Annu1989 (34) 211-229 Brassica Crops

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Ann. Rev. Entomol. 1989. 34:211-29

ENTOMOLOGYOF OILSEEDBRASSICACROPS1Robert J. LambAgriculture CanadaResearch Station, Winnipeg,ManitobaR3T2M9,CanadaINTRODUCTIONHumans ave used vegetable oil pressed from the seeds of the Brassica plantsknown s rape for thousands of years (29, 95), but only during the lastyears have oilseed Brassica crops become nternationally important. Thesmall spherical seeds are harvested and crushed to separate the oil, whichmakes up approximately 40%of the seed by weight, from the remainingmeal. Theoil has been used as a lamp uel, as a lubricant, and in the chemicalindustry, but now s most often used in cookingor the production of food. Themeal contains approximately 40%protein by weight. It has been used as anorganic fertilizer, but now s most often blended in animal feed. Sales of thisbyproduct contribute substantially to the value of the crop.-The expandeduse of the crop, and the consequent ncrease in production,occurred whenplant breeders transformed he chemistry of the seed. First thefatty acid compositionof the oil was modified to reduce the level of erucicacid, and then the levels of glucosinolates in the meal were reduced. Erucicacid was a potential health hazard for humansconsuming the oil (28).Glucosinolates n the meal were repellent or toxic for some arm animals (97).The transformed crop is commonlyeferred to in Europe as 00 rape, signify-ing the goal of reducing he levels of erucic acid and glucosinolates n the seedto zero. In Canada it is knownas Canola. Knowledge f the effects onherbivory of secondaryplant compoundsuch as the glucosinolates is criticalto our understandingof herbivore-plant interactions (54, 81, 11 l). Glucosino-lates are antifeedants for manypolyphagous erbivores, but most insect pests

IThe author is employed y the CanadianFederal Government, nd by reason of this fact, thecopyright coveting this paper is ownedby the Crown n tight of Canada.

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212 LAMBof Brassicacrops use glucosinolates or their fission products as attractants orfeeding stimulants (32).The dramatic change that occurred in the secondary plant compounds foilseed Brassica crops interests entomologists and chemicalecologists. Also,the enormous ncrease in production offered a new environment or insects,which provided entomologists with the opportunity to determine the ecologi-cal characteristics of a new insect community.Nevertheless, entomologistsare interested in oilseed rape primarily because they need to control the manyinsect pests that competewith humans or the crop. The genusBrassica is inthe family Cruciferae, and many f the pests that attack the oilseed Brassicacrops also attack other cruciferous vegetables such as cabbageand radish. Thecommon ests of crucifers were described by Bonnemaison(12) beforeoilseed rape became an important crop in Europe and North America.Rape is an agricultural analog of soybeans, although the two herbaceouscrops are in different plant families. Both produceseeds that yield vegetableoil and high-protein meal, and these competeon international markets. Bothare grownover large areas in North America, Europe, and other parts of theworld, although the oilseed Brassica crops grow better than soybeans incooler, shorter-season climates and less well at higher temperatures. Theplants differ in architecture, but both are sown n dense stands and grow o asimilar height. Both crops harbor complex nsect communities. In general,insect communities n crops have received less attention than insect com-munities in natural habitats (111). As the entomology f the oilseed Brassicacrops becomes better known, a comparison of the insect communityof rapewith the more thoroughly studied soybean community (58, 123) willinstructive.This review emphasizes he relationships between nsect pests and the cropand the agricultural impactof pests on the crop. Thus, it focuses on the plant,its growth, responses to damage 80), and the way nsects interact with theplant to affect the agricultural commodity, apeseed.The Taxonomy and Origins of the Oilseed Brassica CropsAlthoughagriculture and the food industries treat rapeseed as a single com-modity, the crop is a compositeof seed from two or three species (29, 95).Asia, Brassica campestris and Brassica juncea are widely grown, but inEurope, Canada, NewZealand, and Australia, Brassica napusand to a lesserextent B. campestris are grown. These species are closely related. Brassicajuncea probably arose as a natural hybrid betweenB. campestris and Brassicanigra, and B. napusarose as a natural hybrid ofB. campestris and Brassicaoleracea. The latter species is the source of most of our cruciferous vegeta-bles. Brassica carnpestris is native throughout Europe, central Asia, and theNear East and was probably domesticated as a source of oil in central Asia or




