larvae do not carry debris (Terry, 1905; Perkins, 1913) and are marked bright pink, red, yellow and green. The cocoons are egg-shaped and quite hard. Larvae feed on sugarcane aphids, the THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 187 233 Figs 230-236 Apertochrysa. 230, A. madegassa; 231-236, A. umbrosa. 230, fore wing; 231, apex of d abdomen, lateral; 232, apex of 2 abdomen, lateral; 233, d gonarcus complex, lateral; 234, d gonapsis, ventral; 235, 2 spermatheca, lateral; 236, 2 subgenitale, caudal. | sugarcane leafhopper Perkinsiella saccharicida Kirkaldy and caterpillars of the geometrid moth genus Scotorythra. Adult gut contents include insect remains. _ Genus APERTOCHRYSA Tjeder _ Apertochrysa Tjeder, 1966: 480 [as subgenus of Chrysopa Leach]. Type species: Chrysopa umbrosa Navas, by original designation. Apertochrysa Tjeder; Hdlzel, 1973b: 341 [as subgenus of Anisochrysa Nakahara]. [Raised to genus by Tsukaguchi, 1985: 505.] DISTRIBUTION. Palaearctic, Palaeotropics. The genus includes 16 species which are scat- tered throughout the range with a somewhat discontinuous distribution. _ DiaAGnosis. Adult. Small to medium lacewings, fore wing (Fig. 230) 10-13 mm; ground colour green. Head often marked with red stripe on frons, gena, scape or vertex; palps long, narrow, tapered apically; labrum indented; mandibles broad, asymmetrical with basal tooth on left mandible; galea quite narrow; vertex raised; head width : eye width = 2.1—2.5 : 1); antenna longer or shorter than fore wing; flagellar segments 3 times as long as broad, setae arranged in four rings. Pronotum unmarked or with red lateral stripe; dorsal setae long, pale; meso- and metanotum unmarked or with pale median stripe. Legs unmarked, with setae long and pale or short and dark; claws with or without basal dilation. Fore wing unmarked; costal area narrow at base; costal setae short or long, inclined; pterostigma un- marked; Sc and R widely separated; basal Sc crossvein 0.12—0.24 mm.; im narrow, ovate; Rs sinuate; gradates in two parallel series, basal inner gradate not meeting Psm; veins sometimes crassate in d; c, shorter or equal to c,. Abdomen (Figs 231, 232) with setae long, sparse; callus cerci ovate, 27-36 trichobothria; ectoprocts fused dor- sally; d: sternite 8+9 fused, more or less elongate; microtholi absent (but present in large numbers in A. edwardsi); 2: sternite 7 straight apically. 188 GENITALIA 6 (Figs 233, 234). Tignum absent; gonapsis small, basically T-shaped; gonarcus long, narrow; median plate absent; entoprocessus with median projection; arcessus narrow, taper- ing apically, with dorsal striations; pseudopenis absent; gonosaccus long; gonosetae few, short; spinellae absent; gonocristae, when present, positioned in central clump at apex of sternite 8+9. GENITALIA ¢ (Figs 235, 236). Praegenitale absent; subgenitale bilobed apically, tapering basally; spermatheca narrow; ventral impression deep; vela large; duct long, sinuous. Larva. Abdomen broadly fusiform, humped; thoracic tubercles long, cylindrical; metanotum with row of dorsal bristles; lateral abdominal tubercles cylindrical; pair of latero-dorsal setae, arising from short projections, on tergites 6 and 7; dorsal abdominal setae hooked apically; debris packet small. REMARKS. Apertochrysa is apparently closely related to Mallada from which it is indistinguish- able on external characters. However, males of Apertochrysa can be distinguished from Mallada by the absence of a tignum, and the presence of the T-shaped gonapsis, and digitiform entopro- cessus, which are not found in Mallada. In females of Apertochrysa the subgenitale characteristically tapers basally. The costal setae are longer in Apertochrysa than in most species of Mallada and in several species the wing veins of males are crassate. BIOLOGY. In some specimens examined the adult gut contents included traces of insect remains (e.g. setae, lepidopterous scales). New (1981b) recorded that adults of A. edwardsi (Banks) fed on Psylla (Psyllidae) and Brevicoryne (Aphididae) in the laboratory. A. edwardsi is the only species assigned to Apertochrysa in which the larva has been des- cribed (Boros, 1984). It does not differ in any significant characters from the larvae of Mallada species. Genus ATLANTOCHRYSA Hoizel Atlantochrysa Holzel, 1970: 48 [as subgenus of Anisochrysa Nakahara]. Type species: Chrysopa atlantica McLachlan, by original designation. [Raised to genus by Aspock et al., 1980: 379.] DISTRIBUTION. Madeira, Canary Islands. The genus includes three species which are restricted to the eastern Atlantic islands of Madeira and the Canaries. S. J. BROOKS & P. C. BARNARD DIAGNOSIS. Adult. Medium sized lacewings, fore wing (Fig. 237) 13-16 mm; ground colour olive green. Head marked with stripe on gena and X- shaped mark between the antennae which extends along the front of the vertex and across the frons below the antennae; palps tapered; labrum in- dented; mandibles broad, asymmetrical, with tooth on left mandible; galea short; vertex raised; head width : eye width = 2.1-2.3 : 1; antenna about same length as fore wing; flagellar segments about twice as long as wide, with setae arranged in four rings. Pronotum marked with narrow, black, transverse stripe; setae long, black; mesonotum marked with black stripes; metanotum unmarked. Legs unmarked; setae short, black; claws with basal dilation. Fore wing membrane unmarked, but with small black spot at base of costa; length : breadth = 2.7—2.9 : 1; costal area narrow at base; costal setae very short, inclined; stigma unmarked; Sc long; Sc and R widely separated; basal Sc crossvein 0.20—-0.28 mm; im broad, ovate; basal fork of M acute, vein slightly thickened; m5 long, narrow; Rs slightly sinuate; median radial crossveins sinuate; gradates in two parallel series, basal inner gradate usually meeting Psm; c; about same length as c,. Abdomen (Figs 238, 239) marked with black spot on each tergite; setae long, sparse; callus cerci ovate, trichobothria 32-40; ectoprocts fused dorsally, with slight apical invagination; d: microtholi present on sternites; sternite 8+9 fused; 2: sternite 7 straight apically; short apical suture between tergite 9 and ectoproct. GENITALIA 6 (Figs 240-242). Tignum arcuate with short, median process; gonapsis short, broad arcuate; gonarcus long, arcuate; median plate absent; entoprocessus narrow, elongate; arcessus broad, with apical tooth and dorsal striations; additional hooked structure positioned below arcessus, deeply bifurcate basally; gonosaccus with few short gonosetae; gonocristae and spin- ellae absent. GENITALIA ¢ (Figs 243, 244). Praegenitale absent; subgenitale bilobed apically, tapering and slightly extended basally; spermatheca narrow; ventral impression moderate; vela moderate; duct long, coiled. Larva. Abdomen broadly fusiform; thoracic tubercles small, spherical with short setae; ab- dominal latero-dorsal tubercles absent; dorsal setae short, sparse; no debris carried. REMARKS. When Holzel (1970) described Atlanto- chrysa he stated that it could be distinguished from Mallada Navas because it lacked a gonapsis. However, it is now apparent that a small gonapsis is, in fact, present. Nevertheless, males of Atlantochrysa may be distinguished from Mallada THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 189 [IX Zee 243 Figs 237-244 Atlantochrysa atlantica. 237, fore wing; 238, apex of d abdomen, lateral; 239, apex of 2 abdomen, | lateral; 240, 3 tignum, dorsal; 241, d gonapsis, ventral; 242, ¢ gonarcus complex, dorso—lateral (right entoprocessus not figured); 243, 2 subgenitale, ventral; 244, 2 spermatheca, lateral. | by the presence of a hooked structure positioned below the arcessus which is never found in Mallada and which rarely occurs in other chryso- pid genera. In addition, the basal inner gradate meets Psm, the radial crossveins are sinuate and the adult gut contents were found to include insect remains; no species of Mallada has yet been found to be carnivorous. The larval morphology of Atlantochrysa also differs considerably from that | of Mallada since the larvae of Mallada are debris carriers and so have long, hooked setae and cylindrical thoracic tubercles whereas the larvae of Atlantochrysa, which do not carry debris, have short setae and short, spherical tubercles. Mallada, Atlantochrysa and Anomalochrysa McLachlan are the only genera in the Chrysopidae in which males have both a tignum and a gonapsis. However, this does not necessarily indicate a close relationship between the genera since this is probably the plesiomorphic character state for the Chrysopini. There are at least 16 genera in the Chrysopini in which males have either a gonapsis or tignum and it is highly unlikely that these structures have arisen independently on several occasions. The tignum, in particular, has a very similar morphology in all the genera in which it occurs, lending weight to the hypothesis that it only arose once in the Chrysopidae. The probable 190 explanation is that one or other of these structures has been lost on different occasions and that their absence is a derived state. Atlantochrysa may be closely related to Cuncto- chrysa H6lzel since they share several apomorphic characters. In males of both genera there is a hooked structure positioned below the arcessus, there are dorsal striations on the arcessus, and microtholi are present on the sternites. How- ever, the tignum and gonapsis are absent in Cunctochrysa, which is probably the derived state, thus establishing that it is a distinct genus. Meleoma species and Plesiochrysa ramburi also possess an additional hooked structure below the arcessus or pseudopenis, which suggests that these taxa may also be related to Atlantochrysa and Cunctochrysa. It is possible that ‘Anisochrysa’ oblonga Hodlzel, described from Nepal, should be included in Atlantochrysa on the basis of the male genitalia which, in addition to possessing a tignum, gonap- sis and dorsal striations on the arcessus, also have narrow, elongate entoprocessus and a hooked structure below the arcessus. However, the species does differ in several ways from the description given for Atlantochrysa above. In A. oblonga the claws are undilated basally; the basal fork of M is oblique and the vein is not thickened; the radial crossveins are straight; the fore wing is narrow (length : breadth = 3.2 : 1); and insect remains were not present in the gut of the one adult male examined. BIOLoGy. The larva of A. atlantica (McLachlan) has been described by Monserrat (1978). The gut contents of adults examined during this study included large quantities of insect remains. Genus AUSTROCHRYSA Esben-Petersen Austrochrysa Esben-Petersen, 1928: 98. Type species: Austrochrysa samoana, by original designation Scoliochrysa Navas, 1929: 365. Type species: Scoliochrysa loriana Navas, by original desig- nation and monotypy. [Synonymized with Chrysopa Leach by Banks, 1937: 149.] Syn. n. DISTRIBUTION. Oriental, Samoa, Fiji. Austrochrysa includes seven described species, most of which occur in Indonesia. There is an undescribed species from Assam in the BMNH collections. DiAGnosis. Adult. Large lacewings, fore wing (Figs 245, 246) 15-18 mm. Head unmarked or with labrum marked brown, scape marked with lateral brown stripe; palps tapered apically; galea S. J. BROOKS & P. C. BARNARD broad; labrum invaginated; mandibles broad, symmetrical with basal tooth on both mandibles; vertex slightly raised; head width : eye width = 2.1-2.4 : 1; scape swollen, elongate; antenna considerably longer than fore wing; flagellar segments about 3 times as long as broad; setae arranged in four rings. Pronotum somewhat elongate, marked with pair of reddish brown or black medio-lateral oval spots; dorsal setae very long, pale; mesoprescutum marked with small brown spot; yellow median stripe along entire length of pteronota. Legs unmarked; setae long, pale; claws with strong basal tooth. Fore wing broad (length : breadth = 2.42.8 : 1), with pale brown suffusion around most crossveins or black spots on dcc, median radial crossveins, base of outer gradates; outer gradates usually black; costal area narrow at base; costal setae long, erect or slightly inclined; pterostigma marked basally with grey spot; Sc long; Sc and R widely separ- ated; basal Sc crossvein 0.2-0.4 mm; median radial crossveins sinuate; im long, narrow, quad- rangular or ovate; Rs very sinuate; gradates in two or three irregular rows, inner gradates arched; inner basal gradate meeting Psm; anal veins slightly crassate in 6; c, shorter or slightly longer than c). Hind wing with gradates or posterior apical margin lightly suffused grey; gradates in two or three series. Abdomen (Figs 247, 248) unmarked; setae long, sparse; ectoprocts with deep apico-dorsal invagination, fused dorsally, fused with tergite 9; atria sometimes enlarged; d: callus cerci ovate, large; trichobothria 20-25, large, widely dispersed; abdomen ‘gaping’ apic- ally (sternite 8+9 and ectoprocts widely separ- ated); ectoprocts pointed apico-dorsally, extended basally to narrow antero-ventral projection; sternite 8+9 fused, deeply excised anteriorly, invaginated apically; microtholi absent; @: callus cerci small, rounded; trichobothria 26-32, not widely dispersed; sternite 7 straight apically. GENITALIA 6 (Figs 249, 250). Tignum, gonapsis and median plate absent; gonarcus long, narrow, arcuate with small median projection and no lateral expansion; arcessus trilobed apically, sometimes with long, membranous, ventral projection; pseudo- penis absent; entoprocessus short, broad, L- shaped; gonosaccus large, circular; gonosetae long, numerous, evenly dispersed; T-shaped sclerite at apex of sternite 8+9 bearing row of prominent black gonocristae, or gonocristae present on sub- apical tubercle on sternite 8+-9; spinellae absent. GENITALIA 2 (Figs 251, 252). Praegenitale absent; subgenitale narrow, bilobed apically, with long basal extension; spermatheca small, narrow; ventral impression shallow or moderate; vela long; duct long, sinuous. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 191 Figs 245-252 Austrochrysa. 245,249, A. samoana; 246, 247, 250-252, A. abnormis; 248, A. javanensis. 245, 246, fore wing; 247, 248, apex of d abdomen, lateral; 249, 250, 3 genitalia, lateral; 251, 2 subgenitale, ventral; 252, 2 spermatheca, lateral. REMARKS. Austrochrysa, which has recently been redescribed by New (1987), may be distinguished by a combination of characters which is rarely found in the Chrysopidae. There is a large medio- lateral spot on the pronotum, a basal tooth on both mandibles, elongate scape and long, often quadrangular, intramedian cell. In females there is a basal extension of the subgenitale and in males the apex of the abdomen is ‘gaping’, there are relatively few, widely dispersed trichobothria, gonocristae are present, the gonarcus is very narrow with no lateral expansion and sternite 8+9 is apically invaginated. Although there are three series of gradates in Austrochrysa and only two in Scoliochrysa, the male and female genitalia are so similar that they 192 S. J. BROOKS & P. C. BARNARD Figs 253-258 Borniochrysa. 253, 254, 256, B. winkleri; 255, 257, 258, B. squamosa. 253, fore wing; 254, 255, apex of 3 abdomen, lateral; 256, d genitalia, dorsal; 257, 2 spermatheca, lateral; 258, 2 subgenitale, caudal. must be regarded as synonymous. Many of the species now included in Austrochrysa have, in the past, been regarded as Leucochrysa McLachlan (Albarda, 1881; Weele, 1909; Esben-Petersen, 1926; Banks, 1931) because of the quadrangular intramedian cell that occurs in both genera. However, the wing venation and genitalia of Austrochrysa and Leucochrysa have little in common which suggests that the two genera are only distantly related. The type of Austrochrysa samoana Esben- Petersen has lost its abdomen; however, the specimen is probably male since the anal veins are slightly crassate. Esben-Petersen (1928) suggested Austrochrysa had affinities with Anomalochysa McLachlan, because of the three rows of gradates which occur in both genera, and Nothancyla Navas which has a quadrangular intramedian cell. However, it is now apparent that any similarities between these genera and Austrochrysa are superficial. Multiple series of gradate crossveins and a quadrangular intramedian cell have arisen on several occasions in unrelated genera in the Chrysopidae and the male genitalic compliments are totally different in the three genera. BIoLoGy. Unknown. No insect remains were included in the gut contents of any adults exam- ined during this study. Genus BORNIOCHRYSA nom. n. Bornia Navas, 1928: 123. Type species: Bornia winkleri Navas, by original designation and monotypy. [Banks, 1931: 381 as synonym of Chrysopa Leach; Kuwayama, 1962: 372 as synonym of Anomalochrysa McLachlan.] [Homonym of Bornia Philippi, 1836: 13.] DISTRIBUTION. Afrotropics, Oriental, Solomon Is. Five species are included in Borniochrysa; one ee THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) species is widespread throughout the Afrotropics, the other four occurring in the Oriental region. DIAGNOsIs. Adult. Medium-sized lacewings, fore wing (Fig. 253) 10-15 mm; ground colour pale green. Head with red or black stripe on gena, red lateral stripe on clypeus, or with entire reddish suffusion; palps tapered; labrum invaginated; mandibles broad, asymmetrical with basal tooth on left mandible; vertex slightly raised; head : eye width = 1.6—2.3 : 1; antenna slightly shorter than fore wing; flagellar segments about 3 times as long as broad, with four rings of setae. Pronotum unmarked or with median yellow longitudinal _ stripe; dorsal setae long or short, pale or dark; meso- and metanotum unmarked or with brown stripe above wing base. Legs unmarked or with black stripe on tibia; setae short, black; claws with basal dilation. Fore wing unmarked or with brown suffusion on crossveins; costal area narrow at base; costal setae long or short, inclined; ptero- stigma unmarked or with brown basal spot; Sc and R widely separated; basal Sc crossvein 0.12-0.20 mm; im quite broad, long, ovate; m2 broad; Rs sinuate; gradates in two or three parallel or slightly divergent rows; basal inner gradate sometimes meeting Psm; veins not cras- sate in d ; c, same length as or slightly shorter than | >. Hind wing with gradates in two parallel rows. Abdomen (Figs 254, 255) unmarked or sternites with brown apical stripe; setae long, sparse; ¢d: callus cerci rounded or ovate, trichobothria 22— 31; sternite 8+9 fused, elongate, with protruding apical membrane bearing microsetae; ectoprocts with narrow, upturned apical projection, bearing a few stout setae at tip with deep apical invagina- tion, fused only at base dorsally; microtholi present on tergites and sternites or absent; short apodeme situated dorsally on ectoproct; 2: callus | cerci ovate, trichobothria 28-29; ectoprocts with slight dorso-apical invagination, extended ven- trally and curving below abdomen; short apical /suture usually present between ectoproct and | tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 256). Tignum, gonapsis and -median plate absent; gonarcus narrow, long, with dorso-lateral horns; entoprocessus long, T-shaped; arcessus tapering, arrowhead-shaped or trifurcate apically; pseudopenis absent; gono- saccus short; gonosetae long or short, numerous, in lateral clump or at apex of arcessus; hypan- drium large, broad; comes absent. GENITALIA 2 (Figs 257, 258). Praegenitale absent; subgenitale broad, bilobed apically; t ) _| spermatheca broad; ventral impression shallow or _ absent; vela large; duct long, sinuous. Larva. B. squamosa (Tjeder) was reared from 193 larvae collected in Zambia (Brooks, unpublished). The larvae were grey with a fusiform, humped abdomen; cylindrical thoracic tubercles bearing long setae; abdominal setae hooked apically, carrying large packet of debris. REMARKS. Borniochrysa was originally described (Navas, 1928) to include those species that pos- sessed three rows of gradate crossveins in the fore wing but only two rows in the hind wing; a character which also occurs in some species of Chrysopodes within the Chrysopini. However, the male genitalia, with elongate ectoprocts bear- ing stout setae at the apex, dorsal horns on the gonarcus and long entoprocessus which bifurcate apically, are also distinctive. These genitalic characters also occur in Chrysopa appendiculata Esben-Petersen and Chrysopa squamosa Tjeder which are, therefore, transferred to Borniochrysa, although they do not have three rows of gradates in the fore wing. Kuwayama (1962) synonymized Borniochrysa with Anomalochrysa because both genera possess more than two rows of gradates; however, multiple rows of gradates appear to have arisen independently in the Chrysopidae several times and since the male genitalia of Borniochrysa and Anomalochrysa show few similarities the synonymy is unjustified. BIOLOGY. Unknown. Genus BRINCKOCHRYSA Tjeder Neda Navas, 1933: 106. Type species: Neda decaryella Navas, by original designation. {[Homonym of Neda Mulsant, 1850: 274.] Syn. n. Brinckochrysa Tjeder, 1966: 360 [as subgenus of Chrysopa Leach]. Type species: Chrysopa peri Tjeder, by original designation. [Raised to genus by Holzel, 1970: 51.] DISTRIBUTION. Old World. This wide-ranging Old World genus includes 16 described species with several more undescribed in the BMNH collections. Most of the species are concentrated in the Afrotropics. D1AGnosis. Adult. Small to medium lacewings, fore wing (Fig. 259) 7-15 mm; ground colour bright green. Head marked with red stripes on post-ocular region, vertex and frons; palps tapered apically; labrum indented; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width = 1.8-2.4 : 1; antenna longer than fore wing; flagellar segments 3 times as long as broad, with 194 S. J. BROOKS & P. C. BARNARD Figs 259-264 Brinckochrysa. 259, 261, 263, 264, B. amseli; 260, 262, B. alfieri. 259, fore wing; 260, apex of 3 abdomen, lateral; 261, apex of 2 abdomen, lateral; 262, d genitalia, lateral; 263, 2 subgenitale, caudal; 264, 9 spermatheca, lateral. four rings of setae. Pronotum unmarked or with red lateral stripe; dorsal setae long, pale; meso- and metanotum unmarked or with red spots on scutum. Legs unmarked; setae long, dark or pale; stridulatory structure present on inner surface of hind femur; claws with or without basal dilation. Fore wing very narrow (length : breadth = 3.0— 3.8 : 1), pointed apically; unmarked; costal area narrow at base; costal setae short, inclined apically; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.16—0.24 mm; im long, narrow, ovate; Rs straight or slightly sinu- ate; gradates in two parallel series; basal inner gradate not meeting Psm; veins not crassate in d ; c; slightly shorter or longer than c,. Abdomen (Figs 260, 261) with red dorsal markings; setae long and sparse; callus cerci ovate, trichobothria 23-27; stridulatory structure present laterally on sternite 2; d: ectoprocts not fused dorsally, with short, broad apical projection and ventral exten- sion overlapping sternites 8 and 9; sternites 8 and 9 very narrow, not fused, with distinct narrow suture; ventral apodeme (= solimere of Tjeder, 1966) with long apical, toothed hook; @: ecto- procts with apical invagination, greatly extended ventrally and curving below abdomen with small gap between apices; apical suture present be- tween tergite 9 and ectoproct; sternite 7 with deep apical indentation. GENITALIA 6 (Fig. 262). Tignum, gonapsis and median plate absent; gonarcus long, narrow; entoprocessus narrow, sometimes with lateral extension, long or short; arcessus narrow, taper- ing apically; pseudopenis absent; gonosaccus short; gonosetae few, short in lateral clump; gonocristae present at apical and ventral extrem- ities of ectoprocts and on membrane at apex of sternite 9. GENITALIA 2 (Figs 263, 264). Praegenitale absent; subgenitale narrow, bilobed at apex; spermatheca narrow; ventral impression deep; vela long; duct long. Larva. Abdomen fusiform, flattened; tubercles rudimentary or undeveloped; setae not hooked; row of setae absent from metanotum; debris not carried. REMARKS. Brinckochrysa is the only genus in the Chrysopidae in which a stridulatory structure is present in all species. A stridulatory structure also occurs in a few species of Meleoma and in one species of Chrysocerca (Brooks, 1987). Males of Brinckochrysa can be distinguished by the very narrow sternites 8 and 9 and the overlapping ventral extension of the ectoprocts; females by the marked indentation at the apex of sternite 7. The genus may be related to Chrysoperla Steinmann and Peyerimhoffina Lacroix which also have narrow wings, a short intramedian cell and widely spaced, sinuous, costal crossveins at the base of the fore wing. In most Brinckochrysa species the fore wing is less than 11 mm but we have seen an undescribed Madagascan species (in the Muséum National d’Histoire Naturelle, Paris) in which the forewing is 15 mm. In females of this species the indentation at the apex of sternite 7 is unusually broad and the apical margins of the sternite are folded outwards THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 195 Figs 265-272 Ceraeochrysa. 265, C. albatala; 266, 268, 270, C. discolor; 267, 271, C. cincta; 269, C. cubana. 265, fore wing; 266, apex of d abdomen, lateral; 267, apex of 2 abdomen, lateral; 268, 269, ¢ gonarcus complex, dorsal; 270, 3 gonapsis, lateral; 271, 2 subgenitale, ventral; 272, 2 spermatheca, lateral. so that they project ventrally to form a lobe on either side of the indentation. BioLoGy. The eggs and larva of B. kintoki (Okamoto) have been described by Tsukaguchi (1979). The eggs are laid singly or in widely spaced groups of 7-14. From their flattened form, it is likely that the larvae conceal themselves in crevices. Adults are attracted to light and can be found resting on the underside of the leaves of Catalpa bignonioides Walter. Adams (1959) des- cribed the putative larva of B. scelestes (Banks) from Palau, Micronesia. His description agrees closely with that given by Tsukaguchi for kintoki, and differs from most other known chrysopid larvae, so it seems likely that the larva was correctly assigned to B. scelestes. Genus CERAEOCHRYSA Adams Ceraeochrysa Adams, 1982b: 70. Type species: Chrysopa_ cincta Schneider, by original designation. DISTRIBUTION. Nearctic, Neotropics. This genus includes 40 described species and, in addition, several undescribed species are present in the BMNH collections. They are widely dis- tributed throughout North and South America and the West Indies. C. cincta is widely distributed from Florida to Argentina and the Galapagos Islands. Ceraeochrysa is the dominant Neotropical chrysopine genus in numbers of individuals and numbers of species. D1AGNosis. Adult. Medium-sized lacewings, fore wing (Fig. 265) 9-15 mm; ground colour pale green. Head unmarked or with red suffusion and red stripe on scape; palps tapered; galea narrow; labrum indented or straight; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width = 2.3-2.7: 1; antenna slightly longer than fore wing, sometimes black; flagellar segments about 3 times as long as broad; setae arranged in four rings. Pronotum unmarked or with red lateral stripe; dorsal setae long, pale; micropoculae absent; meso- and metanotum unmarked. Legs unmarked; setae short or long, dark or pale; claws with basal 196 dilation. Fore wing unmarked but radial cross- veins dark; costal area narrow at base; costal setae short, inclined; stigma unmarked, long; Sc and R widely separated; basal Sc crossvein 0.20-0.32 mm; im narrow, ovate; Rs slightly sinuate; gradates in two, closely apposed parallel series; basal inner gradate meeting Psm; veins not crassate in 6; c, shorter than c>. Abdomen (Figs 266, 267) unmarked; setae long, sparse; callus cerci ovate, trichobothria 24-33; ectoprocts usually with deep dorso-apical invagination and fused dorsally; ectoprocts fused with tergite 9; d: microtholi absent; apodeme of tergite 9 often with strong ventral hook; sternite 8+9 fused, some- times elongate, dorsally curved; some species with apical elongation of ectoprocts; 9: sternite 7 straight apically. GENITALIA ¢ (Figs 268-270). Tignum absent; gonapsis elongate, narrow, bifurcate apically; median plate large with median horns; ento- processus absent but lateral gonarcus horns often well developed; parameres absent; gonarcus long, narrow often with gonocornua, in cubana-species group gonarcus with massive dorsal expansion; arcessus long or short, narrow tapering apically often with median hook and lateral lobes; pseudo- penis absent; gonosaccus short; gonosetae absent, few or numerous and long arranged in lateral clump; gonocristae usually present at apex of sternite 8+9; spinellae absent; atria sometimes expanded. GENITALIA 2 (Figs 271, 272). Praegenitale absent; subgenitale bilobed, sometimes with small median crumena and mounted on large plate, or pointed basally; spermatheca very small, narrow; ventral impression deep; vela very long; duct long, highly coiled. Larva. Abdomen broadly fusiform, humped; thoracic tubercles long, slender; metanotum with row of dorsal setae and chalazae; abdominal tubercles broad, short; latero-dorsal abdominal tubercles absent; latero-dorsal chalazae present on segments 6 and 7; setae hooked apically, usually smooth but plumose in cubana-group; debris packet very large, obscuring most of body. REMARKS. Species of Ceraeochrysa have no out- standingly distinctive external characters, although members of the genus can often be recognized by the presence of a red lateral stripe on the pro- notum or scape and the dark radial crossveins. Males of Ceraeochrysa can be distinguished from other chrysopid genera by the straight, elongate gonapsis and horned median plate, and females by the relatively small spermatheca. In addition, species of the cubana-group possess a grossly enlarged gonarcus and enlarged atria. S. J. BROOKS & P. C. BARNARD Tjeder (1966) suggested that the genus was related to Glenochrysa due to the presence of a gonapsis and enlarged atria in males. However, the presence of a gonapsis in the Chrysopini is plesiomorphic, and enlarged atria have evolved several times in the Chrysopidae; the two genera do not share any other significant characters which might suggest a close relationship. Adams (1982b) proposed that there was a relationship between Ceraeochrysa and Leucochrysa because in both genera the gonarcus has gonocornua, the apex of the arcessus is hooked and has lateral lobes, and the spermathecae are of similar shape. However, it seems unlikely that these genera are closely related since Ceraeochrysa is clearly a member of the Chrysopini, and exhibits none of the critical leucochrysine characteristics. BioLoGy. Larvae of several species of Ceraeo- chrysa have been described including C. lineati- cornis Fitch (Smith, 1921; 1926b; Muma, 1957), C. bicarnea Banks (Smith, 1926b), C. bimaculata McCloud (Smith, 1922; 1926b), C. cubana Hagen (Muma, 1959), C. damiensis Smith (Smith, 1932) and C. lateralis Guérin (Smith, 1921; 19265; Muma, 1957). Adult gut contents do not include insect remains. Genus CERATOCHRYSA Tjeder Musola Navas, 1929: 367. Type species: Musola impar Navas, by original designation and monotypy. [Homonym of Musola Roewer, 1927: 398 (Arachnida). ] Ceratochrysa Tjeder, 1966: 352 [as subgenus of Chrysopa Leach]. Type species: Chrysopa ceratina Navas, by original designation. [Raised to genus by Barnard & Brooks, 1984: 361.] DISTRIBUTION. Mauritius and Madagascar. The genus includes three species. C. antica (Walker) is widely distributed throughout tropical Africa, C. ceratina is restricted to southern Africa and C. disparilis (Navas) is known only from Madagascar. Afrotropics, DIAGNOSIS. Adult. Large green lacewings, fore wing (Fig. 273) 14-22 mm; ground colour pale green. Head unmarked or marked with red stripe on frons, vertex and scape; palps tapered; labrum deeply emarginated; mandibles broad, asym- metrical with basal tooth on left mandible; galea broad; head width : eye width = 2.1—2.5 : 1; vertex slightly raised; antenna 1.5 times length of fore wing; flagellar segments almost 3 times as long as broad; setae arranged in four rings. Pronotum longer than broad, sometimes with red lateral | | THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 197 SS SSssea9) 7 Figs 273-279 Ceratochrysa. 273-275, 277-279, C. antica; 276, C. ceratina. 