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Zeitschrift/Journal: Stapfia

Jahr/Year: 1999

Band/Volume: 0060

Autor(en)/Author(s): New Timothy R.

Artikel/Article: Neuroptera und Biological Control (Neuropterida). Neuropteren und biologische Schädlingbekämpfung (Insecta: Neuropterida) 147-166 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

Neuroptera and Biological Control (Neuropterida)

Timothy R. NEW

Abstract pidae, predominantly Chrysoperla spp. These developments are outlined and dis- Abstract: Members of three fami- cussed to provide a broad perspective of lies of Neuroptera (Coniopterygidae, current uses of lacewings in integrated Hemerobiidae, Chrysopidae) have been pest management and how uses might used in biological and integrated control, expand in the future. mainly of field crop pests, and in augmen- Key words: lacewings, snakef- tation programmes to increase the impacts lies, Chrysopidae, Hemerobiidae, Coniop- of natural enemies. Most emphasis has terygidaepest management, predators, been in use of a limited range of Chryso- pesticides.

Stapfia 60, zugleich Kataloge des OÖ. Landesmuseums. Neue Folge Nr. 138 (1999). 147-166

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Introduction tory species of the above groups: only 23 were Neuroptera, and these totalled 743 individuals Most neuropteroid insects are predators, of the overall nearly 124000 predators collec- both as larvae and adults. Although other fee- ted. More broadly, SUNDERLAND et al. (1996) ding habits occur in the lacewings, the above cited predator richness of around 400 genera- uncritical generalisation has led to considera- list species in cereals in Britain, and of 600- ble interest in using terrestrial lacewings as 1000 species in cotton and soybean crops in biological control agents able to devour pest the United States. Nevertheless, Neuroptera insects and reduce their populations on a wide remain one of the few groups which, in prac- variety of field, orchard and glasshouse crops. tical IPM terms, may be considered 'aphido- They thus vie with Coccinellidae and other phagous'. They have thus attracted attention groups of predatory insects as popular biologi- for use against a wide variety of small prey cal control agents in many parts of the world. arthropods, in a great variety of agricultural, Considerable research has been undertaken to orchard and forestry crops. determine and quantify their influences, and This account is a broad survey and evalua- how they may be manipulated for enhanced tion of the use of lacewings in biological con- impacts on pest populations. As with many trol, with some discussion of future directions other insect groups, such 'applied relevance' and their incorporation into increasingly has been the major stimulus to research and effective integrated pest management pro- the single most important factor leading to grammes. The main topics reviewed are: increased biological understanding of neurop- teroids, and of the subtle but far-reaching bio- 1. The taxa of lacewings useful in biologi- logical differences between closely related cal control. taxa. 2. The rationale, principles and practice of biological control. Many different neuropteroid groups have 3. The place of lacewings in integrated been implicated as important predators in pest management. various ecosystems, because of their abundan- ce, and their seemingly close and consistent 4- The production and manipulation of associations with particular habitats and/or lacewings for pest management. prey taxa, coupled with their voracity and high fecundity. However, in practice members Why these taxa? of only three families of Neuroptera have pro- ved more broadly amenable to such asses- Early records of lacewings as predators sment and employable in pest management emphasised the voracity and behaviour of lar- programmes. Indeed, the lacewings currently vae, in particular. The conical pits formed by of interest as biological control agents compri- certain myrmeleontoid larvae, in which they se only limited spectra of 3 of the 17 recent lie buried except for the jaws exposed awaiting families of Neuroptera, namely Coniopterygi- the arrival of insects such as wandering ants dae, Hemerobiidae and Chrysopidae, with by which they subdue by dragging under the sub- far the greatest attention given to the last- strate, are known widely as one of the classic named, the green lacewings (NEW 1975, examples of insect predators. Likewise, chryso- CANARD et al. 1984, MCEWEN et al. 1999). In pid larvae camouflaged by the debris from past consequence, we know far more of the biology captures are evocative as 'aphis-lions', paralle- of these taxa than of most other Neuroptera. ling the 'antlions' above. Such examples led to Neuroptera are not one of the more diver- broad assumption that most lacewings could se groups of predators in most agricultural be valuable as predators in crop protection, systems, and can be far outnumbered by groups but this has not been substantiated. Thus, ant- such as Araneae, Carabidae, Staphylinidae, lions and their relatives tend to occur in rela- Coccinellidae, Syrphidae and some Heterop- tively natural habitats, often not associated tera as generalist predators. Thus, in an exten- with vegetation or with the insects that feed sive survey of arable land in Switzerland, on it. Despite their considerable abundance DUELLI & OBRIST (1995) collected 599 preda- and diversity in areas such as the more arid

148 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at regions of southern Africa (leading MAN'SELL likely to be utilised in the foreseeable future. & ASPÖCK, 1990, to comment on their impor- As ASPÖCK et al. (1991, see also H. ASPÖCK tance as significant predators) and Australia, 1991) noted, further investigation of the pre- such areas are usually not those used for inten- datory role of Raphidioptera is needed, becau- sive crop production. The long life cycles of se their large numbers in some environments many such taxa render them somewhat non- suggest that they could be significant. Recent responsive to rapid changes in prey spectrum surveys in Italian vineyards (PANTALEON'I and density. As importantly, though, many are 1990, LETARDI 1994) showed the snakefly 'ambush predators' which do not actively Parainocellia bicolor (COSTA) to be abundant, search for prey but await it coming to them. and it may have some value as a predator on In contrast, many members of Coniop- such longlived plants. However, simple num- terygidae, Hemerobiidae and Chrysopidae are bers do not prove a regulatory role for preda- associated primarily with vegetation (someti- tors on prey, and there is no quantitative evi- mes of restricted subsets, such as coniferous or dence for values of Raphidioptera as biological broadleaved trees, or trees versus low vegetati- control agents. Although larvae of some Bero- on: MONSERRAT & MARIN 1996) and feed thidae feed on termites (JOHNSON & HAGEN almost exclusively on small herbivorous 1981), their general scarcity renders their like- arthropods representing groups which also ly use in control insignificant. However, alt- contain pest taxa, and for which they actively hough there is no quantitative evidence to search. Lacewing larvae can move considera- support the suggestion, it is possible that local ble distances to seek prey. Unfed hatchling high densities of Myrmeleontidae could be larvae of Conwentxia pineticola can travel more inimicable with biological control program- than 40m and survive for more than 10 hours mes through devouring beneficial predatory before feeding, for example (FLESCHNER ants. 1950). Likewise, newly hatched chrysopid lar- vae can survive for up to 72h without food, or Taxonomic introduction to up to 9d if given water (SUNDBY 1966), and the families involved may search actively for prey over most of this time. As such, they are to some extent 'pread- Coniopterygidae, dusty-wings, are an iso- apted' for manipulative use in similar environ- lated lineage of Neuroptera and are the smal- ments and to exploit related prey. Most of the lest members of the Neuroptera. Most species species of interest frequent the temperate have forewings only 2-3 mm long, and they zones with strongly seasonal climates, those are recognised by the reduced venation and regions used most intensively for diverse, the wings and body covered by white or grey- intensive agriculture. They tend to have well- ish wax. The global fauna was monographed defined patterns of voltinism, with one to by MEINANDER (1972), who later (MEINANDER several generations a year, and can respond 1990) provided an augmented checklist of numerically to increased prey density within a world species. Coniopterygidae are widespread season. All can at times be abundant in their but, as for the other families treated here, natural habitats, and may show well defined many of the genera and species have more peaks of abundance, conferring synchronicity restricted distributions. Hemerobiidae, brown with predominant prey species. Within a habi- lacewings, and Chrysopidae, green lacewings, tat a sequence of different lacewing species are also widespread. These two families have can peak at different times of the growing sea- traditionally been treated as closely related, son for considerable collective effect on lon- but U. ASPÖCK (1992) has noted that this glived pests or those present for extended peri- supposed alliance may be based on inadequate ods. appraisal of characters. Sporadic advocacy for use of other neur- Recent taxonomic surveys have done opteroids in biological control, such as for Psy- much to clarify the integrity and relationships chopsidae (TlLLYARD 1919) and Raphidiop- of the genera in each family. OSWALD (1993) tera, has not led to any constructive adoption reviewed the genera of Hemerobiidae, and and, in general, most other groups are not BROOKS & BARNARD (1990) elucidated the

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features and relationships of the mass of potential biological control agents is probably chrysopid genera earlier masquerading under rather more limited than popularly supposed, the name 'Chrysopa', and defined the limits of and becomes more so if we exclude the nume- this genus effectively for the first time. Most rous rare and ecologically specialised taxa earlier literature on Chrysopidae includes whose use would necessitate excessive mani- generic combinations incompatible with these pulation of their normal biology - in as far as modern definitions, and care is needed in its this can be predicted. However, other com- interpretation, even in relatively well known mentators take a broader view, that the poten- faunas. tial for lacewings in biological control has Higher groupings, subfamilies, of the three scarcely been tapped, and that many more families are generally clearly definable, though taxa will eventually be found useful. the relationships between some of them are Within the range of taxa noted above, the unclear. Many structural characters are used vast majority of attention has devolved on one for definition at subfamily and genus levels, small group of chrysopids, sometimes referred and include many features of wing venation to simply as 'the green lacewing', namely the and body. Genitalic arrangement can be of cri- sibling species of the Chrysoperla camea (STE- tical importance, and many female chrysopids PHENS) group. Because of the taxonomic and can still be placed only tentatively to genus if biological confusion which has persisted wit- unaccompanied by the more distinctive males. hin this group, it is noted separately here, to Genitalic features are of almost universal alert readers to the inherent complexities importance for separating species. which can occur in many groups of natural The three families are all diverse; Coniop- enemies and hinder clear interpretation of terygidae comprises around 450 described spe- their use in pest management. cies, Hemerobiidae about 550 species, and Chrysopidae contains around 1200 described The Chrysoperla camea problem species. The genera of current interest in bio- logical control were reviewed by NEW (1999). The taxonomic and other interpretative Briefly, these include 7 genera of Coniop- challenges of this complex of presumed sibling terygidae, 5 of Hemerobiidae and about 12 of taxa are by no means solved. The precise num- Chrysopidae, but those included simply reflect ber of taxa included in the complex is still historical opportunism and implication (often unclear. "C. cornea" was long considered to be based on only a single species in a genus) rat- polymorphic and variable in northern tempe- her than the outcome of more comprehensive rate regions, and complex series of names have surveys of the total utility of each family. been applied both in Europe and North Ame- Thus, whereas both main subfamilies of Coni- rica to account for this variability. Examinati- opterygidae are represented, all Chrysopidae on of a wide range of characters, such as included belong to one tribe (Chrysopini) of courtship songs, adult and larval morphology, the largest subfamily (Chrysopinae). Most of enzyme polymorphisms, variation in diapause the taxa implicated represent large genera or regime and other ecophysiological features those represented in northern temperate regi- have confirmed the presence of at least six ons where interest in biological control was members of the group in Europe alone initiated and where their incidence in associa- (THIERRY et al. 1998). Many areas have at tion with putative pest taxa on crops has been least two sympatric species (DUELLI 1996), and more thoroughly documented. The almost the notion of different taxa occurring together total lack of knowledge of the biology of most is now replacing the concept of a single, varia- tropical lacewings implies strongly that other ble entity (THIERRY & ADAMS 1992). suitable candidates could be revealed in due The taxonomic details of this complex are course. However, some taxa appear not to relevant here only in relation to the inadver- have potential applied value. Species of the tent confusions that have occurred in the past, large chrysopid genus ludochrysa PRINCIPI, for and their wider relevance to confusion bet- example, apparently feed as larvae on ant bro- ween exotic and native biological control od as inquilines, so that the spectrum of agents. In retrospect, early attempts at intro-