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OILSEED BRASSICA ENTOMOLOGY 213adjacent northwestern India. Brassica napus does not occur in wild pop-ulations and was probably domesticated in southern Europe. The speciesdiffer morphologically nd chemically, and they growat different rates; thesedifferences complicate he study of pests associated with the crops. Further-more, the annual forms of B. napusand B. campestris are sown n the springin most of Canada nd in northern Europe, but in central and southern Europewinter-dormant, biennial forms are sown in the late summer 29). Thediffering phenologies of the two forms and the three species affect thesynchronies of insect life histories with the crop.THE PESTSThe most serious pests of the oilseed Brassicacrops are members f the orderColeoptera, particularly in Europe and North America (Table 1). On theIndian subcontinent and in NewZealand aphids cause severe damage(67,108), although in India pests from other orders are also important (10).Asia, insecticides are applied to control aphids and the diamondbackmoth,Plutella xylostella, in oilseed Brassica rops (72), but relatively little has beenpublished on their pest status. Similarly, little has been publishedon the pestsof oilseed Brassicacrops in South America,Australia, and Africa. Previouslyunrecognizedpests are still being identified, For example, ecent research onLygus bugs in Canada ndicates that these bud and seed feeders may causesignificant losses, although he subtle nature of the damage as caused t to beoverlooked (R. A. Butts & R. J. Lamb, unpublished data).Many f the important pests are as cosmopolitan s the crucifers themselves(Table 1). They maynot be important pests wherever they occur, however.The flea beetle Phyllotreta cruciferae is not usually a significant pest inEurope (7), although it is the dominant lea beetle pest of oilseed Brassicacrops in Canada (19) and in India (10). Root maggots, Delia spp., aresignificant pests of rape only in parts of Alberta, Canada 47) and in Finland(118), althoughDelia radicum s a widespread est of other crucifers through-out most of Europe and North America (12).In Europe and Asia, where the use of cruciferous crops is an ancientpractice, most of the insect pests of oilseed Brassica crops are cruciferspecialists. Some f the pests in North America, such as flea beetles, rootmaggots, and diamondbackmoth, are also crucifer specialists; these pestswere introduced from Europe or Asia and becamepests of cruciferous vegeta-bles before oilseed Brassica crops were introduced. In North America, nativepests that are not crucifer specialists have also adapted quickly to the newcrop, as has one native crucifer specialist. The noctuid Mamestraonfiguratawas the first pest to devastate the crop over a wide area in Canada 129).had previously been knownonly as an occasional pest of noncruciferous




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214 LAMB

Tal~|e ! Important pests of oilseed Brassica cropsPlant Plantorgan form

Pesl attacked attacked Area Ref.HenfipteraLygus elisusLygus lineolarisBagrada cruciferarumHomopteraBrevicoryne brassicaeLipaphis erysimi

ColeopteraMeligethes aeneusPhyllotreta cruciferaePhyllotreta striolataPhyllotreta undulataPsylliodes chrysocephalaEntomoscelis americanaCeutorhynchus assimilisCeutorhynchus quadridens

LepidopteraPieris brassicaeDiacrisia obliquaMammestra onfigurataPlutella xylostella

DipteraDasineura brassicaePhytomyzahorticolaDelia radicum

HymenopteraAthalia proxima

Bud, flower, pod, Annual Canada aseed

Bud, flower, pod, Annual Canada aseedStem, leaf, pod Annual India 10

Leaf, stem, bud, Annual, biennial NZ/Australia 67flower, podLeaf, stem, bud, Annual India 107, 108flower, pod

Flower Annual, biennial Europe 90, 122Leaf, stem, pod, root Annual Canada, India 10, 19, 65Leaf, stem, pod, root Annual Canada, Europe 7, 19Leaf, stem Annual Europe 7, 118Stem, leaf Biennial Europe 5, 112Leaf Annual Canada 41, 42Pod Annual, biennial Europe 37, 44Stem Annual, biennial Europe 45, 98,138

Leaf Annual India 10Leaf Annual India 10Leaf, pod Annual Canada 1~-, 129Leaf Annual India, Canada, 10, 72, 93NZ/Australia

Pod Annual, biennial Europe 44, 112,136Leaf Annual India 10, 101Root Annual, biennial Canada, Europe 47, 73,118

Leaf Annt~al India 10, 53aR. A. Butts & R. J. Lamb, npublished ata.bp. xylostella = P. maculipennis.Also dentified as Phytomyzatricornis.