273, fore wing; 274, apex of d abdomen, lateral; 275, apex of 2 abdomen, lateral; 276, ¢ genitalia, dorsal; 277; 2 subgenitale, ventral; 278, 2 subgenitale, lateral; 279, 2 spermatheca, lateral. stripe or yellow median stripe; meso- and meta- notum unmarked or entirely black. Legs un- marked; setae short, pale; claws with basal dilation. Fore wing narrow (length : breadth = 3.1-3.4 : 1); unmarked but stigma sometimes marked black; costal area narrow at base; costal setae short, inclined; Sc and R well separated; basal Sc crossvein 0.24-0.32 mm; radial crossveins sinuous; im ovate, long, narrow; veins sometimes crassate in 6; gradates in two parallel series, at least twice as many outer gradates than inner, outer gradates black; basal inner gradate meeting Psm; c, shorter than c>. Abdomen (Figs 274, 275) sometimes marked with red lateral spots or yellow median stripe; setae long, sparse; callus cerci ovate, trichobothria 35-45; ectoprocts with deep apical invagination, not fused dorsally, fused with tergite 9; d: tergite 9 broad, rounded basally, sternites 8 and 9 not fused; microsetae present throughout all sclerites in some species; micro- tholi absent; 2: sternite 7 straight apically, sometimes thickened. GENITALIA 6 (Fig. 276). Tignum and gonapsis absent; gonarcus long, narrow with dorsal sail-like 198 structure; median plate large, bifurcate; ento- processus extremely elongate with pair of apical teeth; arcessus absent; pseudopenis narrow; gonosaccus with numerous long gonosetae; gono- cristae and spinellae absent. GENITALIA ¢ (Figs 277-279). Praegenitale absent; subgenitale bilobed apically, mounted on large sclerotized plate bearing setae; spermatheca narrow; ventral impression deep; vela small; duct short. Larva. Abdomen broadly fusiform, moderately humped; thoracic tubercles spherical; row of setae present on meso- and metanotum but chalazae absent; small dorso-lateral tubercles present on abdominal segments 1, 6 and 7; abdominal setae smooth, hooked apically; small packet of debris present. REMARKS. Ceratochrysa can be distinguished from other chrysopid genera by the small number of inner gradates compared with outer ones and, in the male, by the extraordinary length of the entoprocessus, which are usually visible protrud- ing beyond the apex of the abdomen, and the presence of the sail-like structures on the gonar- cus. Females are characterized by the sclerotized plate which supports the subgenitale. Ceratochrysa is apparently closely related to Chrysopa Leach, since males of both genera have a pseudopenis, a broad, basally rounded ectoproct and a persistent suture between sternites 8 and 9. There are also close similarities to Plesiochrysa in which, like Ceratochrysa, there is a pair of mid-dorsal horns on the gonarcus, microsetae on the abdominal sclerites of males and a dorsal suture between the ectoprocts. These characters are rarely en- countered in other chrysopine genera. Other shared characters include a long, narrow pro- thorax and antennae which are longer than the fore wing. The larva of Ceratochrysa is similar in appearance to the putative larva of C. ramburi Schneider, which has been described by Adams (1959), further suggesting a relationship between the two genera. BioLoGy. The larva of C. antica has been described by Barnard & Brooks (1984). It has been found to prey on the cassava mealybug Phenacoccus manihoti Matile-Ferrero which was recently introduced into west Africa. Genus CHRYSEMOSA nom. n. Mesochrysa Navas, 1936a: 169. Type species: Mesochrysa stigmata Navas, by original desig- nation and monotypy. [Homonym of Meso- chrysa Navas, 1927a: 55.] S. J. BROOKS & P. C. BARNARD DISTRIBUTION. Afrotropics, Middle East. Chrysemosa includes 10 described species: three from the Middle East, the remaining seven from the Afrotropics. DIAGNOsIs. Adult. Small lacewings, fore wing (Fig. 280) 9-10 mm; ground colour brown or green. Head usually extensively marked with red or dark brown stripes; palps tapered apically; labrum straight apically; vertex raised; head width : eye width = 1.8-1.9 : 1; antenna pale; as long as or slightly longer than fore wing; flagellar segments about 3 times as long as broad; setae short, dark, arranged in four rings. Pronotum marked with broad, brown lateral stripe; dorsal setae short, dark; meso- and metanotum marked with brown lateral stripe. Legs marked with brown annulations on tibia and femora; setae short, dark; Ist tarsal segment elongate in hind leg; claws with basal dilation. Fore wing narrow (length : breadth = 2.9-3.0 : 1); marked with two dark brown spots on dcc, sometimes unmarked; costal area narrow at base; costal setae short, inclined; costal cells almost as broad as long; stigma marked with brown spot; Sc long; Sc and R widely separated; basal Sc crossvein 0.16— 0.20 mm; im narrow, ovate; Rs sinuate; gradates in two parallel series; basal inner gradate not meeting Psm; c, about same length as c>. Hind wing with stigma marked dark brown. Abdomen (Figs 281, 282) with extensive black markings; setae long, sparse; trichobothria 27-29; ecto- procts with broad dorso-apical invagination, separated by wide suture dorsally, fused with tergite 9; d: callus cerci ovate; sternite 8+9 fused; 2: sternite 7 straight apically. GENITALIA 6 (Figs 283, 284). Tignum and median plate absent; gonapsis narrow, arcuate; ento- processus short, fused apically to each other dorso-basally of arcessus; parameres absent; gonarcus long, arcuate, dorsal horns absent; arcessus narrow, tapering apically, bifurcate basally without dorsal striations; pseudopenis absent; gonosaccus small; gonosetae long, numer- ous; gonocristae, spinellae absent. GENITALIA @ (Figs 285, 286). Praegenitale absent; subgenitale bilobed apically, with V- shaped basal indentation; spermatheca narrow; ventral impression broad, moderately deep; vela moderate; duct long, sinuous. REMARKS. Most species of Chrysemosa can be recognized by the prominent black spot on dcc in the fore wing and the broad dorsal suture between the ectoprocts. Males of the genus are character- ized by the presence of a gonapsis and the short, apically fused entoprocessus. In females there | | THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 199 285 280 Figs 280-286 Chrysemosa. 280, 281, 283, 284, C. jeanneli; 282, 285, 286, C. andresi. 280, fore wing; 281, apex of d abdomen, lateral; 282, apex of 2 abdomen, lateral; 283, d gonarcus complex, lateral; 284, d gonapsis, ventral; 285, 2 spermatheca, lateral; 286, 2 subgenitale, ventral. is a V-shaped indentation at the base of the subgenitale. The species now included in Chrysemosa were all previously placed in Suarius Navas following Tjeder (1966). Tjeder grouped all the Afrotropical chrysopine species which lacked a tignum and gonapsis in Suarius. Although this probably is the apomorphic condition for the Chrysopini, never- theless, the tignum and gonapsis appear to have been lost independently several times in the tribe and the resulting group appears to be para- phyletic. However, Tjeder recognized that the jeanneli group of species (his group c) differed from the rest of Suarius because, unlike the other species then included in the genus, the ento- processus were narrow and fused apically to each other dorsal of the arcessus. He also noted that they each possessed what he referred to as an apical apodeme of sternite 8+9. However, it is more likely that this structure is homologous with a gonapsis since an apodeme does not occur in this position in any other genus of Chrysopidae. The holotype of C. stigmata, the type species of Chrysemosa, was deposited in Hamburg Museum but was destroyed during the Second World War (Strimpel, in litt.). Nevertheless, from Navas’s (1936a) description it is possible to deduce that Chrysemosa stigmata Navas is referable to the jeanneli group of species. C. stigmata is a small species, the fore wing is only 9 mm, with only one inner and two outer gradate crossveins. A dark brown mark is described as being situated between the oblique apical branches of the cubitus and the proximal posterior vein (i.e. in dcc) in the fore and hind wing and there is also a brown spot at the base of the stigma. However, Navas included stigmata in a new genus because of the unusual form of the 291 S. J. BROOKS & P. C. BARNARD Figs 287-293 Chrysocerca. 287, 289, 290, 292, 293, C. perturbata; 288, 291, C. sp. indet.; 287, fore wing; 288, apex of 3 abdomen, lateral; 289, apex of 2 abdomen, lateral; 290, d gonarcus complex, lateral; 291, d gonapsis, ventral; 292, 2 spermatheca, lateral; 293, 9? subgenitale, caudal. intra-median cell, which he compared with Nesochrysa Navas. The third median cell was simple without a looping posterior branch, neither ovate nor quadrangular. In all the jeanneli-group species that we have examined cell im is ovate. Indeed, the form of im in Nesochrysa grandidieri is unique in the Chrysopidae. Therefore, the condition that Navas described for Chrysemosa may be aberrant or restricted to one species (as in Nesochrysa) but in any case it is not of generic significance. It therefore seems to be a reasonable supposition to include the jeanneli group species in Chrysemosa. BioLoGy. Unknown. No insect remains were found in the guts of any of the adults examined during this study. Genus CHRYSOCERCA Weele Chrysocerca Weele, 1909: 75. Type species: Chrysocerca jacobsoni Weele, by original designation and monotypy. [Synonymized with Nineta Navas by Lacroix, 1924: 571; reinstated by Tjeder, 1966: 345] Pseudochrysa Okamoto, 1914: 55. Type species: Pseudochrysa formosana Okamoto, by mono- typy. [Synonymized by Kuwayama, 1966: 137.] DISTRIBUTION. Uganda, Oriental region. Five species have been described in Chrysocerca and there is a further undescribed species in the BMNH collections. Only one species is known from the Afrotropical region. DIAGNOsIS. Adult. Small to medium lacewings, fore wing (Fig. 287) 9-13 mm; ground colour pale green. Head unmarked or with red or black markings on gena, clypeus, frons and scape; mandibles broad, symmetrical, with basal tooth on both mandibles; galea broad; palps tapered; labrum indented; vertex raised; head width : eye width = 2.0-2.6 : 1; antenna considerably longer than fore wing; flagellar segments at least 3 times as long as broad. Pronotum unmarked; setae long, pale; mesonotum unmarked or with large red/ black spot above wing base; metanotum un- marked. Legs unmarked; setae long, pale; stridu- latory structure sometimes present at apex of hind femur; claws with or without basal dilation. Fore wing narrow or broad (length : breadth = 2.5-3.0: 1); unmarked or with red suffusion at base and THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) inner gradates with black suffusion; costal area narrow at base; costal setae long erect; stigma unmarked or slightly darkened; Sc and R widely separated; basal Sc crossvein 0.16—0.24 mm; im narrow, ovate, short; 1st Psm crossvein apical of im oblique; Rs sinuate; gradates in two divergent or parallel rows, basal inner gradate meeting Psm; veins not crassate in 6; c, shorter than cp. Abdomen (Figs 288, 289) unmarked; setae long, sparse; callus cerci very large, ovate; tricho- bothria 22-32; d: stridulatory structure some- times present on lateral membrane of 2nd segment; ectoprocts not fused dorsally, produced apically into long cerci; sternite 8+9 fused, usually with median apical projection; tergites and sternites sometimes with microtholi; 2: ecto- procts with deep apical invagination, not fused dorsally, narrow suture present between ecto- procts and tergite 9; sternite 7 straight apically. GENITALIA 6 (Figs 290, 291). Tignum and median plate absent; gonapsis broad, arcuate with apical teeth; gonarcus long, narrow; entoprocessus absent; arcessus with deep pincer-like apical bifurcation; pseudopenis and parameres absent; gonosaccus very long, protruding from apex of abdomen; gonosetae numerous, long, evenly distributed; spinellae and gonocristae absent. GENITALIA @ (Figs 292, 293). Praegenitale absent; subgenitale bilobed apically, tapering basally; spermatheca narrow; ventral impression moderate; vela large; duct short, curved. REMARKS. Males of Chrysocerca can be readily distinguished by the long cerci at the apex of the abdomen, the deeply bifid arcessus and the presence of a broad, arcuate gonapsis with apical teeth. The long antennae, red/black spot on the mesonotum and large callus cerci are also distinc- tive and are useful for assigning unassociated females to this genus. Dr S. Tsukaguchi and Dr S. Takagi have kindly provided us with information on the type of Pseudochrysa formosana Okamoto which has enabled us to confirm the synonymy of Pseudo- _ chrysa with Chrysocerca. BioLoGy. Adults examined in this study did not have insect remains in the gut contents. Genus CHRYSOPA Leach Chrysopa Leach, 1815: 138. Type species: Hemerobius perla L., by subsequent desig- nation by ICZN, 1954: 3. Aeolops Billberg, 1820: 95. Type species: Hemerobius chrysops L., by subsequent desig- nation by Tjeder, 1966: 351. [Synonymized by Tjeder, 1966: 351.] 201 Emerobius Costa, 1834: 72. Type species: Hemerobius chrysops L., by monotypy. Syn. n. Melanops Doumerc, 1861: 192. Type species: Chrysopa parvula Doumerc, by monotypy. [Synonymized by Leraut, 1980: 242.] Chrysopisca McLachlan, 1875: 23. Type species: Chrysopisca minuta McLachlan, by monotypy. Syn. n. Cintameva Navas, 1914c: 214. Type species: Cintameva venulosa Navas, by original desig- nation. [Synonymized by Smith, 1932: 581.] Minva Navas, 1920b: 288. Type species: Minva punctata Navas, by original designation and monotypy. Syn. n. Polyphlebia Navas, 1936b: 88. Type species: Polyphlebia punctata Navas, by original desig- nation and monotypy. Syn. n. Metachrysopa Steinmann, 1964: 264 [as subgenus of Chrysopa Leach]. Type species: Chrysopa septempunctata Wesmael [= Chrysopa pallens Rambur], by original designation. [Synonymized by Tjeder, 1966: 351.] Nigrochrysopa Steinmann, 1964: 264 [as subgenus of Chrysopa Leach]. Type species: Chrysopa formosa Brauer, by original designation. [Syn- onymized by Tjeder, 1966: 351.] Parachrysopa Séméria, 1983: 310. Type species: Hemerobius pallens Rambur, by original desig- nation and monotypy. Syn. n. DISTRIBUTION. Holarctic, including 27 species in the western Palaearctic, 15 species in the eastern Palaearctic and 11 Nearctic species. C. pallens is particularly widespread throughout the Palaearctic region. DIAGNOsIs. Adult. Medium-sized lacewings, fore wing (Fig. 294) 9-19 mm. Head usually exten- sively marked with black spots or stripes; ground colour olive or pale green; palps tapered apically; labrum with deep or slight emargination; man- dibles broad, deeply excised, asymmetrical with basal tooth on left mandible; galea short; vertex slightly raised; head width : eye width = 2.64.9: 1, head broad; antenna shorter than fore wing; flagellar segments about twice as long as broad, setae arranged in four rings. Pronotum unmarked or with extensive black markings; dorsal setae usually short, black; meso- and metanotum marked with black spots or unmarked. Legs unmarked; setae usually short, black; claws with or without basal dilation. Fore wing often quite broad (length : breadth = 2.43.0 : 1), oval, unmarked; basal costal area narrow; costal setae short, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.20-0.52 mm.; im ovate, broad or narrow, occasionally absent; Rs straight or slightly sinuate; gradates in two parallel 202 S. J. BROOKS & P. C. BARNARD PN TS Soe cae a) ‘- ang an 4 < any x A AB Figs 294-300 Chrysopa. 294-297, 299, 300, C. perla; 298, C. pallens. 294, fore wing; 295, apex of d abdomen, lateral; 296, apex of 2 abdomen, lateral; 297, ¢ genitalia, lateral; 298, d tignum, lateral; 299, 2 subgenitale, caudal; 300, 2 spermatheca, lateral. rows, basal inner gradate usually meeting Psm; veins not crassate in 6; c, slightly longer or shorter than c,. Abdomen (Figs 295, 296) marked black or unmarked; setae short, coarse, sparse; callus cerci rounded or ovate, trichobothria 23-47; ectoprocts fused dorsally, not completely fused with tergite 9; d: ectoprocts with deep dorso-apical invagination; tergite 9 broad, rounded basally; sternites 8 and 9 not fused, widely separ- ated; sternites 2-8 with microtholi; apodeme on tergite 9 arcuate, apodeme on sternites 8 and 9 usually with long, dorsally projecting apical tooth; 2: sternite 7 straight apically; ectoprocts with slight dorso-apical invagination. GENITALIA 6 (Figs 297, 298). Tignum usually absent or small, cylindrical or disc-shaped in some species; gonapsis and median plate absent; ento- processus broad basally with dorsal horn; arcessus absent; pseudopenis arcuate, tapering apically; gonarcus long, narrow, arcuate, sometimes with pair of short median horns; gonosaccus short, globular, paired; gonosetae very long, numerous, THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) in lateral clump; gonocristae usually present in large lateral group at apex of sternite 9. GENITALIA @ (Figs 299, 300). Praegenitale absent; subgenitale bilobed apically, sometimes with small median projection; spermatheca broad; ventral impression shallow to moderate, narrow; vela short, duct long or short, sinuous. Larva. Abdomen fusiform, not humped; thoracic tubercles large, spherical, bearing long setae; row of setae absent from metanotum; setae smooth, not hooked apically; abdominal tubercles spheri- cal; dorso-lateral tubercles present on each abdominal segment; debris absent. REMARKS. Chrysopa is a distinctive genus and can be recognized by several characters. The adults of all species are carnivorous, they generally have broad, oval forewings, the head and thorax are more or less marked black and the eyes are small. In both sexes the ectoproct and tergite 9 are only partially fused and in males sternites 8 and 9 are not fused and the apodeme on tergite 9 is arcuate. In the male genitalia a pseudopenis is present, the gonosetae are very long and arranged in lateral clumps, and the entoprocessus have dorsal horns. Chrysopa shares several synapomorphies with Plesiochrysa and Ceratochrysa, suggesting a close relationship between these genera. The most striking of these are the basally rounded ectoproct and arcuate apodeme in the male, with the presence of a pseudopenis and dorsal horns on the gonarcus. A close relationship with Nineta and Tumeochrysa is also suggested by the relatively small eyes, and sutures between sternites 8 and 9 and between the ectoprocts and tergite 9. Electrophoretic studies by Bullini et al. (1983) suggested that C. viridana Schneider was not closely related to walkeri McLachlan, formosa Brauer and dorsalis Burmeister which were also examined during the study. C. viridana also differs morphologically from the latter species. The head is marked only with a black spot on the gena in viridana but is extensively marked in the walkeri-group. In males of viridana the gono- cristae are absent but a large clump is present in the walkeri-group. Chrysopisca was described by McLachlan (1875) to include C. minuta McLachlan which differed from other species of Chrysopas.1. in that it did not have an intramedian cell. However, this study has revealed that in all other important characters C. minuta resembles Chrysopa s.str. It could be argued that, on the basis of the absence of im, Chrysopisca should be retained at least as a subgenus of Chrysopa, but examination of the _ male genitalia of C. minuta has shown that this species is merely a small form of C. sogdianica. 203 There seems, therefore, to be no justification in recognizing Chrysopisca as a valid genus. It was not possible to trace the type species of Minva Navas but from the original description it is apparent that M. punctata Navas is almost certainly a synonym of C. minuta and so must also be regarded as a synonym of Chrysopa. The holotype of the type species of Polyphlebia, P. punctata Navas, was deposited in Rabat, Morocco but unfortunately we have been unable to examine it. Nevertheless, from Navas’s (1936b) description it is possible to synonymise the genus with Chrysopa. The fore wing of P. punctata is broad and rounded apically, im ovate and the basal inner gradate meeting Psm. The head is marked with seven black spots, similar to C. formosa Brauer, and there is a black lateral stripe on the pronotum, all of which support the synonymy with Chrysopa. However, the reason Navas originally described Polyphlebia as a new genus was that it possessed an irregular median series of gradate crossveins. We have not seen any Chrysopa species with a median row of gradates, but this condition has arisen indepen- dently several times in the Chrysopidae and does not appear to have much taxonomic significance. However, this aspect of the wing venation in Polyphlebia does resemble Rexa and it is possible that the genus should be included with that taxon although the other characters mentioned above do not occur in Rexa. In some species, including C. pallens (Rambur) and C. nigricornis Burmeister, the wings are long nd relatively narrow and males possess a small tignum. However, this is not sufficient reason to erect a new genus to accommodate these species since such characters are probably plesiomorphic and the species share many apomorphies with Chrysopa. Therefore, Metachrysopa Steinmann is maintained as a synonym of Chrysopa. Para- chrysopa Séméria is an objective synonym of Metachrysopa, because it shares the same type species. Séméria (1983) argued that because the chromosomal complement of C. pallens Rambur [= angustipennis Stephens] (2n = 10) differed from that of the rest of the genus (2n = 12), ‘pallens’ should have distinct generic status. He used the term ‘twin-genus’ to indicate the close relationship between the taxa. However, until the chromosomes of more than just a few European genera have been examined it seems premature to erect new taxa based solely on this kind of evidence. BIOLOGY. The larvae of many species have been described including C. abbreviata Curtis (Killington, 1937; Gepp, 1983), C. chi Fitch 204 ee 305 S. J. BROOKS & P. C. BARNARD 307 Figs 301-307 Chrysoperla. 301, fore wing; 302, apex of d abdomen, lateral; 303, apex of 2 abdomen, lateral; 304, 3 gonarcus complex, dorsal; 305, ¢ tignum, dorsal; 306, 2 spermatheca, lateral; 307, 2 subgenitale, caudal. (Smith, 1922), C. commata Kis & Ujhelyi (Gepp, 1983), C. dorsalis Burmeister (Alderson, 19115; Killington, 1937; Fraser, 1945; Gepp, 1983), C. flaviceps Brullé (Monserrat, 1982); C. formosa Brauer (Principi, 1947; Gepp, 1983), C. hungarica Klapalek (Gepp, 1983); C. intima McLachlan (Tsukaguchi, 1978), C. majuscula Banks (Clancy, 1946), C. nigra Okamoto (Tsukaguchi, 1978), C. nigricornis Burmeister (Clancy, 1946; Toschi, 1965; Tauber & Tauber, 1972), C. nigricostata Brauer (Brauer, 1850; Gepp, 1983), C. oculata Say (Smith, 1921), C. perla L. (Killington, 1937; Canard, 1971; 1973; 1976; Gepp, 1983), C. phyllochroma Wesmael (Killington, 1937; Gepp, 1983), C. quadripunctata Burmeister (Smith, 1921; 1922; Toschi, 1965), C. pallens (Rambur) (Killington, 1937; Principi, 1940; Gepp, 1983), C. viridana Schneider (Principi, 1954; Gepp, 1983) and C. walkeri McLachlan (Gepp, 1983). The gut contents of adults examined during this study included large quantities of insect remains. Genus CHRYSOPERLA Steinmann Chrysoperla Steinmann, 1964: 260 [as subgenus of Chrysopa Leach]. Type species: Chrysopa carnea Stephens by original designation. [As subgenus of Anisochrysa Nakahara by Hélzel, 1970: 51; raised to genus by Séméria, 1977: 238.] ———————————————————— —————————V7V————_ OOOO ee | THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) DISTRIBUTION. World-wide. This large genus is distributed throughout the world with 12 species in the Afrotropics, 3 species in Madagascar, 7 species in the western Palaearctic, 5 species in the eastern Palaearctic, 5 Oriental and Indian species, 5 Australian species, 5 species in the Nearctic and 8 in the Neotropics. The genus includes several particularly common species, especially C. carnea (Stephens) in the Palaearctic, C. plorabunda (Fitch) in the Nearctic, C. congrua (Walker) in the Afrotropics, and C. externa (Hagen) in the Neotropics. DIAGNosis. Adult. Medium lacewings, fore wing (Fig. 301) 9-14 mm; ground colour pale green. Head unmarked or with red and sometimes brown stripes on gena, clypeus, frons and vertex; palps tapered; labrum indented; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width = 1.9-2.8: 1; antenna longer or shorter than fore wing; flagellar segments 2-3 times as long as broad; setae in four rings. Pronotum with yellow median stripe or red lateral spots; dorsal setae long, pale or short, dark; meso- and metanotum with yellow median stripe or unmarked. Legs unmarked; setae long or short, pale or dark; claws with or without basal dilation. Fore wing narrow (length : breadth = 2.9-3.8 : 1); unmarked; costal area narrow at base; costal setae short, inclined or infrequently long, erect; basal costal crossveins sinuous; stigma unmarked; Sc and R well separated; basal Sc crossvein 0.12—0.20 mm; im narrow, ovate; Ist Rs crossvein usually meeting Psm well distad of apex of im; Rs sinuous; gradates in two parallel series, basal inner gradate meeting Psm; veins not crassate in 6; c; usually shorter than c>; cz broad, rounded apically; posterior marginal crossveins parallel. Hind wing narrow (length : breadth = 2.9-3.7 : 1). Abdomen (Figs 302, 303) unmarked or with median yellow stripe; setae long, sparse; callus cerci narrowly ovate, trichobothria 23-34; ectoprocts with slight apical invagination; 6d: Sternite 8+9 fused, with apical lip; microtholi absent; 2: apex of sternite 7 straight. GENITALIA 6 (Figs 304, 305). Tignum arcuate; gonapsis and median plate absent; entoprocessus absent or very short; arcessus narrow, tapering and often recurved apically; pseudopenis absent; gonarcus long, narrow and arcuate; gonosaccus short; gonosetae long, few, in lateral clump; spinellae present or absent. GENITALIA 2 (Figs 306, 307). Praegenitale absent; subgenitale bilobed apically, slightly extended basally; spermatheca narrow; ventral impression shallow or deep; vela long or short; duct long or short, sinuous. 205 Larva. Abdomen narrow, fusiform, not humped; thoracic and abdominal tubercles small, spherical; setae short, smooth, not hooked apically; trans- verse row of metanotal setae absent; latero-dorsal chalazae, bearing single seta, present but in- distinct; debris not carried. REMARKS. Chrysoperla may be distinguished from other Chrysopini genera by the narrow fore and hind wings with short, inclined costal setae and short im. However, in C. nyerina (Navas) the wings are broad, costal setae long and erect, and im long. In the male genitalia of all species a tignum is present but gonapsis absent and there is usually a distinct lip at the apex of sternite 8+9. However, a few species, including C. satilota (Banks) and C. krakatauensis Tsukaguchi from Australia and the Oriental region, appear to form a discrete species group in which the apex of the abdomen is ’gaping’, the ectoproct and tergite 9 are caudally compressed, the callus cerci very narrow, there is no apical lip on sternite 8+9 but a lateral lobe is present, the gonarcus is short, and there is a broad wing-shaped plate, fused to the apodemes, at the apex of sternite 8+9. The morphology of the apical abdominal segments in these species is therefore similar to that of the Mallada boninensis-group and may indicate a close relationship, since such structures do not occur elsewhere in the Chrysopidae. The apical plate in sternite 8+9 of the satilota-group may be homologous with the gonapsis. Chrysoperla is the only genus in the Chrysopidae in which spinellae occur. These structures appear to be modified microsetae which are distributed ventrally on the gonosaccus of some species. In C. insulata (Fraser), from Réunion, the tignum is absent in males. However, the apex of sternite 8+9 is lipped and there are spinellae on the gonosaccus, which are autapomorphic characters for Chrysoperla, so inclusion of this species in the genus is justified. Chrysoperla and Peyerimhoffina are probably closely related since in both genera the wings are unusually narrow, the basal costal crossveins are sinuous and the posterior marginal crossveins are parallel. In most chrysopid genera the basal costal crossveins are straight and the posterior marginals are divergent. BIoLoGy. The larvae of many species of Chrys- operla have been described including C. carnea (Stephens) (Smith, 1922; Killington, 1937; Neumark, 1952; Putman, 1956; Fleshner & Scriven, 1957; Muma, 1959; Toschi, 1965; Tsukaguchi, 1977; Gepp, 1983), C. commanche (Banks) (Tauber, 1974), C. congrua (Walker) (Brettell, 1982); C. downesi (Smith) (Putman, 206 1937), C. externa (Hagen) (Tauber, 1974), C. harrisii (Fitch) (Tauber, 1974; Tauber & Tauber, 1974), C. interrupta (Schneider) (Smith, 1921; Muma, 1959), C. lanata (Banks) (Ru et al., 1975), C. pudica (Navas) (Tjeder, 1966; Brettell, 1982), C. rufilabris (Burmeister) (Putman, 1937; 1956; Hydorn & Witcomb, 1972; Tauber, 1974) and C. zastrowi (Esben-Petersen) (Barnes, 1975). The guts of adults examined during this study did not contain insect remains although Smith (1922) recorded adult C. rufilabris (Burmeister) feeding on aphids in captivity and there are records of C. carnea (V. F. Eastop, pers. comm.) and C. externa and C. asoralis (E. Nunez, pers. comm.) feeding on aphids. Adults of several species of Chrysoperla are known to hibernate during the winter in temperate regions when they undergo a change from green to brown. Chrysoperla is the only chrysopid genus known to diapause in the adult stage of the life-cycle. Henry (1983) has shown that, by vibrating their abdomens, some morphologically identical popu- lations of Chrysoperla produce different substrate- borne calls. These populations will not mate with each other and Henry terms them ‘acoustical species.’ Genus CHRYSOPIDIA Navas Chrysopidia Navas, 1910a: 54. Type species: Chrysopidia nigrata Navas, by monotypy. [Synonymized with Anomalochrysa McLachlan by Kuwayama, 1962: 372; reinstated by Holzel, 19717 57] DISTRIBUTION. Eastern Palaearctic. Fourteen species have been described in this genus. DIAGNos!Is. Adult. Medium-sized lacewings, fore wing 12-17 mm; ground colour pale green. Head unmarked or with red or brown stripes on the scape, gena or vertex; palps tapered apically, marked with black dorsal stripe; labrum straight or indented; mandibles broad, symmetrical or asymmetrical; vertex raised; toruli small; head width : eye width = 1.8-2.8 : 1; scape more or less elongate; antenna often marked red/brown basally; flagellar segments 3 times as long as broad; setae long, pale, arranged in four rings. Pronotum slightly elongate; dorsal setae long, pale. Legs unmarked; setae long, pale; claws basally dilated. Fore wing: costal area narrow at base; costal setae long, slightly inclined or erect; Sc long; Sc and R widely separated; im narrow, ovate, often short; Rs straight or sinuous; grad- ates in two or three parallel or divergent rows; S. J. BROOKS & P. C. BARNARD basal inner gradate meeting Psm; veins not crassate in 6; c; shorter than c>. Hind wing often with black suffusion on apical posterior margin. Abdomen usually unmarked; setae long (especially at apex of ectoprocts), sparse; sternites long, narrow; callus cerci ovate or rounded; dc: setae at apex of ectoprocts long, coarse; atria grossly enlarged in some species; sternite 8+9 fused, elongate apically; 2: ectoprocts slightly invagi- nated dorso-apically. REMARKS. Species of Chrysopidia can be recog- nized by the presence of an elongate scape and long, slightly inclined costal setae. Males of the genus are characterized by the elongation of sternite 8+9 and the long, coarse apical setae on the ectoprocts. Subgenus ANACHRYSA Hoilzel Anachrysa H6lzel, 1973b: 356 [as subgenus of Chrysopidia Navas]. Type species: Chrysopidia (Anachrysa) elegans WHodlzel, by original designation. DISTRIBUTION. Nepal, India (Assam). There are two described species included in the subgenus. DIAGNOsIS. Adult. Medium-sized lacewings, fore wing (Fig. 308) 15-17 mm. Head marked with dark brown stripe on gena and scape; labrum deeply indented; head width : eye width = 1.8- 2.0 : 1; mandibles asymmetrical with small tooth on left mandible; antenna shorter than fore wing. Pronotum with brown medio-lateral spot; meso- and metanotum unmarked. Fore wing rounded apically; marked with brown spot on dcc, gradates marked black; costal setae long, inclined apically; pterostigma marked with small basal red-brown spot; basal Sc crossvein 0.32-0.36 mm; radial crossveins not sinuate; Rs straight; gradates in three parallel series; dcc open at margin. Hind wing with gradates arranged in three parallel series. Abdomen (Figs 309, 310) with callus cerci rounded; trichobothria 30-32; 6d: ectoprocts deeply invaginated apico-dorsally, not fused dorsally, short apical suture between ectoproct and tergite 9; apodemes long, straight; 9: sternite 7 pointed apically. GENITALIA 6 (Figs 311-313). Weakly sclerotized; tignum narrow, long, slightly curved with short median projection; gonapsis straight, very narrow, hardly sclerotized; median plate absent; ento- processus short, thorn-shaped, deeply bifurcate apically; parameres absent; gonarcus long, arcu- ate; arcessus narrow, elongate, tapering apically; THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 207 311 os 312 313 Figs 308-315 Chrysopidia (Anachrysa). 308, 309, 311-313, 315, C. (A.) elegans; 310, 314, C. (A.) sp. indet. 308, fore wing; 309, apex of d abdomen, lateral; 310, apex of 2 abdomen, lateral; 311, d gonarcus complex, lateral; 312, 3 tignum, dorsal; 313, d gonapsis, ventral; 314, 2 subgenitale, caudal; 315, 2 spermatheca, lateral. gonosaccus short, covered in numerous micro- setae; gonosetae, gonocristae and spinellae absent. GENITALIA ¢ (Figs 314, 315). Praegenitale absent; subgenitale bilobed; spermatheca narrow; ventral impression very deep; vela large, straight; duct long, sinuous. REMARKS. Anachrysa can be distinguished from subgenus Chrysopidia by the larger eyes, small brown stigmatic spot and straight radial cross- veins, which are sinuate in Chrysopidia s.str. In males of Anachrysa, gonosetae are absent but a tignum and gonapsis are present. In Chrysopidia S.Str. gonosetae are present but a tignum and gonapsis are lacking. Despite these differences Anachrysa is clearly closely related to Chrysopidia because both groups share several synapomorphies such as the setose abdominal apex, rounded ectoprocts, elongate sternite 8+9 and long costal setae. BioLoGy. Unknown. No insect remains were discovered in the gut contents of adults examined during this study. Subgenus CHRYSOPIDIA Navas DIAGNosIs. Adult. Medium-sized lacwings, fore wing (Fig. 316) 13-16 mm. Head unmarked or with red stripe on scape, gena, between scape and vertex. Head width : eye width = 2.3-2.5 : 1; mandibles asymmetrical with small tooth on left mandible; antenna as long as or longer than fore wing; often marked red/brown basally. Pronotum slightly elongate; unmarked or with brown lateral spot or red median spot; mesonotum unmarked or with red lateral spot on prescutum; metanotum unmarked. Fore wing oval (length : breadth = 2.7-3.1 : 1); unmarked with gradates black, costal and radial crossveins black at each end; costal 208 Figs 316-321 Chrysopidia (Chrysopidia) nigrata. 316, fore wing; 317, apex of ¢ abdomen, lateral; 318, apex of ? abdomen, lateral; 319, d genitalia, lateral; 320, 2 spermatheca, lateral; 321, 2 subgenitale, caudal. setae long, slightly inclined or erect; stigma unmarked; basal Sc crossvein 0.16—0.28 mm; radial crossveins in apical half sinuate; Rs straight or sinuous; gradates in three parallel or divergent rows. Hind wing often with black suffusion on apical posterior margin; gradates in three parallel rows. Abdomen (Figs 317, 318) with callus cerci ovate; trichobothria 21-25; ectoprocts fused dorsally, fused with tergite 9; d: atria grossly enlarged in some species; apodemes absent; ectoprocts with deep dorso-apical invagination, rounded basally (without hinge), lobate apically; 2: sternite 7 straight apically. GENITALIA 6 (Fig. 319). Tignum, gonapsis and S.J. BROOKS & P. C. BARNARD median plate absent; entoprocessus short, narrow, triangular, bifurcate basally; parameres absent; gonarcus long, narrow, arcuate without horns; arcessus long, narrow, tapering apically, strongly curved ventrally; gonosaccus large; gonosetae long, numerous, in central clump; pseudopenis, gonocristae and spinellae absent. GENITALIA 2 (Figs 320, 321). Praegenitale absent; subgenitale bilobed; spermatheca narrow; ventral impression shallow; duct long or short, sinuous; vela long. REMARKS. The subgenus Chrysopidia is dis- tinguished from Chrysotropia by the presence of THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 209 326 three gradate series, and from Anachrysa by the lack of a tignum and gonapsis. BioLoGy. Unknown. The gut contents of adults examined during this study did not include insect remains. Subgenus CHRYSOTROPIA Navas stat. n. Chrysotropia Navas, 1911: 12. Type species: Chrysopa lacroixi Navas [= ciliata Wesmael], | by monotypy. [As subgenus of Chrysopa Leach | by Holzel, 1967: 36; raised to genus by Hélzel, | 1973b: 354.] DISTRIBUTION. East ' Oriental. One species, C. ciliata (Wesmael), is wide- and west Palaearctic, 329 Figs 322-329 Chrysopidia (Chrysotropia) ciliata. 