150 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at auction such as the release of putative C. ear- major agricultural and environmental weeds ned from the Indian subcontinent into North are exotic species and use of specific herbivor- America to control spotted alfalfa aphid in the es from their areas of origin is a key compo- 1950s (ADAMS in THIERRY & ADAMS 1992) nent of control. Likewise, many insect pests are impossible to analyse, as the released spe- (such as the majority of aphid species in Aus- cies was not differentiated from the very simi- tralia) are exotic species, colonising only after lar C. plorabunda (the 'lead species' of the their host plants had entered cultivation, and complex in North America). Such differentia- use of imported natural enemies is a focus for tion, indeed, might not then have been possi- integrated control. ble. Of greater concern is the current practice 2. Some natural enemies, particularly pre- in Europe of distributing insectary stocks dators, are likely to be already present in the widely across international boundaries. Thus, areas where a pest occurs. It may be possible to insectary stocks from northern Italy have fre- foster, intensify and augment their effects on quently been exported for release in countries the pest population, thereby using native spe- in more northern parts of Europe under the cies rather than imported control agents. The- guise of C. cornea. Some of those releases are se native taxa can then be reared in captivity undoubtedly of species alien to the new areas, in large numbers for augmentative release, undetected because of lack of critical appraisal usually on a periodic basis to match the cha- of the various taxa possibly present. It is racteristics of annual cropping systems. This doubtful whether the nature of some such approach can avoid the problems of introdu- introductions can ever be clarified but, in cing yet further exotic species into possibly order to avoid further confusion, careful docu- sensitive environments, and use of native spe- mentation of the origin of biocontrol stocks cies of predators is widely forecast to increase purported to be of the same species is needed, in the future (WAAGE 1992). PARRELLA et al. together with responsible deposition of vou- (1992) also noted the current and future cher specimens for future reference. values of this strategy. The two approaches are not necessarily Biological control mutually exclusive, in that once an exotic spe- cies becomes established and naturalised as Widespread global concern over the use of part of the regular fauna it can be manipulated pesticides, ranging from problems of pest resi- in similar ways to a native species. stance to side-effects on non-target organisms Massive benefits have occurred from both and dissemination and accumulation in the approaches in the past, and are likely to con- wider environment, during the last few deca- tinue as biological control is increasingly a key des have stimulated development of increa- component of many pest management strate- singly sophisticated management strategies gies. However, the practice of classical biolo- against a wide variety of agricultural, orchard gical control has also engendered considerable and other arthropod pests. A predominant concerns and debate in recent years, over the component of such integrated pest manage- safety of the agents used. Introduction of any ment programmes is the use of parasitoids, pre- exotic species (that is, one new to the site of dators and pathogens affecting the pest, introduction) imposes a duty of care for the collectively its 'natural enemies'. Two major receiving environment. This normally invol- contexts have been pursued, both diverse, but ves assurance of specificity - so that an intro- which are very different in their ecological duced herbivore will not stray from its target implications: weed to eat native or other desirable plant 1. "Classical biological control" involves species, or an introduced predator also attack the importation and release of natural enemies sensitive native taxa. Debate has accelerated from a pest's native range to attack it in areas following HOWARTH'S (1983, 1991) implicati- where it is exotic. Pest status often arises in on (albeit without quantitative scientific evi- part because of release from the factors which dence) that large numbers of endemic Hawai- limit population numbers in a natural, balan- ian insects may have become extinct due to ced context. Thus in Australia many of the introduced parasitoids and predators which

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have spread from crops into natural ecosy- over quarantine issues associated with intro- stems on these remote islands. NAFUS (1994) duced agents (COULSON et al. 1991). Despite implied a similar situation for Guam, and such calls for 'group screening' procedures and appraisals have led to much wider debate over exemptions for additional members of pre- the environmental safety of classical biologi- viously tested taxa, there seems little sound cal control agents. The broader credibility of alternative to a species-by-species evaluation classical biological control has suffered accor- of risk. dingly, and many authors (such as SAMWAYS 1988, 1994; SANDS 1997) have stressed the Properties of desirable predators need for balanced appraisal of 'acceptable for biological control risk'; as SANDS noted "The success and envi- An ideal insect predator for biological ronmental benefits from introduction of nar- control of a given insect pest should be (1) rowly specific exotic agents are often overloo- specific or near-specific to that pest for prey; ked with more public attention given to the (2) respond rapidly in numbers to that prey impact by generalist predators on native spe- when it is encountered, and should be vora- cies". cious (VAN EMDEN 1966, included parameters There is an important dichotomy in the of high appetite, high level of activity and practice of classical biological control as used high abundance: efficient killing and a high against plant weeds and arthropod pests. Scre- rate of capture success are implicit); (3) be ening of herbivores introduced against weeds able to detect the prey in low densities and is well-established, with sound protocols track it effectively in the environment, whilst which do much to guarantee the specificity, not dispersing to colonise neighbouring ecosy- hence the environmental safety, of the agents. stems; (4) be climatically adapted to the full Simplistically, any tendency for the herbivore (or near complete) geographical range of the to eat other unrelated plant taxa is likely to prey, and naturally synchronised in time and lead to its rejection from further considerati- space with the prey; (5) not be unduly disrup- on. Plants of conservation concern may be ted by other environmental cues from efficient incorporated routinely into the spectrum discovery and exploitation of the prey; (6) be tested. In contrast, screening of predators tolerant/resistant to pesticides employed in against pest arthropods is much less advanced the same IPM programmes, and (7) be gene- and, until recently, has often been very super- rally compatible with other natural enemies ficial or generalised; basic protocols are still for potential combined use. In addition, prac- being developed as the need to do so becomes tical aspects such as amenability to economi- apparent. Such screening is difficult but until cal mass rearing, perhaps using artificial diets, this is done, widespread suspicions of'risk' will and inundative and augmentative release inevitably persist (GlLLESPlE & NEW 1998). without undue dispersal from the release site Risk assessment for classical biological control become important in enhancing and manipu- is difficult, not least because an exotic species lating the agent for greater efficiency. continues to evolve and adapt in its new envi- Few, if any, species fulfil the above deside- ronment, and may change its host or prey rata absolutely, and the science and credibility spectrum and tolerances to a wide range of of biological control devolves on selecting the environmental factors, but is a universal need most suitable agents from a variety of subopti- in such cases. SlMBERLOFF & STILING (1996) mal taxa available. Indeed, in many situations emphasised that specific biological control some of the above 'ideals' may not be so; for projects should not be assumed to be inno- example, lack of extreme prey specificity con- cuous until substantial effort has been made to fers practical advantages such as use of the support this assumption. Research and respon- same species against a variety of pest species, sible regulation play important roles in this and allows use of other foods for subsistence complex arena, with the primary responsibili- during periods when the preferred prey is scar- ty of minimising collateral damage from intro- ce or absent, thus providing for effective reser- duced organisms. Wide consultation is occur- voir populations of the predator to be main- ring with increasing frequency, as is debate tained between growing seasons of the target

152 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at crop. Polyphagous predators may be of far and even from the same species on one crop to more practical use than narrowly specific another crop or different environment. Howe- ones. Thus, PREE et al. (1989) advocated the ver, it is important to distinguish between an use of C. cornea on the basis of its wide host introduced predator species (for which specifi- plant and prey ranges. In Australia, Mailada city may be needed for effectiveness and to signatus is promoted as a predator of aphids, avoid undesirable side effects) and exploiting mites, whitefly and Lepidoptera (HÖRNE et al. native predators occurring naturally in the 1999). As with chemical pesticides, strict spe- area of operation and which may be polypha- cificity can restrict marketing opportunity gous as part of the local community of indige- and, in view of the relatively high costs of nous consumers available for manipulation for mass rearing predators, a variety of target pests IPM. and crops is valuable to diversify the applicati- The criteria deemed important may on of the species. However, in some cases depend also on the context for control. The polyphagous predators may eat other natural prime aim of many biological control projects enemies and thereby interfere with other on crops is for short term control during the aspects of 1PM. PRINCIPI & CANARD (1984) life of that crop, with the whole operation cited larvae of C. cornea eating nitidulid and during this period of a few months or less, and coccinellid predators of date palm scale {Par- repeated in its entirety the next season. The latoria blanchardi) in palm groves. Larvae may prime needs may then be for the predator to also eat aphids containing parasitoid mum- be an effective coloniser and build up numbers mies (TREMBLAY 1980). Cannibalism can also rapidly (LUFF 1983). Alternatively, biological be problematical, with virtually all polypha- control may be viewed as more permanent, gous species likely to eat conspecifics, and with the establishment of agents providing cannibalism likely to increase when other prey long term benefit. is scarce (DUELLI 1981). A functional difference between these C. camea larvae can develop to maturity approaches is often that introduction of a clas- on a diet of conspecific eggs, and this was vie- sical biological control agent is expected to wed by BAR & GERLING (1985) as a means to regulate its target prey by density-dependent facilitate survival when other prey is scarce. action, and should therefore be specific, whe- Thus, under some circumstances, cannibalism reas augmentation of naturally occurring pre- can be adaptive (CANARD & DUELLI 1984). dators is viewed as a short term measure. The Sibling cannibalism is one (indirect) means to latter is therefore not usually intended to convert maternal tissue into offspring tissue. maintain the pest below economic threshold Instead of producing a few large eggs, a female levels, but as a 'therapeutic' control of tem- lacewing can produce many small eggs, and porarily high prey numbers (NORDLUND cannibalism result in a few well-fed larvae. In 1996). Specificity is therefore not required for conditions where larval food is abundant, predation to be an effective method of sup- many more may survive. Ability to colonise pression in the field. new areas might be advantageous in tracking a Most use of lacewings is indeed in such low density, dispersed pest. Problems and augmentative releases, involving aspects of potential conflicts in IPM can thus arise when mass production and repeated releases with the same characteristics which render an potential for genetic alteration or various introduced predator a useful, adaptable biolo- forms of behavioural modification using gical control agent essentially coincide with semiochemicals. Species such as C. camea are those which would render a species a dange- well adapted to short duration habitats, such rous invasive exotic from other points-of- as annual cropping systems. Annual crops are view. frequently highly disturbed habitats, monocul- Many of the features of a given predator tures which are very unstable, and with inade- can be elucidated only by detailed study of the quate natural chemical defences against pest species, and only very limited generalisation attack. The biological variability within the should be made by extrapolation from data on 'camea complex' may allow for selection of other species, from species to higher category, appropriate forms suitable for different crops