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OILSEED BRASSICA ENTOMOLOGY 215crops (124). Similarly, Lygusbugs have a wide host range in North America(39, 109); although they are associated primarily with noncruciferous crops,they nowalso attack Canola (R. A. Butts & R. J. Lamb,unpublished data).Another native pest, the chrysomelid beetle Entomoscelis americana, ex-panded its host range from native crucifers to cruciferous vegetables andsubsequently to Canola (124).Pest Damage in Relation to Crop Growth and StructureInsects attack the roots, stems, leaves, flower beds, flowers, pods, and seedsof oilseed Brassica plants (Table 1). The impact on yield of the various kindsof damage s best understood in the context of plant growth and the com-ponents of yield.PLANTGROWTHilseed Brassica crops have small seeds (2.0--5.5 g per1000 seeds), which are usually sownat a depth of 2-3 cm (29). A paircotyledons emerges4-7 days after seeding. The seedling quickly produces hefirst pair of true leaves and then a rosette of broader leaves. At this stage, abiennial crop enters dormancy ntil spring. The annual crop continues to thenext growthstage, whichconsists of rapid elongation of the stem, often to aheight of 1 m or more 116). As the stem elongates, flower buds are producedfirst in a raceme t the tip of the main tem and subsequentlyon side branches,or axillary racemes. The pale-yellow, four-petaled flowers soon open, be-ginning at the base of each raceme 78). When flower is fertilized, its petalsdrop and a cylindrical pod (silique) forms. Each mature pod contains 15-40spherical seeds, which vary in color from yellow to dark browndependingonthe cultivar. Aspods form, the leaves begin to senesce, first at the base of theplant. Podson different parts of a plant matureat different times, so the ripestmayopen and drop their seed prior to harvest. The duration of the growthperiod maybe as short as 70-80days for B. campestris, but this period variesconsiderably depending on the species, cultivar, growth lbrm, and environ-ment (25, 56, 79). Harper & Berkencamp48) have provided a numericalfor the growth stages of B. napus.The yield of the crop depends on plant density, the number of podsproducedper plant, the numberof seeds per pod, and the average weight of aseed.PLANTDENSITY ilseed Brassica crops effectively compensate in seedyield for variation in plant density (40, 79). Reductions n plant density from200 o 40 plants per mz result in a loss in yield of only 20% r less (79). Plantsin sparse populations grow larger and produce more pods and more seeds perplant than densely seeded crops. Thus, pests that kill seedlings, such as theflea beetles, maynot have as great an impacton yield as the plant mortality in




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216 LAMBa damaged ield indicates. The yield loss resulting from seedling mortalitydependson the proportion of the seedlings killed and also on the distributionof mortality. Damageo surviving seedlings, however, mayhave a greaterimpact on yield than the loss of seedlings. For example,an insecticide appliedfor flea beetle control, which permits a higher seedling survival, results inlower yields than one that is less effective at preventingmortality but protectsthe surviving seedlings for a longer time (62).ROOTSMinor insect damage o roots that does not interfere with theirtransport and anchoring functions maynot be important. In Canada, larvae ofPhyllotreta spp. are abundant oot feeders (19), yet only about 5%yield lossresults from densities of 1600 larvae per m2 (17). Root maggots,Delia spp.,commonlyunnel the surface of rape taproots, sometimessevering the rootand killing plants (73), but the impact of these pests on yield has not beenquantified. Fusarium ungi invade the tunnels and maybe a factor in plantmortality (46).STEMStems provide an important photosynthetic surface, especially whenseeds are developing (99), in addition to supporting leaves and transportingnutrients. Danmgeo leaf petioles and stems by Psylliodes chrysocephalareduces yield (30), but the nature of the damagen relation to the functionsstems has not been described. One consequence of damage o stems by P.chrysocephalas higher than normal evels of winter plant mortality in fieldswith heavy infestations (18).LEAVEShe photosynthetic activity of leaves produces muchof the assimi-late neededby oilseed Brassica plants until they flower. Later, the stems andthen the pods gradually replace the leaves as the important photosyntheticorgans (99). Thus, defoliation at the beginning of flowering may beimportant source of loss, but leaf loss becomes ess significant as the cropmatures (26, 94, 117). In Canada, the bertha armyworm,M. configurata, isthe major defoliating pest. It prefers leaves to stems or pods and growsmorerapidly and larger when fed leaves (13). However, larvae consume94%their food in the last two instars (8), which are reached at the end of theflowering period (15). By this stage, leaf feeding by larvae has little impactyield; however, he larvae also attack pods, which does result in yield loss(13).