322, fore wing; 323, galea, dorsal; 324, mandibles, dorsal; 325, apex of 6 abdomen, lateral; 326, apex of 2 abdomen, lateral; 327, ¢ genitalia, lateral; 328, 2 subgenitale, ventral; 329, 2 spermatheca, lateral. spread in Europe with a further species from Nepal and a third from the Philippines. D1AGnos!s. Adult. Medium-sized lacewings, fore wing (Fig. 322) 12-15 mm. Head unmarked or with red stripe on vertex and scape; galea (Fig. 323); mandibles symmetrical with tooth on both (Fig. 324); head width : eye width = 2.62.8 : 1; antenna slightly shorter than fore wing. Pronotum unmarked or with yellow median stripe and red lateral spot; meso- and metanotum unmarked or with yellow median stripe. Fore wing broad (length : breadth = 2.6—2.8 : 1), unmarked; costal setae long, erect or slightly inclined; stigma unmarked, sometimes thickened in <6, pale brown; basal Sc crossvein 0.16—0.24 mm; radial crossveins sinuate, oblique; Rs sinuate; gradates in two parallel or divergent rows. Hind wing with 210 posterior apical margin often with grey suffusion. Abdomen (Figs 325, 326) unmarked or with median yellow stripe and red lateral spots; callus cerci ovate; trichobothria 16-24; ¢: ectoprocts with deep apical invagination, dorso-ventrally flattened; 2: sternite 7 straight apically. GENITALIA 6 (Fig. 327). Tignum, gonapsis and median plate absent; entoprocessus short, bifur- cate basally; arcessus narrow, tapering apically, swollen basally; pseudopenis absent; gonarcus long, narrow; gonosaccus short; gonosetae short, numerous; spinellae and gonocristae absent. GENITALIA @ (Figs 328, 329). Praegenitale absent; subgenitale bilobed apically; spermatheca narrow or broad; ventral impression shallow; vela long; duct short, sinuous. Larva. Abdomen broadly fusiform, humped; thoracic tubercles very long, narrow; setae very long, smooth; metanotum with row of long setae; mid-abdominal tubercles short, very broad; abdominal dorso-lateral tubercles absent; abdomi- nal setae hooked dorsally; large debris packet carried covering entire dorsum. REMARKS. It is evident from the close similarities in male and female genitalia and even in the head and wing markings that Chrysotropia and Chrysopidia are closely related. They could almost be considered as synonymous, but because there are only two series of gradates in Chryso- tropia, rather than three as in Chrysopidia, and the mandibles are symmetrical in Chrysotropia but asymmetrical in Chrysopidia we have decided to treat them as distinct subgenera. BIioLoGy. The larva of C. ciliata Wesmael has been described (Killington, 1937; Gepp, 1983). Insect remains were not present in any of the adult guts that were examined during this study. Genus CHR YSOPODES Navas Chrysopodes Navas, 1913b: 329. Type species: Chrysopodes canudasi Navas, by monotypy Orlandisa Navas, 1914a: 112. Type species: Orlandisa jubilosa Navas, by original desig- nation and monotypy. [Synonymized with Chrysopodes Navas by Adams & Penny, 1986: 422.] Ancylochrysa Navas, 1928: 129. Type species: Ancylochrysa nevermanni Navas, by mono- typy. [Synonymized with Chrysopodes Navas by Adams & Penny, 1986: 422.] DISTRIBUTION. Neotropics. Thirty-two described species are included in the genus, distributed throughout the Neotropical region. S. J. BROOKS & P. C. BARNARD DIAGNnosIs. Adult. Medium-sized lacewings, fore wing 10-17 mm; ground colour green or brown. Head often marked with red stripe on gena, clypeus, scape and with postocular spot; labrum indented or straight; vertex raised; head width : eye width = 1.9-2.8 : 1; flagellar setae arranged in four rings; scape sometimes elongate. Fore wing: costal area narrow at base, stigma unmarked; Rs sinuate; basal inner gradate usually meeting Psm. Abdominal setae long, sparse; ectoproct fused dorsally and fused with tergite 9; d (Figs 334, 344): callus cerci ovate; apodeme projecting apically from tergite 9; sternite 8+9 fused; micro- tholi absent; 2 (Figs 335, 345): sternite 7 straight apically. GENITALIA ¢ (Figs 336, 346). Tignum, gonapsis, median plate and entoprocessus absent; arcessus weakly sclerotized, triangular, often domed, with apical hook, pair of medio-lateral sclerotized rods, and dorsal microsetae; parameres absent; gonarcus usually short, broadly expanded later- ally sometimes with narrow lateral horn; pseudo- penis absent; gonosaccus short; gonosetae absent or few, short; spinellae absent. GENITALIA @ (Figs 337, 338, 347, 348). Praegeni- tale absent; subgenitale bilobed apically; sperma- theca tall, cylindrical; ventral impression very deep; vela very long, coiled; duct long, sinuous. REMARKS. Chrysopodes is a complex genus but can be recognized by examination of the male and female genitalia. In males the apodeme in tergite 9 is extended apico-ventrally and protrudes beyond the apex of the abdomen. The shape of the arcessus is also distinctive since it is weakly sclerotized with a prominent apical hook, sclerot- ized lateral rods and often bears dorsal micro- setae. The large spermatheca is distinctive in females of Chrysopodes since it is hardly con- stricted at the junction with the vela which itself is very long and coiled and there is a very deep ventral impression. Chrysopodes may be related to Chrysopiella, males of which also have dorsal microsetae on the arcessus. Subgenus CHR YSOPODES Navas DIAGnosis. Adult. Medium sized lacewings, fore wing (Fig. 330) 12-16 mm. Head often marked with red stripe on gena, clypeus, scape and with postocular spot; galea narrow (Fig. 332); palps elongate (Fig. 15); labrum indented; mandibles scythe-like, symmetrical with no basal tooth (Fig. 331); head width : eye width = 1.9-2.3 : 1; vertex with numerous small indentations; antenna as long as fore wing; flagellar segments 3 times as THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 211 i 2 Zaw@oeaane AN TRS =. ea —— 332 Figs 330-338 Chrysopodes (Chrysopodes). 330, C. (C.) jubilosa; 331, 332, C. (C.) circumfusa; 333-335, 337, 338, C. (C.) canudasi; 336, C. (C.) sp. indet. 330, fore wing (from Kimmins); 331, mandibles, dorsal; 332, galea, dorsal; 333, apex of d abdomen, lateral; 334, apex of 2 abdomen, lateral; 335, 3 genitalia, ventral; 336, d gonapsis, lateral; 337, 2 spermatheca, lateral; 338, 2? subgenitale, caudal. long as broad. Pronotum marked with yellow median stripe; dorsal setae long, pale; meso- and metanotum marked with median yellow stripe. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.7-2.8 : 1); sometimes marked with faint greyish suffusion on posterior margin and around cross- veins; costal setae long, erect; Sc and R close or widely separated; Sc long; basal Sc crossvein 0.16—0.32 mm; im ovate, sometimes quadrangu- lar; medial radial crossveins oblique, sinuate; gradates in two series (sometimes with additional irregular median series), either parallel or diver- gent; anal veins often slightly crassate in 3; c, shorter than c>. Hind wing narrow (length : breadth = 3.1-3.6 : 1). Abdomen (Figs 333, 334) unmarked or with median yellow stripe. . Abdomen (Figs 344, 345) brown or unmarked; trichobothria 23-39; ectoprocts with slight dorso- apical invagination. REMARKS. Although species of Chrysopodes s.str. and Neosuarius are easily distinguished from each other externally, their genitalia are almost identical. This suggests that the two taxa are very closely related and therefore justifies their treat- ment as subgenera rather than distinct genera. Neosuarius seems to be less specialized than Chrysopodes and still retains many plesiomorphic characters which serve to distinguish it from Chrysopodes. These include: broad, asymmetri- cal mandibles with a basal tooth on the left mandible; broad, tapering palps; narrow wings with short setae; straight radial crossveins and unmodified apical abdominal sternite. However, unlike Chrysopodes the subgenitale is slightly extended in females of Neosuarius. BioLoGy. Unknown. No insect remains were present in the guts of any adults examined during this study but the guts of several specimens of C. | | | | | THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 218 341 Figs 339-348 Chrysopodes (Neosuarius). 339, C. (N.) porterina; 340, 341, 344-348, C. (N.) collaris; 342, 343, C. (N.) divisa. 339, fore wing; 340, galea, dorsal; 341, 342, mandibles, dorsal; 343, galea, dorsal; 344. apex of d abdomen, lateral; 345, apex of 2 abdomen, lateral; 346, d genitalia, dorsal; 347, 2 subgenitale, caudal; 348, 2 spermatheca, lateral. (N.) nigripilosa (Banks) were found to be full of pollen grains. Genus CUNCTOCHRYSA Hélzel Cunctochrysa HOlzel, 1970: 47 [as subgenus of Anisochrysa Nakahara]. Type species: Chrysopa albolineata Killington, by monotypy. [Synony- mized with Suarius Navas by Zeleny, 1971: 176; as subgenus of Chrysopa Leach by Tjeder, 1972: 23; raised to genus by Aspéck et al. , 1980: 271.] DISTRIBUTION. Afrotropics, east and west Palaearctic. Cunctochrysa includes four described species: one from southern Africa, two from western Europe and one from Nepal. DIAGNosIs. Adult. Small to medium lacewings, fore wing (Fig. 349) 9-14 mm; ground colour pale green. Head with black or red stripe on gena, clypeus; palps tapered; galea broad; labrum straight; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head 352 S. J. BROOKS & P. C. BARNARD Figs 349-354 Cunctochrysa albolineata. 349, fore wing; 350, apex of d abdomen, lateral; 351, apex of 2 abdomen, lateral; 352, d genitalia, lateral; 353, 2 subgenitale, ventral; 354, 2 spermatheca, lateral. width : eye width = 2.0-2.7 : 1; antenna as long as fore wing; flagellar segments 2-3 times as long as broad; setae in four rings. Pronotum with median yellow stripe; dorsal setae long or short, black or pale; meso- and metanotum with yellow median stripe. Legs unmarked; setae short, black; claws with or without basal dilation. Fore wing narrow (length : breadth = 3.0 : 1); unmarked, gradate veins dark; costal area narrow at base; costal setae short, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.12—0.24 mm; im narrow, ovate, occasionally rectangular; Rs sinuate; gradates in two parallel series, basal inner gradate meeting Psm; veins not crassate in 3; c; about same length as c,. Abdomen (Figs 350, 351) unmarked; setae short, sparse; callus cerci ovate; trichobothria 21—33; d¢: ectoprocts slightly flat- tened caudally, with dorso-apical invagination sometimes pronounced; sternite 8+9 fused, sometimes elongate; sternites sometimes with microtholi; ¢: ectoprocts with slight apical invagination and apical suture between ectoprocts and tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 352). Tignum, gonapsis and median plate absent; entoprocessus long, taper- ing, ventrally curved; arcessus large, broad, ‘axe head’-shaped in lateral view with large ventral hook and dorsal striations; pseudopenis absent; gonarcus long, narrow; gonosaccus short; gono- setae long, numerous, evenly dispersed; gono- cristae and spinellae absent. GENITALIA 2 (Figs 353, 354). Praegenitale absent; subgenitale bilobed apically with basal crumena; spermatheca narrow or broad; ventral impression variable; vela moderate; duct long or short, sinuous. Larva. Abdomen fusiform, humped; thoracic and abdominal tubercies spherical; transverse row of setae on meso- and metanotum; setae long, smooth, hooked apically; small latero-dorsal tubercle present on abdominal segment 6 but absent from rest of abdomen; small packet of debris carried. REMARKS. Species of Cunctochrysa can only be recognized with certainty from the male genitalia THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) PAIS) in which the arcessus is characteristically ‘axe head’-shaped when viewed laterally and bears a ventral hook. However, a ventral hook also occurs in males of Atlantochrysa McLachlan and Mallada oblonga Holzel, suggesting a close relationship between Cunctochrysa and these taxa, which differ mainly in the possession of a tignum and gonapsis (which is the plesiomorphic condition, see ‘Discussion of taxonomic charac- ters’, p. 122). Males of Meleoma and Plesiochrysa ramburi (Schneider) also have a ventral hook to the arcessus but these latter taxa are undoubtedly more distantly related since they do not share any other significant characters with Cunctochrysa. H6lzel’s (1973b) figures of C. opipara clearly show it to be a Cunctochrysa from the shape of the arcessus and the presence of a ventral hook in the male genitalia. However, he also figures a small cruciform gonapsis and, for this reason includes the species in Apertochrysa Tjeder. C. opipara is present in the BMNH collections but we have been unable to find a gonapsis in the male which suggests that H6lzel’s specimen may have been aberrant. BIOLOGY. The larva of C. albolineata (Killington) has been described (Killington, 1937; Gepp, 1983). The gut contents of adults examined in this study did not include insect remains. Genus EREMOCHRYSA Banks Eremochrysa Banks, 1903: 158. Type species: Chrysopa punctinervis McLachlan, by original designation. Lolochrysa Banks, 1950: 59 [as subgenus of Eremochrysa|. Type species: Eremochrysa hageni Banks, by original designation. Syn. n. DISTRIBUTION. Nearctic, West Indies. Of 19 species included in Eremochrysa, 17 occur in the western U.S.A., one species occurs in Canada and another in Cuba. DiAcnosis. Adult. Small lacewings, fore wing 6-11 mm. Labrum straight or indented; mandibles narrow; vertex raised; flagellar segments 2-3 times as long as broad; setae arranged in four rings. Pronotum unmarked or with dark brown lateral stripe; meso- and metanotum unmarked or with broad, black lateral stripe. Legs with setae short, dark claws without basal dilation. Fore wing unmarked, narrow (length : breadth = 2.9-3.6 : 1); costal area narrow at base; costal setae short inclined; stigma unmarked; Sc short; Sc and R widely separated; basal Sc crossvein 0.16-0.32 mm; im ovate; Rs straight; veins not crassate in d; c, shorter than c>. Hind wing without inner gradate series. Callus cerci ovate; trichobothria 20-30; ectoprocts with slight or deep dorso-apical invagination, fused dorsally; d: sternite 8+9 fused, or separated by narrow suture; sternites without microtholi; small, ovate sclerot- ized area, bearing setae present on membrane below tergite 9; 2: setae short, sparse; apex of sternite 7 straight; short apical suture between ectoproct and tergite 9. GENITALIA 6. Tignum absent; gonapsis narrow, hooked apically; median plate absent; entopro- cessus broad, short, positioned dorsally; para- meres absent; gonarcus long, narrow; pseudopenis absent; gonosaccus short; gonocristae and spinel- lae absent. GENITALIA @. Praegenitale absent; subgenitale at apex of long membrane, bilobed apically, slightly extended ventrally. REMARKS. This genus of small lacewings can be recognized by the absence of an inner gradate series in the hind wing, Rs is straight with broad radial cells and the mandibles are narrow. Males are characterized by the narrow, hooked gonapsis and arcessus; the short, dorsally positioned ento- processus; the small sclerotized plate below the ectoprocts; and the blunt lobe at the apex of the ectoprocts. In females the subgenitale is situated at the apex of a long membrane and the basal lobe of the subgenitale projects ventrally. Banks (1950) erected the subgenus Lolochrysa to accommodate those species which had a broad vertex, long pale setae on the pronotum, uni- formly brown veins, and an elongate, upturned apex of sternite 8+9 in males. On examination of the species assigned to Lolochrysa the vertex was not found to be any broader than Eremochrysa species and sternite 8+ 9 did not differ significantly in shape between species of the two subgenera. In addition to the above characters, Lolochrysa species have a row of teeth at the apex of the gonapsis, there is a short suture between sternites 8+9 and the gonosetae are numerous. However, none of these characters is sufficient to regard these species as anything more than a species group and accordingly Lolochrysa is synonymized with Eremochrysa. Subgenus CHRYSOPIELLA Banks stat. n. Chrysopiella Banks, 1911: 344. Type species: Chrysopa_ sabulosa Banks, by original designation DISTRIBUTION. Nearctic. Four species are included in Chrysopiella, all from the western U.S.A. 359 S. J. BROOKS & P. C. BARNARD Figs 355-362 Eremochrysa (Chrysopiella). 355, E. (C.) pallida; 356, 359, E. (C.) sabulosa; 357, 358, 360-362, E. (C.) brevisetosa. 355, fore wing; 356, apex of ¢ abdomen, lateral; 357, apex of 2 abdomen, lateral; 358, d gonarcus complex, lateral; 359, d gonapsis, lateral; 360, 361, 2 subgenitale, ventral; 362, 2 spermatheca, lateral. DiAGNosIs. Adult. Small lacewings, fore wing (Fig. 355) 10-11 mm; ground colour green. Head long, narrow; marked with black spots on gena, clypeus, scape and vertex; mandibles narrow, symmetrical either with small basal tooth on both mandibles or with tooth lacking in both man- dibles; galea narrow, elongate; head width : eye width = 2.9-3.3 : 1, eyes very small; antenna as long as fore wing. Pronotum unmarked or with dark brown median stripe; dorsal setae short, black; meso- and metanotum unmarked or with black longitudinal stripe. Legs unmarked. Fore wing unmarked, narrow (length : breadth = 2.9: 1); basal Sc crossvein 0.24—-0.28 mm; Rs straight, widely separated below R, particularly below stigma; gradates in one series, inner gradate series absent. Abdomen (Figs 356, 357) unmarked; ectoprocts deeply invaginate apically, not fused dorsally; trichobothria 20-24; ¢: abdominal setae short, sparse, often with longer setae inter- spersed; ectoproct with short, broad ventral lobe; ectoprocts fused with tergite 9; setae at apex of sternite 8+9 stout, reclinate. GENITALIA 6 (Figs 358, 359). Arcessus long, narrow with dorsal microsetae at base; gonosetae numerous, long. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 217 Figs 363-372 Eremochrysa (Eremochrysa) punctinervis. 363, fore wing (from Kimmins); 364, apical segment of maxillary palp, lateral; 365, galea, dorsal; 366, mandibles, dorsal; 367, apex of d abdomen, lateral; 368, apex of abdomen, lateral; 369, ¢ gonarcus complex, lateral; 370, ¢ gonapsis, lateral; 371, 2 subgenitale, caudal; 372, 2 spermatheca, lateral. GENITALIA ¢ (Figs 360-362). Spermatheca large; ventral impression shallow; vela moderately long; duct short, sinuous. REMARKS. The male and female genitalia of Chrysopiella and Eremochrysa have many charac- ters in common that are not found elsewhere in the Chrysopidae and this suggests that they are closely related. However, rather than synonymize them, we have decided to treat them as distinct subgenera because there are several characters which are not shared. In Chrysopiella the man- dibles are symmetrical, the galea is narrow, the eyes are small (head width : eye width 3.0: 1), the inner gradates are absent from the fore wing and microsetae are present on the dorsum of the arcessus. However, in Eremochrysa the man- dibles are asymmetrical, the galea_ broad, the eyes larger (head width : eye width = 2.42.9 : 1), an inner gradate series is present in the fore wing and microsetae are absent from the arcessus. BioLoGy. Unknown. Adams & Garland (1981) noted that adult Chrysopiella species were pollen- feeders and that the guts of C. brevisetosa Adams & Garland contained pollen resembling Atriplex (Chenopodiaceae). Pollen was also found in the guts of C. brevisetosa and C. sabulosa (Banks) during this study. 218 Subgenus EREMOCHRYSA Banks D1AGNosIs. Adult. Small lacewings, fore wing (Fig. 363) 6-9 mm; ground colour brown. Head extensively marked with black or red/brown spots and stripes; palps narrow (Fig. 364); galea broad (Fig. 365); mandibles narrow, asymmetrical with basal tooth on left mandible (Fig. 366); head width : eye width = 2.3—2.9 : 1; antenna shorter than fore wing. Pronotum with dark brown lateral stripe; dorsal setae short, black or long, pale, coarse, erect; meso- and metanotum marked with broad, black lateral stripe. Legs marked with black spot at apex of hind femur; 1st tarsal segment elongate. Fore wing very narrow (length: breadth = 3.2-3.6: 1); basal Sc crossvein 0.16— 0.32 mm; gradates in two parallel series, basal inner gradate not meeting Psm. Abdomen (Figs 367, 368) with black dorsal markings; tricho- bothria 21-30; ectoprocts with slight or deep dorso-apical invagination, fused dorsally; d: most setae very short and dense with a few longer setae interspersed; pair of microsetae at base of each seta except on ectoproct and tergite 9; sternite 8+ 9 fused or sometimes with short suture, elongate, tapering apically, deeply bifid apically; setae at apex of sternite 8+9 long, coarse, recli- nate in most species. GENITALIA 6 (Figs 369, 370). Arcessus long, narrow, tapering apically to strong hook, bifur- cate basally; gonosetae long, few or numerous, in lateral clump. GENITALIA @ (Figs 371, 372). Spermatheca minute, broad; ventral impression deep; vela short; duct very short. Larva. Abdomen broadly fusiform, humped; thoracic tubercles short, cylindrical, bearing very long setae; abdominal tubercles spherical; abdominal latero-dorsal tubercles present on tergites 6 and 7; abdominal setae hooked apically; debris carried. BIoLoGy. The larva of E. punctinervis (McLachlan) has been described by Smith (1926a). No insect remains were found in the guts of any adults examined during this study. Genus GLENOCHRYSA Esben-Petersen Glenochrysa Esben-Petersen, 1920: 518. Type species: Glenochrysa typica Esben-Petersen, by original designation. [As subgenus of Chrysopa Leach by Tjeder, 1966: 410; reinstated as genus by New, 1980: 32.] DISTRIBUTION. Afrotropics, Oriental, N. Australia, Samoa. S. J. BROOKS & P. C. BARNARD There are 16 described species and subspecies included in this widely dispersed genus. Four species occur in southern and west Africa, three in the Indian subcontinent, three in the Oriental region, five in Australia (all in Queensland), and one in Samoa. D1AGnNosis. Adult. Small to medium lacewings, fore wing (Fig. 373) 9-13 mm; ground colour pale green. Head with extensive black markings; palps tapered; labrum slightly indented or straight; mandibles broad, asymmetrical with basal tooth on left mandible; vertex steeply raised; head width : eye width = 2.0—2.5 : 1; antenna shorter or as long as fore wing; flagellar segments about 3 times as long as broad; setae arranged in four rings. Pronotum usually marked with dark brown spot in each corner; dorsal setae long, pale, sometimes with clump of long coarse setae; mesonotum with dark brown spots; metanotum with broad dark brown marking or unmarked. Legs unmarked or with black median annulation; setae long, pale; claws with basal dilation. Fore wing narrow (length : breadth = 2.8-3.2 : 1); extensively marked with dark brown and brown- ish yellow shading around crossveins and usually with iridescent pustules; costal area narrow at base; costa slightly convex in basal half; costal setae short, inclined; stigma marked dark brown; Sc very short; Sc and R widely separated; basal Sc crossvein 0.16—-0.36 mm; im narrow, ovate; Rs sinuate; gradates in two parallel rows, basal inner gradate usually not meeting Psm; veins not crassate in d; c, about same length as c); basal branch of Cu, recurrent. Hind wing with dark brown markings, especially in basal half; stigma marked dark brown. Abdomen (Figs 374, 375) often with dorsal dark brown markings; setae long, sparse; callus cerci rounded; trichobothria 23-35; ectoprocts fused with tergite 9; d: ecto- procts broadly elongate apically, with deep apical invagination; atria large or small; sternite 8+9 fused, elongate; 2: ectoprocts with slight apical invagination; sternite 7 straight apically. GENITALIA 6 (Figs 376, 377). Tignum and median plate absent; gonapsis small, indistinct, rod- shaped; entoprocessus long, narrow; arcessus broad, elongate, pointed or trifurcate apically; pseudopenis absent; gonarcus long, narrow with upturned median projection; gonosaccus very long, sometimes protruding from apex of ab- domen; gonosetae long, numerous, evenly dis- persed; gonocristae present in lateral and median rows; spinellae absent. GENITALIA 2 (Figs 378, 379). Praegenitale absent; subgenitale bilobed apically with long basal lobe; spermatheca narrow; ventral THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 219 375 Figs 373-379 Glenochrysa 373, G, typica; 374-376, 378, 379, G. splendida; 377, G. principissa. 373, fore wing (from Kimmins); 374, apex of d abdomen, lateral; 375, apex of 2 abdomen, lateral; 376, d gonarcus complex, lateral; 377, 3 gonapsis, ventro-lateral; 378, 2 spermatheca, lateral; 379, 2? subgenitale, ventral. impression shallow or absent; vela short; duct long, coiled. REMARKS. Glenochrysa is one of the most dis- tinctive genera of the Chrysopini. It is easily recognized by the extensive dark and pale brown wing markings and iridescent pustules, caused by embossed parts of the wing membrane. In addition, Sc is short, the basal branch of Cup is recurrent, the basal half of the fore wing costal margin is slightly convex and the anterior part of the vertex markedly raised. In the male genitalia the gonocristae and deeply invaginated ectoprocts are distinctive, as is the upturned median projec- tion of the gonarcus. In females the long basal lobe of the subgenitale is characteristic of the genus. Tjeder (1966) included two American species, C. bimaculata McClendon and C. cubana Hagen, in Glenochrysa because the male genitalic com- plement was similar, although they lacked wing markings. These species have now been shown to belong to Ceraeochrysa Adams (q.v.). BrioLoGy. Unknown. The gut contents of adults examined during this study did not include insect remains. Genus HIMALOCHRYSA Holizel Himalochrysa H6lzel, 1973b: 367. Type species: Himalochrysa modesta Hodlzel, by original designation and monotypy. S. J. BROOKS & P. C. BARNARD Figs 380-387 Himalochrysa. 380, 384, H. bhandarensis; 381-383, 385-387, H. modesta. 380, fore wing; 381, apex of 6d abdomen, lateral; 382, apex of 2 abdomen, lateral; 383, 5 gonapsis, ventral; 384, d gonarcus complex, lateral; 385, 6 gonarcus complex, dorsal; 386, 2 subgenitale, ventral; 387, 2 spermatheca, lateral. Nepalochrysa H6lzel, 1973b: 347 [as subgenus of Anisochrysa Nakahara]. Type species: Anisochrysa (Nepalochrysa) bhandarensis Holzel, by original designation and monotypy. Syn. n. DISTRIBUTION. Nepal. The genus includes two species. DIAGNOsIs. Adult. Medium lacewings, fore wing (Fig. 380) 15-16 mm; ground colour pale green. Head marked with small red or black spot on gena; palps tapered apically; labrum with deep, narrow indentation; mandibles broad, asymmetri- cal, with basal tooth on left mandible; vertex raised; toruli small; head width : eye width = 1.9-2.2 : 1; scape squared; antenna unmarked, shorter than fore wing; flagellar segments about twice as long as broad; setae short, dark, arranged in four rings. Pronotum with red latero-median stripe in apical half, unmarked or with yellow, median longitudinal stripe; dorsal setae long, pale; mesonotum marked with red spot on pre- scutum or yellow median stripe; metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing length : breadth = 2.8-3.1 : 1; unmarked, gradates black; costal area narrow at base; costal setae quite short, inclined; stigma unmarked; Sc long; Sc and R widely separated; basal Sc crossvein 0.28— 0.32 mm; radial crossveins sinuate in apical half of wing; im ovate (sometimes quadrangular), narrow, long or short; Rs straight; gradates in two or four parallel series; basal inner gradate meeting Psm; inner gradate series extended basally in species with multiple series; veins not crassate in 3; c; slightly shorter than c>. Hind wing with THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 221 gradates black; gradates arranged in two or four parallel series. Abdomen (Figs 381, 382) un- marked; setae long, sparse; callus cerci ovate; trichobothria 25-34; ectoprocts slightly invaginate apico-dorsally, fused dorsally; d: ectoprocts fused with tergite 9; microtholi present on tergites and sternites or absent; sternites 8+-9 fused, short; apodemes straight; 2: sternite 7 straight apically; short apical suture present between tergite 9 and ectoprocts. GENITALIA 6 (Figs 383-385). Tignum and median plate absent; gonapsis absent or large with broad median lobe and shorter, rounded lateral lobes bearing small tubercles on inner face; entoproces- sus short, broad, toothed or tapering apically with ventral lobe; parameres absent; gonarcus long, arcuate with rounded, apico-lateral lobes and broad lateral expansion; arcessus short, narrow, hooked apically, with pair of long, curved sub- apical ventral horns; gonosaccus large with broad median tooth below arcessus; gonosetae few, short, medially placed; gonocristae and spinellae absent. GENITALIA @ (Figs 386, 387). Praegenitale absent; subgenitale bilobed apically, basally slightly elongate, tapering; spermatheca narrow; ventral impression shallow; vela long; duct short, sinuous. REMARKS. Species of Himalochrysa may be dis- tinguished by the complex male genitalia. The arcessus with paired ventro-apical horns, the gonarcus with the broad, rounded latero-apical lobes and the median tooth on the gonosaccus situated below the arcessus are unique in the Chrysopini. Other significant characters shared by species of Himalochrysa include the black gradate crossveins, sinuate radial crossveins, the short arcessus and the entoprocessus which taper apically and have a ventral lobe. H6lzel (1973b) regarded Nepalochrysa as dis- tinct from Himalochrysa and, indeed, there are some characters which are not shared by the two taxa. For example, a gonapsis is present in Himalochrysa but absent in Nepalochrysa, micro- tholi are absent in Himalochrysa but present in Nepalochrysa, the arcessus has a pair of dorsal horns in Himalochrysa which are absent in Nepalochrysa, there are four series of gradates in Himalochrysa but only two in Nepalochrysa, and the intramedian cell is long (and sometimes quadrangular) in Himalochrysa but short in Nepalochrysa. However, apart from the presence or absence of the gonapsis, none of the above characters appears to have much value in separat- ing chrysopid genera since they have frequently been lost or gained within other genera in the family. Therefore, the striking synapomorphic characters which occur in the male genitalia seem to fully justify the synonymy of Himalochrysa and Nepalochrysa. Holzel (1973b) noted that males of Himalo- chrysa are unusual amongst the Chrysopini in the apparent possession of parameres, a character usually confined to the Italochrysini. However, re-examination of the male genitalia has revealed that what Hdlzel took to be parameres is an enlarged gonapsis. BioLoGy. Unknown. There were no insect re- mains in the gut contents of any of the adults examined during this study. Genus KOSTKA Navas Kostka Navas, 1913b: 319. Type species: Kostka nacaratus Navas, by original designation and monotypy. DISTRIBUTION. Sumatra. Only one species has been described, and is known from only two specimens. DIAGNOsIs. Adult. Large lacewings, fore wing (Fig. 388) 22-23 mm. Head unmarked; palps tapered apically; labrum slightly indented; vertex raised; eyes large, head width : eye width = 1.9-2.0 : 1; antennae broken in all specimens examined; flagellar segments about twice as long as broad; setae arranged in four rings. Pronotum elongate, marked with dark lateral spot; setae long, pale; meso- and metanotum unmarked. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing broad (length : breadth = 2.42.6: 1); marked with several large dark spots; costal area narrow at base; costal setae long, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein at level of or slightly basal of m,—m) crossvein; im broad, quadrangu- lar; radial crossveins sinuate; Rs abruptly curved towards R in apical half of wing; gradates in two divergent series; inner gradate series irregular; veins not crassate in 6; c, shorter than c); posterior margin very broad. Hind wing broad (length : breadth = 2.8-2.9 : 1); unmarked. Abdomen (Figs 389, 390) unmarked, very long; setae long, sparse; callus cerci rounded or slightly ovate; trichobothria 29-33; ectoprocts with shal- low apical invagination, not fused dorsally; ecto- procts fused with tergite 9; 5: sternites 8 and 9 not fused, sternite 9 broad, extending beyond apex of ectoprocts; short setae scattered on all sclerites except tergite 9 and ectoproct; setae very dense at apex of sternite 9; sternites 7 and 8 very short; 222 S. J. BROOKS & P. C. BARNARD Figs 388-393 Kostka nacaratus. 388, fore wing (from Kimmins); 389, apex of 2 abdomen, lateral; 390, apex of ? abdomen, lateral; 391, ¢ genitalia, lateral; 392, 2 spermatheca, lateral; 393, 2 subgenitale, ventral. apodemes absent; atria large; ectoprocts rounded basally, not hinged; 2: apex of sternite 7 slightly convex. GENITALIA 6 (Fig. 391). Tignum, gonapsis and median plate absent; entoprocessus very long, broad; arcessus long, narrow, tapering, trifurcate apically; pseudopenis absent; gonarcus long, narrow; gonosaccus very short; gonosetae long, numerous in median clump; parameres absent; spinellae and gonocristae absent. GENITALIA @ (Figs 392, 393). Praegenitale absent; subgenitale bilobed, slightly exten- ded basally; spermatheca very narrow; ven- tral impression very deep, broad, occupying most of spermatheca; vela long; duct long, sinuous. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 223 REMARKS. Kostka can be recognized by the broad wings with a few large spots, irregular inner gradate series and quadrangular intramedian cell. Males of Kostka are unusual in having sternites 8 and 9 not fused and an elongate sternite 9 and in females the very narrow spermatheca and heart- shaped subgenitale are distinctive. It is difficult to suggest a possible sister-group of Kostka, although the basal position of the Sc crossvein possibly indicates ankylopterygine affinities. Similarly, the lateral spot on the elong- ate pronotum and the sinuous radial crossveins suggest a relationship with Austrochrysa Esben- Petersen. BioLoGy. Unknown. Adult gut contents do not include insect remains. Genus MALLADA Navas Mallada Navas, 1925a: 24. Type species: Mallada stigmatus Navas, by monotypy. Anisochrysa Nakahara, 1955: 145. Type species: Anisochrysa paradoxa Nakahara, by monotypy and original designation. [Synonymized with Chrysopa Leach by Adams, 1959: 25; as sub- genus of Chrysopa by Tjeder, 1966: 416; reinstated as full genus by Holzel, 1970: 46; synonymized with Mallada by Adams, 1975: 172.] Triadochrysa Adams, 1978b: 294 [as subgenus of Mallada|. Type species: Mallada (Triadochrysa) triangularis Adams, by original designation and monotypy. Syn. n. DISTRIBUTION. World-wide, but absent from much of the Neotropics. This is the largest genus of the Chrysopidae with at least 122 described species and, doubtless, many more remain to be described. At present 29 species are known from the Afrotropics, 8 from Madagascar and neighbouring islands, 31 species from the western Palaearctic, 8 from the eastern Palaearctic, 19 species from the Indian subcon- tinent, 10 Oriental species, 12 species from Australia and the nearby Pacific islands and a further 5 from south-western Canada, western U.S.A. and Mexico. Two species are particularly widespread: M. boninensis (Okamoto) occurs throughout the Afrotropics and Indo-Malaysia, and M. basalis (Walker) is distributed from Australia to Japan and the Pacific Islands. DIAGNosIs. Adult. Small to medium lacewings, fore wing (Figs 394, 402) 7-16 mm; ground colour pale green, sometimes brown. Head sometimes unmarked or with red, brown or black markings on gena, clypeus, scape, vertex; labrum straight or invaginated; galea broad; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width = 1.9-2.9: 1; antenna slightly longer or shorter than fore wing; flagellar segments 2-3 times as long as broad; setae arranged in four rings. Pronotum unmarked or with yellow median stripe or red or brown stripes or spots; dorsal setae long or short, pale or black; meso- and metanotum marked or unmarked; calciform organs (cuticular glands) present on pronotum and central region of mesonotum in 6. Legs unmarked; setae long or short, pale or dark; claws with or without basal dilation. Fore wing narrow or broad (length : breadth = 2.5-3.2 : 1); unmarked or sometimes with dark suffusion around crossveins or spots; costal area narrow at base; costal setae short, inclined, occasionally long, erect; stigma un- marked or sometimes pale brown; Sc and R widely separated; basal Sc crossvein 0.16— 0.32 mm; im short, ovate; Rs straight or slightly sinuate; gradates in two, occasionally three, parallel series, basal inner gradate not usually meeting Psm; veins often crassate in 6 ; c, shorter than c). Hind wing with Sc and R fused and stigma thickened, pale brown in 6 of some species. Abdomen (Figs 395, 396, 404) unmarked or with red or black markings; setae long, sparse; callus cerci ovate; trichobothria 25-42; ectoprocts with slight apical invagination, fused dorsally, fused with tergite 9; d: microtholi usually absent; sternite 8+9 fused; 2: sternite 7 straight or convex apically. GENITALIA 6 (Figs 397-399, 403). Tignum arcuate or T-shaped, absent in some individuals; gonapsis variable, often three-pronged, absent in some individuals; median plate absent; ento- processus short, straight, occasionally absent; gonarcus long, narrow; arcessus narrow, tapering apically, often with dorsal striations and basal invagination; pseudopenis absent; gonosaccus short; gonosetae few, short; spinellae and gono- cristae usually absent or weakly developed. GENITALIA 2 (Figs 400, 401). Praegenitale absent; subgenitale bilobed apically; spermatheca broad or narrow; ventral impression deep or shallow; vela long or short; duct long or short, sinuous. Larva. Abdomen broad, fusiform, humped; thoracic tubercles elongate, cylindrical; row of setae on metanotum; setae long, smooth or plumose; latero-dorsal abdominal tubercles absent; paired dorsal setae on abdominal seg- ments 5—7; small packet of debris carried. REMARKS. Mallada is a very diverse genus but the 224 S. J. BROOKS & P. C. BARNARD Figs 394401 Mallada. 394, 398, M. boninensis; 395-397, 399-401, M. basalis; 394, fore wing; 395, apex of d abdomen, lateral; 396, apex of 2 abdomen, lateral; 397, d tignum, ventral; 398, ¢ gonapsis, ventral; 399, d gonarcus complex, lateral; 400, 2 subgenitale, caudal; 401, 2 spermatheca, lateral. following characters are shared by most species. The basal inner gradate crossvein does not meet Psm, Rs is straight or only slightly sinuate, a tignum and gonapsis are present in the male genitalia, there are very few gonosetae and the arcessus has dorsal striations and a deep basal invagination. However, in some species, such as M. inotata (Walker) and a second undescribed Australian species, the gonapsis and/or tignum may be absent in certain individuals. Adams & Penny (1987) have suggested that some male genitalic components may become more heavily sclerotized as the insect matures. However, this seems unlikely to be the case in this instance since the gonapsis or tignum may be absent in indivi- duals which are apparently otherwise mature and have the gonarcus and other genitalic structures heavily sclerotized. It seems that some Mallada species are prone to have a variable male genitalic complement; of course, this can lead to errors in generic placement of some species. Triadochrysa Adams differs from Mallada only in the possession of three gradate series. How- ever, from this study it is now clear that a third series of gradate crossveins has arisen indepen- dently within several chrysopid genera and that this character does not have much taxonomic significance. Moreover, Tsukaguchi (1985) has shown that M. babai Kuwayama, which has three series of gradates, is conspecific with M. formosana Matsumurae which has only two rows of gradates. Therefore, there seems to be no justification for regarding Triadochrysa as a distinct subgenus of Mallada. Within Mallada there seems to be a distinct species group which has an Austro-Indonesian THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 225 ond SSlsiaiaiet | art sella ol 403 Figs 402-404 Mallada prasina. 402, fore wing; 403, d gonapsis, ventral; 404, apex of d abdomen, lateral. origin, and comprises adamsi, basalis, boninensis, desjardinsi, dierli, dispar, flaveola, flavostigma, khandalensis, morota, nea, noumeana, scolius, serrandi, signata and tripunctata. In these species the basal inner gradate crossvein meets Psm, c> is relatively long and broad and the radial crossveins in the apical half of the fore wing are usually sinuous or oblique. In males, the apex of the abdomen is gaping; the ectoprocts are compressed caudally and sternite 8+9 is elongate; there is a lateral lobe on sternite 8+-9; the gonapsis is scale- like; there are few gonosetae which often have an elongate, swollen base; in most species veins Sc and R in the hind wing are closely apposed or fused and the stigma is thickened. BIoLoGy. The biologies and larvae have been described in several species including M. boninen- sis (Okamoto) (Tsukaguchi, 1977; Brettell, 1979; New, 1984), M. burgeonina (Navas) (Tjeder, 1966); M. clathrata (Schneider) (Principi, 1956), M. flavifrons (Brauer) (Lacroix, 1925; Killington, 1937; Principi, 1956; Gepp, 1983), M. innotata (Walker) ((Boros, 1984); M. inornata (Navas) (Lacroix, 1925; Gepp, 1983), M. prasina (Burmeister) (Withycombe, 1923; Principi, 1956; Gepp, 1983), M. ventralis (Curtis) (Withycombe, 1923; Gepp, 1983), M. basalis (Walker) (Adams, 1959), M. cockerelli (McCloud) (Smith, 1926b), M. madestes (Banks) (Mehra, 1966; Dessart, 1973), M. microphya (= basalis) (McLachlan) (Terry, 1908; Zimmerman, 1957), M. signata (Schneider) (Boros, 1984); M. traviata (Banks) (Boros, 1984); M. tripunctata (McLachlan) (Boros, 1984). In most chrysopid species occurring in temper- ate regions it is the pupal stage which overwinters; however, in Mallada it is the third instar larva which goes through the winter (Séméria, 1977; Barnard, Brooks & Stork, 1986). Duelli & Johnson (1981) observed males of M. basalis during courtship vibrate their wings so vigorously that they hit the substrate and pro- duced an easily perceptible sound. Thickening of the hind wing pterostigma probably enhances sound production and protects the wing from damage. The adult gut contents do not contain insect remains. Genus MELEOMA Fitch Meleoma Fitch, 1855: 786. Type species: Meleoma signoretii Fitch, by monotypy. DISTRIBUTION. Nearctic, Neotropics. There are 26 described species of Meleoma with seven in the south-western U.S.A. and two species from the south-eastern U.S.A. An addi- tional 14 species occur in Central America (mainly Mexico) and there are three more species from northern South America. S. J. BROOKS & P. C. BARNARD Figs 405-413 Meleoma. 405, 410, M. signoretii; 406-409, 411-413, M. emuncta. 405, fore wing (from Kimmins); 406, apex of d abdomen, lateral; 407, apex of 2 abdomen, lateral; 408, d gonarcus complex, lateral; 409, d tignum, dorsal; 410, 3 gonapsis, ventral; 411, 2 spermatheca, lateral; 412, 2 subgenitale, ventral; 413, 2 subgenitale, caudal. D1AGnosis. Adult. Medium to large lacewings, fore wing (Fig. 405) 11-21 mm. Head unmarked or with red or black stripes on frons, clypeus, gena and vertex; palps tapered apically; labrum indented; vertex raised; head broad, head width : eye width = 2.7-3.9 : 1; frons of 6 often ornamented with horns and cavity; scape some- times elongate; antenna shorter than fore wing, widely separated at base; flagellar segments 1.5— 2.0 times as long as broad; setae arranged in four rings. Pronotum unmarked or with yellow median stripe and red lateral stripe; dorsal setae long or short, pale or dark; meso- and metanotum unmarked or with yellow median stripe and red lateral stripe. Legs unmarked; setae usually short, dark; hind femur sometimes with stridulatory file; claws usually with basal dilation. Forewing narrow (length : breadth = 3.0-3.3 : 1); un- marked; costal area narrow at base; costal setae short, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.24—0.32 mm; im narrow or broad, ovate; Rs sinuate; gradates in two parallel series, basal inner gradate usually meeting Psm; veins usually not crassate in d; c; shorter than c>. Abdomen (Figs 406, 407) un- marked; setae long, sparse; callus cerci ovate; THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 227 trichobothria 28-39; ectoprocts sometimes deeply invaginated apically, fused dorsally, sometimes fused to tergite 9; lateral stridulatory structure sometimes present on 2nd sternite; ¢: ectoproct usually somewhat flattened; microtholi absent; sternite 8+9 fused; ?: sternite 7 straight apically. GENITALIA 6 (Figs 408-410). Tignum present or absent; gonapsis W-shaped, lateral wings ex- panded, bearing gonocristae; arcessus broad, often with dorsal striations; entoprocessus short, projecting ventrally; pseudopenis short, pointed apically, upturned; parameres absent; gonarcus long, narrow; gonosaccus short; gonosetae usually numerous, long; gonocristae usually absent; spinellae absent. GENITALIA 2 (Figs 411-413). Praegenitale absent; subgenitale bilobed apically, often with short median projection, basal lobe similar to praegeni- tale, and broad basal expansion; spermatheca variable; ventral impression deep; vela long or short; duct short, sinuous. Larva. Abdomen narrow, fusiform; thoracic tubercles short, spherical; abdominal latero-dorsal tubercles absent; setae not hooked apically; debris absent or present only in small amounts. REMARKS. Meleoma is very heterogenous but some external characters are shared by most species. The head is usually broad and the antennae are shorter than the fore wing and widely separated at the base. Males often have an intricate head ornamentation or a stridulatory structure and in the male genitalia there is a broad arcessus with upturned pseudopenis. A similar structure occurs in Atlantochrysa, Cunctochrysa and Plesiochrysa ramburi, which suggests that Meleoma may be distantly related to these taxa. Tauber (1969) stated that a tignum (= transverse arch) is present in all species; however, we have been unable to find one in dolicharthra Navas and signoretii Fitch. In the female the basal lobe resembling a praegenitale is an outgrowth of the membranous tube rather than an extension of the base of the subgenitale itself, and is therefore not a true praegenitale. A crumena is not present. The head is narrow in both sexes of those species which do not have frontal ornamentation in males. However, the head is broad even in females of those species in which the males have frontal ornamentation. The frons and clypeus are marked with a broad transverse stripe in species without frontal ornamentation but unmarked in those with ornamentation. BIOLOGY. Tauber (1969) described a complex courtship display in which the male abdomen is vibrated and the wings held out while the female advances and inserts her mandibles into the frontal cavity of the male prior to mating. Some species, which do not have this frontal ornamen- tation in the male, have been found to possess a stridulatory structure in both sexes (Adams, 1962; Tauber, 1969; Brooks, 1987) which is presumably used to produce songs during courtship. The larvae have been described in several species including M. arizonensis Banks (Tauber, 1969), M. cavifrons Banks (Toschi, 1965), M. colhuaca Banks (Tauber, 1969), M. comata Banks (Toschi, 1965), M. dolicharthra Navas (Tauber, 1969), M. emuncta Fitch (Putman, 1932; 1937; Tauber, 1969), M. hageni Banks (Tauber, 1969), M. pipai Tauber (Tauber, 1969), M. kennethi Tauber (Tauber, 1969); M. schwarzi Banks (Tauber, 1969) and M. signoretii Fitch (Putman, 1932; 1937; Tauber, 1969). The gut contents of adults examined during this study did not include insect remains. Genus NINETA Navas Nineta Navas, 1912: 98. Type species: Hemerobius flavus Scopoli, by original designation. [As subgenus of Chrysopa Leach by Banks, 1940: 187; Hélzel, 1965: 92; reinstated as genus by Tjeder, 1966: 345.] Parachrysa Nakahara, 1915: 121. Type-species: Nothochrysa olivacea Gerstaecker, by original designation and monotypy. [Synonymized by Kuwayama, 1962.] DISTRIBUTION. West and east Palaearctic, Nearctic. Nineta is restricted to the Holarctic with seven species occurring in the western Palaearctic, four species in the eastern Palaearctic, one un- described species from the Nilgiri Hills, southern India and two species known from western U.S.A. N. vittata Scopoli is the most widely distributed species and occurs throughout the Palaearctic region. DIAGNOsIs. Adult. Large lacewings, fore wing (Fig. 414) 16-22 mm. Head usually unmarked or with red lateral stripe on vertex and scape or broad dark brown band across frons, front of vertex and between antennae; palps tapered apically; galea long (Fig. 415), broad with large apical papilla; mandibles broad, symmetrical with small basal tooth on each mandible (Fig. 416); labrum almost straight or with deep indentation; vertex flat; head width : eye width = 2.7-3.0: 1 (2.4: 1 in Nilgiri species), head broad; scape sometimes elongate; antenna shorter than fore wing; flagellar segments 1.5—2.0 times as long as broad; setae arranged in four rings. Pronotum 228 S. J. BROOKS & P. C. BARNARD NN 416 415 420 Figs 414-422 Nineta. 414-416, 418, 420-422, N. vittata; 417, 419, N. flava. 414, fore wing; 415, galea, dorsal; 416, mandibles, dorsal; 417, apex of d abdomen, lateral; 418, apex of 2 abdomen, lateral; 419, 5 genitalia, lateral; 420, 2 spermatheca, lateral; 421, 2 subgenitale, ventral; 422, 2 subgenitale, caudal. marked with red or dark brown lateral spot or stripe; dorsal setae short, dark (long, pale in Nilgiri species); meso- and metanotum unmarked or with red or dark brown spot. Legs unmarked; setae short, dark (long, pale in Nilgiri species); claws with basal dilation. Fore wing unmarked; narrow or ovate (length : breadth = 2.7-3.0: 1); costal area narrow at base; costal setae usually very short, inclined (long, erect in Nilgiri species); stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.4-1.3 mm; radial crossveins sinuate; im narrow, ovate; Rs sinuate; gradates in THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 229 two parallel or slightly divergent series; basal inner gradates extended basally, sometimes meet- ing Psm; veins sometimes crassate in 3; costa sometimes concave medially; c,; shorter than c>. Abdomen (Figs 417, 418) elongate, unmarked; setae long, sparse; ectoprocts deeply invaginated apically, fused dorsally, not fused with tergite 9; 6: additional very short, coarse setae present at least on sclerites 3—5; setae long, coarse at apex of ectoprocts and sternite 9; microtholi present or absent; callus cerci rounded or ovate; tricho- bothria 25-32; ectoprocts elongate apically; ster- nite 8 and 9 not fused; sternite 9 elongate, curved dorsally with apical swelling; 9: setae long, coarse at apex of sternite 7; callus cerci rounded; trichobothria 34—37; sternite 7 convex and slightly projecting apically. GENITALIA 6 (Fig. 419). Tignum, gonapsis and median plate absent; entoprocessus straight; gonarcus long, narrow with median horns; arces- sus short, L-shaped; pseudopenis absent; gono- saccus short; gonosetae numerous, long or short, in circular median clump; gonocristae and spinel- lae absent. GENITALIA 2 (Figs 420-422). Praegenitale absent; subgenitale bilobed apically; ventral impression absent or very shallow; vela long; duct long, sinuous; spermatheca narrow. Larva. Abdomen fusiform, not humped; prothor- acic tubercles hardly developed; other thoracic and abdominal tubercles very short, spherical; setae short, smooth, not hooked; latero-dorsal abdominal tubercles absent; debris not carried. REMARKS. Species of Nineta may be distinguished from most other Chrysopini by their large size, and usually by their lack of head markings and elongate scape. The apex of the male abdomen is also very distinctive with the elongate ectoprocts and upturned tip of sternite 9. The genus appears to be very closely related to Tumeochrysa Needham, since both genera share many apomorphies. Similarities in the male genitalia and abdominal apex are particularly striking. However, Tumeochrysa differs from Nineta in that there are three gradate series, dcc is short and closed before the posterior margin of the fore wing, the costal margin of the hind wing is convex and the scape is grossly enlarged in all species, not just narrowly elongate as in some Nineta species. There is a geographical overlap in the ranges of the two genera in India and China but no species in this region show intermediate characters. Therefore, it seems justifiable to continue to regard them as distinct genera. The possession of short abdominal setae and median horns on the gonarcus in males of Nineta is reminiscent of Ceratochrysa Tjeder and there are also similarities with Chrysopa Leach, species of which also have broad heads, a suture between sternites 8 and 9 and the ectoprocts and tergite 9 incompletely fused. Navas (1912) suggested that Nineta was similar to Chrysocerca Weele because males of both genera have apical extensions of the ectoprocts. However, these two genera do not appear to be closely related since they share no significant characters and even the cerci are morphologically quite different. BIoLoGy. The larvae of several species of Nineta have been described including N. carinthiaca (Holzel) (Gepp, 1983), N. flava (Scopoli) (Alderson, 1911a; Killington, 1937; Canard, 1983; Gepp, 1983), N. guadarramensis (Pictet) (Gepp, 1983), N. inpunctata (Reuter) (Gepp, 1983), N. pallida (Schneider) (Brauer, 1867; Gepp, 1983) and N. vittata (Wesmael) (Killington, 1937; Gepp, 1983). Adult gut contents do not include insect remains. Genus PARACHRYSOPIELLA gen. n. Type species: Eremochrysa argentina Banks. DISTRIBUTION. Neotropics. The genus includes two species, one from Argentina and an undescribed species, present in the BMNH collections, from Chile. DiAGnosis. Adult. Small lacewings, fore wing (Fig. 423) 6-11 mm; ground colour green. Head long, narrow; marked with dark brown stripe on gena, clypeus, vertex, scape; labrum slightly invaginated; mandibles narrow, symmetrical with basal tooth on each mandible (Fig. 424); galea long, narrow (Fig. 425); palps long, narrow; vertex slightly raised; head width : eye width = 3.0-4.9 : 1, eyes very small; pedicel elongate; antenna as long as fore wing; flagellar segments 3 times as long as broad; setae short, dark, arranged in four rings. Pronotum marked with dark brown longitudinal stripe; dorsal setae short, coarse, black; meso- and metanotum marked with longi- tudinal median brown stripe. Legs unmarked; setae short, dark; claws with basal dilation. Fore wing oval (length : breadth = 2.62.8 : 1); unmarked; costal area narrow at base; costal setae short, inclined; costal cells broad; Sc short; Sc and R widely separated; basal Sc crossvein 0.24 mm; im ovate; Rs straight; inner gradates absent; veins not crassate in 6; c, slightly longer than c,. Hind wing with inner gradates absent. Abdomen (Figs S.J. BROOKS & P. C. BARNARD Figs 423-431 Parachrysopiella sp. indet. 423, fore wing; 424, mandibles, dorsal; 425, galea, dorsal; 426, apex of d abdomen, lateral; 427, apex of 2 abdomen, lateral; 428, d gonapsis, ventral; 429, ¢ gonarcus complex, lateral; 430, 2 spermatheca, lateral; 431, 2 subgenitale, ventral. 426, 427) unmarked; setae coarse, short, sparse; microtholi absent; trichobothria 20-34; ecto- procts deeply invaginated apico-dorsally, not fused dorsally, fused with tergite 9; ¢: ectoprocts extended apically with short, very coarse setae at apex; sternite 8+9 fused, elongate; apodeme on sternite 8+9 curving ventrally; callus cerci ovate; ?: callus cerci rounded; sternite 7 straight apically. GENITALIA 6 (Figs 428, 429). Tignum absent; gonapsis short, broad, bifurcate; median plate absent; entoprocessus elongate with dorsal median tooth; parameres absent; gonarcus long, arcuate; arcessus short with strong apical hook; pseudo- penis absent; gonosaccus very short; gonosetae absent or very few, short; gonocristae and spinellae absent. GENITALIA @ (Figs 430, 431). Praegenitale absent; subgenitale bilobed apically with long basal extension; spermatheca large, narrow; ventral impression moderate; vela long; duct short, sinuous. REMARKS. Species of Parachrysopiella super- ficially resemble Eremochrysa (subgenus Chry- sopiella) which also lacks inner gradates in the fore wing. However, the male and female geni- talia differ considerably which suggests that they are not congeneric. In males of Eremochrysa, unlike Parachrysopiella, the ectoprocts are not THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 231 extended apically, dorsal microsetae are present on the arcessus, long gonosetae are present, there is a small ovate sclerite situated below the ectoprocts, the gonapsis is long and narrow and the entoprocessus are broad and not toothed. In females of Eremochrysa the subgenitale is not extended basally. In addition, the claws are undilated and the pedicel is not elongate in Eremochrysa. Nevertheless, the two genera do share several synapomorphies, such as the narrow galea, narrow symmetrical mandibles, narrow palps and absence of inner gradates, which suggest a close relationship. BIoLoGy. Unknown. Pollen grains were present in the guts of adults examined during this study, but no insect remains. Genus PEYERIMHOFFINA Lacroix Peyerimhoffina Lacroix, 1920: 83. Type species: Peyerimhoffina pudica Lacroix, by monotypy. Tjederina Hdlzel, 1970: 48. Type species: Chrysopa gracilis Schneider, by original designation and monotypy. [Synonymized with Apertochrysa Tjeder by Zeleny, 1971: 175; as subgenus of Chrysopa Leach by Principi, 1977: 326; re- instated as genus by Aspock et al., 1980: 261.] Syn. n. DISTRIBUTION. Eastern and western Europe, Morocco. At present, two described species are included in Peyerimhoffina but further investigation will probably reveal that they are synonymous. DiAGnosis. Adult. Small lacewings, fore wing (Fig. 432) 9-10 mm; ground colour green. Head marked with dark brown stripe on gena, scape and lateral red stripe below antennae; palps squared apically; labrum straight; vertex raised; head width : eye width = 3.0-3.3 : 1; antenna shorter than fore wing; flagellar segments about twice as long as broad; setae arranged in four rings. Pronotum marked with yellow median stripe; dorsal setae short, dark; meso- and metanotum marked with yellow median stripe. Legs un- marked; setae short, dark; claws without basal dilation. Fore wing very narrow (length : breadth = 3.54.0 : 1); unmarked; costal area narrow at base; costal setae short, inclined; basal costal crossveins sinuous; stigma long, thickened, marked pale brown; Sc and R widely separated; basal Sc crossvein 0.16—0.24 mm; im narrow, ovate; 1st Rs crossvein meets im at or just basal or proximal of apex; mp elongate; Rs straight; gradates in two parallel series; basal inner gradate meeting Psm; inner gradates more numerous than outer grad- ates; veins not crassate in d; c; same length as c); posterior marginal crossveins parallel. Hind wing narrow (length : breadth = 3.2-3.6 : 1); un- marked; inner gradates more numerous than outer gradates; costal margin convex. Abdomen (Figs 433, 434) unmarked or with yellow dorsal median stripe; setae quite short and sparse; callus cerci large, ovate; trichobothria 26-28; ectoprocts very narrow, pointed apico-dorsally; ectoprocts without dorsal invagination, fused dorsally, fused with tergite 9; d: microtholi absent; sternite 8+9 fused, short; apodemes linear; 9: sternite 7 straight apically. GENITALIA 6 (Figs 435, 436). Tignum absent; gonapsis V-shaped; median plate absent; ento- processus short, T-shaped; parameres absent; gonarcus long, narrow, arcuate, lateral lobes large; arcessus short, narrow, tapering to apical hook; gonosaccus small; gonosetae short, few arranged in lateral clump; gonocristae and spinel- lae absent. GENITALIA 2 (Figs 437, 438). Praegenitale absent; subgenitale bilobed apically; spermatheca very narrow, flattened; ventral impression shal- low; vela quite long; duct short, sinuous. Larva. Abdomen narrow, fusiform; thoracic and abdominal tubercles hardly developed; setae short; no debris carried. REMARKS. Peyerimhoffina is a very distinctive genus and has several synapomorphic characters suggesting that it is highly derived. Species may be distinguished by the narrow wings, inner gradates out-numbering the outer ones and the narrow, pointed ectoprocts in both sexes. However, the narrow fore wing, short intramedian cell and sinuous basal costal crossveins are reminiscent of Chrysoperla and these characters do not occur widely in the rest of the Chrysopidae so may indicate a close relationship. Principi (1977) and Aspock et al. (1980) figured a tignum in males of P. gracilis but this was not shown by Kis et al. (1970). We have been unable to find a tignum in any of the males examined in this study. Although we have synonymized Tjederina Ho6lzel with Peyerimhoffina, we have been unable to examine the type specimen of P. pudica Lacroix (the type species of Peyerimhoffina) because it could not be located in the Muséum National d’Histoire Naturelle, Paris. From Lacroix’s (1920) description it is evident that P. pudica shares many characters with Tjederina which are not found elsewhere in the Chrysopidae. The ptero- stigma is long, thickened and pale brown, there are more inner than outer gradates, the wings are 232 S. J. BROOKS & P. C. BARNARD Figs 432-438 Peyerimhoffina gracilis. 432, fore wing; 433, apex of ¢d abdomen, lateral; 434, apex of @ abdomen, lateral; 435, d gonapsis, ventral; 436, d gonarcus complex, lateral; 437, 2 spermatheca, lateral; 438, 9 subgenitale, ventral. very narrow, the basal costal crossveins are sinuous, the first Rs crossvein meets Psm just proximal of the apex of im, the ectoprocts are pointed dorso-apically, the callus cercus is rela- tively large and the species is associated with the conifer Abies numidica. Lacroix commented on the lack of Sc crossveins in the pterostigmal region: these are present in Tjederina but are often difficult to see. BIoLoGy. The larva of P. gracilis has been described by Gepp (1983) and is said to be associated with conifers. No insect remains were found in the guts of adults examined during this study. Genus PLESIOCHRYSA Adams stat. n. Plesiochrysa Adams, 1982a: 28 [as subgenus of Chrysopa Leach]. Type species: Chrysopa brasiliensis Schneider, by original designation. DISTRIBUTION. Neotropics, Oriental, Australasia. The genus includes 23 described species and several undescribed ones are in the BMNH collec- tions. Five species occur in the Neotropics and the rest are widely distributed from Australia and the Pacific Islands, through Malaysia to southern India and the Seychelles. Plesiochrysa ramburi (Schneider) is common and occurs through-out Austro-Malaysia and most of Micronesia. DIAGNOSIS. Adult. Medium to large lacewings, fore wing (Fig. 439) 13-18 mm; ground colour pale green. Head unmarked or with red or yellow markings on gena, clypeus, frons, vertex, mark- ings sometimes black; palps tapered apically; mandibles broad, asymmetrical with basal tooth on left mandible; galea variable (Figs 440, 441); labrum indented; vertex slightly raised; head width : eye width = 1.9-2.6 : 1; antenna slightly longer or shorter than fore wing; scape as long as broad; flagellar segments 2-3 times as long as broad; setae arranged in four rings. Pronotum sometimes elongate with numerous micropocu- lae; dorsal setae coarse, pale; marked with yellow median stripe, red lateral spot or stripe, some- times with black markings; meso- and metanotum unmarked or marked with yellow median stripe or red lateral spots, sometimes with black markings. Legs unmarked; setae long or short, pale or dark; claws with basal dilation, occasionally undilated. Fore wing often narrow (length : breadth = 2.7— 3.3: 1); unmarked, occasionally with brown THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 233 Figs 439-448 Plesiochrysa. 439, 440, P. oceanica; 441, 445, P. ramburi; 442, 444, 446, P. brasiliensis; 443, 447, 448, P. elongata. 439, fore wing; 440, 441, galea, dorsal; 442, apex of ¢ abdomen, lateral; 443, apex of 2 abdomen, lateral; 444, 3 gonarcus complex, dorsal; 445, d gonarcus complex, caudal; 446, d tignum, dorsal; 447, 2 subgenitale, caudal; 448, 2 spermatheca, lateral. : : shading on basal crossveins; costal area narrow at base; costal setae short, inclined; stigma un- marked; Sc long; Sc and R widely separated; basal Sc crossvein 0.20-0.36 mm; radial crossveins straight in Old World species, sinuous in New World species; im ovate, narrow or broad; Rs sinuate; gradates in two, occasionally three, parallel or divergent series; basal inner gradate usually meeting Psm; veins rarely crassate in 0; c, shorter than c); cp rounded apically, broad. Hind 234 wing narrow (length : breadth = 2.8-3.6 : 1); unmarked; gradates in two (occasionally three) series. Abdomen (Figs 442, 443) unmarked or with yellow mid-dorsal stripe or brown markings; setae long, sparse, occasionally dense; callus cerci rounded or ovate; trichobothria 28-40; ectoprocts fused dorsally, with or without suture between ectoproct and tergite 9; d: additional short setae often present on sclerites; microtholi present or absent; ectoprocts broad, basally rounded, deeply invaginated apically; sternites 8 and 9 not fused or incompletely fused with short suture present; apodeme on tergite 9 arcuate; ?: ectoprocts slightly invaginated dorso-apically; sternite 7 straight apically. GENITALIA 6 (Figs 444-446). Tignum narrow, linear in Neotropical species, absent in Old World species; gonapsis and median plate absent; entoprocessus elongate, usually Y-shaped (bifur- cating basally); arcessus absent; parameres absent; gonarcus long, arcuate with paired median horns; pseudopenis elongate, arcuate, tapering apically (in P. ramburi additional up-turned ventral hook bearing setae and bilobed apex is present); gonosaccus large, globular; gonosetae numerous, long, usually projecting inwards, in lateral clump; spinellae absent; gonocristae usually absent but small subapical patch present in ramburi. GENITALIA @ (Figs 447, 448). Praegenitale absent; subgenitale bilobed apically sometimes with basal crumena; spermatheca narrow; ventral impression shallow; vela short; duct long or short, curved, sometimes broad basally. Larva. Abdomen broadly fusiform; thoracic tubercles spherical; setae smooth; row of setae absent from metanotum; laterodorsal abdominal tubercles absent except from tergites 6 and 7; small packet of debris carried. REMARKS. Plesiochrysa may be recognized by its large size, elongate prothorax and long, narrow wings. The genus is obviously closely related to Chrysopa Leach since in males the apodeme on tergite 9 is arcuate, a pseudopenis is present, dorsal horns are present on the gonarcus and the gonosetae are long, numerous and in lateral clumps. However, it differs from Chrysopa Leach in that the head width : eye width is greater, males sometimes possess a tignum, the head markings are generally red not black, the entoprocessus are elongate and not horned, there are short micro- setae on all sclerites, the gut contents show no signs of insect remains and the larvae lack latero- dorsal tubercles. All these characters justify the elevation of Plesiochrysa to a genus. Plesiochrysa is also probably closely related to Ceratochrysa Tjeder which share a similar wing shape and S. J. BROOKS & P. C. BARNARD venation, elongate prothorax and males of which also have additional short abdominal setae. There are four distinct species groups within Plesiochrysa: the Neotropical species; the Old World species including oceanica; ramburi; and an undescribed species from the Solomon Islands. In the Neotropical species the head is marked with narrow red stripes; the wings are narrow; the radial crossveins are sinuous; sternites 8 and 9 are incompletely fused; the ectoproct and tergite 9 are not completely fused; a tignum is present; the short, additional setae on the male abdomen are sparse. The Old World species differ from the others in having slightly broader wings; the head is marked with large red or yellow spots; the intramedian cell is broad; the short setae on the male abdomen are dense; sternites 8 and 9 are entirely separate; the ectoprocts and tergite 9 are completely fused in males and females. In P. ramburi the head and thorax have black markings; the claws are undilated; sternites 8 and 9 are entirely separate; the ectoprocts and tergite 9 are completely fused; an additional ventral hook is present in the male genitalia (Fig. 445); a small patch of gonocristae is present subapically on sternite 9; the gonosetae do not project inwards. In the undescribed species from the Solomon Islands the head is unmarked; the eyes large (head width : eye width = 1.9: 1); thorax is marked with black; the gradates are arranged in three series in the fore and hind wing; the basal crossveins are shaded brown; abdominal setae are dense; micro- tholi are present in males. BIOLOGY. The larvae of two species of Plesiochrysa have been described: P. lacciperda (Kimmins) (Mehra, 1966) and P. ramburi (Schneider) (Adams, 1959). No insect remains were present in the guts of any of the adults examined during this study, although P. paessleri is reported to be an insect feeder (E. Nunez, pers. comm.). Genus REXA Navas Rexa Navas, 1920b: 289. Type species: Rexa lordina Navas, by original designation and monotypy. Eurochrysa Esben-Petersen, 1925: 67. Type species: Chrysopa corsica Hagen, by original designation. [Synonymized by Hédlzel, 1973c: 78.