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and regions. In annual crops, augmentative are based commonly on flimsy evidence, releases of natural enemies are viewed increa- mainly co-incidence and the undoubted vora- singly as 'inundative' to cause immediate mor- city of many lacewing larvae. Numerous early tality, so that measurement of mortality is a records (summarised by NEW 1975) show that critical aspect of monitoring success (OBRYCK1 individual larvae of all three families may con- et al. 1997). Enhancing the effects of released sume up to several hundred prey individuals as predators through optimal release strategies they develop. Many such records include and local environmental manipulation is an impressively high numbers, but most do not integral part of such exercises. differentiate between 'prey killed' and 'prey eaten'; partial consumption of prey is frequent Lacewings as 'ideal predators' among insect predators (NEW 1991), and many of the problems of interpreting voracity Relatively few Neuroptera appear to be are noted by PRINCIPI & CANARD (1984). specific in their choice of prey species, but From the practical viewpoint of reducing pest opportunistically take a variety of prey taxa numbers, of course, 'killing without eating' is a (NEW 1986). Limitations may be more on the perfectly satisfactory outcome! NEUMARK grounds of physical comparability and ease of (1952) reported two larvae of Chrysoperla car- capture, rather than on strict taxonomic gro- nea as devouring 6487 and 4645 eggs of Mat- unds. Thus, lacewings are only rarely reported sucoccus, respectively. C. plorabunda and as predators of Thysanoptera in Europe (RlU- several related species in California can eat DAVETS 1995), probably because thrips are more than 200 cotton aphids (BURKE & MAR- 'tough'. However, CHANG (1998) reported lar- TIN 1956). Larvae of smaller hemerobiids and vae of C. plorabunda (FITCH) to consume more coniopterygids are similarly voracious. Data thrips than aphids and suggested that they for Coniopterygidae, summarised by Lo VERDE might have value in controlling them in some & MONSERRAT (1997), clearly imply the like- field crops in North America. Most prey ly value of some species as control agents but, reported commonly are relatively soft-bodied again, much of the information on this family taxa, such as aphids, other Homoptera, mites needs corroboration and further research. or small caterpillars, with some lacewings They are generally deemed likely to be impor- apparently associated reasonably consistently tant in biological control of insect pests (TAU- with one or more of the above groups, which BER & ADAMS 1990), but are 'usually overloo- collectively contain a high proportion of her- ked because of their small size' (PENNY et al. bivorous crop pests. Relatively few complete 1997). surveys of the prey spectrum of lacewing spe- cies have been made, and the relationships Attributes of both larva and adult stages between habitat specificity and food specifici- are relevant in evaluating the likely impacts of ty are commonly not clear. For example, spe- lacewings on prey populations. Whereas both cies of Coniopterygidae or Hemerobiidae asso- stages are predators in Coniopterygidae, most ciated consistently with either coniferous or Hemerobiidae and some Chrysopidae, adults broadleaved trees in the northern hemisphere of some Chrysopidae (including many of tho- (MONSERRAT & MARIN 1996) are clearly se predominantly valued in pest control) feed restricted to developing on prey species on on honeydew or nectar, thus requiring diffe- those tree groups and thus correspondingly rent food sources from their larvae. limited in distribution. But whether habitat There have been relatively few comparati- restriction mediates prey restriction or the ve trials on the suitability of particular prey converse is not easy to determine. In some species to particular lacewing species, with other cases, sporadic records of prey associati- their influences on development, fecundity on have been translated into broader dogma and general fitness reported. CANARD'S (1970, without critical reappraisal. 1972) classic study of the suitability of five Most lacewing taxa implicated as useful aphid species as prey for Chrysopa perla larvae predators are simply those which have been indicated some important effects on adult bio- studied, or which have been found with the logy. Thus, although Megoura viciae supported pest species of concern. Their practical values larval development and was readily eaten,

154 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at resulting adults had very low fecundity, with However, polyphagy is a complex state to males sterile. Many such effects were summa- appraise. It can enable a predator to extend rised by PRINCIPI & CANARD (1984), showing and diversify its habitats and ecological influ- influences of food on duration and extent of ences. Food quality1, though, can vary greatly chrysopid development, differential mortality, not only between prey species (so that some and adult performance. Some prey can be prey are 'essential food' for particular predator toxic, even though they are acceptable to species in that they are adequate to support lacewing larvae. The main conclusion is that reproduction, whereas others are 'alternative studies of food utilisation and acceptability food' which support development but not may need very careful conduct as 'information reproduction), but also within a prey species, published on the feeding habits of Chrysopi- depending on its own nutritional regime. dae in nature is often imprecise' (PRINCIPI & Influence of suitability of a particular prey spe- CANARD 1984). cies to a predator can vary seasonally. In a bro- ad appraisal of predators' diets in arable ecosy- Two Australian Hemerobiidae compared stems, SUNDERLAND et al. (1996) suggested by NEW (1984) differed considerably in their that 'It seems likely that, in general, more relative response to Psyllidae and cabbage than one prey type is needed by predators to aphids (Brevicoryne brassicae) as prey. The maximise fitness'. Polyphagy allows for aug- generalist Micromus tasmaniae WALKER perfor- mentation of limited supplies of high quality med well on both prey species, but the more foods to ensure adequate energy and nutrients specialised arboreal Drepanacra binocula to grow, develop and reproduce. (NEWMAN) appeared to be much the better adapted to native psyllid prey. However, labo- Searching behaviour by lacewing larvae is ratory trials of the sort reflected in such studies efficient, with details of search pattern chan- may not always adequately represent the field ging once prey is encountered during extensi- situation in which a variety of prey species ve wandering, to more intensive patterns. may coexist, and may overlook some other The complexity of the plant substrate may aspects of the interaction (FRAZER & GILBERT 'guide' how predators search in particular 1976). In general, 'specialist' predators may be environments. Plant structure may also affect expected to be adapted in various ways to the susceptibility of prey, by providing refuges their prey of choice whereas generalist species in which they can hide. C. camea larvae are may show less distinct differences on different more effective in reducing populations of Rus- prey species. Many of the lacewings used com- sian wheat aphid (Diuraphis noxia (MORDVIL- monly in biological control fall into the latter KO)) on Indian rye-grass (Oryzopsis hymenoi- category. des) than on crested wheat grass (Agropyron Prey specificity can be based on a broad desertorum), and MESSINA et al. (1995, 1997) variety of genetically controlled traits (TAU- attributed this in part to the narrower blades BER & TAUBER 1987), including (a) choice of of the former, so that lacewings could then oviposition site, (b) adult food requirements rarely pass an aphid without contacting it. for mating, (c) large egg size, (d) specialisati- Their capture rate was therefore higher, but ons of larval behaviour, (e) enhanced larval also led to many aphids contacted falling from development and (0 univoltinism. TAUBER &. the plant. On crested wheat grass, fewer TAUBER showed that considerable genetic aphids were contacted and fewer were dislod- variation underlies the feeding differences bet- ged, so contacts were more likely to result in ween the monophagous Chrysopa sbssonae capture (CLARK & MESSINA 1998). (which feeds only on a single aphid species, Diet substantially influences egg produc- Prociphilus tesselatus (FlTCH)) and the genera- tion by lacewings, and the size of eggs in par- list C. quadripunctata. Young larvae of C. qua- ticular species may be related to prey specifici- dripunctata suffered high mortality in presence ty (TAUBER & TAUBER 1987). Fecundity of all of this aphid (ALBUQUERQUE et al. 1997), and three families is usually in the order of hun- clear species-specific defensive and feeding dreds, and can extend much higher in some behaviour patterns occur in these two sister- species. With rapid growth rates in the warmer species (MlLBRAlTH et al. 1993) parts of the year, some species can respond