BUDSANDFLOWERSecause oilseed Brassica plants may abort up to50-60% f their flower buds and flowers, they can compensate or damage othese organs by retaining buds or flowers that wouldotherwise be aborted (78,115, 135)..Low or moderate levels of damage o buds and flowers caused by




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OILSEED BRASSICA ENTOMOLOGY 217pollen beetles, Meligethesspp. (90, 113, 115, 122, 137), E. americana 41),and Lygus bugs (R. A. Butts & R. J. Lamb, unpublished data) have littleimpact on yield. In fact, such damagemaybenefit the plant by reducingshading of the photosynthetic surfaces by flowers (117). Although suchbenefit has not been confirmed, low levels of bud damage y Meligethes spp.have been shown o increase yield slightly (90). There is a limit to the extentto which compensation an occur, however, and yield loss resulting from highlevels of Meligethes damage o buds and flowers is commonn Europe (90,113,122). Inability to compensateully is in part a result of damageo the tipsof the mainshoot and side branches rather than just a loss of buds (90). Thedamage robably also causes a loss of assimilates destined for seed produc-tion, because somedamaged lots yielded less than protected control plotsalthough the former had more pods (90).PODSDamage to young pods may be compensated in the same way asdamage o buds and flowers (135). However,as plants and pods mature andseeds develop, the plants capacity to compensate or pod loss declines (78,135). Furthermore, pods are important photosynthetic organs later in the lifeof the plant (99). Thus, loss of expandedpods as occurs when plants areattacked by bertha armyworman cause a substantial yield loss (14).SEEDSFeeding damage to developing seeds may be compensated for byreduced natural abortion of seeds (94) and possibly by an increase in the sizeof surviving seeds, although neither has been documented.Seeds are some-times larger in defoliated plants (26), but this type of compensation as notbeen observedconsistently (94, 135). Plants do appear to compensate or seeddamageby larvae of the weevil Ceutorhynchus assimilis and by the podmidge, Dasineurabrassicae, but through an increase in the numberof pods(115). However, his compensation maybe for loss of pods resulting frompod damage aused by adult weevils rather than for seed loss to larvae. Thelevel of compensation ften is not sufficient to prevent yield loss. Determin-ing the degree of compensation for seed loss is difficult because of theintimate relationship between seed growth and pod development (94). Fur-thermore, in Europe D. brassicae depends on pod damage by adult C.assimilis for oviposition sites (112).Damage hat kills plants or damage hat has immediate effects on thenumberof racemes, flowers, pods, or seeds is more easily quantified thandamage hat has lasting effects on the growth or developmentof the crop.Many f the pests that cause well-defined damage o particular plant organsalso have effects that cannot be explained in terms of the physiologicalcharacteristics of plant growth and components f yield. For example, theyield losses causedby flea beetles (62) are not all due to the seedling mortality




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218 LAMBthey cause, nor do the losses of buds and flowers by Meligethes spp. fullyaccount for the loss in yield (90). Similarly, aphid damage as a complexseries of effects on the plant that are not yet fully understood 67).