| DISTRIBUTION. Mediterranean. The genus includes three described species. DIAGNosIs. Adult. Medium sized lacewings, fore wing (Fig. 449) 13-14 mm; ground colour olive THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 235 Figs 449-455 Rexa. 449, 451, 454, 455, R. lordina; 450, 452, 453, R. raddai. 449, fore wing; 450, apex of ¢ abdomen, lateral; 451, apex of 2 abdomen, lateral; 452, d gonapsis, dorsal; 453, d gonarcus complex, lateral; 454, ° spermatheca, lateral; 455, 2 subgenitale, caudal. : green. Head with black stripe on gena or red suffusion, red stripe on vertex; palps truncate, flattened apically; labrum deeply invaginated; | vertex raised; toruli small; head width : eye width = 2.3-2.7 : 1; antenna shorter than fore wing; flagellar segments twice as long as broad, first segment elongate; setae arranged in four rings. Pronotum with broad yellow median longitudinal stripe or red lateral stripe; dorsal setae long or short, pale or dark, coarse; meso- and metanotum with yellow longitudinal median stripe. Legs unmarked; setae long or short, pale or dark; claws without basal dilation. Fore wing broad (length : breadth = 2.5 : 1), oval; unmarked; costal area 236 narrow at base; costal setae quite short, inclined; stigma unmarked; Sc long; Sc and R widely separated; basal Sc crossvein 0.32 mm; im long, quadrangular; Rs straight; veins not crassate in 6; gradates in three irregular series or two parallel series; basal inner gradate not meeting Psm; c; shorter than c; cp broad, squared apically; posterior margin broad; posterior marginal cross- veins parallel, forked from just proximal of mid- wing. Hind wing very broad (length : breadth = 2.5 : 1); gradates in three irregular series or two parallel series. Abdomen (Figs 450, 451) unmarked; setae quite short, sparse; tricho- bothria 32-34; ectoprocts with slight dorso-apical invagination, fused dorsally, fused with tergite 9; atria small; ¢: microtholi absent; callus cerci ovate; sternite 8+9 fused, broad, short; 9: callus cerci rounded; sternite 7 straight apically. GENITALIA 6 (Figs 452, 453). Tignum absent; gonapsis broad with pair of strong submedian teeth; median plate absent; entoprocessus broad, triangular, strongly toothed at apical angle; arcessus trifurcate at apex with pair of dorsal teeth basally; pseudopenis absent; gonarcus long, narrow, arcuate; gonosaccus short; gonosetae few, short, positioned in central clump below arcessus; spinellae and gonocristae absent; hypan- drium small. GENITALIA 2 (Figs 454, 455). Praegenitale absent; subgenitale bilobed apically with median projection, lobes widely separated, with very long basal extension; spermatheca very narrow; ventral impression very deep, broad; vela long; duct long, sinuous. Larva. Abdomen humped; jaws shorter than head; thoracic tubercles broad and long, bearing many long, smooth setae with apical hooks; meso- and metanotum with transverse row of setae; abdominal tubercles situated laterally on seg- ments 2-7; abdominal setae hooked apically; debris carried. REMARKS. Rexa may easily be distinguished from other Chrysopini genera by the quadrangular intramedian cell, straight radial sector, broad posterior margin with posterior marginal cross- veins forked from mid-wing and irregular gradate series. The female genitalia are distinctive with the long basal extension of the subgenitale. Rexa may be related to Himalochrysa because of the following shared characters: in the male the arcessus is trilobed apically with dorsal horns, the entoprocessus is toothed apically with a ventral lobe and a gonapsis present; in the female the subgenitale has a ventral extension. The description of Eurochrysa given by Esben- Petersen (1925) agrees closely with that of Rexa S. J. BROOKS & P. C. BARNARD Navas and there can be little doubt that they are synonymous. However, Esben-Petersen (1925) noted that there were two syntypes of Chrysopa corsica Hagen, the type-species, in the de Selys collection, Institut Royal des Sciences Naturelles de Belgique, Brussels. At present, there are two specimens in the de Selys collection over a label reading ‘Chrysopa corsica Hagen’. However, these specimens are Mallada flavifrons (Brauer), and were determined as such by Tjeder in 1967. Hagen’s (1864) description was very detailed and clearly does not refer to the above specimens but does correspond with Esben-Petersen’s diagnosis, so it must be assumed that these are not the types and that they were placed over the ‘corsica’ label after Esben-Petersen examined them. However, there are apparently no other specimens in the de Selys collection which could be the syntypes of C. corsica. BIioLoGy. The biology of Rexa lordina Navas has been investigated in south-west France by Canard & Labrique (1989). In that region the species was associated with the oleaceous bush Phillyrea angustifolia (L.) which supported nymphs of the jumping plantlouse Euphyllura olivina (Costa). Adults were on the wing from May to early July and they had a glycophagous diet. Eggs were laid singly, and the life cycle was univoltine. The gut contents of adults examined in this study did not contain insect remains. Genus SUARIUS Navas Suarius Navas, 1914a: 73. Type species: Suarius walsinghami Navas, by original designation and monotypy. [As subgenus of Chrysopa Leach by Tjeder, 1966: 372; reinstated as genus by Holzel, 1970: 51.] Vasquezius Navas, 1914a: 75. Type species: Vasquezius alisteri Navas, by original desig- nation and monotypy. [Synonymized with Suarius by Hélzel, 1980: 169.] Prochrysopa Tjeder, 1936: 18. Type species: Prochrysopa mongolica Tjeder, by monotypy. [Synonymized with Suarius by Tjeder, 1966: 372; as subgenus of Suarius by Hélzel, 1970: 51.] Syn. rev. DISTRIBUTION. Eastern and western Palaearctic. Suarius includes 17 species and subspecies described from the western Palaearctic region and five species from the eastern Palaearctic. DiAGnosis. Adult. Small lacewings, fore wing (Fig. 456) 6-12 mm; ground colour brown or green. Head usually extensively marked with red THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 237 Figs 456-461 Suarius. 456, S. nana; 457, 459, S. walsinghami; 458, 460, 461, S. caviceps. 456, fore wing (from Kimmins); 457, apex of d abdomen, lateral; 458, apex of 2 abdomen, lateral; 459, ¢ genitalia, caudal (right entoprocessus not shown); 460, ? subgenitale, ventral; 461, 2 spermatheca, lateral. or black stripes; palps tapered; galea short, broad; labrum slightly indented; mandibles broad, asym- metrical with basal tooth on left mandible; vertex raised; head width : eye width = 1.9-2.6 : 1; antenna as long as fore wing; flagellar segments about 3 times as long as broad; setae arranged in four rings. Pronotum marked with broad, brown lateral stripe; dorsal setae short, dark; meso- and metanotum unmarked or with brown lateral stripe. Legs unmarked or with tibial or femoral annulations; setae short, dark; 1st tarsal segment elongate, especially in hind leg; claws without basal dilation. Fore wing quite narrow (length : breadth = 2.6-3.0: 1); unmarked or marked with numerous small black spots; costal area narrow at base; costal setae very short, inclined; underside of R bearing dense, thickened setae (‘scales’) in males of some species; costal crossveins widely spaced, costal cells almost as long as broad; stigma usually unmarked; Sc and R widely separated; basal Sc crossvein 0.16—-0.28 mm; im ovate, broad; Rs straight; few, if any, radial crossveins forking at apex; gradates in two parallel series; inner basal gradate not meeting Psm; R sometimes crassate in 3; c,; about same length as c>. Hind wing with stigma unmarked; dorsal side of R crassate with thickened setae (‘scales’) in ¢ of some species. Abdomen (Figs 457, 458) with extensive black markings or unmarked; setae long, sparse but often coarse, short; callus cerci rounded or ovate; trichobothria 25—33; ectoprocts with broad dorso- apical invagination, fused dorsally or separated by deep groove; d: ectoprocts and tergite 9 fused; sternite 8+9 fused; 2: sternite 7 straight apically; short apical suture present between ectoproct and tergite 9. GENITALIA 6 (Fig. 459). Tignum, gonapsis and median plate absent; entoprocessus long, often broadening apically, not fused apically; para- meres absent; gonarcus long, arcuate with pair of dorsal horns; arcessus narrow, arcuate, trifurcate apically; pseudopenis absent; gonosaccus very small; gonosetae short, few; gonocristae and spinellae absent. GENITALIA @ (Figs 460, 461). Praegenitale absent; subgenitale bilobed apically, elongate basally; spermatheca narrow; ventral impression broad, quite deep; vela moderate length; duct long, sinuous. REMARKS. Species of Suarius may be distin- guished from other chrysopids by their distinctive genitalia. In males, a tignum and gonapsis are absent, the arcessus is trifurcate apically and has 238 dorsal striations, the entoprocessus are long, broad and not fused apically, the gonarcus has a pair of dorsal horns and there are a few short gonosetae. In addition, the wings are unmarked or marked with numerous small spots, the claws are undilated and the female subgenitale is extended basally. Hdlzel (1970) re-established Prochrysopa Tjeder as a subgenus of Suarius to include those species in which males do not possess dense thickened setae (or ‘scales’) on the underside of the radius in the fore wing and the upper surface in the hind wing and in which the radius is not thickened. However, since the subgenus is based solely on the absence of a single character it must fall as a synonym of Suarius. The type of S. walsinghami Navas has aberrant venation so, as Tjeder (1966) points out, the description and figure of the wings given by Navas (1914a) are misleading and do not provide suitable characters on which to base the genus. BIOLOGY. Unknown. No insect remains were found in the guts of any adults examined during this study. Genus TUMEOCHRYSA Needham Tumeochrysa Needham, 1909: 204. Type species: Tumeochrysa indica Needham, by original designation and monotypy Chrysoplecta Navas, 1910a: 55. Type species: Chrysoplecta immaculata Navas, by monotypy. [Synonymized by Banks, 1940: 187.] DISTRIBUTION. Eastern Palaearctic. The seven described species of Tumeochrysa are restricted to the highlands of India, Nepal and eastern China, with one species known from Taiwan. DIAGnosIs. Adult. Large lacewings, fore wing (Fig. 462) 19-25 mm; ground colour green. Head unmarked; palps tapered apically; labrum in- dented; vertex slightly raised; head width : eye width = 2.42.9 : 1, head broad; scape grossly enlarged, elongate; antenna shorter than fore wing; flagellar segments about 1.5 times as long as broad; ist and 2nd antennal segments fused; setae very short, arranged in four rings. Pronotum unmarked; setae very short, dark; meso- and metanotum unmarked. Legs unmarked; setae short, dark; hind coxa swollen; claws basally dilated. Fore wing narrow; unmarked; costal area narrow at base; costal setae very short, inclined; stigma unmarked; Sc and R widely separated; basal Sc crossvein 0.28—-0.41 mm; im ovate, S. J. BROOKS & P. C. BARNARD narrow; Rs straight or sinuate; radial crossveins sinuate; gradates in three or four parallel series; inner gradates extended basally, not meeting Psm; veins not crassate in d; c, shorter than c); dcc short, closed before posterior margin. Hind wing with three gradate series; costa convex. Abdomen (Figs 463, 464) unmarked; elongate; trichobothria 35, densely packed; ectoprocts fused dorsally; 5: setae long, sparse with short, dense setae on tergites 4-5 and very long, erect setae on tergites 7-8; microtholi absent; callus cerci ovate; ectoprocts with deep, broad apical invagination and blunt apical projection, fused with tergite 9; sternites 8 and 9 not fused; sternite 9 elongate, curved dorsally, with large apical tubercle; 2: setae long, dense; callus cerci rounded; ectoprocts with slight apical invagina- tion and incompletely fused with tergite 9 ; sternite 7 straight apically. GENITALIA 6 (Fig. 465). Tignum and gonapsis absent; median plate absent; entoprocessus short, broad; parameres absent; gonarcus long, narrow, arcuate with median horns; arcessus L-shaped, tapering apically; pseudopenis absent; gono- saccus small; gonosetae long, several; gonocristae and spinellae absent. GENITALIA 2 (Figs 466, 467). Praegenitale absent; subgenitale bilobed; spermatheca narrow; ventral impression absent; vela long; duct long, sinuous. REMARKS. Species of Tumeochrysa can be dis- tinguished by the swollen scape, distal cubital cell closed at wing margin, gradates arranged in three or four rows and convex costa in the hind wing. The male genitalia and abdominal apex are very similar to Nineta which suggests a close affinity between these genera since these characters do not occur in the rest of the Chrysopidae. On the basis of the irregular and additional gradate series Needham (1909) suggested that Tumeochrysa was allied to Anomalochrya McLachlan. However, it has now been shown that this character has arisen independently on several occasions in the Chrysopidae and is of little value in assessing possible generic affinities. BioLoGy. Unknown. There were no insect re- mains in any of the guts of adults examined during this study. Genus UNGLA Navas gen. rev. Ungla Navas, 1914b: 224. Type species: Ungla annulata Navas, by original designation and monotypy. [Synonymized with Suarius Navas by Adams, 1975: 169.] THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 239 gil ‘SS SSS 7 [LLEEE (3 Figs 462-467 Tumeochrysa. 462, T. sp. indet.; 463-467, T. indica. 462, fore wing (from Kimmins); 463, apex of 3 abdomen, lateral; 464, apex of 2 abdomen, lateral; 465, d genitalia, lateral; 466, 2 spermatheca, lateral; 467, 2 subgenitale, ventral. DISTRIBUTION. Southern Neotropics. The genus includes four described species that occur in Argentina and Peru. D1AGnosis. Adult. Medium-sized lacewings, fore wing (Fig. 468) 11-13 mm; ground colour pale green. Head marked with brown stripe on gena, vertex and between antennae; palps tapered apically; labrum emarginate; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width =1.8-2.4: 1; scape about as broad as long; antenna about as long as fore wing; flagellar segments about twice as long as broad; flagellar setae arranged in four rings. Pronotum marked with brown spots; dorsal setae short, black; meso- and metanotum 468 _.. AST PUY 469 S. J. BROOKS & P. C. BARNARD 472 Figs 468-473 Ungla binaria. 468, fore wing; 469, apex of ¢ abdomen, lateral; 470, apex of 2 abdomen, lateral; 471, 6 genitalia, dorsal; 472, 2 subgenitale, ventral; 473, 2 spermatheca, lateral. unmarked. Legs unmarked; setae short, black; claws undilated basally. Fore wing unmarked; narrow (length : breadth = 2.9-3.1 : 1); costal area narrow, undilated basally; costal setae short, inclined; stigma unmarked; Sc long; Sc and R widely separated; im short, broad, ovate; mz, long, narrow; Rs sinuate; gradates in two, closely apposed, parallel series; basal inner gradate not meeting Psm; veins not crassate in 6; c, slightly shorter than c). Hind wing narrow (length : breadth = 3.3-3.4 : 1). Abdomen (Figs 469, 470) unmarked; setae long, sparse; callus cerci ovate; trichobothria 29-30; ectoprocts deeply invagi- nated dorso-apically, fused dorsally, fused with tergite 9; d: ectoprocts rounded basally, not hinged; sternite 8+9 fused, elongate; setae at apex of sternite 8+9 short, coarse; atria enlarged; apodeme in tergite 9 absent; @ : sternite 7 pointed apically. GENITALIA 6 (Fig. 471). Tignum, gonapsis, median plate, entoprocessus and parameres absent; gonarcus arcuate with long sinuous lateral horn; arcesuss narrow, tapering apically; pseudo- penis absent; gonosaccus large, globular; gono- setae long, numerous, arranged in lateral clump; gonocristae absent. GENITALIA @2 (Figs 472, 473). Praegenitale absent; subgenitale bilobed apically, slightly extended basally; spermatheca short, narrow; ventral impression moderate; vela long; duct long, sinuous. REMARKS. Externally species of Ungla are similar to Neosuarius Adams & Penny, but they may be THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 241 distinguished by examination of the male and female genitalia. In males of Ungla the apodeme in tergite 9 is absent, the ectoproct is rounded basally, sternite 8+-9 is elongate, the gonosaccus is large and bears numerous gonosetae, and the arcessus is well sclerotized. In Neosuarius the apodeme in tergite 9 protrudes from the apex of the abdomen, the ectoprocts are extended basally, sternite 8+9 is short, the gonosaccus is short and gonosetae are usually absent, and the arcessus is weakly sclerotized and has dorsal microsetae and lateral rods. In females of Ungla the spermatheca is short and pill-box-shaped but in Neosuarius it is large and hardly constricted at the junction with the vela. In his redescription of C. argentina Navas, Adams (1975) described entoprocessus in the male genita- lia. However, it is apparent from his figures and examination of U. binaria that true entoprocessus, which freely articulate with the gonarcus, are absent in the genus. What Adams interpreted as entoprocessus are lateral horns on the gonarcus which are present in many chrysopine genera (some of which also have entoprocessus) and may act as an attachment for the gonosaccus. Adams (1975) incorrectly stated (citing Article 1 of the International Code of Zoological Nomenclature) that the name Ungla Navas is unavailable because the holotype of the type- species is a composite specimen. However, this is a misinterpretation of Article 1 and, according to Article 17, the name is available. In the same paper Adams synonymized the genus with Suarius Navas, species of which also lack a tignum and gonapsis in the male genitalia, and because, like C. squamosa Tjeder, the ectoprocts are not hinged basally. However, it is probable that the tignum and gonapsis have been lost independently in the Chrysopidae on several occasions and the absence of these structures does not necessarily indicate a close relationship between taxa. In addition, C. squamosa has now been shown to belong to Borniochrysa so it would seem that the synonymy with Suarius is unjustified. Moreover, Ungla does not show any of the apomorphies of Suarius such as the dorsal suture between the ectoprocts, a trifurcate arcessus and dorsal horns on the gonarcus. In several chrysopid genera the ectoprocts are rounded basally, such as Borniochrysa, Chrysopidia, Chrysocerca, and it is possible that Ungla is related to these genera, although this seems to be a tenuous link since they share no other apomorphies. BIOLOGY. Unknown. The gut contents of adults examined during this study did not include insect remains. Genus YUMACHRYSA Banks Yumachrysa Banks, 1950: 51 [as subgenus of Chrysopa Leach]. Type species: Chrysopa apache Banks, by original designation. [Syn- onymized with Suarius Navas by Tjeder, 1966: 372; raised to genus by Adams & Penny, 1986: 121.] DISTRIBUTION. Western U.S.A., Mexico At present the genus includes three described species and there is one undescribed species in the BMNH collections. DIAGNOosIs. Adult. Small lacewings, fore wing (Fig. 474) 8-12 mm; ground colour green or brown. Head unmarked or marked with brown stripes on frons, vertex, scape; palps tapered apically; labrum indented; vertex raised; toruli small; head width : eye width = 2.1-2.5 : 1; antenna as long as fore wing; flagellar segments about twice as long as broad; setae arranged in four rings. Pronotum quite broad; dorsal setae long or short, fine or coarse; unmarked or marked with broad brown lateral stripe; meso- and metanotum marked brown laterally. Legs un- marked or with brown spots and stripes on femur and tibia; setae short, dark; claws with basal dilation. Fore wing narrow (length : breadth = 3.0-3.3 : 1); unmarked, crossveins dark; costal setae short; costal area narrow basally; costal cells as broad as long; stigma unmarked; Sc long; Sc and R widely separated; basal Sc crossvein 0.28-0.32 mm; im narrow, ovate; Rs straight; gradates in two parallel series; basal inner gradate meeting Psm; veins not crassate in d ; c, shorter or same length as c>. Hind wing narrow (length : breadth = 3.0-3.8 : 1). Abdomen (Figs 475, 476) marked extensively with brown; setae long, sparse; callus cerci ovate; trichobothria 21-31; d: sometimes with additional short setae sparsely scattered on sclerites; very long, coarse dorsal setae present at apex of ectoprocts; microtholi sometimes present on tergites and sternites; ectoprocts deeply invaginated apico-dorsally, fused dorsally, fused with tergite 9, slightly elongate apically; sternite 8+9 fused, projecting dorsally at apex; ?: ectoprocts slightly invaginate apico-dorsally, incompletely fused with tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 477). Tignum, gonapsis and median plate absent; entoprocessus narrow, greatly extended ventrally, bearing dorsal hooks; arcessus long, swollen medially, tapering apically; parameres absent; gonarcus short, broad, broadly expanded laterally; gonosaccus large; gonosetae numerous, long, in lateral clump; gonocristae in large apical group, with small lateral clump; spinellae absent. 242 S. J. BROOKS & P. C. BARNARD 475 Figs 474-479 }=Yumachrysa. 474, 475, 477, Y. apache; 476, 478, Y. sp. indet. 474, fore wing; 475, apex of d abdomen, lateral; 476, apex of 2 abdomen, lateral; 477, 3 genitalia, lateral; 478, 2 subgenitale, ventral; 479, 2 spermatheca, lateral. GENITALIA @ (Figs 478, 479). Praegenitale absent; subgenitale broad, bilobed with squared basal extension; spermatheca small; vela long, curved; ventral impression deep; duct long, highly coiled. Larva. Abdomen broadly fusiform, distinctly humped; thoracic tubercles cylindrical, long; row of plumose setae present on metanotum; abdomi- nal dorso-lateral tubercles absent except from tergites 6 and 7; setae hooked, long; debris carried. REMARKS. Species of Yumachrysa can be dis- tiguished by examination of the male or female genitalia. In males, the arcessus is long and swollen medially and the entoprocessus are narrow, greatly extended ventrally and bear strong dorsal hooks. In females the small sperma- theca has a relatively long duct and vela with a deep ventral impression and a square basal extension to the subgenitale. Tjeder (1966) linked Yumachrysa with Suarius because the male genitalia of both genera lack a tignum and gonapsis. However, although this is probably the advanced condition, it appears to have arisen independently several times in the Chrysopidae and so does not necessarily indicate a close relationship between the two genera. The characters of the male genitalia of Yuma- chrysa suggest an affinity with Chrysopa Leach. In both genera gonocristae are present, sternite 9 has a subapical dorsal projection, the entoprocessus bear dorsal hooks, the gonosetae are long, numerous and arranged in a lateral clump, and the ectoprocts are deeply invaginated apico-dorsally. Although, unlike Chrysopa, a pseudopenis is not present in Yumachrysa, the arcessus is arcuate and not closely attached to the gonarcus and may be intermediate between a true arcessus and fully detached pseudopenis. BIOLOGY. The larvae and aspects of the biolog of Y. apache (Banks) and Y. yuma (Banks) have been described by Tauber (1975). No insect remains were present in any of the adult guts examined during this study. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 243 Tribe LEUCOCHRYSINI Adams Leucochrysini Adams, 1978a: 211. Type genus: Leucochrysa McLachlan. DISTRIBUTION. Nearctic, Neotropics. DiAGnosis. Adult. Large lacewings, fore wing > 15 mm. Palps tapered apically (Fig. 501); galea broad (Fig. 500); mandibles broad, asymmetrical with tooth on left mandible (Fig. 499); vertex raised; toruli small; eyes large (head width : eye width = 1.6-2.4 : 1); antenna at least 1.5 times longer than fore wing; flagellar segments at least 3 times as long as broad; setae arranged in four rings. Pronotum narrow. Claws with narrow basal dilation. Fore wing marked with black spot on stigma; basal Sc crossvein 0.16—-0.60 mm; im rectangular or broadly ovate; Psm curves towards costa at wing apex before fusing with outer gradates; a few additional gradates distal to outer gradate series; c,; shorter than c); c, squared apically, narrow. Abdominal setae long, sparse; trichobothria < 35; d: microtholi usually present. GENITALIA 6. Tignum, gonapsis and entoproces- sus absent; gonarcus broad, short, transverse with gonocornua; arcessus broad, short with median hook and usually with blunt lateral lobes; para- meres absent; gonosetae absent or few. GENITALIA 2. Praegenitale absent; subgenitale often extended, downcurved basally, and mounted on a broad membranous plate; crumena present; spermatheca narrow; ventral impression deep; vela very long, sinuous. REMARKS. Leucochrysini includes Berchmansus Navas, Cacarulla Navas, Gonzaga Navas, Leuco- chrysa McLachlan (including the subgenus Nodita Navas), Neula Navas, Nuvol Navas and Vieira Navas. Adams (1978a) stated that the Apochry- sinae are closely related to the Leucochrysini. However, there seems to be little substantiating evidence for this as the genitalia and wing venation of the two taxa do not share any signifi- cant apomorphic characters. Conversely, there are several characters which suggest a close relationship between the Leucochrysini and Belonopterygini. The male genitalia of these tribes are quite similar. The gonarcus is broad, transverse and hardly arcuate, and bears lateral gonocornua but entoprocessus are absent. This condition is probably apomorphic within the Chrysopidae since the plesiomorphic condition, which occurs in the rest of the family and also in the Hemerobiidae, is for a narrow, strongly arcuate gonarcus with entoprocessus and no gonocornua. Another link between the tribes is suggested in the female genitalia. The extended and downcurved basal part of the subgenitale in the Leucochrysini might be the precursor of the praegenitale which occurs as a synapomorphy in most of the belonopterygine genera. If this basal section became detached from the rest of the subgenitale and migrated basally it would be termed a praegenitale, and this condition is closely approached in several species of Leucochrysa. A crumena is present in the basal part of the subgenitale in the Leucochrysini and is also present in the praegenitale of the Belonopterygini but is absent from the belonop- terygine subgenitale, which further supports this theory. Genus BERCHMANSUS Navas Berchmansus Navas, 1913b: 327. Type species: Berchmansus adumbratus Navas, by original designation and monotypy. DISTRIBUTION. Trinidad. Three species are included in Berchmansus from Panama and Brazil. Central and South America, DIAGNosIs. Adult. Medium-sized lacewings, fore wing (Fig. 480) 13-15 mm. Head marked with red or black stripes on gena, frons, clypeus and vertex; palps elongate or tapered (in elegans); labrum indented; galea short, broad; mandibles broad, asymmetrical with basal tooth on left mandible; vertex raised; head width : eye width = 1.9-2.4 : 1; scape elongate or as broad as long; antenna twice length of fore wing or same length; flagellar segments about 3 times as long as broad, with setae in four rings. Pronotum with red lateral stripe or unmarked; setae long, pale; meso- and metanotum unmarked or entirely black. Legs unmarked or with black annulation on tibiae; setae long, pale; tarsi very short in adumbratus; claws with or without basal dilation. Fore wing narrow or broad (length : breadth = 2.4-3.3 : 1); lightly suffused on base of Rs and on posterior marginal crossveins or with numerous large black spots; costal area narrow at base; costal setae long, inclined; Sc long; stigma marked with small brown spot; Sc and R widely separated; basal Sc crossvein 0.2—0.4 mm; basal R crossvein proximal of Rs; im ovate; Rs sinuous; gradates in two parallel series; basal inner gradate meeting Psm; veins not crassate in 6; c; slightly longer or shorter than c); cp squared apically; c3; present in adumbratus; dcc very broad in adumbratus. Hind wing narrow (length : breadth = 3.3-3.4 : 1); marked with numerous brown spots or faint suffusion. Abdomen (Figs 481, 482) unmarked or S. J. BROOKS & P. C. BARNARD WM (e225 HH Figs 480-485 Berchmansus. 480, 482, 484, 485, B. adumbratus; 481, 483, B. elegans. 480, fore wing (from Kimmins); 481, apex of d abdomen, lateral; 482, apex of 2 abdomen, lateral; 483, d genitalia, ventral; 484, 2 subgenitale, ventral; 485, 2 spermatheca, lateral. with extensive black markings; setae long, sparse; sternites with microtholi in 6; callus cerci rounded, trichobothria 23-33; sternites large; ectoprocts fused dorsally with slight apical invagi- nation; sternite 8+-9 fused in 6; sternite 7 straight apically in 2; ectoproct and tergite 9 fused. GENITALIA 6 (Fig. 483) (B. elegans only). Tignum, gonapsis, median plate and entoproces- sus absent; parameres absent; gonarcus long, narrow with short gonocornua; arcessus broad, short with apical tooth; additional arcuate struc- ture, sometimes with pair of median horns, situated below gonarcus, attached laterally to ectoproct; pseudopenis absent; gonosaccus, gono- cristae, spinellae absent. GENITALIA 92 (Figs 484, 485). Praegenitale absent; subgenitale bilobed apically; spermatheca very broad or narrow; ventral impression very deep; vela moderate or very long, highly coiled or simply curved; duct long, coiled. REMARKS. Berchmansus can be distinguished from all other chrysopid genera by the position of the basal radial crossvein which leaves the radius before the origin of Rs, and by the arcuate structure, situated ventrad of the gonarcus complex, in the male genitalia. The two species examined in this study (B. adumbratus Navas and B. elegans Guérin) differ considerably in many respects as noted in the generic description. Unfortunately, it has not been possible to examine a male of B. adumbratus so it is unclear at present which characters in the male genitalia are of generic significance. Until the male of adumbratus is described it seems reasonable to group the species in the same genus on the basis of the ee THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 245 489 Figs 486-491 Cacarulla maculipennis. 486, fore wing; 487, apex of d abdomen, lateral; 488, apex of 2 abdomen, lateral; 489, d genitalia, lateral; 490, 2 subgenitale, ventral; 491, 2 spermatheca, lateral. synapomorphy of the basally positioned radial crossvein. However, it is unsatisfactory to base the genus solely on this venational character, particularly in view of the many characters not shared by the two species, and it is possible that the male genitalia of adumbratus will show that the species are not congeneric. BioLoGy. Unknown. The guts of adults examined during this study did not contain insect remains. Genus CACARULLA Navas Cacarulla Navas, 1910b: 479. Type species: Allochrysa maculipennis Banks, by original designation and monotypy. DISTRIBUTION. Colombia, Peru. Only one species is known. D1AGnosis. Adult. Large lacewings, fore wing (Fig. 486) 20-21 mm. Head marked with brown stripe on labrum, clypeus, gena, scape, base of antenna; palps tapered; labrum straight; vertex slightly raised, small; head broad, head width : eye width = 1.8-1.9 : 1; scape elongate; antenna about 1.5 times length of fore wing; flagellar segments about 3 times as long as broad with setae arranged in four rings. Pronotum elongate; marked with median lateral spot; setae long, pale; meso- and metanotum marked with black spots. Legs unmarked; setae long, pale; claws with narrow basal dilation. Forewing length : breadth = 2.7-3.0: 1; marked with numerous black spots; 246 costal area narrow at base; costal setae long, slightly inclined; stigma long, marked with black spot; Sc very long, curving round wing apex; Sc and R well-separated; basal Sc crossvein 0.36— 0.48 mm; im very narrow, long, quadrangular; Rs straight, with two origins; Rs cells in two rows; gradates in three series, inner gradates greatly extended basally, not meeting Psm; a few extra gradates distal to outer gradates; veins not crassate in d; c, much shorter than c; c) squared apically. Hind wing marked with faint shading on posterior marginal crossveins; Rs with one origin; Rs cells not doubled; gradates in two parallel series, greatly extended basally. Abdomen (Figs 487, 488) marked with black dorsal spots; setae long, sparse; callus cerci rounded; trichobothria 26-33; sternites large; ectoprocts with apical invagination, dorsal suture absent; ectoproct and tergite 9 fused; d: sternite 8+9 fused; microtholi present on sternites; 9: sternite 7 straight apically. GENITALIA 6 (Fig. 489). Tignum, gonapsis, median plate, entoprocessus and parameres absent; gonarcus long, narrow, with lateral horns and very large ventro-lateral expansion; arcessus tapering to apical hook with latero-apical lobes; pseudopenis absent; gonosaccus, gonosetae, gonocristae and spinellae absent. GENITALIA @ (Figs 490, 491). Praegenitale absent; subgenitale bilobed apically, extended basally, with small basal crumena; spermatheca very small, narrow; ventral impression very deep; vela long, curved; duct long, sinuous. REMARKS. This striking genus may be distin- guished from other chrysopid genera by the very long antennae; the fore wings which are marked with numerous black spots; cell im which is narrowly quadrangular; the double origin of Rs; Rs cells in two rows; and three rows of gradates in the fore wing with two in the hind wing. BIOLOGY. Unknown. Genus GONZAGA Navas Gonzaga Navas, 1913b: 317. Type species: Gonzaga torquatus Navas, by original desig- nation and monotypy. [Synonymized with Allochrysa Banks by Banks, 1915: 624; re- instated as valid genus by Banks, 1944: 32.] DISTRIBUTION. Neotropics. The seven species are distributed throughout South and Central America and the West Indies (Cuba). DIAGNOsIs. Adult. Medium to large lacewings, S. J. BROOKS & P. C. BARNARD fore wing (Fig. 492) 14-21 mm; ground colour green. Head entirely dark brown or marked with black stripes on frons, clypeus, vertex, scape; palps tapering; labrum deeply indented; vertex raised; mandibles broad, right mandible un- toothed; galea broad, apical papilla prominent; head width : eye width = 1.6—-1.7 : 1, eyes very large; antenna longer than fore wing; flagellar segments 3 times as long as broad; setae arranged in four rings. Pronotum entirely dark or un- marked; dorsal setae long, pale; meso- and metanotum usually with extensive black mark- ings. Legs unmarked; setae long, pale; claws with or without basal dilation. Fore wing broad (length : breadth = 2.3-2.6 : 1); extensively marked with large black spots; costal area narrow at base; costal setae long erect; stigma marked with large black spot; stigmal area between C and Sc often broad; Sc and R widely separated; basal Sc crossvein 0.36—-0.44 mm; im long, quadrangu- lar; Rs straight or sinuate; gradates in two parallel rows, inner basal gradates usually extended basally, not meeting Psm; Psm crossveins often extending beyond outer gradate series; veins not crassate in 6; c, shorter than cy; cp narrow, squared apically. Hind wing marked with black spots. Abdomen (Figs 493, 494) marked entirely black on dorsum; setae long, sparse; callus cerci prominent, rounded; trichobothria 23-39; ster- nites enlarged; ectoprocts fused dorsally; 3d: dense microtholi present on sternites 2-8; ecto- procts with slight apical invagination, incom- pletely fused with tergite 9 and with short ventral lobe; sternite 8+9 fused; short teeth on dorsal margin of sternite 9; 2: ectoprocts with deep apical invagination, fused with tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 495). Tignum, gonapsis and median plate absent; arcessus narrow with dorsal striations and membranous lateral lobes, tapering to apical tooth; parameres absent; gonarcus short, broad, transverse with broad lateral lobes and lateral horns bearing a small submedian tubercle; pseudopenis absent; gonosaccus very small; gonosetae absent or few, long; gonocristae and spinellae absent. GENITALIA 2 (Figs 496, 497). Praegenitale absent; subgenitale bilobed apically, mounted on large plate, basal half curved ventrally at nght angles, bearing median crumena; spermatheca narrow; ventral impression deep; vela long; duct long, coiled. REMARKS. Gonzaga was placed in the Leuco- chrysini by Adams (1978a). The genus appears to be closely related to Leucochrysa McLachlan with which it shares many characters. Indeed, some THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 247 ane 497 496 Figs 492-497 Gonzaga. 