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rapidly to abundance of suitable prey. Adult Increased compatibility between different diet is also highly influential, with dramatic predator species may be achieved if they inter- responses found in two species of Chrysoperla act in ways which serve to reduce the like- by CARVALHO et al. (1996). Females fed on lihood of competitive effects between them, high quality diet (yeast plus honey, 1:1) laid and increase their collective influences on a an average of 2304 ± 188 eggs (C. extema, n = prey species. Some chrysopid larvae have been 6, oviposition period 84-5 ± 10.5 days) and shown to secrete an oviposition-deterring 2160 ± 159 eggs (C. mediterranea, n = 6, ovi- pheromone (ODP), which deters females from position period 103.3 ± 8.6 days). The poorest laying eggs at sites searched previously by lar- food tested, pure honey, in contrast led to out- vae. The effects of this can extend beyond puts of only 25 ± 6 and 94 ± 20 eggs, respec- intraspecific influences, to interactions of dif- tively for the two species. Measures of repro- ferent species and genera, so that the chryso- ductive performance in lacewings must be pid ODPs are also allomones (RuziCKA 1997a, related clearly to both larval prey suitability b 1998, RUZICKA &. HAVELKA 1998). Substra- and adult diet. Protein and carbohydrate are tes contaminated by chrysopid ODP deterred both needed by adults of such species for good a ladybird, C. septempunctata, from laying. reproduction, but measuring the efficiency of The ODP of C. oculata SAY persists for food utilisation is difficult. several weeks and is relatively stable up to about 140° C. Unfed first instars, as well as Single or multiple predator later stages, mark the sites they search for prey species? (RuziCKA 1994). The ramifications of this The question of whether to use single spe- behaviour are complex and remain to be inve- cies of natural enemy in a given pest control stigated in detail. Predators may, for example, operation, or a combination of species, be able to avoid others which would eat them remains controversial, with one key argument if prey were scarce, so that there may be being whether different predators may interact advantage in spreading offspring more evenly in an 'additive way' (CHANG 1996) to enhan- between patches of aphids. This would be ce control, or interfere with each other to achieved by females searching for ODP-free reduce overall control success. Such influen- sites. In practical terms, this might lead to a ces for generalist predators may include inter- higher efficiency of aphid control. Conversely, specific cannibalism. Support for both view- as RUZICKA & HAVELKA (1998) noted, a points exists, and there is probably no univer- strong female response to ODPs (or ODAs) sal answer in view of the great variety of pre- might favour growth of aphid colonies which datory insects and strategies (NEW 1991). survive the 'first wave' of predators but are still Some cases of complementarity may be rea- protected by those deterrents for some time. sonably clearcut: in field crops in southeastern Females of C. perla, C. commata and C. Australia a single common hemerobiid camea tested by RuziCKA (1998) all strongly {Micromus tasmaniae) is mainly aphidopha- avoided oviposition on substrates contamina- gous and a manipulable chrysopid (Mallada ted by their own larvae or the other species in signatus) feeds easily on lepidopterous larvae, this group. prey not favoured by Micromus. The two spe- Nevertheless, intraguild predation may be cies may thereby complement each other for responsible for low numbers of lacewings multi-pest contexts. found after some commercial releases (ROSEN- CHANG (1996) undertook laboratory trials HEIM et al. 1993). Intraguild predation is very to determine the larval behaviour of C. plora- common (POLIS et al. 1989). The egg-stalks of bunda and the ladybird Coccinella septempunc- chrysopids have commonly been presumed to tata whilst feeding on bean aphids {Aphis fabae confer protection from other predators, but SCOPOLI), but found no clear evidence of the eggs are indeed vulnerable to ladybirds interactions between these species, probably (LUCAS 1998) and others. The effects of pre- reflecting that the larvae tend to occur in dif- dators may vary also depending on whether ferent places on the plant so that they rarely one or several species of prey are present. Pre- encountered each other. sence of alternative prey may simply provide a

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distraction, especially if it is more easily acces- quently tested for insecticidal effects (CROFT sible than the target species. Conversely, the 1990); indeed, it has been adopted as a 'repre- presence of one herbivore species can increase sentative predator' in such studies, simply the susceptibility of a second species - for because of the extent of background knowled- example by presenting a resource attractive to ge and the easy availability of the lacewing for predators and increasing their abundance and trials. GRAFTON-CARDWELL & HOY (1985a) feeding intensity in a habitat patch. Trials on enumerated laboratory trials on 128 different the interaction between the bird cherry-oat pesticides. Many of these were difficult to eva- aphid (Rhopaiosiphum padi (L.)) and the Russi- luate because of the different methods and an wheat aphid (Diuraphis noxia) suggested protocols used, and differences in the stages or that C. plorabunda can control the latter on age classes tested. However, all stages were range grasses, but its effectiveness may be highly tolerant of many of the pesticides eva- hampered by presence of such other cereal luated. High susceptibility to field dosage rates aphids. Presence of R. padi may have reduced of most carbamates and organophosphorous predation on D. noxia because R. padi is the insecticides prompted a survey of interspecific more accessible species or because its faster variability, as a prelude to investigating deve- growth rate caused it to outnumber the other lopment of resistance in the laboratory. species on plants that initially received equal Other extensive trials on C. camea from ratio of the two species (BERGESON & MESSI- Californian alfalfa fields were summarised by NA 1997). HOY (1990). Eggs, larvae and adults from four colonies were screened with six pesticides Integrated pest management (carbarylmethoxyl [carbamates], permethrin, Integration of natural enemies with insec- fenvalerate [pyrethroids], diazinon, phosmet ticide use and other methods of suppression is [organophosphates]). Populations from the a priority in developing integrated pest mana- four locations responded with significant dif- gement, so that themes such as insecticide ferences to all six chemicals; one population tolerance and resistance become central to consistently had highest mortality, one the development of practical programmes. A wide lowest, and the other two were intermediate. variety of pesticides used in agriculture also Differences in survival corresponded generally affect beneficial insects, and knowledge of with pesticide usage in the alfalfa. This survey these effects is clearly important in designing was important in suggesting that populations pest management regimes. Early studies of of C. camea responded to local pesticide sel- integration of chrysopids with pesticide use ection pressures, and implied that geographi- were reviewed by BlGLER (1984), who noted cal differences in tolerance reflect past selec- the apparent advantages of Chrysopidae for tion pressures (PREE et al. 1989). IPM as reflected in their physiological and HOY (1990) summarised detailed earlier ecological tolerances to a variety of pesticides studies (GRAFTON-CARDWELL & HOY 1985a, then widely used. 1985b, 1986) in which this intraspecific varia- bility and trials for selection for insecticide a. Chemical pesticides. resistance were discussed in more detail. Sel- The effects of pesticides on arthropod ection response with carbaryl led to a rapid natural enemies of agricultural pests have and high level of resistance; the resistant been reviewed extensively (THEILING & strain could not be killed after the third selec- CROFT 1988, HOY 1990), but some lacewings tion by a wide range of concentrations of the had earlier attracted attention because of their pesticides, and the resistance may be determi- apparently natural tolerances, based on labo- ned by one or few major genes (GRAFTON- ratory trials with larvae. Some generalities CARDWELL & HOY 1986). Changes in fitness thereby started to emerge - such as the like- due to laboratory rearing were compared bet- lihood that Chrysopidae are more tolerant ween the 'base' and 'resistant' colonies, to than Hemerobiidae to pyrethroid insecticides determine any differences in performance. (CROFT 1990). C. camea has figured highly in Larvae and pupae of the base colony had hig- such trials, as one of the insects most fre- her survival rates than the resistant strain, but

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duration of development did not differ signifi- wing generation (tebufenozide), and (4) daily cantly between the two colonies. The fecundi- oviposition rate reduced (diflubenzuron). ty of resistant females was slightly the higher, Literature on the sublethal effects of pesti- but percentage hatch of eggs did not differ. cides continues to expand. The IGRs noted Methods for screening the various stages were above represent the recent trend toward using developed by these authors. more slowly acting pesticides with a more sel- ective mode of action. They differ in their pre- In addition to intraspecific variations and cise modes of action: fenoxycarb is a juvenile changes in tolerance to pesticides, tolerance hormone analogue, diflubenzuron is a chitin may differ greatly between species (ROUSCH synthesis inhibitor, and tebufenozide is an 1995) but, as ROUSCH noted, this does not ecdysone based moulting inducer. This variety usually influence biological control in practice suggests the likelihood that IGRs will be because primary choice of agents is mainly by found to have a substantial array of undesira- other criteria. In contrast, pesticide resistance ble effects on natural enemies, and clarifying is of greater relevance because it implies grea- these is urgent in helping dispel the illusion of ter variation which allows some individuals to safety such chemicals have fostered. HAT- tolerate doses which are normally lethal. T1NGH & TATE'S (1995) studies in South Afri- Assumptions that particular pesticides or ca demonstrated clearly that IGRs can interfe- groups of pesticides are 'safe' for use in con- re with biological control by coccinellids. A junction with particular groups of natural ene- second chitin synthesis inhibitor, triflumuron, mies are based commonly simply on the lack applied topically to adult C. carnea did not of conspicuous lethal effects. However, a wide affect fecundity or longevity at doses tested by variety of sublethal effects, many of them dif- SENIOR et al. (1998), but egg and larval deve- ficult to enumerate and evaluate, may also cast lopment were both substantially retarded. doubt on the values of some such chemicals. First instars were the most heavily affected. For example, insect growth regulators (IGRs) An effective dose can be obtained by larvae have been presumed generally to be compa- simply by tarsal contact (SHUJA et al. 1998). tible with IPM, and have been defended as Severe effects, with mortality of up to 100 %, such (HATTINGH 1996), from the reasoning of chitin synthesis inhibitors on C. camea that since they interfere with specific metabo- were noted also by BlGLER & WALDBURGER lic pathways of pests they are more selective to (1994), although earlier studies by WILKINSON natural enemies than conventional insectici- et al. (1978) had shown no effects on adults or larvae, but substantial pupal mortality. VOGT des (CROFT 1990). They often interfere with (1994) also found substantial effects on larvae natural enemy development and their biologi- and pupae of this species. cal control capacity. Following documentati- on that IGRs can have severe detrimental Case by case evaluation is needed to assess effects on populations of some beneficial coc- effects of IGRs, as for any other group of pesti- cinellids, RUMPF et al. (1998) showed that cides. Sensitive measures of sublethal exposu- they can have more severe effects on Micro- res to insecticides, such as toxin-specific rate mus tasmaniae than organophosphorous insec- of head acetylcholinesterase (AChE) inhibiti- ticides and a pyrethroid they tested. Effects of on, may be of some value in studies of impac- three IGRs (fenoxycarb, diflubenzuron, tebu- ts of particular chemicals on lacewings. fenozide) were compared with those of two Activity of AChE was inhibited much more organophosphar.es (methyl parathion, azin- rapidly in M. tasmaniae than in C. camea lar- phos-methyl) and a pyrethroid (permethrin) vae, corresponding to higher tolerance of the for life table parameters determined for adults latter in mortality tests (RUMPF et al. 1997), reared from treated larvae and controls. The and again confirming the dangers of trying to effects noted included (1) a higher proportion generalise about insecticide effects on larger of female lacewings (diflubenzuron), (2) redu- taxonomic groups. ced longevity (fenoxycarb, diflubenzuron), (3) total number of eggs redued for one generati- b. Biological pesticides. on (fenoxycarb, diflubenzuron) and the follo- Protocols for testing for side effects of

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microbial pest control agents have been deve- (1998) revealed much higher mortality in lar- loped mainly because of use of specially vae reared on Bt-fed lepidopterous larvae (62 engineered pesticides, such as forms of BaciUus %) compared with Bt-free prey (37 %). 'Sick' thuringiensis. CROFT (1990) used C. camea as a prey may also cause nutritional deficiencies in representative model predator to discuss for- predators, but HlLBECK et al.'s study suggested mulation of standard protocols for evaluation. that reduced fitness of C. camea was indeed For adults, ingestion and direct contact, associated with Bt. More limited trials by because of direct penetration of the integu- SALAMA et al. (1982) had earlier suggested ment (by fungi) was recommended for tests, that C. camea larvae had prolonged develop- whereas the wider activities of larvae sugge- ment and reduced food intake when fed on sted need for three categories of tests: (a) topi- Spodopiera larvae fed on a diet containing Bt, cal, with larvae (with introduced prey) tested but those larvae were fed the test diet for only in cages which have been sprayed with the seven days, and then returned to normal, pesticide and allowed to dry to leave residue; healthy prey. (b) feeding larvae on prey sprayed with the Tests of the effects of nuclear polyhedrosis pesticide, and (c) feeding larvae on prey virus from a noctuid moth on larvae of C. cor- which have ingested the pesticide. For the last nea (HASSAN & GRONER 1977) revealed no two trials, feeding for seven day periods was influences on larvae, or on fecundity of resul- recommended. ting adults (or viability of their eggs) when The entomophagous fungus Verticillium larvae were sprayed directly, fed on infected lecanii has been used successfully as a microbi- diet, or exposed to residual film of the sprays. al insecticide against aphids in greenhouses, and has recently been tested for effects on C. c. Cultural controls. camea as an important predator in the same Considerations of larval and adult beha- environments. Larvae of C. camea were trea- viour of lacewings can affect their use in con- ted with the fungus by immersion, and fungus- junction with 'cultural controls' such as inter- treated aphids (A. craccivora) were fed to third cropping, and changing tillage regimes in instar larvae. Feeding capacity of larvae was crops to help conserve natural diversity. decreased by both treatments. Likewise, both Counts of chrysopid eggs (species, singular or direct infection and feeding on fungus-infe- plural, not specified or separated) on cotton in sted aphids increased duration of the larval Nicaragua interplanted with corn, beans or and pupal stages; and feeding on infected weeds (SCHULTZ 1988) revealed some signifi- aphids reduced the proportion of adults that cant numerical differences. Significantly fewer emerged, due to contamination within the eggs were found on cotton interplanted with cocoon. Searching capacity of larvae was also corn or weeds than in monocultures. In this impaired by infection (SEWIFY & EL ARNAOU- case, crop diversification was apparently asso- TY 1998). ciated with reduced numbers of beneficial ins- Under laboratory conditions, larvae of the ects, contrary to the results of many other, closely related C. kolthoffi were highly suscep- similar appraisals, and corroboration of this tible to Metarhizium anisopliae, the green mus- would be of considerable general interest. cardine fungus, one of the most widely used fungi in microbial control (VENTURA et al. 1996). Manipulating lacewings for biolo- Commercial Bacillus thuringiensis (Bt) ins- gical control ecticides have been used predominantly against lepidopterous pests, and transgenic The foregoing has confirmed that lace- crop plants incorporating endotoxins from the wings, at least of a few generalist taxa, can bacterium are now being introduced. The play very positive roles in a variety of pest safety of genetically engineered crop plants of control contexts, but predominantly for short- this nature is currently of considerable con- term manipulation to control seasonal pests cern. Prey fed with Bt may cause death of pre- on field and glasshouse crops. The biological dators. Trials with C. camea by HlLBECK et al. basis of their values is reasonably well under-

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stood, and underpins also the ways in which the mode of delivery and the release rate and the lacewings can be manipulated for enhan- timing) were evaluated in California ced value in integrated pest management. Use vineyards in trials against leafhopper pests of C. camea and its close allies, in particular, (DAANE & YOKOTA 1997). Chrysopids are has led to development of commercial mass- usually released as eggs, with the caveats of rearing operations in North America and losses from cannibalism during shipment and Europe, which have spawned similar operati- any period of delay before deployment, but ons elsewhere for other species of potential DAANE & YOKOTA noted the possible advan- value. Much of this draws directly on the tages of larval release because the voracious pioneer work by HAGEN and his colleagues in third instar stage would be reached more California, by which the use of artificial diets, quickly, and predatory pressure increased and of attractant principles for field manipula- rapidly in the field. This might also help to tion were established as a solid basis for later reduce intraguild predation in some circum- development and refinement. stances. In Californian cotton fields, the dominant predatory arthropods other than However, some taxonomic confusion is lacewings are Heteroptera. Suspicion of preda- still apparent in the precise species used. In a tion by the bugs on lacewing eggs was raised recent survey, O'NEILL et al. (1998) reared ins- by the great differences in field abundances of ects sold in the USA by three commercial ins- chrysopid eggs (common) and larvae (rare). ectaries as C. camea; although their sample ROSENHEIM et al. (1993) believed that decrea- was small, all individuals reared were C. rufila- sed survival of lacewing larvae was primarily bris, a species not usually placed in the camea due to predation by Heteroptera. It occurred complex. Although C. rufilabris is indeed sup- despite prey being abundant. plied extensively by commercial insectaries in the United States (HUNTER 1997), its biology However, releases of eggs by DAANE & differs considerably from that of C. camea. As YOKOTA (1997) gave very low hatchability noted earlier, such confusions can have far- (28.4 ± 12 % of eggs), and this high mortality reachng consequences in evaluation of control was attributed tentatively to high temperatu- success, and in broader conservation issues. As res in the vineyards, coupled with a relatively with most aspects of mass-rearing of natural long pre-hatching period because freshly laid enemies, adequate quality control is vital. eggs were used. Point-of-hatch eggs may be a Additional point-of-sale evaluations by more suitable release stage in increasing survi- O'NEILL and his colleagues showed the impor- val, despite prior risks of cannibalism from tance of posting at a correct, critical stage of 'early hatchers'. Release of eggs gave no diffe- development. Chrysopids are posted as eggs in rences in leafhopper numbers when compared most operations, usually with a supply of food to untreated control plots, but significant (such as Sitotroga eggs) to sustain larvae as reductions occurred from larval releases and they hatch, but hatching of larvae en route high survival of third instars. can lead to considerable cannibalism. The Various systems have been developed for percentage alive on receipt was generally in releasing lacewings in the field. Those tested the range of 60-80 %, but addition of supple- by DAANE & YOKOTA (1997) ranged from 6 mentary food is clearly advantageous, and 175 to 1 235 000 eggs or larvae/hectare, but rapid release of the agents may also be neces- showed no correlation between numbers sary to avoid further deaths from starvation or released and prey density. cannibalism in confined surroundings. Most The production and manipulation of such releases are to open field systems (DAANE chrysopids, in particular, for augmentative et al. 1997), notwithstanding the values also control has achieved a high level of success. of lacewings in glasshouses (SCOPES 1969), The stages of a commercial operation are (1) where both C. camea and wholly predatory mass rearing, with use of artificial or semiarti- species (such as Chrysopa perla) may be useful. ficial diets for larvae and adults, (2) holding The precise method of release can also adults and harvesting the eggs for sale and have far-reaching consequences. Three aspec- colony maintenance, (3) optimising releases ts of this (the developmental stage released, and (4) monitoring and maintaining the

160 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at released populations in the vicinity of the concentrate lacewings in particular areas of crop. Early developments in this field were crops, with sprays of artificial honeydew con- described by TULISALO (1984). Some hemero- taining acid hydrolysed L-tryptophan effective biids also show considerable promise for this in increasing numbers of C. camea (HAGEN et approach, with mass rearing for Micromus al. 1976, VAN EMDEN & HAGEN 1976). Refi- angulatus in Europe described by STELZL &. nement of this approach is continuing (McE- HASSAN (1992). WEN et al. 1994), Mass rearing must be cost effective, and Capitalising on lacewing lifecycles in bio- considerable efforts continue to refine the pro- logical control necessitates understanding of cess, depending increasingly on artificial diets any diapause regime and the factors which and progressive automation of the production control seasonal development. This is also process. Again, most work has been underta- relevant in designing optimal regimes for stor- ken on species of Chrysoperla. NORDLUND & age of reared material without loss of repro- MORRISON (1992) identified major areas for ductive performance and general viability. attention as larval food composition and pre- TAUBER et al. (1993) showed that C. camea sentation, adult feeding and oviposition effec- adults can be maintained at low temperatures ts, mechanised egg collection and de-stalking, for around six months without significant mechanised larval rearing units, preceding reduction in fertility or fecundity. CHANG et field application. Details of all these stages are al. (1995) extended that study to show that readily available, but the main biological inte- storage under short day (8 hour light: 16 hour rest is in understanding and exploiting the ins- dark) conditions at 5°C , with a carbohydrate ects' feeding habits and patterns of develop- and protein diet, gave post-storage adults ment and movement in relation to producing whose reproductive performance equaled that large numbers efficiently and enhancing their of 'normal' unstored adults. The implications effects in the field. of this were seen as reducing investment in facilities and labour without loss of product The use of artificial diets for lacewings is quality. CHANG et al. proposed a protocol by no means new, extending back some 50 which could be tailored both for short-term years, with HAGEN's (1950) studies of protein and long-term maintenance of the lacewing. requirements in adult diet still of fundamental Considerable variation in the life history traits importance. These early diets were liquid- of this species suggested the need for different based, but recent trends to develop more solid storage strategies for different populations diets for chrysopids, to better resemble the (TAUBER et al. 1997b). texture of their normal prey (COHEN 1998, COHEN & SMITH 1998) appear likely to impro- C. extema, some populations of which do ve both quality and quantity of the insects not diapause, is restricted to relatively warm produced. However, even simple 'artificial conditions, but trials on storage at 10 ° C sho- honeydew' (commonly a mixture of sucrose, wed considerable possibility for keeping stocks yeast and water) can facilitate larval develop- for release (TAUBER et al. 1997a). Adults sur- ment and be of practical importance in allo- vived well for at least four months, though wing C. carnea to develop at low prey densi- oviposition rates declined from two to four ties. months in storage. Adult diets for such glyciphagous species Dispersal of liberated predators from the have incorporated yeast hydrolysates and release site can obviate any desirable longer sugars to produce high reproductive activity term benefit. As DUELLI (1980) showed, C. and, again, those diets derived from HAGEN's camea undertakes migratory flights soon after work (for example, HAGEN & TASSAN 1966) emergence and may thus leave the areas whe- to enhance protein content have been used re it is needed. Use of semiochemicals, as abo- frequently since that time. However, particu- ve, is important in helping to maintain popu- lar constituents of adult diet may be critical in lations in crop environments, but MCEWEN et affecting fecundity (MCEWEN et al. 1995), and al. (1998) also described the use of overwinte- in field manipulation through their attractant ring chambers for adult C. camea to help sta- properties. Food sprays have been employed to bilise populations. These chambers provide