492, 493, 495, G. torquata; 494, 496, 497, G. nigriceps. 492, fore wing (from Kimmins); 493, apex of ¢ abdomen, lateral; 494, apex of 2 abdomen, lateral; 495, ¢ genitalia, dorsal; 496, 2 spermatheca, lateral; 497, 2 subgenitale, ventral. species placed in Leucochrysa, such as L. risi which has extensive dark brown wing markings, appear to be very closely related to Gonzaga. However, males of Gonzaga share several syna- pomorphies, which do not occur in Leucochrysa, and which suggest that the genus is distinct. In Gonzaga there is a short lobe at the apex of the ectoprocts, there is no suture between sternites 8+9 and the arcessus is narrow with dorsal striations. Gonzaga may be distinguished from other chrysopid genera by the very long antennae, conspicuous black wing and head markings and the rectangular intramedian cell. Banks (1944) noted that the costal stigmal area is usually twice as broad as the subcostal stigmal area in Gonzaga but in Leucochrysa the costal stigmal area is rarely twice as broad as the subcostal stigmal area. However, there are so many exceptions to this in each genus that the character is practically worthless. BIOLOGY. Unknown. Genus LEUCOCHRYSA McLachlan Leucochrysa McLachlan, 1868: 208. Type species: Chrysopa varia Schneider, by original designation. Protochrysopa Kolbe, 1888: 174. Type species: Chrysopa insularis Walker, by monotypy. [Synonymized by McLachlan, 1894.] Allochrysa Banks, 1903: 143. Type species: Chrysopa virginica Fitch, by original desig- nation. [Synonymized by Navas, 1917.] DISTRIBUTION. Nearctic, Neotropics. There are 160 described species in Leucochrysa. DIAGNOsIs. Adult. Medium to large lacewings, fore wing 10-25 mm; ground colour green. Head with palps tapered apically (Fig. 501); labrum indented; galea broad (Fig. 500); mandibles broad, asymmetrical with small tooth on left mandible (Fig. 499); vertex raised; head width : eye width = 1.82.3: 1, eyes large; antenna longer than fore wing; flagellar segments 3 times as long as broad; setae arranged in four rings. Pronotum with dorsal setae long, pale. Legs unmarked; setae long, pale; claws with basal dilation. Fore wing with costal area narrow at base, costal setae long, slightly inclined; Sc and R widely separated; veins not crassate in ¢; c, shorter than c); cz narrow, squared apically. Abdomen (Figs 502, 503, 513, 514) unmarked or with dorsal black spots; setae long, sparse; callus cerci rounded or 248 S. J. BROOKS & P. C. BARNARD Figs 498-511 Leucochrysa (Leucochrysa). 498-507, L. (L.) varia; 508, 509, L. (L.) dolichocera; 510, 511, L. (L.) longicornis. 498, fore wing (from Kimmins); 499, mandibles, dorsal; 500, galea, dorsal; 501, maxillary palp, dorsal; 502, apex of d abdomen, lateral; 503, apex of 2 abdomen, lateral; 504, ¢ genitalia, dorsal; 505, d arcessus, ventral; 506, 509, 510, 2 subgenitale, lateral; 507, 508, 2 subgenitale, ventral; 511, 2 spermatheca, lateral. ovate; ectoprocts invaginated apically, fused dorsally; ectoproct fused with tergite 9; d: sternites 8 and 9 separated by narrow suture; @?: sternite 7 straight apically. GENITALIA 6. Tignum, gonapsis, median plate, entoprocessus and parameres absent; pseudo- penis absent. GENITALIA 2. Praegenitale absent; subgenitale bilobed apically. BIOLoGy. Larvae of both subgenera of Leuco- chrysa are debris-carriers. Adults examined during this study did not have insect remains included in the gut contents. Subgenus LEUCOCHRYSA McLachlan DISTRIBUTION. Nearctic, Neotropics. There are 41 described species in Leucochrysa s.str. which has its centre of distribution in the Neotropics. The majority occur in central and THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 249 south America, with a few in the West Indies and in southern U.S.A. DIAGNOSIS. Adult. Large lacewings, fore wing (Fig. 498) 15-25 mm; ground colour green. Head unmarked or with red or black markings on gena, clypeus, labrum, vertex, post-ocular region; labrum deeply indented; galea broad; head width : eye width = 1.8—2.1 : 1; antenna longer than fore wing. Pronotum unmarked or with red lateral spot; slightly elongate; meso- and metanotum unmarked or with red/brown spots. Fore wing usually unmarked or with shading on some veins; stigma marked with black basal spot; basal Sc crossvein 0.36—0.60 mm; im quadrangular, broad; Rs almost straight; gradates in two parallel series, inner gradates extended basally, not meeting Psm in most species, outer gradates black; Psm crossveins often extending beyond outer gradates. Hindwing stigma marked black. Trichobothria 23-38; 3: sternites enlarged; microtholi present. GENITALIA 6 (Figs 504, 505). Arcessus broad, short, with apical hook and membranous lateral lobes; gonarcus usually broad, short, transverse, usually with submedian projections; gonosaccus absent or very small; gonosetae absent or few, short; gonocristae and spinellae absent. GENITALIA @ (Figs 506-511). Subgenitale often greatly extended and recurved basally; sperma- theca broad or narrow, sometimes tubular; ventral impression usually deep; vela usually long, sometimes highly coiled; duct long or short, sinuous or coiled. REMARKS. The male and female genitalia and wing venation of Leucochrysa s.str. are very similar to those of Nodita and Gonzaga, suggest- ing a close relationship between these taxa. However, species of Gonzaga may be readily distinguished by the prominent black wing spots which are absent in most Leucochrysa and Nodita species. Also, unlike Leucochrysa and Nodita, males of Gonzaga possess a short lobe at the apex of the ectoprocts, sternites 8+9 are completely fused and the arcessus has dorsal striations. In females of Gonzaga the base of the subgenitale is not as extended or elaborated as in the other two taxa. The wing venation of Gonzaga and Leucochrysa is very similar, in particular both have a quadrangular intramedian cell. However, this does not necessarily indicate a close relation- ship since this is probably the plesiomorphic condition in the Chrysopidae. Species of Leucochrysa s.str. can usually be distinguished from those of Nodita by the quad- rangular intramedian cell and straight radial sector. In Nodita cell im is ovate or triangular and Rs is sinuous, although these characters are not always reliable (Adams, 1977). Nevertheless, the two taxa are apparently very closely related since the genitalia are virtually indistinguishable and for this reason we have decided to treat them as subgenera. However, in the females we have been able to detect a slight difference in the extent of the modification of the subgenitale between the two subgenera. In Nodita the subgenitale is often extended basally and the basal half curved ventrally, sometimes at right angles. In some Leucochrysa species this elaboration is taken a stage further and the basal half is greatly extended and only membranously attached to the rest of the subgenitale. Sometimes the base is completely re- curved on itself so that it resembles a praegenitale. Adams (1978a) suggested that the Apochrysinae were an off-shoot of Leucochrysa, but there seems to be little evidence to support this view since the Apochrysinae lack many of the leucochrysine apomorphies. BIioLoGy. Larvae of Leucochrysa are debris- carriers (Canard & Principi, 1984). Adults exam- ined during this study did not have insect remains included in the gut contents. Subgenus NODITA Navas stat. n. Nodita Navas, 1916: 21. Type species: Nodita ramosi Navas, by PRESENT DESIGNATION. Lachlanita Navas, 1929: 369. Type species: Lachlanita mainerina Navas, by original desig- nation and monotypy. Syn. n. DISTRIBUTION. Nearctic, Neotropics. Only five species of Nodita are known from the U.S.A. compared with 114 species distributed throughout South America. D1AGnosis. Adult. Medium to large lacewings, fore wing (Fig. 512) 10-23 mm. Head marked with red or brown spots and stripes on gena, frons, vertex and scape; vertex slightly raised; head width : eye width = 1.9-2.3 : 1; scape usually slightly elongate; antenna at least 1.5 times length of fore wing. Pronotum marked with red lateral stripe or spot; mesonotum marked with red/ brown spots or entirely suffused; metanotum unmarked. Fore wing rounded apically; un- marked or with a few small black spots, Rs often black medially; stigma usually marked with small basal black spot; basal Sc crossvein 0.8—1.4 mm; im broad, triangular (occasionally narrow, ovate); Rs sinuate; gradates in two parallel series; basal inner gradate usually meeting Psm. Hind wing of larger species with posterior margin usually suffused dark grey; Rs often marked black 250 S. J. BROOKS & P. C. BARNARD Figs 512-518 Leucochrysa (Nodita) intermedia. 512, fore wing (from Kimmins); 513, apex of d abdomen, lateral; 514, apex of 2 abdomen, lateral; 515, 3 genitalia, lateral; 516, 2 subgenitale, lateral; 517, 2 subgenitale, caudal; 518, 2 spermatheca, lateral. medially. ¢: sternites small, with or without microtholi; trichobothria 22-31; 9: trichobothria 25-34. GENITALIA 6 (Fig. 515). Arcessus short with median apical tooth, usually with lateral pro- jections; gonarcus short, broad with lateral projections; gonosaccus short; gonosetae, spinel- lae and gonocristae absent. GENITALIA @ (Figs 516-518). Subgenitale often extended and downcurved basally, with basal papilla, mounted on broad, membranous struc- ture; spermatheca short, broad; ventral impres- sion deep; vela very long; duct long, coiled. REMARKS. Species of Nodita can be distinguished by their long antennae, the black spot on the stigma, the broadly triangular intramedian cell and the sinuate radial sector. Nodita is very similar externally to Leucochrysa s.str. but in Leucochrysa the intramedian cell is quadrangular, Rs is straight, and Rs and the posterior margin of the wing are unmarked. However, these charac- ters are not always reliable and Adams (1977) noted that some species, such as Leucochrysa risi Navas, could be assigned equally well to either group. The male and female genitalia of Nodita and Leucochrysa are almost indistinguishable (see above), but because of the venational differences we have decided to recognize Nodita as a sub- genus of Leucochrysa, rather than as a synonym. Navas (1916) made Nodita available when he described two species in the genus, although he did not include a description of Nodita itself. Instead he merely noted that it was the same as Leucochrysa sensu Banks nec McLachlan. No type species was designated. He later (Navas, 1917) made a formal description of Nodita and designated Chrysopa intermedia Schneider as type species. However, since this species was not THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 251 Fig 519 Nuvol umbrosus, fore wing. included in the original description this desig- nation is invalid. BIOLOGY. The larva of N. floridana Banks was described by Smith (1926b), but the only informa- tion given was that it was a debris-carrier. N. oenops has recently been partially figured and described by Adams (1987). This species is also a debris-carrier and is shown to have extremely elongate prothoracic tubercles. Genus NEULA Navas Neula Navas, 1917: 280. Type species: Neula mesana Navas, by original designation and monotypy. DISTRIBUTION. Colombia. This genus is monotypic. DiAGNosis. Adult. Large lacewings, fore wing 25 mm; ground colour yellowish green. Head marked with red stripe on vertex; antenna longer than fore wing, marked black at base. Thorax marked with red lateral stripe. Legs unmarked; claws with basal dilation. Fore wing unmarked; broad; costal area narrow at base; Sc long; stigma marked with black basal spot; im quadrangular; Rs sinuate; gradates in three parallel series; inner gradates greatly extended basally; basal inner gradate not meeting Psm; Psm extending slightly beyond base of outer gradate series. Hind wing with gradates in three series; posterior margin with brownish suffusion. GENITALIA. Unknown. REMARKS. It has not been possible to trace any specimens of this genus. From the long antennae, long, broad fore wing, black spot on the stigma and upturned Psm extended beyond the base of the outer gradates it is evidently a leucochrysine genus, probably related to Leucochrysa. How- ever, the presence of three gradate series is distinctive and should make it easy to recognise if any specimens are subsequently discovered. BIOLOGY. Unknown. Genus NUVOL Navas Nuvol Navas, 1916: 24. Type species: Nuvol umbrosus Navas, by original designation and monotypy. DISTRIBUTION. Neotropics (Brazil). This genus is monotypic. DiaGnosis. Adult. Large lacewings, fore wing (Fig. 519) 17 mm. Head marked with black lateral stripe on vertex and black postocular spot. Pronotum marked with three longitudinal black stripes; setae long. Legs unmarked; claws basally dilated. Fore wing marked with five, faint yellow- ish brown transverse stripes radiating from centre of wing; costal area narrow at base; costal setae short, inclined; stigma marked with small black spot; Sc and R widely separated; R greatly extended apically, curving posteriorly around wing apex; branching veinlets at apex of Sc and R unforked, marked black; im short, broadly ovate; Rs almost straight; gradates arranged in two parallel series; outer gradates closely aligned; inner gradates extended basally, not meeting Psm; c, about as long as c; cp broadening apically; dcc closed before posterior margin of wing. Hind wing marked similarly to fore wing. REMARKS. The type specimen of N. umbrosus is apparently lost. However, we are grateful to Prof. P. A. Adams for providing a photograph of a specimen of umbrosus which is in the collection of the University of Sao Paulo Museum of Zoology. Unfortunately, we have been unable to examine the actual specimen which seems to be the only extant example of this species. The specimen is in poor condition, lacking the left fore and hind ZOD, wings, antennae and abdomen, although the right fore and hind wings are in good condition leaving no doubt about the identity of the specimen. Nuvol can be readily distinguished from other chrysopid genera by the long apical extension of the radius, which curves posteriorly at the wing apex, running parallel to the costa. In addition, there are seven unbranched veinlets at the apex of the radius. In other Chrysopidae (except the Apochrysinae) the radius is shorter, with fewer apical veinlets, most of which are branched. The close alignment of the outer gradate series is also an unusual feature of Nuvol, since in most Chrysopidae the outer gradates are arranged step- wise, although this character is paralleled in the Apochrysinae and in Vieira. The short, almost triangular intramedian cell and the unusual black longitudinal stripes on the head and pronotum of Nuvol are also similar to Vieira and suggest that these two genera may be closely related. Nuvolis clearly a member of the Leucochrysini, with which it shares several venational characters, and the characters described above indicate its generic validity, but further speculation on the generic affinities of Nuvol must await the dis- covery of additional specimens with their ab- domens intact so that the genitalia can be examined. BIOLOGY. Unknown Genus VIEIRA Navas Vieira Navas, 1913a: 152. Type species: Leuco- chrysa leschenaulti Navas, by original desig- nation and monotypy. DISTRIBUTION. French Guiana, Surinam, Amazonia. This genus is known from only two species. D1aAGnosis. Adult. Large lacewings, fore wing (Fig. 520) 21-24 mm. Head marked with black stripes on gena, frons, laterally on vertex; mandibles broad, asymmetrical with basal tooth on left mandible; palps entirely black, tapered apically; labrum emarginate; vertex flattened; toruli small; head width : eye width = 1.6-1.8 : 1, eyes very large; antenna shorter than fore wing; flagellar segments 2 times as long as broad; setae short, in four rings. Pronotum broad; marked with black lateral stripe; setae very long, black, coarse; mesonotum with black stripes along sutures; metanotum marked with black lateral stripe. Legs long; unmarked; setae very long, pale; claws with small basal dilation. Fore wing broad (length : breadth = 2.42.6 : 1); extensively marked with large pale brown spots and stripes; costal setae S.J. BROOKS & P. C. BARNARD long, erect; costal area narrow at base, widening abruptly at level of Sc crossvein; costal crossveins highly sinuate at level of Rs base; Sc long; stigma marked with brown spot; Sc and R widely separ- ated; basal Sc crossvein 0.32—0.36 mm; radial crossveins sinuate; im short, broad, triangular; m long, narrow; first Psm crossvein oblique; Rs very sinuate; gradates in two divergent rows; basal inner gradate meeting Psm, not extending basally; veins not crassate in 6; c; and cy about same length; c. broad, squared apically; dcc open at wing margin, anterior corner adjacent to c) not Psm. Hind wing broad (length : breadth = 2.7-3.1 : 1), falcate apically; marked with large pale brown spot on mid Rs and on pterostigma. Abdomen (Figs 521, 522) marked with pair of longitudinal submedian and lateral black stripes; setae long, sparse (shorter on tergites than sternites); callus cerci ovate; trichobothria 40-46; ectoprocts with slight dorso-apical invagination, fused dorsally, fused with tergite 9; tergite 9 short basally, not hinged; sternites enlarged; spiracles large; d: microtholi present on sternites; sternite 8+9 fused; 2: sternite 7 straight and setose at apex. GENITALIA 6 (Fig. 523). Tignum, gonapsis and median plate absent; parameres absent; gonarcus short, broad, transverse, with long lateral horn tapering to apical tooth; arcessus short, broad with strong apico-median hook and blunt lateral lobes bearing 2 short gonosetae situated laterally below arcessus; gonosaccus short; gonocristae and spinellae absent. GENITALIA @ (Figs 524-526). Praegenitale absent; subgenitale bilobed apically with median pro- jection, extended and ventrally curved basally; spermatheca long, narrow; vela large; ventral impression deep, narrow; duct long, coiled. REMARKS. Vieira may be distinguished by its large broad fore wings which are extensively marked with pale brown spots and streaks and have very sinuate radial and basal costal crossveins. It is the only leucochrysine genus in which the antennae are shorter than the fore wing. BIoLoGy. Unknown. Insect remains were not included in the gut contents of adults examined during this study. Subfamily NOTHOCHRYSINAE Navas Notochrysinos Navas, 1910a: 59. Type-genus: Nothochrysa McLachlan. Nothochrysini Navas, 1913b: 303. Dictyochrysinae Esben-Petersen, 1918: 26. Type- genus: Dictyochrysa Esben-Petersen. [Syn- onymized by Adams, 1967: 219.] THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 253 = 1 ee DE aa Z Witter \ yrs MM, S\N a — EO Ge an ae AN" Figs 520-526 Vieira leschenaulti. 520, fore wing (from Kimmins); 521, apex of 2 abdomen, lateral; 522, apex of 2 abdomen, lateral; 523, d genitalia, dorsal; 524, 2 spermatheca, lateral; 525, 2 subgenitale, lateral; 526, 2 subgenitale, ventral. Nothochrysinae Navas; Adams, 1967: 219. width : eye width = 2.54.7 : 1; eyes small; palps rounded apically (Fig. 547); toruli large; vertex domed; pedicel hardly constricted medially; antennae shorter than fore wing; flagellar seg- DIAGNOsIs. Adult. Small to large lacewings, fore ments narrow, at least twice as long as broad, wing 6-23 mm. Head narrow, elongate; head often short; setae arranged in 5-6 rings. Claws DISTRIBUTION. Afrotropical, western Palaearctic, Australia, western Canada and U.S.A. 254 usually without basal dilation. Fore wing quite narrow (length : breadth = 2.6-3.2 : 1); costal area narrow; costal setae short, inclined; basal Sc crossvein relatively distal (0.76-1.68 mm); im broad, variable in shape; 1st Rs crossvein meets im in basal half; R and Cu hardly swollen basally; M sometimes forked after m3; tympanal organ absent; Rs arises distally; c; often considerably shorter than c); Psm and Pcu widely separated; Psm short, fusing with inner gradates; gradates arranged in two or three rows; jugal lobe large; veins not crassate in ¢6. Abdomen short; setae short, sparse, often absent from intersegmental membrane; trichobothria 20-39; d: ectoprocts with slight dorso-apical invagination; sternites 8+9 fused; @: ectoproct and tergite 9 not completely fused. GENITALIA 6. Tignum, gonapsis, median plate, gonocristae and spinellae absent; gonarcus arcu- ate; entoprocessus present, often long and well developed. GENITALIA @. Praegenitale absent; subgenitale bilobed apically, extended basally; spermatheca large. REMARKS. The Nothochrysinae may be readily distinguished from the rest of the Chrysopidae by the short pseudomedian vein which is continuous with the inner gradates. In the rest of the Chrysopidae Psm meets the outer gradates. Adams (1967) established that the Notho- chrysinae are the most primitive group of the Chrysopidae on the grounds that they retain many archaic features; our analysis confirms that they are the least derived subfamily. The most ad- vanced group of genera within the Nothochrysinae appears to include Nothochrysa and the two Australian genera Dictyochrysa and Triplochrysa. Unlike other nothochrysine genera Psm is dis- tinct, c, long, the claws are basally dilated, 1A is forked and no pollen was found in the gut contents of adults. Hypochrysa lacks these apomorphies but shares two other apomorphies with the group: in males the tergites and ectoprocts are fused, and microtholi are present. The remaining three genera are probably the most primitive in the subfamily, although we have been unable to find an apomorphic character to link them. The South African genera Kimochrysa and Pamochrysa both have Sc and C fused basal to the stigma in the hind wing which suggests a close relationship. In Pamochrysa and Pimachrysa tergite 8 is extended laterally and the spiracle is located on the tergite rather than on the lateral membrane. These are the only two genera in the entire Chrysopidae which exhibit this character. However, it would appear to be plesiomorphic since it occurs S. J. BROOKS & P. C. BARNARD widely in the Neuroptera (e.g. Hemerobiidae, Myrmeleonidae, Rapismatidae) and in the Raphidioptera and so does not necessarily suggest a close relationship between these two genera. Key to the genera of the Nothochrysinae 1 Gradate crossveins arranged in three or more series (Fig. S67) edad cident soso he dP ew oes eee 2 533) .y. i fo See Se elle eee & 3 2 Gradates arranged in 5-7 irregular series (Fig. 527); 1A forked..... DICTYOCHRYSA Esben-Petersen — Gradates arranged in three parallel series (Fig. 567); 1A unforked......... TRIPLOCHRYSA Kimmins 3 Scand C fused basad of pterostigma in hind wing . .4 — ScandCnotfusedin hindwing ................. 5 4 Cell im rhomboidal, about as long as broad (Fig. SSA) head Sie oe Meee re PAMOCHRYSA Tjeder — Cellim triangular or pentagonal, about twice as long as broad (Fig. 539)........ KIMOCHRYSA Tjeder 5 Cell im narrow, quadrangular (Fig. 545); large species, fore wing > 14 mm; 1A forked in fore Wilts theses gees NOTHOCHRYSA McLachlan — Cell im broad, subtriangular (Fig. 533); small species, fore wing < 10 mm; 1A not forked in fore WAND. ws < oka S Siee cuelgyess tes =) s etre eee 6 6 2A and 3A not fused in fore wing (Fig. 561) PIMACHRYSA Adams — 2A and 3A fused apically in fore wing (Fig. 533) HYPOCHRYSA Hagen Genus DICTYOCHRYSA Esben-Petersen Dictyochrysa Esben-Petersen, 1917: 214. Type species: Dictyochrysa fulva Esben-Petersen, by original designation and monotypy. DISTRIBUTION. Australia and Tasmania. Three species have been described from this region. DIAGNOsIS. Adult. Medium-sized lacewings, fore wing (Fig. 527) 14-17 mm; ground colour brown. Head unmarked or marked with brown arcuate stripe on front of vertex; palps truncate, rounded apically; labrum indented; vertex flattened; toruli large; head broad (head width : eye width = 2.6— 3.9 : 1); scape as long as broad; pedicel as long as broad, hardly constricted medially; antenna longer than fore wing; flagellar segments about 2.5 times as long as broad; setae arranged in six rings. Pronotum broad; marked with numerous dark spots and stripes; dorsal setae short, dark, THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 255 WLLL S th ipi Figs 527-532 Dictyochrysa. 527, D. fulva; 528-532, D. peterseni. 527, fore wing (from Kimmins); 528, apex of 3 abdomen, lateral; 529, apex of 2 abdomen, lateral; 530, d genitalia, dorsal; 531, 2 spermatheca, lateral; 532, ? subgenitale, ventral. coarse; mesonotum with black markings; meta- notum entirely black or unmarked. Legs un- marked or with black annulations; setae short, dark; pair of tibial spurs present; claws without basal dilation. Fore wing unmarked; basal half with numerous rounded shallow depressions in some species; slightly pointed apically; costal area narrow at base but abruptly expanding subbasally; costal setae very short, coarse, inclined; costa indented immediately proximal to stigma; Sc quite long but not extending beyond pterostigma; Sc and R widely separated; basal Sc crossvein 1.48-2.12 mm; M not fusing basally with R; tympanal organ absent; im not apparent but M forked; Rs straight; gradates arranged into S—7 irregular series, wing highly reticulated; veins not crassate in 6; c, shorter than c); 1A forked. Hindwing with C and Sc fused (or very closely apposed) just basad of pterostigma. Abdomen (Figs 528, 529) unmarked or sclerites marked dark brown; setae short, sparse; callus cerci rounded or ovate; trichobothria 32-54; ectoprocts slightly invaginated dorso-apically, fused dorsally; atria small; ¢: ectoproct fused with tergite 9, narrowly extended basally; microtholi absent or present on tergites; sternites 8+9 fused; 2: ectoproct not fused with tergite 9; sternite 7 straight apically. GENITALIA 6 (Fig. 530). Tignum, gonapsis and median plate absent; entoprocessus long, spoon- shaped; gonarcus arcuate with short medio-lateral projections; arcessus long, narrow, broadening apically with narrow dorsal groove; pseudopenis, gonocristae and spinellae absent; gonosaccus short, gonosetae short, numerous in lateral clump. GENITALIA ¢ (Figs 531, 532). Praegenitale absent; subgenitale extended basally, bilobed apically with small median projection; ventral impression broad, deep; vela very short; duct short, curved, highly setose in apical half; sperma- theca broad. Larva (Ast instar only). Abdomen fusiform, not humped; abdominal tubercles hardly developed; chalazae absent; dorsal thoracic setae and ab- dominal setae not hooked apically; debris not carried. REMARKS. Dictyochrysa is readily distinguishable from other nothochrysine genera by the highly reticulated venation in the discal area of the wings. BIOLOGY. The first instar larva of Dictyochrysa fulva Esben-Petersen has been described by New S. J. BROOKS & P. C. BARNARD Figs 533-538 Hypochrysa elegans. 533, fore wing (from Kimmins); 534, apex of 2 abdomen, lateral; 535, apex of ? abdomen, lateral; 536, d genitalia, dorsal; 537, 2 subgenitale, ventral; 538, 2 spermatheca, lateral. (1981a). The eggs are stalked and laid singly. None of the adults examined had insect remains included in the gut contents. Genus HYPOCHRYSA Hagen Hypochrysa Hagen, 1866: 377. Type species: Chrysopa nobilis Schneider, by monotypy. Hypochrysodes Leraut, 1980: 243 [replacement name for Hypochrysa Hagen; unnecessary replacement name; Oswald, 1987: 225.] DISTRIBUTION. Southern Europe, Argentina. There are only two species known in Hypochrysa and only females are known of H. viridula Adams, the Argentinian species. DIAGNOSIS. Adult. Small lacewings, fore wing (Fig. 533) 9-10 mm; ground colour brown. Head marked with black median stripe and stripe on gena; palps rounded apically, slightly elongate; labrum indented; vertex domed; toruli large; eyes small (head width : eye width = 4.44.7 : 1); scape as broad as long; antennae dark brown, shorter than fore wing; flagellar segments 3 times as long as broad; setae arranged in five rings. Pronotum unmarked or marked with median and lateral black stripe; dorsal setae few, long, pale; meso- notum unmarked or marked with black stripes along sutures; metanotum unmarked. Legs with black stripes on tibiae and femora; setae short, dark; claws without basal dilation. Fore wing unmarked; rounded apically; costal area narrow at base; costal setae very short, inclined; stigma suffused with pale brown; Sc long; Sc and R widely separated; basal Sc crossvein 1.16— 1.40 mm; im broad, triangular; M forking after m3; Rs straight; gradates in two parallel series; inner gradates not fused with Psm; c, at most half as long as cz; 2A and 3A fused apically; 1A not forked; dcc broad, open at margin. Abdomen (Figs 534, 535) unmarked or with dark brown THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 257 markings; setae short, sparse; callus cerci ovate; trichobothria 20-27; ectoproct with slight dorso- apical invagination, fused dorsally, fused with tergite 9, not articulated with sternite 8+9; d: microtholi present on all sclerites; sternites 8+9 fused; apodeme greatly elongated beyond apex of ectoproct, bearing stout seta; 2: ectoproct not completely fused with tergite 9; apex of sternite 7 emarginate. GENITALIA 6 (Fig. 536). Tignum, gonapsis and median plate absent; gonarcus heavily sclerot- ized, arcuate, flattened, with lateral horns; entoprocessus large, L-shaped; arcessus rounded, bilobed; pseudopenis, gonosetae, gonocristae and spinellae absent. GENITALIA @ (Figs 537, 538). Praegenitale absent; subgenitale bilobed apically, with or with- out basal extension; spermatheca long, narrow, heavily sclerotized; ventral impression broad, deep; duct short, narrow, sinuous; vela short. Larva. Abdomen very slender, fusiform, not humped; tubercles not at all developed; setae short; no debris carried. REMARKS. Species of Hypochrysa may be dis- tinguished from other nothochrysine genera by 2A and 3A fused apically in the fore wing; in males there is a long ventral extension of the apodeme. Adams (1967) stated that in males of H. elegans sternites 8 and 9 are not fused but moveable. However, in all the male specimens examined during this study sternites 8+9 were fused. BIoLoGy. The larva of Hypochrysa elegans (Burmeister) has been described (Brauer, 1867; Principi, 1956; Gepp, 1983). The guts of adults examined during this study contained pollen grains. Genus KIMOCHRYSA Tijeder Kimochrysa Tjeder, 1966: 254. Type species: Kimochrysa impar Tjeder, by original designation. DISTRIBUTION. Three species are known from southern Africa. DiaAGnosis. Adult. Small lacewings, fore wing (Fig. 539) 6-9 mm. Head unmarked or marked with dark brown on gena, labrum, vertex, antennae, scape, pedicel; palps rounded apically; galea narrow with large apical papilla; mandibles broad, asymmetrical, with basal tooth on left mandible; labrum indented; vertex domed, some- times pitted; toruli very large; eyes small (head width : eye width = 4.1-4.3 : 1); scape squared, bearing numerous short, dorsal setae; pedicel hardly constricted medially; antenna shorter than fore wing; flagellar segments short, 2-3 times as long as broad; setae arranged in six rings. Pteronotum entirely dark brown; dorsal setae on pronotum long, pale. Legs unmarked or marked with dark brown dorsal stripe on femora and tibiae; setae short, black; claws without basal dilation. Fore wing unmarked; rounded apically; costal area narrow at base; costal setae quite short, inclined; stigma suffused pale brown, broad; Sc very short, fused for short distance with C basad of stigma; Sc and R widely separated; basal Sc crossvein 0.44-1.08 mm, sometimes multiple Sc crossveins present; m, very short; mp» very long; im broad, triangular or quadrangular; M forks at or just distal to m3; tympanal organ absent; Rs straight; gradates in two parallel series; c, 2-3 times shorter than c); cy narrow, squared apically; 2A unforked; dcc broad, open at margin. Hind wing with C and Sc fused basad of ptero- stigma. Abdomen (Figs 540, 541) marked brown dorsally; setae short, sparse; trichobothria 23-35; ectoprocts with slight dorso-apical invagination, fused dorsally, not fused with tergite 9; d: callus cerci rounded; microtholi absent; sternites 8+9 short, broad, incompletely fused, dorsal suture present; apodemes absent; ¢: callus cerci ovate; sternite 7 straight apically. GENITALIA 6 (Fig. 542). Tignum, gonapsis and median plate absent; entoprocessus short, broad with median lobe; parameres absent; gonarcus narrow, long, arcuate; arcessus short, broadly triangular with few, very short dorsal setae; gonosaccus short; gonosetae, gonocristae and spinellae absent. GENITALIA 2 (Figs 543, 544). Praegenitale absent; subgenitale bilobed apically, slightly extended and tapering basally; spermatheca long, quite broad; ventral impression deep, tapered abruptly apically; vela very short; duct long, coiled. REMARKS. Kimochrysa impar Tjeder is the only species amongst the Nothochrysinae to have multiple subcostal crossveins in the fore wing. Kimochrysa and Pamochrysa are probably closely related since in both genera the subcosta and costa are fused before the stigma in the hind wing. The two genera can be separated by the lack of the spiracle on the 8th tergite in Kimochrysa and by the shape of cell im. BIoLoGy. Unknown. The gut contents of speci- mens examined in this study contained only pollen grains. 