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adults with a focal retreat, by which overwin- Acknowledgements tering survival could be enhanced considerab- ly; the chambers may also be transportable to I thank Univ. Prof. Dr. Horst Aspöck for provide inocula of adults for fresh sites in his invitation to contribute to this special spring. issue of Stapfia, for his friendship and advice The extensive work summarised above for over many years, and for his comments on an C. camea and its relatives is among the most earlier draft of this essay. wide-ranging and informative bodies of data on any insect predator, and displays the varied uses for generalist predators in biological con- Zusammenfassung trol. As PARRELLA et al. (1992) emphasised, such use of native natural enemies is likely to Bisher sind Arten von drei Familien der become increasingly important in IPM, and Neuroptera - Coniopterygidae, Hemerobiidae marks a determined transition from the impor- und Chrysopidae - in der biologischen Schäd- tation of specialist natural enemies and classi- lingsbekämpfung bzw. im integrierten Pflan- cal biological control. Use of specialist preda- zenschutz eingesetzt worden, vorwiegend zur tors is likely to continue in control of particu- Bekämpfung von Schädlingen von Feldfrüch- lar pests, but the effective integration of C. ten und im Rahmen von Programmen zur camea (s.l.) with a variety of other facets of Steigerung der Einbindung natürlicher Fein- IPM renders it exceptional. Further studies of de. Besondere Aufmerksamkeit wurde einigen the complex, and of other chrysopids will Chrysopiden, vorwiegend Arten der Gattung undoubtedly extend their range of applicati- Chrysoperla, geschenkt. Die gegenwärtigen ons. Entwicklungen werden umrissen, und Mög- lichkeiten für einen vermehrten Einsatz von A number of generalist hemerobiids clear- Neuropteren in der Zukunft werden diskutiert. ly also merit additional attention for enhance- ment and manipulation in crops. The toleran- ce of some hemerobiids to lower temperatures than many chrysopids can withstand can ren- References

der them useful for control purposes at times ALBUQUERQUE G.S., TAUBER M.J. & CA. TAUBER (1997): when chrysopids are not active. Thus, species Life-history adaptations and reproductive costs of Hemerobius in California are active throug- associated with specialization in predacious ins- ects. — J. Anim. Ecology 66: 307-317. hout the year and H. pacificus is the only com- mon active predator in artichoke fields during ASPÖCK H. (1991): Grundlagen des möglichen Einsat- zes von Raphidiopteren in der biologischen the winter (NEUENSCHWANDER 1975, NEUEN- Schädlingsbekämpfung. — Verh. XII. Int. Symp. SCHWANDER et al. 1975, NEUENSCHWANDER & Entomofauna Mitteleuropa. Akad. Wiss. Ukrai- HAGEN 1980; further details in NEW 1988) ne (1991): 26-33. and its use might extend the overall period of ASPÖCK H., ASPÖCK U. & H. RAUSCH (1991): Die Raphidi- opteren der Erde. — Goecke & Evers, Krefeld. effective control by lacewings. M. tasmaniae in Australia and New Zealand might play a simi- ASPÖCK U. (1992): Crucial points in the phylogeny of the Neuroptera (Insecta). pp. 63-73 in CANARD lar role, and other parallels may be sought in M., ASPÖCK H. & M.W. MANSELL (Eds), Current other places. research in Neuropterology, Toulouse. There would appear to be considerable BAR D. & D. GERUNG (1985): Cannibalism in Chryso- potential to manipulate other species of lace- perla camea (STEPHENS) (Neuroptera, Chrysopi- dae). — Israel J. Entomol. 19: 13-22. wings to increase their values as inoculative BERGESON E. & F.J. MESSINA (1997): Resource- versus control agents in a broad variety of pest con- enemy-mediated interactions between cereal trol contexts, and to improve the selection of aphids (Homoptera: Aphididae) on a common species for use in a wide variety of habitats and host plant. — Ann. Entomol. Soc. Amer. 90: 425- climate regimes. 432. BIGLER F. (1984): Biological control by chrysopids: integration with pesticides, pp. 233-245. In: CANARD M., SEMERIA Y. 8 T.R. NEW (Eds), Biology of the Chrysopidae. W. Junk, Dordrecht.

BIGLER F. & M. WALDBURGER (1994): Effects of pesticides

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on Chrysoperla carnea STEPH. (Neuroptera, DAANE K.M., YOKOTA G.Y., ZHENG Y. & K.S. HAGEN Chrysopidae) in the laboratory and semi-field. (1996): Inundative releases of common green — IOBC Bulletin 17:55-69. lacewings (Neuroptera: Chrysopidae) to sup- press Erythroneura variabilis and £. elegantula BROOKS SJ. & P.C. BARNARD (1990): The green lace- (Homoptera: Cicadellidae) in vineyards. — Envi- wings of the world: a generic revision (Neurop- ron. Entomol. 25: 1224-1234. tera: Chrysopidae). — Bull. Brit. Mus. (Nat. Hist.), Entomology Ser. 59: 117-286. DAANE K.M. & G.Y. YOKOTA (1997): Release strategies affect survival and distribution of green lace- BURKE H.R. & D.F. MARTIN (1956): The biology of three wings (Neuroptera: Chrysopidae) in augmenta- chrysopid predators of the cotton aphid. — J. tion programs. — Environ. Entomol. 26: 455- Econ. Entomol. 49: 698-700. 464. CANARD M. (1970): Incidence de la valeur de diverses DUELU P. (1980): Preovipository migration flights in pucerons (Homoptera, Aphididae) sur le poten- the green lacewing Chrysopa carnea (Planipen- tial de multiplication de Chrysopa perla (L.) nia, Chrysopidae). — Behav. Ecol. Sociobiol. 7: (Neuroptera, Chrysopidae). — Ann. Zool.-Ecol. 239-246. anim. 2: 345-355. DUELU P. (1981): Is larval cannibalism in lacewings CANARD M. (1972): Biologie de la reproduction de adaptive? (Neuroptera: Chrysopidae). — Res. Chysopa perla (L.) (Neuroptera, Chrysopidae) en Popul. Ecology 23: 193-209. function de l'alimentation imaginale. — Ann. Zool.-Ecol. anim. 4: 213-227. DUELU P. (1996): The working group 'carnea-com- plex': report on activities, results and cooperati- CANARD M. & P. DUELU (1984): Predatory behaviour of ve projects, pp. 307-311. In: CANARD M., ASPÖCK larvae and cannibalism, pp. 92-100. In: CANARD H. & M.W. MANSELL (Eds), Pure and applied rese- M., SEMERIA Y. S T.R. NEW (Eds), Biology of arch in Neuropterology, Toulouse. Chrysopidae. W. Junk, Dordrecht. DUELU P. & M. OBRIST (1995): Comparing surface CANARD M., SEMERIA Y. & T.R. NEW (Eds) (1984): Biology activity and flight of predatory arthropods in a of Chrysopidae. W. Junk, Dordrecht. 5 km transect, pp. 283-293. In: TOFT S. & W. RIE- DEL (Eds), Arthropod natural enemies in arable CARVALHO C.F., CANARD M. & C. ALAUZET (1996): Com- parison of the fecundities of the Neotropical land. I. Acta Jutlandica 70. green lacewing Chrysoperla externa (HAGEN) FLESCHNER C.A. (1950): Studies on searching capacity and the West Palaearctic Chrysoperla mediter- of the larvae of three predators of the citrus red ranea (HÖLZEL) (Insecta: Neuroptera: Chrysopi- mite. — Hilgardia 20: 233-265. dae). pp. 103-107. In: CANARD M., ASPÖCK H. & FRAZER B.D. & N. GILBERT (1976): Coccinellids and M.W. MANSELL (Eds), Pure and applied research aphids: a quantitative study of the impact of in Neuropterology, Toulouse. adult ladybirds (Coleoptera: Coccinellidae) prey- CHANG G. C. (1996): Comparison of single versus mul- ing on field populations of pea aphids (Homop- tiple species of generalist predators for biologi- tera: Aphididae). — J. Entomol. Soc. Brit. Colum- cal control. — Environ. Entomol. 25: 207-212. bia 73: 33-56.

CHANG G.C. (1998): Chrysoperla plorabunda (Neurop- GILLESPIE R.G. & T.R. NEW (1998): Compatibility of con- tera: Chrysopidae) larvae feed disproportiona- servation and pest management strategies, pp. tely on thrips (Thysanoptera: Thripidae) in the 198-208. In: ZALUCKI M., DREW R.A.I. & G.C. WHITE field. — Canad. Entomol. 130: 549-550. (Eds), Pest management - future challenges, Brisbane. CHANG Y.-F., TAUBER M.J. S CA. TAUBER (1995): Storage of the mass-produced predator Chrysoperla car- GRAFTON-CARDWELL E.E. & M.A. HOY (1985a): Intraspe- nea (Neuroptera: Chrysopidae): influence of cific variability in response to pesticides in the photoperiod, temperature and diet. — Environ. common green lacewing, Chrysoperla carnea Entomol. 24: 1365-1374. (STEPHENS) (Neuroptera: Chrysopidae). — Hilgar- dia 53: 1-31. CLARK T.L. & F.J. MESSINA (1998): Foraging behavior of lacewing larvae (Neuroptera: Chrysopidae) on GRAFTON-CARDWELL E.E & M.A. HOY (1985b): Short- plants with divergent architecture. — J. Ins. term effects of permethrin and fenvalerate on Behavior 11:303-317. oviposition by Chrysoperla carnea (Neuroptera: Chrysopidae). — J. Econ. Entomol. 78: 955-959. COHEN A.C. (1998): Solid-to-liquid feeding: the insi- GRAFTON-CARDWELL E.E. & M.A. HOY (1986): Genetic de(s) story of extra-oral digestion in predaceous improvement of common green lacewing, Arthropoda. — Amer. Entomol. 44: 103-117. Chrysoperla carnea (Neuroptera; Chrysopidae): COHEN A.C. & L.K. SMrm (1998): A new concept in arti- selection for carbaryl resistance. — Environ. Ent- ficial diets for Chrysoperla carnea: the efficacy omol. 15: 1130-1136. of solid diets. — Biol. Control 13: 49-54. HAGEN K.S. (1950): Fecundity of Chrysopa californica COULSON J. R., SOPER R.S. & D.W. WILUAMS (Eds) (1991): as affected by synthetic foods. — J. Econ. Ento- Biological control quarantine: needs and proce- mol. 43: 101-104. dures. — Workshop report. USDA-ARS, Beltsvil- HAGEN K.S., GREANY P., SAWALL E.F. Jr. S R.L. TASSAN le, Maryland. (1976): Tryptophan in artificial honeydews as a CROFT B.A. (1990): Arthropod biological cotrol agents source of an attractant for adult Chrysoperla and insecticides. Wiley, New York. carnea. — Environ. Entomol. 5: 458-468.