258 S. J. BROOKS & P. C. BARNARD Figs 539-544 Kimochrysa. 539, 541, 543, 544, K. africana; 540, 542, K. impar. 539, fore wing; 540, apex of 3 abdomen, lateral; 541, apex of 2 abdomen, lateral; 542, d genitalia, dorsal; 543, 2 spermatheca, lateral; 544, 2 subgenitale, caudal. Genus NOTHOCHRYSA McLachlan Nothochrysa McLachlan, 1868: 195. Type species: Chrysopa fulviceps Stephens, by subsequent designation by Banks, 1903. Nathanica Navas, 1913c: 180. Type species: Hemerobius capitatus Fabricius, by original designation. [Synonymized by Tjeder, 1941: 30.] DISTRIBUTION. Two species are known from the western Palaearctic and one from western U.S.A. DIAGNOsIs. Adult. Medium to large lacewings, fore wing (Fig. 545) 12-23 mm; ground colour brown. Head unmarked or with broad black stripes on vertex and frons; palps rounded, truncate apically (Fig. 547); labrum indented; mandibles broad, asymmetrical with basal tooth on left mandible (Fig. 548); galea broad with large apical papilla (Fig. 546); vertex domed; toruli large; head width : eye width = 2.5-3.2 : 1, head broad; pedicel barely constricted medially; antenna shorter than fore wing; flagellar segments 3 times as long as broad; setae arranged in 5-6 rings. Pronotum marked laterally or entirely dark brown; dorsal setae short, dark or pale; meso- and metanotum marked dark brown. Legs unmarked; setae short, dark; claws with or without basal dilation. Fore wing unmarked; narrow (length : breadth = 2.9-3.2 : 1); costal area narrow at base; costal setae short, inclined; stigma suffused red/ brown; Sc long; Sc and R widely separated; basal Sc crossvein 0.76—-1.80 mm; im rectangular, narrow; Rs sinuate; gradates in two parallel series; crossveins not crassate in 6; c,; shorter than c,; 1A forked. Abdomen (Figs 549, 550) marked entirely brown; setae long, sparse on sternites, shorter on tergites; trichobothria 36-39; ectoprocts slightly invaginated dorso-apically, narrow dorsal suture present; d: microtholi THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 259 547 Figs 545-553 Nothochrysa. 545, N. fulviceps; 546-553, N. capitata. 545, fore wing (from Kimmins); 546, galea, dorsal; 547, apical segment of maxillary palp, dorsal; 548, mandibles, dorsal; 549, apex of d abdomen, lateral; 550, apex of 2 abdomen, lateral; 551, d genitalia, ventral; 552, 2 spermatheca, lateral; 553, 2 subgenitale, ventral. present on all sclerites except sternite 9 and ectoproct; callus cerci ovate; ectoprocts fused with tergite 9; sternites 8+9 fused; sternites broad; atria small; apodemes strongly developed on sternites 14; @: callus cerci rounded; ecto- procts not completely fused with tergite 9; sternite 7 convex apically. GENITALIA ¢ (Fig. 551). Tignum, gonapsis and median plate absent; entoprocessus minute, triangular; gonarcus arcuate; arcessus short, triangular; pseudopenis absent; gonosaccus very short; gonosetae short, in small lateral group; gonocristae, spinellae absent; hypandrium large. GENITALIA @ (Figs 552, 553). Praegenitale absent; subgenitale bilobed apically with broad basal extension; spermatheca long, broad; ventral impression broad, deep; vela absent; duct long, coiled. Larva. Abdomen broadly fusiform, humped; thoracic tubercles weakly developed, bearing two short setae; metanotum with row of setae on chalazae; abdominal tubercles short, spherical with a few short setae; abdominal setae hooked; abdominal latero-dorsal tubercles absent; single latero-dorsal setae with chalazae on abdominal tergites 6 and 7; debris often carried, usually composed of large particles. REMARKS. Nothochrysa and Dictyochrysa are the only nothochrysine genera in which 1A is forked 260 S. J. BROOKS & P. C. BARNARD Figs 554-560 Pamochrysa stellata. 554, fore wing; 555, apex of d abdomen, lateral; 556, apex of 2 abdomen, lateral; 557, 3 arcessus, lateral; 558, d genitalia, dorsal; 559, 2 spermatheca, lateral; 560, 2 subgenitale, ventral. in the fore wing. Dictyochrysa can be distin- guished from Nothochrysa by the dense reticula- tion of the wing venation. Nothochrysa is unusual amongst the Nothochrysinae in having reduced entoprocessus. Adams (1967) showed a suture between sternites 8+9 in N. californica and Psm not meeting the inner gradates but extending slightly beyond them, although the vein stops short of the outer gradates. BIoLoGy. The larvae of all three species of Nothochrysa have been described: N. californica Banks (Toschi, 1965), N. capitata Fabricius (Killington, 1937; Kimmins, 1939; Gepp, 1983) and N. fulviceps Stephens (Killington, 1937; Gepp, 1983). Adults examined during this study did not have insect remains in their gut contents. Genus PAMOCHRYSA Tjeder Pamochrysa Tjeder, 1966: 248. Type species: Pamochrysa stellata Tjeder, by monotypy. DISTRIBUTION. The single included species is known from southern Africa. DIAGNosIs. Adult. Small lacewings, fore wing (Fig. 554) 9-11 mm; ground colour pale green. Head, scape, pedicel marked extensively with black spots and stripes; palps narrow, rounded apically; mandibles broad, asymmetrical with basal tooth on left mandible; galea long, narrow with prominent apical papilla; labrum indented; vertex domed; toruli large; eyes small (head width : eye width = 3.7-4.4 : 1); scape squared; pedicel slightly constricted medially; antenna shorter than fore wing; flagellar segments 3 times as long as broad; setae arranged in five rings. Pronotum marked with black lateral stripes; dorsal setae short, black; meso- and metanotum with black markings. Legs marked with black stripes on femora and tibiae; setae short, dark; claws without basal dilation. Fore wing rounded apically; marked with pale brown shading on gradates; costal area narrow at base; costal setae quite long, inclined; stigma marked with black spots; Sc long; R sinuate below stigma; Sc and R widely separated; basal Sc crossvein 1.4—1.6 mm; im broad, diamond-shaped; M forking after base of m3; Rs straight; gradates in two parallel series; veins not crassate in 6; c, shorter than c7; 1A not forked; dcc broad, open at margin. Hind wing with C and Sc fused just basad of pterostigma. Abdomen (Figs 555, 556) marked along entire length with black dorsolateral stripe; setae short, coarse on apical segments, quite dense; callus cerci ovate; trichobothria 41-44; ectoprocts with slight dorso-apical invagination, fused dorsally, THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 261 Figs 561-566 Pimachrysa. 561, P. nigra; 562-566, P. fusca. 561, fore wing; 562, apex of d abdomen, lateral; 563, apex of 2 abdomen, lateral; 564, ¢ genitalia, dorsal; 565, 2 spermatheca, lateral; 566, 2 subgenitale, ventral. not fused with sternite 9; 6: microtholi absent; sternite 8+-9 fused; apodemes absent; @: spiracle on segment 8 opens on sternite; sternite 7 straight apically. GENTALIA 6 (Figs 557, 558). Tignum, gonapsis and median plate absent; entoprocessus large, linear; parameres absent; gonarcus arcuate, narrow with short, medio-lateral horns; arcessus short, narrow, tapering apically, curved medially at 90°; gonosaccus short; gonosetae few, short, evenly distributed; gonocristae and spinellae absent. GENITALIA @ (Figs 559, 560). Praegenitale absent; subgenitale bilobed apically, tapered slightly basally; spermatheca large, broad; ventral impression very broad, deep; vela very short; duct short, curved. REMARKS. Pamochrysa may be distinguished from other nothochrysine genera by the rhomboidal intramedian cell since im is quadrangular in other Nothochrysinae. The genus is probably closely related to Kimochrysa with which it shares the apomorphy of having Sc and C fused in the hind wing. Pamochrysa and Pimachrysa are the only genera in the Chrysopidae in which a spiracle (on tergite 8 in females) opens on a tergite, in all other genera the spiracles open on the lateral 262 S. J. BROOKS & P. C. BARNARD Figs 567-572 Triplochrysa pallida. 567, fore wing (from Kimmins); 568, apex of 2 abdomen, lateral; 569, apex of ? abdomen, lateral; 570, d genitalia, dorsal; 571, 2 spermatheca, lateral; 572, 2 subgenitale, ventral. membrane. This is a plesiomorphic condition which occurs widely throughout the Neuroptera (e.g. Hemerobiidae and Myrmeleonidae). Pamochrysa, Kimochrysa and Pimachrysa plesiomorphically retain a suture completely separating tergite 9 and the ectoprocts in males. BioLoGy. Unknown. Pollen grains were present in the guts of adults examined during this study and Tjeder (1966) identified Dipsacacae and Compositae pollen grains in the gut contents of specimens that he examined. Genus PIMACHRYSA Adams Pimachrysa Adams, 1956: 67. Type species: Pimachrysa grata Adams, by original designation. DISTRIBUTION. Five species are known from western U.S.A. and northern Mexico. DIAGNOsIs. Adult. Small lacewings, fore wing (Fig. 561) 7-10 mm; ground colour brown. Head marked with brown stripes on labrum, clypeus, frons, vertex, scape; palps broad, rounded apic- ally; labrum with straight apical margin; vertex domed; toruli large; head width : eye width = 2.64.1 : 1, eyes very small; antenna, pale brown, shorter or same length as fore wing; flagellar segments 3 times as long as broad; setae arranged in 6 rings. Pronotum marked with brown lateral stripe; dorsal setae very short, few; meso- and metanotum with brown lateral stripe. Legs with short, black setae; claws without basal dilation. Fore wing narrow (length : breadth = 3.0-3.3: 1); unmarked; costal area narrow at base; costal setae short, inclined; stigma marked with brown spot on each crossvein; Sc long or short; Sc and R widely separated; basal Sc crossvein 1.16—1.52 mm; im long, quadrangular; Rs straight; mm» very long; radial crossveins straight; gradates in two parallel series; crossveins not crassate in 6; c) up to 1.5 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 263 577 Figs 573-578 Chrysaloysia somalica d . 573, fore wing; 574, apex of abdomen, dorsal view; 575, sternite 8 + 9, ventral view (stippling indicates apodemes); 576, ectoproct + tergite 9, lateral view; 577, gonarcus and arcessus, caudal view; 578, arcessus, lateral view. times longer than c,; 1A unforked; dcc open at wing margin, very broad. Abdomen (Figs 562, 563) extensively marked brown; setae very short and sparse; trichobothria 20-25; ectoprocts not fused with tergite 9, fused dorsally, deeply invagi- nated apico-dorsally; d: callus cerci ovate; ster- nites 8 and 9 not completely fused; microtholi present on all sclerites except sternite 8+9 and ectoproct; ?: sternite 7 straight apically; gona- pophyses grossly enlarged, bearing large spoon- shaped setae; spiracle opens on 8th tergite. GENITALIA 6 (Fig. 564). Tignum, gonapsis and median plate absent; entoprocessus narrow, elongate; parameres absent; gonarcus arcuate with shallow curve; arcessus short, narrowing abruptly to apical hook, curved ventrally at base; gonosaccus short; gonosetae absent or few, short, evenly dispersed; gonocristae and spinellae absent. GENITALIA @ (Figs 565, 566). Praegenitale absent; subgenitale with narrow basal extension, bilobed apically; spermatheca broad; ventral impression deep, broad basally, tapering abruptly apically; duct long, coiled; vela absent. REMARKS. Pimachrysa has retained the primitive character of the spiracle opening on the 8th tergite (like Pamochrysa), but it can be distinguished by 264 the narrow wings with the pterostigmatic sub- costal crossveins marked with black. The female is readily distinguished by the spoon-shaped setae on the gonapophyses. BIoLoGy. Larva unknown. The guts of adult specimens of P. nigra Adams contained pollen grains, but insect remains were not present. Dr N. D. Penny (in litt.) states that the adults are on the wing during the winter. Genus TRIPLOCHRYSA Kimmins Triplochrysa Kimmins, 1952a: 69. Type species: Triplochrysa pallida Kimmins, by monotypy and original designation. DISTRIBUTION. Queensland (Australia). Two species are known in the genus. DIAGNOsIS. Adult. Medium-sized lacewings, fore wing (Fig. 567) 12-14 mm; ground colour pale yellowish green. Head unmarked or marked with broad black stripe between antennae; palps narrow, rounded apically; labrum deeply invagi- nated; mandibles broad, left mandible with basal tooth; galea long, narrow with long apical papilla; vertex small, hardly raised; toruli large; eyes small, (head width : eye width = 2.32.9 : 1); scape as broad as long; antenna shorter than fore wing; flagellar segments short, twice as long as broad; setae arranged in 5 rings. Pronotum short, broad; unmarked or marked with small black spot in each corner; dorsal setae long, pale; meso- and metanotum unmarked or with lateral black spot. Legs unmarked; setae short, dark; claws with small median tooth, hardly dilated. Fore wing unmarked; rounded apically; costal area very narrow at base; costal setae short, inclined; stigma unmarked; narrow strip of dense microsetae present on pterostigma between Sc and R; Sc long; Sc and R widely separated; basal Sc cross- vein 1.08—1.12 mm; mp very long; M forked distal to m3; im broadly quadrangular or open apically (apical crossvein absent); Rs straight; gradates in three parallel series; inner gradates extended basally, not fusing with Psm; c; about half as long as co; 1A not forked; 2A and 3A not fused apically; dcc broad, open at margin. Abdomen (Figs 568, 569) unmarked; setae short, sparse; callus cerci ovate; trichobothria 31—35; ectoprocts with slight dorso-apical invagination, fused dor- sally; 3: microtholi absent; tergite 9 and ecto- proct fused; sternite 8+9 fused, broad, short; 9: suture present between ectoprocts and tergite 9; sternite 7 convex apically. GENITALIA 6 (Fig. 570). Tignum, gonapsis and S. J. BROOKS & P. C. BARNARD median plate absent; entoprocessus short, broad; parameres absent; gonarcus short, broad, trans- verse; arcessus long, narrow, rounded apically, strongly curved ventrally basad; gonosaccus short; gonosetae, gonocristae and spinellae absent. GENITALIA @ (Figs 571, 572). Praegenitale absent; subgenitale bilobed apically with tapering basal extension; spermatheca long, narrow or broad; ventral impression deep; vela short; duct very long, narrow, sinuous. REMARKS. Triplochrysa may be readily distin- guished from other nothochrysine genera by the three parallel series of gradate crossveins. The two Australian genera Triplochrysa and Dictyochrysa are the only nothochrysines with more than two gradate series, Dictyochrysa being distinguished by its reticulate venation. BioLocy. Unknown. Adult gut contents do not include insect remains. CHECKLIST OF EXTANT SPECIES OF CHRYSOPIDAE (* indicates material examined) Family CHRYSOPIDAE Schneider Subfamily APOCHRYSINAE Handlirsch, 1908 Genus ANAPOCHRYSA Kimmins, 1952 * africana Kimmins, 1952 voeltzkowi (Weele, 1909) Genus APOCHRYSA Schneider, 1851 *Jeptalea (Rambur, 1842) picteti (Navas, 1911) Genus CLA VERINA Navas, 1913 *beata (Walker, 1858) Genus DOMENECRHUS Navas, 1913 *marianella (Guérin, 1853) *mirifica (Gerstaecker, 1888) *isigillatus Navas, 1913 Genus JOGUINA Navas, 1912 *borneensis Kimmins, 1952 *malayana Banks, 1931 *nicobarica (Brauer, 1864) Genus LAINIUS Navas, 1913 *constellatus Navas, 1913 decoratus Navas, 1930 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) Genus LOYOLA Navas, 1913 *croesus (Gerstaecker, 1893) tripunctata Banks, 1924 Genus NACAURA Navas, 1913 *matsumurae (Okamoto, 1912) minomoana (Nakahara, 1915) Genus NOBILINUS Navas, 1913 *albardae albardae (McLachlan, 1875) *albardae insignitus Navas, 1913 *albardae phantoma (Gerstaecker, 1893) *aurifera (Walker, 1853) bellula (Banks, 1914) coccinea (Brauer, 1864) Genus NOTHANCYLA Navas, 1910 *verreauxi Navas, 1910 Genus OLIGOCHRYSA Esben-Petersen, 1914 *Jutea (Walker, 1853) gracilis Esben-Petersen, 1914 Genus SYNTHOCHRYSA Needham, 1909 *cognata Kimmins, 1953 evanida (Gerstaecker, 1893) *montrouzieri (Girard, 1862) stigma (Girard, 1862) *salomonis (Kimmins, 1951) Subfamily CHRYSOPINAE Schneider, 1851 Tribe ANKYLOPTERYGINI Navas, 1910 Genus ANKYLOPTERYX Brauer, 1864 Ethiochrysa Fraser, 1952 syn. n. Subgenus ANKYLOPTERYX Brauer, 1864 *alluaudi Navas, 1910 *basalis Kimmins, 1952 braueri Banks, 1937 *buttikaferi Weele, 1905 *collarti Navas, 1925 *decorsei Navas, 1910 decormei misspelling *delicata Navas, 1935 delicatula Banks, 1937 *doleschali Brauer, 1864 *fastuosa Navas, 1929 *fraterna Banks, 1939 gracilis Nakahara, 1955 *grata Tjeder, 1966 immaculata Brauer, 1864 *lambillioni Navas, 1929 *Jaticosta Banks, 1939 *nepalensis H6lzel, 1973 *nepheloptera Navas, 1912 *nesiotica Navas, 1913 *nonelli Navas, 1913 *obliqua Banks, 1924 265 octopunctata octopunctata (Fabricius, 1793) octopunctata borneensis Weele, 1909 *octopunctata candida (Fabricius, 1798) octopunctata kisserensis Weele, 1909 *octopunctata punctata (Hagen, 1858) *octopunctata sigillaris Gerstaecker, 1893 *octopunctata trimaculata (Girard, 1859) *octopunctata (Fraser, 1951) comb. n. [Homonym] *overlaeti Navas, 1936 * pallida Banks, 1910 *pallidula Tjeder, 1966 * pellucida Tjeder, 1966 perpallida Banks, 1924 polychlora (Fraser, 1952) comb. n. *polygramma Gerstaecker, 1893 *nervosa Navas, 1913 pusilla Tjeder, 1966 quadrimaculatus (Guérin, 1844) rhodocephala Navas, 1914 rieki New, 1980 scioptera Navas, 1924 splendidissima Gerstaecker, 1884 tanana Fraser, 1952 tesselatus Needham, 1909 *tristicta Navas, 1910 *venusta (Hagen, 1853) Subgenus SENCERA Navas, 1925. stat. n. *anomala (Brauer, 1864) *exquisita Nakahara, 1955 *feae Navas, 1929 *scioneura Navas, 1924 Genus PARANK YLOPTERX Tijeder, 1966 stat. n. *burgeoni (Navas, 1929) comb. n. *elgonica (Navas, 1936) comb. n. *feana (Navas, 1929) comb. n. *maculata (Kimmins, 1939) comb. n. *multipunctata (Fraser, 1951) comb. n. *neavei (Navas, 1913) *verdcourti (Kimmins, 1951) *polysticta (Navas, 1910) comb. n. speciosa (Navas, 1924) comb. n. *tetrasticta (Navas, 1932) comb. n. *walterloti (Navas, 1911) comb. n. waterloti’ misspelling Genus RETIPENNA Brooks, 1986 *burmana Brooks, 1986 chione (Banks, 1940) comb. n. *dasyphlebia (McLachlan, 1894) comb. n. *jrregularis (Navas, 1914) syn. n. grahami (Banks, 1940) comb. n. *hasegawai (Nakahara, 1955) jubingensis (H6lzel, 1973) comb. n. *notata (Navas, 1910) *guttata (Navas, 1930) *longipilis (Banks, 1938) 266 *variegata Brooks, 1986 Genus SEMACHRYSA Brooks, 1983 Indochrysa Banks, 1938 *claggi (Banks, 1937) *contorta Brooks, 1983 *cruciata (Esben-Petersen, 1928) *dammermanni (Esben-Petersen, 1929) *decorata (Esben-Petersen, 1913) *rectoides (Banks, 1939) *hyndi Brooks, 1983 *matsumurae (Okamoto, 1914) *minuta Brooks, 1983 *nigribasis (Banks, 1920) *papuensis Brooks, 1983 *picilabris (Kimmins, 1952) *polysticta Brooks, 1983 *sagitta Brooks, 1983 *wallacei Brooks, 1983 Genus SIGNOCHRYSA gen. n. *bakeri (Banks, 1924) comb. n. boettcheri (Esben-Petersen, 1933) buruensis (Esben-Petersen, 1929) stat. n. & comb. n. *caliptera (Banks, 1920) comb. n. *catenulata (Gerstaecker, 1893) comb. n. *jocaste (Banks, 1940) comb. n. *mira (Navas, 1913) comb. n. ornatissima (Nakahara, 1955) comb. n. rizali (Banks, 1924) comb. n. *signatipennis (Banks, 1910) comb. n. Tribe BELONOPTERYGINI Navas, 1913 Genus ABACHRYSA Banks, 1938 “eureka (Banks, 1931) Genus BELONOPTER YX Gerstaecker, 1863 *arteriosa Gerstaecker, 1863 Genus CALOCHRYSA Banks, 1943 *“extranea (Essben-Petersen, 1917) Genus CHRYSACANTHIA Lacroix, 1923 Nesochrysa Fraser, 1951 syn. n. *esbeniana Lacroix, 1923 “varicella (Fraser, 1951) comb. n. Genus CHRYSALOYSIA Navas, 1927 *somalica Navas, 1927 Genus DYSOCHRYSA Tjeder, 1966 “furcata Tjeder, 1966 *reflexa Tjeder, 1966 Genus EVANOCHRYSA gen. n. *evanescens evanescens (McLachlan, 1869) comb. n. evanescens javanica (Weele, 1909) comb. n. *infecta (Newman, 1838) comb. n. S. J. BROOKS & P. C. BARNARD subcostalis (Navas, 1917) *Jevasseuri (Navas, 1921) comb. n. Genus ITALOCHRYSA Principi, 1946 *aequalis aequalis (Walker, 1853) *aequalis polychroa (Gerstaecker, 1893) *aequalis sumatrana (Albarda, 1881) *aethiopiae (Lacroix, 1925) albescens (Navas, 1932) comb. n. amplipennis Tjeder, 1966 asirensis Holzel, 1980 banksi New, 1980 *bimaculata H6lzel, 1980 *boueti (Navas, 1927) *burgeoni (Navas, 1924) carletoni (Banks, 1939) *chloromelas (Girard, 1862) *cornuta (Navas, 1935) cruciata (Navas, 1935) *cuneata (Navas, 1911) *everetti (Weele, 1909) stat. n. *exilis Tjeder, 1966 *fascialis (Banks, 1910) falcata Tjeder, 1966 *ferruginea (McLachlan, 1869) flavobrunneus Ghosh, 1981 froggatti (Esben-Petersen, 1914) *fulvicornis Kimmins, 1955 gagginoi (Navas, 1929) * gigantea (McLachlan, 1867) *finoti (Navas, 1908) * gillavryi (Navas, 1924) *guerini (Navas, 1911) *henryi (Kimmins, 1938) *impar (Navas, 1912) indigena (Needham, 1909) *insignis (Walker, 1853) stictoneura (Gerstaecker, 1885) insignita (Navas, 1913) xanthocephala (Navas, 1932) *jtalica (Rossi, 1790) lateralis Olivier, 1792) grandis (Thunberg, 1838) * japonica (McLachlan, 1875) modesta (Nakahara, 1955) * jubilaris (Navas, 1924) *Jata (Banks, 1910) lefroyi (Needham, 1909) *limbata (Navas, 1924) lobini Holzel & Ohm, 1982 luddemanni (Navas, 1935) *Judekingi (Weele, 1909) stat. n. ignobilis (Navas, 1913) lyrata Tjeder, 1966 maclachlani (Wallengren, 1875) modesta (Navas, 1935) comb. n. *mozambica (Walker, 1860) THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 267 *neobritanica (Navas, 1913) elizabethae (Navas, 1928) comb. n. *neurodes (Rambur, 1842) * grandidieri Navas, 1910 nigrinervis (Esben-Petersen, 1917) *iNotus (Tjeder, 1966) comb. n. nigrovenosus Kuwayama, 1970 *macrostigma (Gerstaecker, 1894) comb. n. nossibensis (Navas, 1928) *auricollis Navas, 1913 *oberthuri (Navas, 1908) *marginata (Navas, 1912) comb. n. okavangoensis Tjeder, 1966 *marginicollis (Kimmins, 1957) comb. n. pallicornis (Banks, 1920) rubeolus (Tjeder, 1966) comb. n. *peringueyi (Esben-Petersen, 1920) *ruficeps (Tjeder, 1966) comb. n. polemia (Navas, 1916) *seyrigi (Navas, 1934) comb. n. punctistigma (Esben-Petersen, 1918) virgatus (Tjeder, 1966) comb. n. robusta (Needham, 1909) *rufostigma (McLachlan, 1867) maculata (Esben-Petersen, 1912) Genus NODOCHRYSA Banks, 1938 *necrota (Banks, 1920) rugosa Tsukagachi, 1988 Genus OYOCHRYSA Brooks, 1985 *sectoria (Navas, 1925) *ancora Brooks, 1985 serrata Tjeder, 1966 *sanguinea Brooks, 1985 similis Tjeder, 1966 *spadix Brooks, 1985 simplex (Banks, 1920) stigmalis (Navas, 1928) es : a ee mse) elegans Esben-Petersen, 1933 > * * . 4 talaverae (Navas, 1928) kervillei Navas, 1925 *temerata (Navas, 1914) Genus TURNEROCHRYSA Kimmins, 1935 * tibialis (Navas, 1913) *mirifica Kimmins, 1935 *turneri (Kimmins, 1948) soror Tjeder, 1966 Genus STIGMACHRYSA Navas, 1925 *cladostigma (Navas, 1913) uchidae (Kuwayama, 1927) Tribe CHRYSOPINI Schneider, 1851 vansoni Tjeder, 1966 *variegata (Burmeister, 1839) Genus ANOMALOCHRYSA McLachlan, 1883 sordidata (Navas, 1908) *angulicosta Perkins, 1899 *zonata (Navas, 1913) *cognata Perkins, 1899 vartianorum Holzel, 1967 *debilis Perkins, 1899 zulu Tjeder, 1966 nana Perkins, 1899 *frater Perkins, 1899 *fulvescens fulvescens Perkins, 1899 *fulvescens rhododora Perkins, 1899 *haematura Perkins, 1899 *hepatica McLachlan, 1883 proteus Perkins, 1899 *Jongipennis Perkins, 1899 *maclachlani maclachlani Blackburn, 1884 paurosticta Perkins, 1899 deceptor Perkins, 1899 gayi Perkins, 1899 zoe Perkins, 1899 *maclachlani simillima Perkins, 1899 *molokaiensis Perkins, 1899 *montana Blackburn, 1884 ornatipennis Blackburn, 1884 *peles Perkins, 1899 *princeps Perkins, 1899 *raphidioides raphidioides Perkins, 1899 *raphidioides reticulata Perkins, 1899 Genus NESOCHRYSA Navas, 1910 *rufescens McLachlan, 1883 Oviedus Navas, 1913 syn. n. biseriata Perkins, 1899 Madachrysa Navas, 1934 syn. n. *soror Perkins, 1899 Genus NACARINA Navas, 1915 Nadiva Navas, 1919 Goliva Navas, 1920 Rameta Navas, 1920 Mesochrysa Navas, 1927 syn. n. balboana (Banks, 1941) deletangi (Navas, 1920) egena (Navas, 1930) furcata Navas, 1916 megaptera (Navas, 1927) comb. n. *neotropica (Navas, 1913) comb. n. *panchlora (Gerstaecker, 1888) pletorica (Navas, 1919) *sanctiignatii (Navas, 1927) comb. n. sanguinea (Navas, 1920) *titan (Banks, 1915) comb. n. *valida (Erichson, 1848) viridipennis (Alayo, 1968) *wagneri (Navas, 1924) 268 *sylvicola Perkins, 1899 *viridis Perkins, 1899 Genus APERTOCHRYSA Tjeder, 1966 afghanica (Holzel, 1973) anomala (Tillyard, 1917) comb. n. araucariae (Tillyard, 1917) comb. n. crassinervis (Esben-Petersen, 1927) comb. n. *edwardsi (Banks, 1940) comb. n. *eremita (Kimmins, 1955) *ictericus (Esben-Petersen, 1927) comb. n. kichijoi (Kuwayama, 1936) leai (Tillyard, 1917) comb. n. *madegassa (Navas, 1921) comb. n. *metastigma (Tillyard, 1917) comb. n. nautarum (Tillyard, 1917) comb. n. *norfolkensis (Tillyard, 1917) comb. n. * physophlebia (Navas, 1914) comb. n. *umbrosa (Navas, 1914) waitei (Tillyard, 1917) comb. n. Genus ATLANTOCHRYSA Hoizel, 1970 *atlantica (McLachlan, 1882) * pseudoatlantica (Tjeder, 1939) sororcula (Tjeder, 1939) Genus AUSTROCHRYSA Esben-Petersen, 1928 Scoliochrysa Navas, 1929 syn. n. *abnormis abnormis (Albarda, 1881) comb. n. lunigera (Gerstaecker, 1893) *abnormis javanensis (Weele, 1909) comb. n. apoana (Banks, 1937) comb. n. *hexasticha (Gerstaecker, 1893) comb. n. leucoptera (Esben-Petersen, 1926) comb. n. *Joriana (Navas, 1929) comb. n. samoana Esben-Petersen, 1928 Genus BORNIOCHRYSA nom. n. for Bornia Navas, 1928 stat. n. *appendiculata (Esben-Petersen, 1926) comb. n. luzonica (Banks, 1939) comb. n. *solomonis Banks, 1941 *squamosa (Tjeder, 1966) comb. n. *winkleri Navas, 1928 Genus BRINCKOCHRYSA Tjeder, 1966 Neda Navas, 1933 (Homonym) syn. n. *alfierii (Navas, 1926) comb. n. *amseli (H6lzel, 1967) comb. n. *cardaleae (New, 1980) comb. n. *chlorosoma (Navas, 1915) comb. n. *decaryella (Navas, 1933) comb. n. *goniophora (Navas, .1935) comb. n. kintoki (Okamoto, 1919) comb. n. *Jauta (Esben-Petersen, 1927) comb. n. *michaelseni (Esben-Petersen, 1928) nachoi (Monserrat, 1977) naumanni (Holzel, 1982) plagata (Navas, 1929) comb. n. S. J. BROOKS & P. C. BARNARD pulchella Ho6lzel, 1987 scelestes (Banks, 1911) comb. n. *stenoptera (Navas, 1910) *mozambica (Navas, 1931) syn. n. sigwalti (Séméria, 1984) syn. n. *turkanensis (Navas, 1936) peri (Tjeder, 1966) syn. n. Genus CERAEOCHRYSA Adams,1982 acutipuppis Adams & Penny, 1987 *adornata (Lacroix, 1926) *anceps (Navas, 1925) ariasi Adams & Penny, 1987 *arioles (Banks, 1944) binaria (Navas, 1928) *azoguesina (Navas, 1928) comb. n. *berlandi (Navas, 1924) caligata (Banks, 1945) castilloi (Navas, 1913) caucana (Banks, 1910) comb. n. *cincta (Schneider, 1851) *bilineata (Navas, 1913) lafonei (Navas, 1914) incalis (Banks, 1915) bicarnea (Banks, 1920) *advena (Navas, 1922) habana (Navas, 1922) mestiza (Navas, 1924) villosula (Navas, 1924) bessona (Navas, 1924) bina (Navas, 1924) cornuta (Navas, 1925) alternans (Navas, 1933) wollebaeki (Esben-Petersen, 1934) sallei (Banks, 1940) iona (Banks, 1944) *claveri (Navas, 1911) *silvana (Navas, 1913) deficiens (Navas, 1930) sibirica (Navas, 1930) haitiensis (Smith, 1931) adoina (Banks, 1945) inexpectata (Alayo, 1968) *cubana (Hagen, 1861) tolteca (Banks, 1901) albatala (Banks, 1913) venularis (Navas, 1913) *imbecilla (Navas, 1914) epheba (Navas, 1924) seminole (Banks, 1924) freemani (Smith, 1931) Jamaicensis (Banks, 1941) * discolor (Navas, 1914) *effusa (Navas, 1911) everes (Banks, 1920) *furcata (Navas, 1922) *furculata (Navas, 1923) THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 269 gundlachi (Navas, 1924) dimidiata (Navas, 1928) *instabilis (Navas, 1925) iniqua (Navas, 1929) jacobaea (Navas, 1925) impar (Navas, 1929) syn. n. peterseni (Navas, 1929) *undulata (Fraser, 1952) petersenia (Navas, 1931) *atrostrata (Tjeder, 1966) gloriae (Alayo, 1968) *ceratina (Navas, 1910) fairchildi (Banks, 1945) *disparilis (Navas, 1934) falcifera Adams & Penny, 1987 *fiebrigi (Navas, 1913) Genus CHRYSEMOSA nom. n. for Mesochrysa * gradata (Navas, 1913) PENS 1936 — * guatemalteca (Navas, 1914) aoe (Navas, 1915) comb. n. *indicata (Navas, 1913) comb. n. Noe (Tjeder, 1966) comb. n. infausta (Banks, 1945) jeanneli (Navas, 1915) comb. n. josephina (Navas, 1926) *ellenbergeri (Navas, 1921) *angulata (Navas, 1929) laristanus (Holzel, 1982) comb. n. *Jaufferi (Navas, 1922) comb. n. parva (Tjeder, 1966) comb. n. *Jineaticornis (Fitch, 1855) a (Holzel & Ohm, 1982) comb. n. puncticornis (Fitch, 1855) simillima (Tjeder, 1966) comb. n. parvula (Banks, 1903) sodomensis (Holzel, 1982) comb. n. columbiana (Banks, 1903) ae Navas, 1936 stichoptera (Navas, 1914) umbralis (Navas, 1933) comb. n. michaelmuris Adams & Penny, 1987 Genus CHRYSOCERCA Weele, 1909 montoyana (Navas, 1913) nigripes Adams & Penny, 1987 *placita (Banks, 1908) intacta (Navas, 1912) *forreri (Navas, 1913) rafaeli Adams & Penny, 1987 reddyi Adams & Penny, 1987 Pseudochrysa Okamoto, 1914 formosana (Okamoto, 1914) jacobsoni Weele, 1909 *nigrivultuosa (Kimmins, 1955) *perturbata (Banks, 1931) comb. n. timorina Handschin, 1935 *reducta (Banks, 1944) Genus CHRYSOPA Leach, 1815 *rochina (Navas, 1915) Aeolops Billberg, 1820 sanchezi (Navas, 1924) Melanops Doumerc, 1861 scapularis (Navas, 1914) Chrysopisca McLachlan, 1875 syn. n. *silvanoi (Navas, 1916) comb. n. Cintameva Navas, 1914 *smithi (Navas, 1914) Minva Navas, 1920 syn. n. poeyi (Navas, 1924) Polyphlebia Navas, 1936 syn. n. neotropica (Navas, 1929) Metachrysopa Steinmann, 1964 squalidens Adams & Penny, 1987 Nigrochrysopa Steinmann, 1964 tenuicornis Adams & Penny, 1987 Parachrysopa Séméria, 1983 syn. n. * valida (Banks, 1895) *abbreviata abbreviata Curtis, 1834 bimaculata (McClendon, 1901) immaculata Stephens, 1836 *limitata (Navas, 1913) chlorophanus Ratzeburg, 1844 *longicella (Navas, 1913) decora Evans, 1847 *breviata (Banks, 1915) germanica (Esben-Petersen, 1913) *lioni (Navas, 1927) abbreviata caerulescens Bianchi, 1931 damiensis (Smith, 1931) abbreviata maclachlaniola Bianchi, 1931 wolcotti (Smith, 1931) *albicornis Fitch, 1855 *altaica Holzel, 1967 Genus CERATOCHRYSA Tjeder, 1966 altaiensis H6lzel, 1980 Musola Navas, 1929 syn. n. replacement name for quadripunctata *antica (Walker, 1853) Steinmann, 1968 *nesaea (Navas, 1911) *astarte Holzel, 1967 inaequalis (Navas, 1912) *chazaudi Navas, 1922 *ducissa (Navas, 1914) *chi Fitch, 1855 *regina (Navas, 1914) upsilon Fitch, 1855 *yuilleti (Navas, 1914) *chlorophana Burmeister, 1839 pooana (Navas, 1922) latipennis Schnieder, 1851 270 chlorophana Walker, 1853 xanthocephala Fitch, 1855 bipunctata Fitch,1855 transmarina Hagen, 1861 *coloradensis Banks, 1895 *commata Kis & Ujhelyi, 1965 curdica H6lzel, 1967 *dasyptera McLachlan, 1872 minima Keljander, 1881 lubischewi (Navas, 1933) depressa Steinmann, 1968 schamona HoOlzel, 1971 “dorsalis Burmeister, 1839 pini Brauer, 1850 bifidilinea Costa, 1884 ypsilon Costa, 1884 *dubitans McLachlan, 1887 venulosa (Navas, 1914) excepta Banks, 1911 fezzanina (Navas, 1932) comb. n. *flaviceps (Brullé, 1840) *formosa Brauer, 1850 burmeisteri Schneider, 1851 jJapana Okamoto, 1919 pyreaea (Navas, 1930) bicristata Tjeder, 1936 fuscostigma Esben-Petersen, 1933 hummeli Tjeder, 1936 *“hungarica Klapalek, 1899 *intima McLachlan, 1893 fracta Navas, 1910 silens Steinmann, 1971 kansuensis Tjeder, 1936 *lezeyi Navas, 1910 minuta (McLachlan, 1975) syn. n. *nierembergi Navas, 1908 nigra Okamoto, 1919 nigrescens H6lzel & Ohm, 1986 *nigricornis Burmeister, 1839 colon Fitch, 1855 erythrocephala Banks, 1898 majuscula Banks, 1906 crotchi Banks, 1938 *nigricostata Brauer, 1850 heydenii Schneider, 1851 nigrovenosa Pongracz, 1912 *fastigiata Navas, 1914 cosmia Navas, 1918 laburdensis Lacroix, 1924 *mediata (Navas, 1924) *neuralis (Navas, 1926) ingens Steinmann, 1964 *oculata Say, 1849 euryptera Burmeister, 1839 illepida Fitch, 1855 omikron Fitch, 1855 mississipiensis Fitch, 1855 S.J. BROOKS & P. C. BARNARD assimilis Banks, 1898 rubicunda Navas, 1913 *pallens (Rambur, 1838) septempunctata Wesmael, 1841 mauriciana (Rambur, 1842) nobilis Brauer, 1850 *cognata McLachlan, 1867 *centralis McLachlan, 1875 robusta (Gerstaecker, 1893) ricciana Navas, 1910 puncticollis Navas, 1915 parvula (Doumerc, 1861) *perla L., 1758 chrysops (L., 1758) viridis (Retzius, 1783) cancellatus (Schrank, 1802) reticulata Leach, 1815 maculata Stephens, 1836 fallax Navas, 1913 nigriceps Okamoto, 1914 nothochrysodes (Navas, 1935) *perplexa McLachlan, 1887 kreyembergi (Navas, 1933) persica Holzel, 1966 *phyllochroma Wesmael, 1841 tenella Brauer, 1850 pusilla Brauer, 1850 labbei Navas, 1910 magnicauda Tjeder, 1936 electra Holzel, 1965 *pleuralis Banks, 1911 punctata (Navas, 1920) comb. n. punctata (Navas, 1936) comb. n. *quadripunctata Burmeister, 1839 sulphurea Fitch, 1855 sichelii Fitch, 1855 *quettana (Navas, 1930) *regalis Navas, 1915 dorsalis Navas, 1904 nec Burmeister sapporensis Okamoto, 1914 *separata Banks, 1911 *slossonae Banks, 1924 *sogdianica McLachlan, 1875 *minuta (McLachlan, 1875) syn. n. nadali Navas, 1913 euprepia Navas, 1916 indiga Navas, 1916 *harterti Navas, 1929 asiatica Steinmann, 1971 thibetana McLachlan, 1887 *viridana Schneider, 1845 geniculata Pictet, 1865 peterseni Navas, 1910 galaica Navas, 1927 *clypealis Navas, 1931 *collina Navas, 1934 zelenyi Steinmann, 1964 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 271 *walkeri McLachlan, 1893 italotis (Banks, 1910) novempunctata Navas, 1912 krakatauensis Tsukagachi, 1988 xanthocephala Navas, 1916 *Jindana (Navas, 1924) comb. n. * maquilingi (Banks, 1937) comb. n. Genus CHRYSOPERLA Steinmann, 1964 replacement name for inconspicua (Navas, *acutella (Navas, 1933) comb. n. 1926) anpingensis (Esben-Petersen, 1913) comb. n. * mediterranea (H6lzel, 1972) *arequipae (Navas, 1929) comb. n. *meloui (Navas, 1924) comb. n. *asoralis (Banks, 1915) comb. n. *mutata (McLachlan, 1898) “australis (New, 1980) comb. n. expurgata (Tjeder, 1949) *carnea carnea (Stephens, 1836) *nepia (Navas, 1911) comb. n. affinis (Stephens, 1836) *nigriciana (Navas, 1931) comb. n. microcephala (Brauer, 1850) *nyerina (Navas, 1933) comb. n. vulgaris (Schneider, 1851) oblita (Holzel, 1973) lampropter (Stein, 1863) *oscillans (Navas, 1922) comb. n. lucasina (Lacroix, 1912) *otalatis (Banks, 1910) comb. n. pillichi (Pongracz, 