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HAGEN K.S. S R.L. TASSAN (1966): The influence of pro- LUFF M.L. (1983): The potential of predators for pest tein hydrolysate of yeasts and chemically defi- control. — Agric, Ecosystems, Environment 10: ned diets upon the fecundity of Chrysopa car- 159-181. nea STEPHENS (Neuroptera). — Vesl. csl. zool. MANSELL M.W. & H. ASPÖCK (1990): Post symposium Spol. 30: 219-227. neuropterological excursions, pp. 287-298. In: HASSAN S.A. & A. GRONER (1977): Die Wirkung von MANSEU. M.W. & H. ASPÖCK (Eds), Advances in Kernpolyedern (Baculovirus sp.) aus Mamestra Neuropterology. — Department of Agricultural brassicae auf Trichogramma cacoeciae (Hymen- Development, Pretoria. optera, Trichogrammatidae) und Chrysopa car- MCEWEN P.K., JERVIS M.A. & N.A.C. KIDD (1994): Use of nea (Neuroptera, Chrysopidae). — Entomopha- sprayed L-tryptophan solution to concentrate ga 22: 281-288. numbers of the green lacewing Chrysoperia car- HATTINGH V. (1996): The use of insect growth regula- nea in olive tree canopy. — Ent. Exp. et Appl. tors - implications for IPM with citrus in sou- 70: 97-99. thern Africa as an example. — Entomophaga MCEWEN P.K., NEW T.R. & A.E. WHITTINGTON (Eds) 41: 513-518. (1999): Lacewings in the crop environment (in HATTINGH V. & B. TÄTE (1995): Effects of field-weathe- press).

red residues of insect growth regulators on MCEWEN P.K., SHUJA A. & L. SENIOR (1998): Conservati- some Coccinellidae (Coleoptera) of economic on of the common green lacewing (Chrysoperia importance as biocontrol agents. — Bull. Ent. carnea s.l.) (Neuroptera, Chrysopidae) to control Res. 85: 489-493. crop pests. — Acta Zool. Fenn. 209: 153-156.

HILBECK A., BAUMGARTNER M., FRIED P.M. S F. BIGLER MEINANDER M. (1972.): A revision of the family Coni- (1998): Effects of transgenic Bacillus thuringien- opterygidae (Planipennia). — Acta Zool. Fenn. sis corn-fed prey on mortality and development 136: 1-357. time of immature Chrysoperia carnea (Neurop- MEINANDER M. (1990): The Coniopterygidae (Neurop- tera: Chrysopidae). — Environ. Entomol. 27: tera, Planipennia). A checklist of the species of 480-487. the world, descriptions of new species and HORNE P.A., NEWT.R. & D. PAPACEK (1999): Preliminary other new data. — Acta Zool. Fenn. 189: 1-95. notes on Mallada signatus (Chrysopidae) as a MESSINA F.J., JONES T.A. & D.C. NIELSON (1995): Host predator in field crops in Australia. In: MCEWEN plant affects the interaction between the Russi- P.K, NEWT.R. & A.E. WHITTINGTON (Eds), Lacewings an wheat aphid and a generalist predator, in the crop environment (in press). Chrysoperia carnea. — J. Kansas Entomol. Soc. HOWARTH F.G. (1983): Classical biological control: 68: 313-319. panacea or Pandora's box? — Proc. Hawai'ian MESSINA F.J., JONES T.A. & D.C. NIELSON (1997): Host Entomol. Soc. 24: 239-244. plant effects on the efficacy of two predators HOWARTH F.G. (1991): Environmental aspects of classi- attacking Russian wheat aphids (Homoptera: cal biological control. — Ann. Rev. Entomol. 36: Aphididae). — Environ. Entomol. 26: 1398-1404. 485-509. MILBRAITH L.R., TAUBER M.J.S CA. TAUBER (1993): Prey HOY M.A. (1990): Pesticide resistance in arthropod specificity in Chrysopa: an interspecific compari- natural enemies: variability and selection res- son of larval feeding and defensive behavior. — ponses, pp. 203-236. In: ROUSCH R.T. S B.E. TABAS- Ecology 74: 1384-1393. HNIK (Eds), Pesticide resistance in arthropods. — MONSERRAT V.J. & F. MARIN (1996): Plant substrate spe- Chapman S Hall, London. cificity of Iberian Hemerobiidae (Insecta: Neur- HUNTER CD. (1997): Suppliers of beneficial organisms optera). — J. Nat. Hist. 30: 775-787. in North America. — California Environmental NAFUS D.R. (1994): Extinction, biological control and Protection Agency, Sacramento. insect conservation on islands, pp. 139-154. In: JOHNSON J.B. S K.S. HAGEN (1981): A neuropterous lar- GASTON K.J., NEW T.R. & M.J. SAMWAYS (Eds), Per- va uses an allomone to attack termites. — Natu- spectives on insect conservation. — Intercept, re 289: 506-507. Andover.

LETARDI A. (1994): Dati sulla distribuzione italiana di NEUENSCHWANDER P. (1975): Influence of temperature Megaloptera Sialidae, Raphidioptera Inocelli- on the immature stages of Hemerobius pacifi- idae e Planipennia Mantispidae, con particolare cus. — Environ. Entomol. 4: 215-220. riferimento all'ltalia centrale (Neuropteroidea). NEUENSCHWANDER P. & K.S. HAGEN (1980): Role of the — Boll. Soc. Entomol. Ital. 125: 199-210. predator Hemerobius pacificus in a non-insec- ticide treated artichoke field. — Environ. Ento- Lo VERDE G. S V.J. MONSERRAT (1997): Nuovi dati sui mol. 9: 492-495. Coniopterygidae Siciliani. — Naturalista Sicil. 21: 57-66. NEUENSCHWANDER P., HAGEN K.S. & R.F. SMITH (1975): Predation on aphids in California's alfalfa fields. LUCAS E. (1998): How do ladybirds {Coleomegilla — Hilgardia 43: 53-78. maculata /eng; [Coleoptera; Coccinellidae]) feed on green lacewing eggs {Chrysoperia rufilabris NEUMARK S. (1952): Chrysopa carnea STEPH. and its [Neuroptera: Chrysopidae])? — Canad. Ento- enemies in Israel. — llanoth, Agricultural Stati- mol. 130: 547-548. on Bull. 1: 1-127.

164 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

NEW T.R. (1975): The biology of Chrysopidae and PRINCIPI M.M. & M. CANARD (1984): Feeding habits, pp. Hemerobiidae (Neuroptera) with reference to 76-92. In: CANARD M., SEMERIA Y. & T.R. NEW (Eds), their usage as biocontrol agents: a review. — Biology of Chrysopidae. W. Junk, Dordrecht. Trans. R. Entomol. Soc. Lond. 127: 115-140. ROSENHEIM J.A., WILHOIT L.R. & CA. ARMER (1993): Influ- NEW T.R. (1984): Comparative biology of some Aus- ence of intraguild predation among generalist tralian Hemerobiidae. pp. 153-166. In: GEPP J., insect predators in the suppression of a herbi- ASPOCK H. & H. HOLZEL (Eds), Progress in world's vore population. — Oecologia 96: 439-449. Neuropterology, Graz. RIUDAVETS J. (1995): Predators of Frankliniella occi- NEW T.R. (1986): A review of the biology of Neurop- dentalis and Thrips tabaci. —Wageningen Agri- tera Planipennia. — Neur. int.. Supplement I: 1- cultural University Papers 95-1: 43-87. 57. ROUSCH R.T. (1995): Genetic variation in natural ene- NEW T.R. (1988): Neuroptera. pp. 249-258. In: MINKS mies: critical issues for colonization in biological A.K. & P. HARREWIJN (Eds), Aphids. Their biology, control, pp. 263-288. In: MACKAUER M., EHLER L.E. natural enemies and control. Vol 2B. — Elsevier, & J. ROLANDE (Eds), Critical issues in biological Amsterdam. control. — Intercept, Andover.

NEW T.R. (1991): Insects as predators. — New South RUMPF S., HETZEL F. & C. FRAMPTON (1997): Lacewings Wales University Press, Kensington. (Neuroptera: Hemerobiidae and Chrysopidae) NEW T.R. (1999): Introduction to the systematics and and integrated pest management: enzyme distribution of Coniopterygidae, Hemerobiidae activity as biomarker of sublethal insecticide and Chrysopidae. In: MCEWEN P.K., NEW T.R. & exposure. — J. Econ. Entomol. 90: 102-108. A.E. WHITTTINGTON (Eds), Lacewings in the crop RUMPF S., FRAMPTON C. S D.R. DIETRICH (1998): Effects of environment, (in press) conventional insecticides and insect growth NORDLUND D.A. (1996): Biological control, integrated regulators on fecundity and other life-table pest management and conceptual models. — parameters of Micromus tasmaniae (Neurop- Biocontrol News and Information 17: 35-44. tera: Hemerobiidae). — J. Econ. Entomol. 91: 34-40. NORDLUND D.A. & R.K. MORRISON (1992): Mass rearing of Chrysoperla species, pp.427-439. In: ANDERSON RUZICKA Z. (1994): Oviposition-deterring pheromone T.E. & N.C. LEPPLA (Eds), Advances in insect rea- in Chrysopa oculata (Neuroptera: Chrysopidae). ring for research and pest management. — — Eur. J. Entomol. 91: 361-370. Westview Press, Boulder. RUZICKA Z. (1997a): Persistence of the oviposition- OBRYCKI J.J., LEWIS L.C. & D.B. ORR (1997): Augmenta- deterring pheromone in Chrysopa oculata tive releases of entomophagous species in (Neur: Chrysopidae). — Entomophaga 42: 107- annual cropping systems. — Biol. Control 10: 112. 30-36. RUZICKA Z. (1997b): Recognition of oviposition-deter- O'NEILL R.J., GILES K.L., OBRYCKI J.J., MAHR D.L., LEGASPI ring allomones by aphidophagous predators J.C. & K. KATOVICH (1998): Evaluation of the qua- (Neuroptera: Chrysopidae, Coleoptera: Cocci- lity of four commercially available natural ene- nellidae). — Eur. J. Entomol. 94: 431-434. mies. — Biol. Control. 11: 1-8. OSWALD J.D. (1993): Revision and cladistic analysis of RUZICKA Z. (1998): Oviposition deterring semiochemi- the world genera of the family Hemerobiidae cals in insect predators, pp 461-462. In: BRUNN- (Insecta: Neuroptera). — J. New York Entomol. HOFER V. & T. SOLDÄN (Eds), Book of Abstracts, Soc. 101: 143-299. Vlth European Congress of Entomology, Ceske Budejovice. PANTALEONI R. (1990): Un nuovo ausilario nel vigneto: Parainocellia bicolor (COSTA). — Inform. Fitopat. RUZICKA Z. & J. HAVELKA (1998): Effects of oviposition- 40: 39-43. deterring pheromones and allomones on Aphi- dolectes aphidimyza (Diptera: Cecidomyiidae). PARRELLA M.P., HEINZ K.M. & L. NUNEY (1992): Biological — Eur. J. Entomol. 95: 211-216. control through augmentative releases of natu-

ral enemies: a strategy whose time has come. — SALAMA H.S., ZAKI F.N. S A.F. SHARABY (1982): Effect of Amer. Entomol. 38: 172-179. Bacillus thuringiensis BERL. on parasites and pre-

PENNY N.D., ADAMS P.A. S LA. STANGE (1997): Species dators of the cotton leafworm Spodoptera lit- catalog of the Neuroptera, Megaloptera and toralis (BOISD.). — Z. Angew. Entomol. 94: 498- Raphidioptera of America north of Mexico. — 504. Proc. Calif. Acad. Sei. 50: 39-114. SAMWAYS M.J. (1988): Classical biological control and Pous G.A., MYERS C.A. & R.D. HOLT (1989): The ecolo- insect conservation: are they compatible? — gy and evolution of intraguild predation, Environ. Conservation 15: 348-354. potential competitors that eat each other. — Ann. Rev. Ecol. Syst. 20: 297-330. SAMWAYS M.J. (1994): Insect conservation biology. — Chapman & Hall, London. PREE D.J., ARCHIBALD D.E. a R.K. MORRISON (1989): Resi- stance to insecticides in the common green lace- SANDS D.P.A. (1997): The 'safety' of biological control wing Chrysoperla carnea (Neuroptera: Chrysopi- agents: assessing their impact on beneficial and dae) in southern Ontario. — J. Econ. Entomol. other non-target hosts. — Mem. Mus. Victoria 82: 29-34. 56:611-615.

165 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

SCHULTZ B.B. (1988): Reduced oviposition by green THIEUNG K.M. & B.A. CROFT (1988): Pesticide side effec- lacewings (Neuroptera: Chrysopidae) on cotton ts on arthropod natural enemies: a database intercropped with corn, beans or weeds in Nica- survey. — Agric. Environ. 21: 191-218. ragua. — Environ. Entomol. 17: 229-232. THIERRY D. & P.A. ADAMS (1992): Round table discussi- SCOPES N.E.A. (1969): The potential of Chrysopa car- on on the Chrysoperla carnea (STEPHENS) complex nea as a biological control agent of Myzus per- (Insecta: Neuroptera: Chrysopidae). pp. 367-373. sicae on glasshouse chrysanthemums. — Ann. In: CANARD M., ASPÖOC H. & M.W. MANSELL (Eds). Appl. Biol. 64: 433-439. Current research in Neuropterology, Toulouse.

SENIOR L.J., MCEWEN P.K. & N.A.C. KIDD (1998): Effects THIERRY D., CLOUPEAU R., JARRY M. & M. CANARD (1998): of the chitin synthesis inhibitor triflumuron on Discrimination of the West Palaearctic Chryso- the green lacewing Chrysoperla carnea (STE- perla STEINMANN species of the carnea STEPHENS PHENS) (Neuroptera, Chrysopidae): influence on group by means of claw morphology (Neurop- adult potentialities and offspring. — Acta Zool. tera: Chrysopidae). — Acta Zool. Fenn. 209: 255- Fenn. 209:227-231. 262.

SEWIFY G.H. & A. EL ARNAOUTY (1998): The effect of the TILLYARD R.J. (1919): Studies in Australian Neuroptera. entomopathogenic fungus Verticillium lecanii No. 6. The family Psychopsidae, with descripti- (ZIMM.) Viegas on mature larvae of Chrysoperla ons of new genera and species. — Proc. Linn. carnea STEPHENS (Neuroptera, Chrysopidae) in Soc. New South Wales 43: 750-786. the laboratory. — Acta Zool. Fenn. 209: 233- TUUSALO U. (1984): Mass rearing techniques, pp. 213- 237. 220. In: CANARD M., SEMERIA Y. & T.R. NEW (Eds), SHUJA A., SENIOR L.J., MCEWEN P.K. & P.A. LANGLEY Biology of Chrysopidae. — W. Junk, Dordrecht. (1998): Effects of the chitin synthesis inhibitor, TREMBLAY E. (1980): Su di un caso di predatismo da triflumuron, on the common green lacewing parte di Chrysopa formosa BR. (Neuroptera) a Chrysoperla carnea (STEPHENS): influence of resi- carico di imenotteri braconidi endoparassiti di dence time. — Acta Zool. Fenn. 209: 239-242. afidi. — Boll. Soc. Entomol. Ital. 112: 189-191.

SIMBERLOFF D. & P. STILING (1996): How risky is biologi- VAN EMDEN H.F. (1966): The effectiveness of aphido- cal control? — Ecology 77: 1965-1974. phagous insects in reducing aphid populations, STELZL M & M.A. HASSAN (1992): Über die Zucht von pp. 227-235. In: HODEK I. (Ed.), Ecology of aphi- Micromus angulatus STEPH. (Neuropteroidea, dophagous insects, Prague. Hemerobiidae), einer neuen Nützlingsart zur VAN EMDEN H.F. S K.S. HAGEN (1976): Olfactory reac- Bekämpfung von weichhäutigen Schadarthro- tions of the green lacewing Chrysopa carnea to poden in Gewächshäusern. —J. Appl. Entomol. tryptophan and certain breakdown products. — 114: 32-37. Environ. Entomol. 5: 469-473.

SUNDBY R.A. (1966): A comparative study of the effi- VENTURA M.A., RIBEIRO C. & V. GARCIA (1996): Suscepti- ciency of three predatory insects Coccinella sep- bility of third instar larvae of the green lace- tempunctata L. (Coleoptera, Coccinellidae), wing Chrysoperla kolthoffi (NAVAS) to the ento- Chrysopa carnea STEPHENS (Neuroptera, Chryso- mopathogenic fungus Metarhizium anisopliae pidae) and Syrphus ribesii L. (Diptera, Syrphi- (METSCHNIKOFF) Sorokin var. anisopliae TULLOCH in dae) at two temperatures. — Entomophaga 11: the laboratory (Insecta: Neuroptera: Chrysopi- 395-404. dae). pp. 241-249. In: CANARD M., ASPOCK H. S SUNDERLAND K.D., BlLDE T., DEN NlJYS L.J.M.F., DlNTER A., M.W. MANSELL (Eds), Pure and applied research in Neuropterology, Toulouse. HEIMBACH U., LYS J.A., POWELL W. & S. TOFT (1996): Reproduction of beneficial predators and parsi- VOGT H. (1994): Effects of pesticides on Chrysoperla toids in agroecosystems in relation to habitat carnea STEPH. (Neuroptera, Chrysopidae) in the quality and food availability. In: Boou C.J.H. & field and comparison with laboratory and semi- L.J.M.F. DEN NIJYS (Eds), Athropod natural ene- field results. — IO8C Bulletin 17: 71-82. mies in arable land. II. — Acta Jutlandica 71: pp. 117-153. WAAGE J.K. (1992): Biological control in the year 2000. pp. 329-340. In: KADIR A.A.S.A. & H.S. BAR- TAUBER CA. & P.A. ADAMS (1990): Systematics of the LOW (Eds), Pest management and the environ- Neuropteroidea: present status and future ment in 2000. — CAB International, Walling- needs. — Va Polytech. Inst. State Univ. Coll. ford. Agric. Life Sei. Inf. Ser. 90: 151-164. WILKINSON J.D., BIEVER K.D., IGNOFFO CM., PONS W.J., TAUBER CA. & M.J. TAUBER (1987): Food specificity in MORRISON R.K. & R.S. SEAYS (1978): Evaluation of predacious insects: a comparative ecophysiolo- diflubenzuron formulations on selected insect gical and genetic study. — Evol. Ecol. 1: 175-186. parasitoids and predators. — J. Georgia Ento-

TAUBER M.J., ALBUQUERQUE G.S. S CA. TAUBER (1997a): mol. Soc. 13: 227-236. Address of the author: Storage of non-diapausing Chrysoperla extema adults: influence on survival and reproduction. Dr Timothy R. NEW, — Biol. Control 10: 68-72.

Department of Zoology, La Trobe TAUBER M.J., TAUBER CA. & J.I. LOPEZ-ARROYO (1997b): University, Life-history variation in Chrysoperla carnea: implications for rearing and storing a Mexican Bundoora, Victoria 3083, Australia population. — Biol. Control 8: 185-190.

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