1913) *lemoulti (Lacroix, 1923) nipponensis (Okamoto, 1914) * phaeocephala (Navas, 1931) comb. n. kurisakiana (Okamoto, 1914) plicata (Tjeder, 1966) *kolthoffi (Navas, 1927) plorabunda (Fitch,1855) angelina (Navas, 1931) pseudographa (Fitch, 1855) *quettana (Navas, 1931) robertsonii (Fitch, 1855) downesi (Smith, 1932) illinoiensis (Shimer, 1865) ferganica (Navas, 1933) *pudica (Navas,1913) renoni (Lacroix, 1933) punensis (Ghosh, 1976) pictavica (Lacroix, 1933) *rufilabris (Burmeister, 1839) sinica (Tjeder, 1936) interrupta (Schneider, 1851) mohave (Banks, 1938) *attenuata (Walker, 1853) lundbladi (Tjeder, 1939) *repleta (Walker, 1853) maderensis (Tjeder, 1939) novaeboracensis (Fitch, 1855) canariensis (Tjeder, 1939) tabida (Fitch,1855) carnea nanceiensis Séméria, 1980 medialis (Banks, 1903) *clypealis (Navas, 1929) comb. n. sanandensis (Ghosh, 1976) comans (Tjeder, 1966) *satilota (Banks, 1910) *comanche (Banks, 1938) comb. n. savioi (Navas, 1933) comb. n. sperryae (Banks, 1943) shansiensis (Kuwayama, 1962) comb. n. *concinna (Holzel, 1974) *socia (Navas, 1936) comb. n. *congrua (Walker, 1853) suzukii (Okamoto, 1919) *concolor (Walker, 1853) triactinata (New, 1980) comb. n. *bequaerti (Navas, 1912) *zastrowi (Esben-Petersen, 1928) *decaryana (Navas, 1934) comb. n. dozieri (Smith, 1931) comb. n. Genus CHRYSOPIDIA Navas, 1910 *exotera (Navas, 1913) comb. n. Subgenus ANACHRYSA Holzel, 1973 *externa externa (Hagen, 1861) *elegans Holzel, 1973 *lanata (Banks, 1910) erato Holzel, 1973 gracina (Navas, 1919) Subgenus CHRYSOPIDIA Navas, 1910 externa cocosensis Adams, 1983 fuscata Navas, 1914 *exul (McLachlan, 1869) comb. n. *ignobilis (Walker, 1860) comb. n. *wollastoni (Navas, 1913) Jiriana Holzel, 1973 *furcifera (Okamoto, 1914) jocasta Holzel, 1973 * galapagoensis (Banks, 1924) comb. n. Junbesiana Holzel, 1973 gujaratensis (Ghosh, 1976) *nigrata Navas, 1910 *harrisii (Fitch, 1855) numerosa Navas, 1914 stenostigma (Navas, 1914) regulata Navas, 1914 incongrua (Navas, 1914) comb. n. remanei Holzel, 1973 *insulata (Fraser, 1957) comb. n. Subgenus CHRYSOTROPIA Navas, 1911 stat. n. iranica (H6lzel, 1967) ‘ciliata (Wesmael, 1841) comb. n. 272 alba auctorum nec L., 1767 kusnezovi (Navas, 1911) lacroixi (Navas, 1911) japonica (Nakahara, 1915) linensis (Navas, 1916) absona (Navas, 1916) melaneura (Navas, 1916) *obliquata (Banks, 1931) comb. n. *orientalis (H6lzel, 1973) comb. n. Genus CHRYSOPODES Navas, 1913 Orlandisa Navas, 1914 Ancylochrysa Navas, 1928 Subgenus CHRYSOPODES Navas, 1913 albopalpis (Banks, 1910) breviata Adams & Penny, 1987 *circumfusa (Burmeister, 1839) comb. n. burmeisteri (Navas, 1929) conisetosa Adams & Penny, 1987 costalis (Schneider, 1851) diffusa (Navas, 1927) duckei Adams & Penny, 1987 gonzalezi (Navas, 1913) indentata Adams & Penny, 1987 inornata (Banks, 1910) *jubilosa (Navas, 1914) laeva (Navas, 1910) *limbata (Navas, 1926) lineafrons Adams & Penny, 1987 mediocris Adams & Penny, 1987 nebulosa Adams & Penny, 1987 nevermanni (Navas, 1928) polygonica Adams & Penny, 1987 *pulchella (Banks, 1910) *geayi (Navas, 1910) *canudasi Navas, 1913 spinella Adams & Penny, 1987 tetifera Adams & Penny, 1987 Subgenus NEOSUARIUS Adams & Penny, 1987 *collaris (Schneider, 1851) *thoracica (Walker, 1853) krugii (Kolbe, 1888) signatalis (Banks, 1911) rufolinea (Banks, 1914) acolhua (Banks, 1949) *divisa (Walker, 1853) *transversa (Walker, 1853) *nobregana (Navas, 1913) hesperina (Banks, 1915) *agatha (Navas, 1925) debilis (Navas, 1926) *uruguaya (Navas, 1927) oglobini (Navas, 1931) *escomeli (Navas, 1922) *figuralis (Banks, 1915) comb. n. *verticalis (Navas, 1929) *flavescens (Blanchard, 1851) S. J. BROOKS & P. C. BARNARD * jaffuelina (Navas, 1918) comb. n. *nigripilosa (Banks, 1924) comb. n. *nosina (Navas, 1913) comb. n. *porterina (Navas, 1910) * poujadei (Navas, 1910) comb. n. *tristella (Navas, 1920) comb. n. Genus CUNCTOCHRYSA Holzel, 1970 *albolineata (Killington, 1935) replacement name for tenella (Schneider, 1851) *baetica (Ho6lzel, 1972) kannemeyeri (Esben-Petersen, 1920) comb. n. ignobilis (Navas, 1921) tarsalis (Navas, 1927) *instabilis (Navas, 1934) *opipara (Holzel, 1973) comb. n. Genus EREMOCHRYSA Banks, 1903 Lolochrysa Banks, 1950 syn. n. Subgenus CHRYSOPIELLA Banks, 1911 stat. n. *brevisetosa Adams & Garland, 1981 *minora Banks, 1935 * pallida Banks, 1911 *sabulosa (Banks, 1897) Subgenus EREMOCHRYSA Banks, 1903 *altilis Banks, 1950 california Banks, 1906 canadensis (Banks, 1911) *digueti Navas, 1911 fraterna (Banks, 1897) *hageni Banks, 1903 israeli Alayo, 1968 *pima Banks, 1950 *pumilis Banks, 1950 *punctinervis (McLachlan, 1869) *rufifrons Banks, 1950 rufina Banks, 1956 *spilota Banks, 1950 * tibialis Banks, 1950 *yosemite Banks, 1950 Genus GLENOCHRYSA Esben-Petersen, 1920 *conradina (Navas, 1910) *franzeni Kimmins, 1952 gloriosa (Navas, 1931) comb. n. *hopkinsi (Esben-Petersen, 1928) comb. n. irregularis (Banks, 1910) marmorata (Needham, 1909) nimbosa (Banks, 1918) *opposita (McLachlan, 1863) * principissa (Navas, 1915) regularis (Banks, 1910) *splendida splendida (Weele, 1909) comb. n. *faceta (Navas, 1912) splendida lucasseni (Weele, 1909) comb. n. splendida timorensis (Weele, 1909) comb. n. tillyardi New, 1980 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) *typica Esben-Petersen, 1920 zeylanica (Banks, 1913) Genus HIMALOCHRYSA Holzel, 1973 Nepalochrysa H6lzel, 1973 syn. n. bhandarensis (H6lzel, 1973) *modesta Holzel, 1973 Genus KOSTKA Navas, 1913 *nacaratus Navas, 1913 Genus MALLADA Navas, 1925 Anisochrysa Nakahara, 1955 Triadochrysa Adams, 1978 syn. n. adamsi (New, 1980) comb. n. *alarconi (Navas, 1915) *alcestes (Banks, 1911) esakii (Esben-Petersen, 1926) alcines (Banks, 1940) comb. n. *alluaudi (Navas, 1930) comb. n. *alticola (Banks, 1931) comb. n. amseli (H6lzel, 1980) ariadne (HGlzel, 1978) astur (Banks, 1937) *atomalis (Navas, 1933) comb. n. *atrosparsa (Tjeder, 1966) *baronissa (Navas, 1921) *basalis (Walker, 1853) *microphya (McLachlan, 1883) jolyana (Navas, 1910) latotalis (Banks, 1910) olatatis (Banks, 1910) formosana (Esben-Petersen, 1913) tagalica (Banks, 1914) skottsbergi (Esben-Petersen, 1924) stigmatus Navas, 1924 *delmasi (Navas, 1927) *delmasinus Navas, 1935 paradoxa (Nakahara, 1955) *bertrani (Navas, 1931) comb. n. *boninensis (Okamoto, 1914) *rutila (Esben-Petersen, 1927) obliqua (Navas, 1929) *burgeonina (Navas, 1936) *caesa (Navas, 1929) comb. n. chailensis (Ghosh, 1977) *chlorella (Navas, 1915) comb. n. *chloris (Schneider, 1851) *clathrata (Schneider, 1845) neglectus (Costa, 1855) nympha (Navas, 1910) riveti (Navas, 1923) cypria (Navas, 1932) cognatella (Okamoto, 1914) nakaharai (Navas, 1915) hoffmanni (Esben-Petersen, 1916) *collartina (Navas, 1932) comb. n. *crassoneura (Weele, 1909) comb. n. darwini (Banks, 1940) comb. n. *decolor (Navas, 1913) comb. n. derbendica (H6lzel, 1967) *desjardinsi (Navas, 1911) comb. n. dierli (H6lzel, 1973) * dispar (Kimmins, 1952) dubia (Holzel, 1973) *duplicata (Navas, 1934) comb. n. *eurycista (Navas, 1914) comb. n. *eurydera (Navas, 1910) comb. n. flaveola (Schneider, 1851) comb. n. * flavifrons (Brauer, 1851) clathrata (Pictet, 1865) *lineolata (McLachlan, 1880) luteola (Navas, 1901) narcissina (Navas, 1910) gallica (Lacroix, 1914) irenaea (Navas, 1915) fiorina (Navas, 1926) cyprina (Navas, 1932) * flavostigma (Esben-Petersen, 1927) comb. n. formosana (Matsumura, 1911) sauteri (Esben-Petersen, 1913) yamamurae (Nakahara, 1915) babai (Kuwayama, 1962) *fortunata (McLachlan, 1882) *genei (Rambur, 1842) * granadensis (Pictet, 1865) clathrata (Pictet, 1865) escudera (Navas, 1908) gunvorae (Tjeder, 1966) hamata (Tjeder, 1966) *handschini (Navas, 1929) collarti (Navas, 1931) *herasina (Navas, 1929) comb. n. *iberica (Navas, 1903) *incerta (Navas, 1936) comb. n. *inclinata (Navas, 1934) comb. n. incrassata (Tjeder, 1966) ingae (Tjeder, 1966) *jgnita (Navas, 1910) comb. n. iota (Navas, 1908) comb. n. *incongrua (Fraser, 1951) comb. n. *innotata (Walker, 1853) comb. n. assimilata (Navas, 1934) *inornata (Navas, 1901) craspedia (Navas, 1915) infecta (Lacroix, 1915) caverina (Navas, 1933) *joannisi (Navas, 1910) comb. n. liberata (Navas, 1935) *khandalensis (Navas, 1931) comb. n. *khandalina (Navas, 1931) comb. n. kibonotoensis (Weele, 1910) kinnaurensis (Ghosh, 1977) *Javata (Navas, 1914) comb. n. Juaboensis (Tjeder, 1966) 274 *Juctuosa (Banks, 1911) macleodi Adams & Garland, 1982 *maculithorax (Kimmins, 1936) comb. n. *madestes (Banks, 1911) comb. n. maghrebinus H6lzel & Ohm, 1984 *makrana (H6lzel, 1966) *melanopis (Navas, 1913) comb. n. mira (H6lzel, 1973) morota (Banks, 1915) comb. n. unicolor (Navas, 1918) herasi (Navas, 1923) *murreensis (Tjeder, 1963) *nea (Navas, 1912) comb. n. *neglecta (Banks, 1931) comb. n. nepalica (Holzel, 1973) *nesophila (Navas, 1920) comb. n. oceanica (Navas, 1914) buxtoni (Esben-Petersen, 1928) *nigra (McLachlan, 1869) comb. n. *notalis (Navas, 1914) comb. n. *noumeana (Navas, 1910) comb. n. nuristana (Holzel, 1982) *nyassalandica (Navas, 1914) *oblonga (H6lzel, 1973) *obvia (Holzel, 1973) opima (H6lzel, 1973) parabola (Okamoto, 1919) *perfecta (Banks, 1895) cockerelli (Banks, 1903) injusta (Banks, 1906) marginalis (Banks, 1906) perpallida (Tjeder, 1966) personata (Navas, 1934) comb. n. pervenosa (Tjeder, 1966) phlebia (Navas, 1927) *pictella (Navas, 1933) comb. n. * picteti (McLachlan, 1880) thoracica (Pictet, 1865) *polyneura (Navas, 1940) comb. n. * prasina (Burmeister, 1839) aspersa (Wesmael, 1841) abdominepunctata (Brauer, 1850) coerulea (Brauer, 1850) ramburii (Costa, 1855) abdominalis (Brauer, 1856) mariana (Navas, 1905) sachalinensis (Matsumura, 1911) *burri (Navas, 1914) caucasica (Navas, 1914) nikkoensis (Okamoto, 1914) *vernalis (Navas, 1926) *pulchrina (Tjeder, 1966) *punctilabris (McLachlan, 1894) comb. n. *rocasolanoi (Navas, 1929) comb. n. *sanvitoresi (Navas, 1913) comb. n. scolius Navas, 1928 sensitiva (Tjeder, 1939) S. J. BROOKS & P. C. BARNARD *serrandi (Navas, 1921) comb. n. *sierra (Banks, 1924) *signata (Schneider, 1851) comb. n. alcatoa (Banks, 1943) *sjoestedti (Weele, 1910) nubilata (Navas, 1910) inopina (Navas, 1914) burgeoni (Navas, 1929) spissinervis (Tjeder, 1966) *subcostalis (McLachlan, 1882) *subcubitalis (Navas, 1901) subflavifrons (Tjeder, 1949) *sumatrensis (Esben-Petersen, 1926) comb. n. *sybaritica (McLachlan, 1875) *tacta (Navas, 1921) horcheri (Esben-Petersen, 1928) teiresias (H6lzel & Ohm, 1982) *tosta (Navas, 1932) comb. n. traviata (Banks, 1940) comb. n. triangularis Adams, 1978 *tripunctata (McLachlan, 1867) *tropica (Hagen, 1858) comb. n. *varians (Kimmins, 1959) comb. n. vartianorum (Holzel, 1973) *venosa (Rambur, 1842) reticulata (Steinmann, 1965) venosella (Esben-Petersen, 1920) distracta (Navas, 1934) * ventralis (Curtis, 1834) aspersa (Schneider, 1851) *venusta (Holzel, 1974) *zelleri (Schneider, 1851) soumainae (Lacroix, 1915) benedictae (Séméria, 1976) Genus MELEOMA Fitch, 1855 *adamsi Tauber, 1969 antennensis Tauber, 1969 arizonensis (Banks, 1903) beardi Tauber, 1969 carapana Adams, 1969 colhuaca Banks, 1949 *dolicharthra (Navas, 1913) cavifrons Banks, 1950 *emuncta (Fitch, 1855) slossonae Banks, 1896 verticalis Banks, 1908 comata Banks, 1950 festivata Adams, 1969 furcata (Banks, 1911) delicata Banks, 1950 hageni Banks, 1949 innovata (Hagen, 1861) kennethi Tauber, 1969 *macleodi Tauber, 1969 mexicana Banks, 1899 nahoa (Banks, 1949) THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 2S) pallida Banks, 1908 *pinalena (Banks, 1958) pipai Tauber, 1969 *poolei Adams, 1969 powelli Tauber, 1969 *rubricosa (Navas, 1913) comb. n. *schwarzi (Banks, 1903) *signoretii (Fitch, 1855) tezcucana (Banks, 1949) *titschacki Navas, 1928 Genus NINETA Navas, 1912 Parachrysa Nakahara, 1915 afghanica HOlzel, 1982 alpicola (Kuwayama, 1956) carinthiaca (Holzel, 1965) *dolichoptera (Navas, 1910) *flava (Scopoli, 1763) subfalcata (Stephens, 1836) *grandis Navas, 1915 gravida (Banks, 1911) *guadarramensis guadarramensis (Pictet, 1865) alvesi (Navas, 1917) guadarramensis principiae (Monserrat, 1980) inpunctata (Reuter, 1894) impunctata (Klingstedt, 1935) reuteri (Tjeder, 1967) nanina (Banks, 1911) * pallida (Schneider, 1845) pomacea (Zakharneko, 1983) *vittata (Wesmael, 1841) proximus (Rambur, 1842) integra (Hagen, 1852) olivacea (Gerstaecker, 1894) inornata (Matsumura, 1911) inornatella (Nakahara, 1914) matsumurana (Navas, 1915) Genus PARACHRYSOPIELLA gen. n. argentina (Banks, 1910) comb. n. Genus PEYERIMHOFFINA Lacroix, 1920 Tjederina H6lzel, 1970 syn. n. * gracilis (Schneider, 1851) comb. n. stenoptila (Schneider, 1851) tricolor (Brauer, 1856) pudica Lacroix, 1920 Genus PLESIOCHRYSA Adams, 1982 stat. n. *armstrongi (Esben-Petersen, 1928) comb. n. *atalotis (Banks, 1910) comb. n. *brasiliensis (Schneider, 1851) *cubana (Navas, 1921) antillana (Navas, 1924) bouvieri (Navas, 1924) *rata (Lacroix, 1926) uribei (Navas, 1927) scalaris (Navas, 1929) yucatanensis (Navas, 1929) cajencis (Navas, 1930) divergens (Navas, 1931) *dussumieri (Navas, 1912) comb. n. *elongata (Navas, 1913) *augusta (Navas, 1913) mariae (Lacroix, 1919) submarginata (Banks, 1918) josephina (Navas, 1930) * gilola (Navas, 1914) comb. n. *impunctata (Navas, 1914) comb. n. *invaria (Walker, 1853) comb. n. *lacciperda (Kimmins, 1956) comb. n. *litorosa (Navas, 1911) comb. n. *oceanica (Walker, 1853) v-rubrum (Brauer, 1865) marcheana (Navas, 1910) ogasawarensis (Okamoto, 1914) paessleri (Navas, 1928) * paraguaria (Navas, 1920) comb. n. *peterseni (Banks, 1924) comb. n. *ramburi (Schneider, 1851) affinis (Rambur, 1842) vicina (Kempny, 1904) jaluitana (Kempny, 1904) neutra (Navas, 1910) *notosticta (Navas, 1913) *deutera (Navas, 1913) *reaumuri (Navas, 1914) *controversa (Lacroix, 1920) *remota (Walker, 1853) comb. n. rubida (Navas, 1929) comb. n. ruficeps ruficeps (McLachlan, 1875) comb. n. procubitalis (Navas, 1912) *ruficeps fervida (Gerstaecker, 1893) comb. n. *scotti (Esben-Petersen, 1927) comb. n. *seurati (Navas, 1922) comb. n. *tahitensis (Navas, 1913) comb. n. *tetrasticta (Navas, 1913) comb. n. hieronyma (Navas, 1917) Genus REXA Navas, 1920 Eurochrysa Esben-Petersen, 1925 corsica (Hagen, 1864) *lordina Navas, 1920 almerai (Navas, 1920) *jordani (Navas, 1929) corsicana (H6lzel, 1965) * raddai (HOlzel, 1966) Genus SUARIUS Navas, 1914 Vasquezius Navas, 1914 Prochrysopa Tjeder, 1936 syn. n. afghana (Holzel, 1967) *alisteri (Navas, 1914) *puparia (Navas, 1914) *caviceps (McLachlan, 1898) fedtschenkoi (McLachlan, 1875) 276 *gobiensis (Tjeder, 1937) kaszabi (Steinmann, 1968) iberiensis Holzel, 1974 iranensis Holzel, 1974 *Jucasi (Navas, 1910) *Juchi (Navas, 1913) pilosella (Navas, 1916) maroccanus Holzel, 1965 mongolica (Tjeder, 1937) gobica (Steinmann, 1965) *nanus (McLachlan, 1893) pretiosa (Gerstaecker, 1894) nymphula (Navas, 1910) nymphulina (Navas, 1915) *egena (Navas, 1940) *nanchanica (Navas, 1927) comb. n. paghmana (H6lzel, 1967) mongolica (Steinmann, 1968) pallidus H6lzel, 1978 ressli Holzel, 1974 storeyi (Navas, 1926) comb. n. *tigridis (Morton, 1921) vanensis (H6lzel, 1967) vartianae (H6lzel, 1967) *walsinghami walsinghami Navas, 1914 *walsinghami orientalis H6lzel, 1978 yasumatsui (Kuwayama, 1962) comb. n. Genus TUMEOCHRYSA Needham, 1909 Chrysoplecta Navas, 1910 *caesarea HOlzel, 1973 cirerai (Navas, 1930) immaculata (Navas, 1910) *indica Needham, 1909 issikii (Kuwayama, 1961) comb. n. *magnifica HOlzel, 1973 *praeclara Holzel, 1973 Genus UNGLA Navas, 1914 stat. n. “argentina (Navas, 1911) comb. n. annulata Navas, 1914 *binaria (Navas, 1923) comb. n. confraterna (Banks, 1913) comb. n. nesotala (Banks, 1944) comb. n. Genus YUMACHRYSA Banks, 1950 stat. n. *apache (Banks, 1938) clarivena (Banks, 1950) yuma (Banks, 1950) Tribe LEUCOCHRYSINI Adams, 1978 Genus BERCHMANSUS Navas, 1913 *adumbratus Navas, 1913 cinctipes (Banks, 1915) *elegans (Guerin, 1844) Genus CACARULLA Navas, 1910 S.J. BROOKS & P. C. BARNARD *maculipennis (Banks, 1910) Genus GONZAGA Navas, 1913 amabilis Navas, 1932 *calliptera Banks, 1944 *nigriceps (McLachlan, 1867) comb. n. notatus Navas, 1929 palliatus Navas, 1929 soroana Alayo, 1968 *torquatus Navas, 1913 Genus LEUCOCHRYSA McLachlan, 1868 Protochrysopa Kolbe, 1888 Allochrysa Banks, 1903 Subgenus LEUCOCHRYSA McLachlan, 1868 *ampla (Walker, 1853) comb. n. annulata (MacGillivray, 1894) arizonica (Banks, 1906) bedoci Navas, 1923 *benoisti Navas, 1933 *benoistina Navas, 1934 bolivari Banks, 1944 boliviana (Banks, 1915) boxi Navas, 1930 brasilica (Navas, 1913) callota Banks, 1915 christophei Banks, 1938 *clara (McLachlan, 1867) *scioptera (Navas, 1913) *colombia (Banks, 1910) claverina Navas, 1927 californica Navas, 1928 cordillera (Banks, 1910) *dolichocera (Navas, 1913) duarte Banks, 1945 ehrhardti Navas, 1929 *erminea Banks, 1945 geminata Navas, 1913 haitiensis Smith, 1931 *ignatii Navas, 1923 insularis (Walker, 1853) virginica (Fitch, 1855) phantasma (MacGillivray, 1894) cerverai Navas, 1924 lestagei Navas, 1922 *Jongicornis (Gray, 1832) loretana Navas, 1935 * magnifica (Banks, 1920) navasi Banks, 1941 antennata Navas, 1921 *negata (Navas, 1913) *singularis Navas, 1913 nigrilabris (Banks, 1915) *notha Navas, 1913 *pretiosa (Banks, 1910) *angrandi (Navas, 1911) *variata (Navas, 1913) delicata Navas, 1925 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 277 reedi Navas, 1919 gloriosa (Banks, 1910) *risi Esben-Petersen, 1933 *gossei (Kimmins, 1940) comb. n. *riveti (Navas, 1913) replacement name for conformis (Walker, serrula Adams, 1979 1853) tavaresi Navas, 1916 *grisoli (Navas, 1912) comb. n. *varia (Schneider, 1851) heriocles Banks, 1944 vegana Navas, 1917 horni Navas, 1932 *phaeocephala Navas, 1929 *hybrida (Rambur, 1842) comb. n. *vigoi (Navas, 1913) indiga Navas, 1928 *vulnerata (Navas, 1913) inquinata (Gerstaecker, 1888) walkerina Navas, 1913 *intermedia (Schneider, 1851) *internata (Walker, 1853) *intermedia (Walker, 1853) comb. n. Subgenus NODITA Navas, 1916 stat. n. [Homonym] Lachlanita Navas, 1929 syn. n. israeli Alayo, 1968 aleura Banks, 1944 kotzbaueri Navas, 1926 alloneura Banks, 1945 laertes Banks, 1945 *alternata (Navas, 1913) *Jafoni (Navas, 1911) comb. n. *amazonica (Navas, 1913) *Jancala Banks, 1944 americana (Banks, 1897) *Jateralis (Navas, 1913) anchietai Navas, 1922 lenora Banks, 1944 antennalis Navas, 1932 longistigma Navas, 1930 antennata (Banks, 1906) *Juctuosa (Banks, 1914) *antica (Navas, 1913) maculata Navas, 1928 *apicalis (Banks, 1915) comb. n. mainerina (Navas, 1929) comb. n. apicata Navas, 1926 marginalis (Banks, 1915) *askanes Banks, 1946 *maronica (Navas, 1915) australis Navas, 1917 [nomen nudum] marquezi (Navas, 1913) azevedoi (Navas, 1913) melanocera Navas, 1916 *camposi Navas, 1933 meridana Navas, 1927 *caucella (Banks, 1910) meteorica (Gerstaecker, 1893) *centralis (Navas, 1913) mexicana (Banks, 1900) ceratica (Navas, 1911) minima (Banks, 1918) *cerverai Navas, 1914 montanola (Banks, 1910) *championi (Navas, 1914) *morenoi Navas, 1934 citri (Ashmead, 1880) *morrisoni (Navas, 1914) comb. n. clepsydra (Banks, 1918) *mortoni (Lacroix, 1926) comb. n. clystera (Banks, 1918) *nativa (Navas, 1911) comb. n. compar (Navas, 1921) *navasi Kimmins, 1940 loyolana Navas, 1925 replacement name for alternata (Navas, 1914) cornesta Banks, 1944 *nesites (Navas, 1913) *cortesi (Navas, 1913) neuralis (Banks, 1910) calverti (Banks, 1914) nevermanni Navas, 1928 cruentata (Schneider, 1851) nictheroyana Navas, 1926 *deminuta (Lacroix, 1926) comb. n. nigrinervis Banks, 1939 diasi Navas, 1922 *nigrovaria (Walker, 1853) dimidia Navas, 1925 notulata Navas, 1924 *diversa (Walker, 1853) comb. n. oenops Adams, 1987 *egregia (Navas, 1913) orthones Banks, 1945 eubule Banks, 1944 pacifica Navas, 1928 *euterpe Banks, 1944 *pallescens Banks, 1946 explorata (Hagen, 1861) *palliceps (McLachlan, 1867) firmini Navas, 1924 panama Banks, 1945 firmini (Navas, 1927) [Homonym] paraquaria Navas, 1929 *floridana (Banks, 1897) *pavida (Hagen, 1861) *fuscinervis (Navas, 1914) platyptera (Gerstaecker, 1888) garridoi Alayo, 1968 *postica (Navas, 1913) gemina Navas, 1929 punctata (Banks, 1903) 278 radiosa (Gerstaecker, 1888) ramosa Navas, 1917 ramosi Navas, 1916 rochana Navas, 1922 rodriguezi (Navas, 1913) rufescens Navas, 1931 salleana (Navas, 1911) senior Navas, 1935 *scurra (Lacroix, 1926) comb. n. *serrei Navas, 1924 stichocera (Navas, 1908) submacula (Banks, 1915) *sulcata (Navas, 1921) *superior (Navas, 1913) surinamensis Banks, 1944 tarini Navas, 1924 texana Banks, 1939 theodori Navas, 1932 theodorina Navas, 1935 vegana Navas, 1925 vieirana (Navas, 1913) vinesi Navas, 1924 ypirangana Navas, 1932 *zapotina (Navas, 1913) zayasi Alayo, 1968 Genus NEULA Navas, 1917 mesana Navas, 1917 Genus NUVOL Navas, 1916 *umbrosus Navas, 1916 Genus VIEIRA Navas, 1913 iridea (Olivier, 1792) comb. n. *Jeschenaulti (Navas, 1911) Subfamily NOTHOCHRYSINAE Navas, 1910 Dictyochrysinae Esben-Petersen, 1918 Genus DICTYOCHRYSA Esben-Petersen, 1917 *fulva Esben-Petersen, 1917 *Jatifascia Kimmins, 1952 *peterseni Kimmins, 1953 Genus HYPOCHRYSA Hagen, 1866 Hypochrysodes Leraut, 1980 *elegans (Burmeister, 1839) nobilis (Schneider, 1851) pernobilis Tjeder, 1967 viridula Adams, 1978 Genus KIMOCHRYSA Tjeder, 1966 *africana (Kimmins, 1937) *impar Tjeder, 1966 raphidioides Tjeder, 1966 Genus NOTHOCHRYSA McLachlan, 1868 Nathanica Navas, 1913 S. J. BROOKS & P. C. BARNARD californica Banks, 1892 *capitata (Fabricius, 1793) *fulviceps (Stephens, 1836) erythrocephalus (Rambur, 1842) Genus PAMOCHRYSA Tijeder, 1966 *stellata Tjeder, 1966 Genus PIMACHRYSA Adams, 1956 *albicostales Adams, 1967 *fusca Adams, 1967 grata Adams, 1956 *intermedia Adams, 1967 *nigra Adams, 1967 Genus TRIPLOCHRYSA Kimmins, 1952 kimminsi New, 1980 * pallida Kimmins, 1952 Genus ‘CHRYSOPA’ incertae sedis Many species of Chrysopidae were originally described in Chrysopa when that genus was ill- defined and much broader in scope than it is now. Although some of those species have now been placed correctly in other genera, a large number still remain incertae sedis. These 239 species are listed below; where possible we have indicated the genus in which the species may eventually prove to belong. aculeata Tjeder, 1966 acuta Hoffmansegg, 1805 adnexa Navas, 1929 adnixa Esben-Petersen, 1913 (? Mallada) adonis Banks, 1937 aegyptiaca Navas, 1915 (? Mallada) alethes Banks, 1940 alobana Banks, 1944 *amabilis Banks, 1938 (? Apertochrysa) annotaria Banks, 1945 annularis Navas, 1921 anomala (Navas, 1929) *antennalis Navas, 1915 (? Mallada) apurina Navas, 1935 argyrea Navas, 1915 atala Brauer, 1865 atrioris Banks, 1920 *aurea Kimmins, 1951 aztecana Banks, 1903 azygota Banks, 1915 bandrensis (Navas, 1929) bandrina Navas, 1935 barberina Navas, 1932 barbouri Navas, 1924 basuto Tjeder, 1966 (? Mallada) batesi Banks, 1946 beccarii Navas, 1929 behni Benthin, 1875 (? Nothochrysa) benaventi (Navas, 1930) bermudezi Navas, 1927 [bimaculata Hagen, 1864 nomen nudum] bineura Navas, 1936 bipunctata Burmeister, 1839 birungana Navas, 1924 (? Mallada) bolivarensis Navas, 1929 bolivari Banks, 1913 bonnini Lacroix, 1919 brevicollis (Rambur, 1842) brevihirta Banks, 1946 bruchi Navas, 1914 buhleri Handschin, 1936 bulbosa (Navas, 1926) bullocki Navas, 1933 caffer Tjeder, 1966 (? Mallada) camposana Navas, 1935 canaria Navas, 1915 (? Mallada) cantonensis Navas, 1931 caprae (Navas, 1929) castalia Banks, 1949 cephalica Navas, 1936 chacranella Banks, 1915 (? Ungla) chemoensis (Navas, 1936) (? Chrysopa s.str.) chusanina Navas, 1933 climacia Navas, 1935 comitissa (Navas, 1914) concinna Banks, 1944 conformis (Rambur, 1842) congolana Navas, 1911 conspersa (Navas, 1929) cornuta Navas, 1926 ricana Navas, 1929 ceratodes Navas, 1932 coronata Navas, 1930 cufrina Navas, 1932 (? Suarius) cymantis Banks, 1944 cymbele Banks, 1933 dahli Navas, 1925 (? Plesiochrysa) *dampfina Navas, 1928 (? Ungla) dancalia Navas, 1931 (? Chrysoperla) darlingtoni Banks, 1938 decarlina Navas, 1924 decaryna Navas, 1924 decolor (Navas, 1936) derota Banks, 1937 deserta Navas, 1912 (? Mallada) devia McLachlan, 1887 dichroa Navas, 1923 diploa Navas, 1935 (? Mallada) distracta Navas, 1930 durandi Lacroix, 1925 (? Chrysoperla) emiliae Lacroix, 1919 estradai Navas, 1924 eudora Banks, 1937 everina Banks, 1946 THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) excelsior Banks, 1937 exterior Navas, 1925 externa Navas, 1924 cubensis Navas, 1927 extranea Navas, 1923 facialis Navas, 1927 fascialis Banks, 1906 favrei Navas, 1935 feana (Navas, 1929) festana Navas, 1932 filosa (Fabricius, 1787) fischerina Navas, 1933 fratercula Banks, 1940 frequens Esben-Petersen, 1913 (? Mallada) gasteria Navas, 1917 gestroi Navas, 1929 279 geyri Esben-Petersen, 1915 (? Brinckochrysa) gialina Navas, 1932 (? Suarius) grandis Navas, 1933 (? Austrochrysa) gratiosa Navas, 1933 (? Chrysopa s.str.) gravesi Navas, 1926 (? Mallada) grazianii Navas, 1932 (? Chrysoperla) guineensis Navas, 1929 hansensis Navas, 1929 healdi Navas, 1926 (? Mallada) hestia Banks, 1918 heudei Navas, 1934 huasanensis Navas, 1915 ifranina Navas, 1936 (? Mallada) ilota Banks, 1915 (? Chrysoperla) inaequata Navas, 1935 incerta Banks, 1895 (? Ceraeochrysa) incisa Banks, 1949 incompleta Banks, 1911 inconspicua Navas, 1914 (? Mallada) *jniqua Navas, 1931 (? Mallada) intemerata Navas, 1934 irrorella Navas, 1935 (? Mallada) isolata Banks, 1913 jacobsom Weele, 1909 (? Apertochrysa) javanica Esben-Petersen, 1913 (? Mallada) julia Navas, 1927 karakurti Rossikova, 1904 kiangsuensis Navas, 1934 kulingensis (Navas, 1936) Jagunensis Navas, 1920 (? Mallada) lambda Navas, 1933 Jaminaris Navas, 1917 lateralis (Guérin, 1844) (? Ceraeochrysa) latithorax Banks, 1913 leptana Banks, 1914 libera Navas, 1928 (? Chrysoperla) lojensis Navas, 1929 lorenzana Navas, 1913 loretensis (Navas, 1931) loriae Navas, 1929 luederwaldti Navas, 1923 280 lurida (Navas, 1930) lybica Navas, 1914 (? Mallada) mainerii Navas, 1929 malayana Esben-Petersen, 1926 marchionissa Navas, 1915 marcida Banks, 1937 margaritina (de Beauvois, 1809) mendocensis Navas, 1918 menetriesi Hagen, 1861 meriani Navas, 1925 (? Plesiochrysa) mesonotalis Esben-Petersen, 1926 (? Chrysocerca) metanotalis Navas, 1924 mexicana Banks, 1901 meyeri Handschin, 1936 mimeuri Navas, 1936 mindanensis Banks, 1937 mosconica Navas, 1931 (? Mesochrysa) nadali Navas, 1913 naesonympha Brauer, 1865 navasi Lacroix, 1914 nicolaina Navas, 1929 (? Brinckochrysa) nigripalpis Banks, 1910 notulata Banks, 1937 nymphodes Navas, 1914 (? Suarius) obesa Navas, 1929 ochracea Albarda, 1881 ophthalmica Navas, 1913 oralis Navas, 1914 orestes Banks, 1911 orientalis Hagen, 1859 (? Mallada) paolii Navas, 1928 parallela Navas, 1931 parishi Banks, 1913 (? Ceraeochrysa) peruviana Navas, 1924 pieli (Navas, 1931) pigmentata Handschin, 1935 plesia Navas, 1918 polonica Lurié, 1897 polyphlebia Navas, 1914 postica Navas, 1936 pucayensis Navas, 1929 puerula Navas, 1921 pullata Banks, 1944 (? Ceraeochrysa) punctithorax New, 1980 (? Apertochrysa) pusilla Schneider, 1851 pygmaea Navas, 1930 (? Suarius) quadornia Banks, 1949 reboredina Navas, 1933 reedina Navas, 1919 reichardti Bianchi, 1931 robusta Banks, 1906 (? Nineta) rossa Navas, 1924 rothschildi Navas, 1915 rotundata Navas, 1929 (? Brinckochrysa) ruizi Navas, 1934 sajanina (Navas, 1928) S. J. BROOKS & P. C. BARNARD sanguinea (Navas, 1928) sansibarica Kolbe, 1897 (? Mallada) sarta Banks, 1914 satoruna Navas, 1922 scalai Navas, 1917 selenia Navas, 1912 (? Mallada) senior (Navas, 1928) sequens Banks, 1943 (? Mallada) serrana Navas, 1927 siderocephala Navas, 1933 sillemi Esben-Petersen, 1935 (? Chrysoperla) silvestrina Navas, 1929 simplex Navas, 1908 smitzi Navas, 1913 (? Mallada) *sobria Navas, 1933 (? Mallada) solaria Navas, 1930 (? Chrysoperla) steinbachi Navas, 1925 sumatrensis (Navas, 1929) (? Austrochrysa) tacorensis Navas, 1934 taikuensis Kuwayama, 1962 (? Cunctochrysa) tenera Navas, 1924 [ternata Hagen, 1861 nomen nudum] tetuanensis (Navas, 1934) (? Chrysopa s.str.) thallina Navas, 1919 thieli Navas, 1929 (? Plesiochrysa) thomasensis Navas, 1929 tibialis Banks, 1914 (? Leucochrysa) torrei Navas, 1924 tortolana Banks, 1949 trifurcata Banks, 1949 tucumana Navas, 1919 valdezi Banks, 1924 (? Signochrysa) *varicosa Navas, 1913 vegeta Navas, 1917 venulosa Navas, 1918 (? Ungla) venulosa Navas, 1923 vilallongai Navas, 1940 replacement name for villalongae Navas, 1935 villica Navas, 1929 virgata Handschin, 1935 virgestes Banks, 1911 viridinervis Jakowleff, 1869 wagneri Esben-Petersen, 1932 (? Chrysopa s.str.) yuanensis Navas, 1932 yuanica Navas, 1932 zina Navas, 1933 zulu Tjeder, 1966 (? Mallada) ACKNOWLEDGEMENTS. This review was based partly on an uncompleted work on the chrysopid genera of the world begun by the late D. E. Kimmins, and we have made use of many of Kimmins’ wing venation drawings in this paper. We are particularly grateful to Prof. P. A. Adams (California State University, Fullerton, U.S.A.), Dr H. THE GREEN LACEWINGS OF THE WORLD: A GENERIC REVIEW (NEUROPTERA: CHRYSOPIDAE) 281 Holzel (Bruckl, Austria), Dr T. R. New (La Trobe University, Victoria, Australia), Mr J. D. Oswald (Cornell University, Ithaca, U.S.A.) and Dr S. Tsukaguchi (University of Osaka, Japan) with whom we have had interesting and productive discussions and correspondence and who have willingly provided us with information on chrysopid taxonomy. We would also like to thank the following for the loan of specimens: Prof. P. A. Adams, Dr H. M. André (Musée Royal de l’ Afrique Centrale, Tervuren, Belgium), Dr Burmeister (ZoGdlogische Sammlung des Bayerischen Staates, Munich, West Germany), Mr R. Danielsson (Zoologiska Institution, Lund, Sweden), Dr O. S. Flint (USNM), Dr P. Grootaert (Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium), Dr K. K. Giinther (Museum fiir Naturkunde der Humboldt-Umiversitat, Berlin), Dr. W. Hogenes (Instituut voor Taxonomische Zoologie, Amsterdam, The Netherlands), Dr H. Holzel, DrN. P. Kristensen (Zoologisk Museum, Copenhagen, Denmark), Mr J. Legrand (Muséum National d’ Histoire Naturelle, Paris, France), Dr M. Mansell (National Collection of Insects, Pretoria, South Africa), Dr N. D. Penny (California Academy of Sciences, San Francisco, U.S.A.), Dr R. Poggi (Museo Civico di Storia Naturale, Genoa, Italy), Dr C. Remington (Peabody Museum, Yale University, New Haven, Connecticut, U.S.A.), Miss M. Schneider (Queensland University, St Lucia, Australia), Dr R. T. Schuh (American Museum of Natural History, New York, U.S.A.), Dr J. van Tol (Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands), Dr S. Tsukaguchi, Mr C. Vogt (Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A.) and Dr V. B. Whitehead (South African Museum, Cape Town, South Africa). 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Abachrysa 163, 165, 177 Aeolops 201 Allochrysa 247 Anachrysa 206, 209 Anapochrysa 132, 134 Ancylochrysa 210, 212 Anisochrysa 223 Ankylopterygini 118, 120, 121, 155 Ankylopteryx 121, 155, 212 Anomalochrysa 185, 189, 192, 193, 238 Apertochrysa 187, 215 Apochrysa 118, 132, 133, 135 Apochrysinae 118, 120, 121, 132, 252 Atlantochrysa 123, 188, 215, 227 Austrochrysa 190, 223 Belonopterygini 118, 120, 121, 122, 123, 163 Belonopteryx 121, 163, 165, 177 Berchmansus 243 Bornia 186, 192, 241 Borniochrysa 192 Brinckochrysa 193 Cacarulla 243, 245 Calochrysa 163, 167, 175, 177 Ceraeochrysa 123, 195, 219 Ceratochrysa 121, 196, 203, 229, 234 Chrysacanthia 163, 168 Chrysaloysia 149, 163, 170, 172 Chrysemosa 198 Chrysocerca 123, 194, 200, 229, 241 Chrysopa 118, 119, 121, 123, 198, 201, 234, 242 Chrysoperla 194, 204, 231 Chrysopidia 186, 206, 207, 210, 241 Chrysopiella 121, 210, 215, 230 Chrysopinae 118, 120, 149 Chrysopini 118, 121, 122, 184 Chrysopisca 201, 203 Chrysopiscini 184 Chrysoplecta 238 Chrysopodes 120, 155, 193, 210 Chrysotropia 123, 208, 209 Cintameva 201 Claverina 132, 136 Cunctochrysa 190, 213, 227 Dictyochrysa 254, 259, 260, 264 Dictyochrysinae 252 Domenechus 132, 137 Dysochrysa 163, 170 Emerobius 201 Eremochrysa 121, 215, 217, 218, 230 Ethiochrysa 155, 157 Eurochrysa 234 Evanochrysa 163, 172, 177 Glenochrysa 169, 196, 218 Goliva 175 Gonzaga 132, 243, 246, 249 Himalochrysa 123, 219, 236 Hypochrysa 254, 256 Hypochrysodes 256 Indochrysa 161 Italochrysa 123, 163, 168, 173, 175, 177, 182 Ttalochrysini 118, 163 Joguina 132, 138 Kimochrysa 254, 257, 261, 262 Kostka 221 Lachlanita 249 Lainius 132, 140 Leucochrysa 118, 123, 192, 196, 243, 246, 2A7, 248, 250, 251 Leucochrysini 118, 122, 243 Lolochrysa 215 Loyola 132, 141 Madachrysa 177, 179 Mallada 123, 186, 188, 205, 215, 223 Melanops 201 Meleoma 123, 215, 225 Mesochrysa (Navas, 1927) 175, 177 Mesochrysa (Navas, 1936) 198 Metachrysopa 201, 203 Minva 201, 203 Musola 196 Nacarina 123, 163, 165, 167, 175 Nacaura 132, 141 Nadiva 175, 177 Nadivini 163, 177 Nathanica 258 Neda 193 Nepalochrysa 220, 221 Neosuarius 211, 212, 240 Nesochrysa (Navas) 163, 170, 172, 177, 180, 183, 200 Nesochrysa (Fraser) 168 Neula 126, 149, 243, 251 Nigrochrysopa 201 Nineta 123, 203, 227, 238 Nobilinus 132, 143 Nodita 177, 243, 249 Nodochrysa 163, 179 Nothancyla 132, 145, 192 Nothochrysa 254, 258 Nothochrysinae 118, 120, 121, 252 Nuvol 126, 149, 243, 251 Oligochrysa 132, 146, 148 Orlandisa 210 Oviedus 172, 177, 179 Oyochrysa 163, 175, 180 Pamochrysa 254, 257, 260, 263 Parachrysa 227 Parachrysopa 201, 203 Parachrysopiella 229 Parankylopteryx 155, 157 Peyerimhoffina 194, 231 Pimachrysa 254, 261, 262 Plesiochrysa 121, 190, 198, 203, 215, 227, 232 Polyphlebia 201, 203 Prochrysopa 236 Protochrysopa 247 Pseudochrysa 200 Rameta 175 Retipenna 155, 159 Rexa 234 Scoliochrysa 190 Semachrysa 155, 161 Sencera 155, 157 Signochrysa 155, 162 Stigmachrysa 163, 179, 183 Suarini 184 Suarius 199, 236, 241, 242 Synthochrysa 132, 147 Tjederina 205, 231 Triadochrysa 223, 224 Triplochrysa 254, 264 Tumeochrysa 203, 229, 238 Turnerochrysa 163, 175, 183 Ungla 238 Vasquezius 236 Vieira 243, 252 Yumachrysa 241 A kenites, Retervwnes b ihe (49 B54) hd Hm cw Bah (ge BOE aie gree a ae eo = ean a ee ws ApPm- Pica U OAs wy 67° bom img , a enya 0 . ke ia wera OS Ze 4 wa ee ya ay eG 1 to¢ 9 A » = 2 ind ‘baat aa — - Ce - bee te, a “eq « ~ —, aor a =. ’ ya ja | hah) Ka i Tipe Atyes (0 1 : haere ls - ies PS 1h 7 “a< . = ® .. i. ‘ . —— _ _— i= ij i —_ 06 = i > i = * a +. »S® eet ad 3 “ (eS ove t- 3, a E ie ; yo Gt Oe ~ .* ¢ a) « 2 ow —ee, am |/ sf i aks Ps 2 tn wl Apes Ue <4 ~~ Fi —onfere fei E. W. CLASSEY Ltd P.O. Box 93 FARINGDON, OXON SN7 7DR, ENGLAND rr Natural History Publishers Booksellers and Library Agents SU REEEEEERenmend_cooeneeeeEneEE Catalogues on request Agents for the publications of the BRITISH MUSEUM (NATURAL HISTORY) SOO Publishers of ENTOMOLOGIST’S GAZETTE An illustrated quarterly journal of Palaearctic Entomology Volume 41 commences January 1990 Current Annual Subscription (post free) £16.00 (US $35.00) | "j 1 The green lacewings of the world: a generic review (Neuroptera: c ht S. J. Brooks & P. C. Barnard — ENTOMOLOGY SERIES Vol. 59, No. 2, September 1990