Froman agronomicpoint of view, the return to the producer depends notonly on the yield, but also on the harvestability and quality of the seed. Pestssuch as Phyllotreta spp. (17, 62), C. assimilis (121), and Meligethesspp. (90)affect the harvestability and quality of the seed by delaying the maturation ofdamaged lants and causing the crop to mature unevenly. Slowing maturationdelays harvest, which increases the probability of frost damagewhere thegrowingseason is short (62). Whenhe crop matures unevenly, seeds of themost mature plants are lost whenpods ripen, open, and drop their seed beforeharvest. Furthermore, the amountof chlorophyll in the harvested seed in-creases because he late-maturing plants are harvested before all their pods areripe, reducing the quality of the crop (14, 62, 90, 121). Insect damage analso reduce the oil content (90, 108, 121) and increase the glucosinolatecontent of seed (68).Brassica Chemistry in Relation to Insect PestsOilseed Brassica plants produce glucosinolates, a group of more than 70compoundsharacteristic of the Cruciferae and a few other plant families (57,97, 130). The glucosinolates are secondary plant compounds 81), andknow ittle of their metabolic function in the Cruciferae (97). Theyor theirfission products, however, mayfunction as antifungal agents (82), feedingdeterrents, or toxins for some nsect herbivores (11, 31) and as antifeedantstoxins for somewild vertebrate herbivores (103) and somefarm animals (97).The glucosinolates are hydrolyzed by myrosinases to yield volatileisothyiocyanates, particularly when plant tissue is damaged 130). Mostinsects that attack the Cruciferae are attracted, stimulated to feed, or stimu-lated to oviposit by allyl isothiocyanate or its parent glucosinolate, sinigrin(33, 34, 51, 77, 86, 119, 120). Although hey are attracted to and confinedcrucifers as hosts, some nsects find certain fission products of glucosinolatestoxic (2). Cruciferousplants probablyevolved glucosinolates and their fissionproducts as defensive chemicals; insects that specialize in attacking crucifersare thought to have evolved ways to minimize the toxic effects of thesechemicals and to have then adopted the glucosinolates as cues for theidentification of their hosts (32).Only a small proportion (approximately 12) of the known lucosinolatesoccur in the genus Brassica (50), but these mayproduce a complexarrayvolatile fission products (27, 59, 132). Interestingly, allyl glucosinolate,which yields the commonly sed insect attractant allyl isothiocyanate (23,34), is not found in B. napusand B. campestris (55, 59, 102). It occurs assignificant componentof the glucosinolates in B. juncea and B. oleracea(102). Nevertheless, many f the insects that attack oilseed mustard and the




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OILSEED BRASSICA ENTOMOLOGY 219cruciferous vegetables and that are attracted to allyl isothiocyanatealso attackB. napusand B. campestris (Table 1; 12). Presumablyother volatiles pro-duced by B. napusand B. campestris serve as attractants and feeding stimu-lants, although these have not yet been identified. The volatile cues producedby oilseed Brassicacrops and related crucifers are likely to be very complex.Not only are many volatile chemicals produced, but the composition ofvolatiles varies rapidly over time with the growthof the plant and to a lesserextent varies with the plant part assayed (27, 55, 59, 102). The analysisvolatiles emanating romBrassicaplants has not yet revealed the diversity ofcompoundsuggested by analysis of extracted plant tissues, but has clearlydemonstrated that the amount and probably the type of volatiles variesconsiderably between ntact and damaged oliage (35, 132). Becauseof theirdiversity, proven biological activity, and limited distribution in the plantkingdom, glucosinolates have been emphasized n research on the chemicalcues for the insects that attack crucifers, but other plant chemicalssuch asflavonol glycosides also stimulate feeding, and cucurbitacins and cardeno-lides are feeding inhibitors for someof these insects (85, 87, 88).Pests of oilseed Brassicacrops such as the aphid Myzuspersicae (84, 104),the bertha armyworm,M. configurata (129), and Lygus bugs (39, 109) arepolyphagous. When eeding on B. napus, M. persicae excretes high con-centrations of glucosinolates, whereas another aphid, Brevicoryne brassicae,which s a crucifer specialist, excretes lower concentrations of these chemi-cals and retains them n its body(133). Brevicorynebrassicae has a glucosi-nolase system, which hydrolyzes glucosinolates (75). The mechanisms hatallow other generalist herbivores that do not exhibit these adaptations toexploit the oilseed Brassica crops are not known. Sinigrin and ethylisothiocyanate are mild feeding deterrents for the polyphagousygus ineolar-is (49), but this species attacks Canola n Canada R. A. Butts & R. J. Lamb,unpublished data). Obviously, not all polyphagousherbivores are deterredfrom attacking oilseed Brassica plants by glucosinolates and their fissionproducts.The glucosinolates and their fission products are antifeedant or toxic fordomesticated animals such as fowl and hogs (97). The presence of glucosino-lates limited the utility of rapeseed mealuntil plant breeders produced ulti-vars with very low levels of the aliphatic glucosinolates in the meal 29). Thecultivars with low glucosinolate levels in the seeds also have lower levels ofthe aliphatic glucosinolates in the young oliage, although the indolyl gluco-sinolates are still present at higher levels (131). Changing he chemicalcomposition of oilseed Brassica crops could conceivably affect theirattractiveness, palatability, or toxicity to crucifer specialists or potentialpolyphagous pests. However, although virtually the whole crop area inCanada 28) and a large part of the crop area in Europehave been convertedlow-glucosinolate types (110), no changes in the insect pests have been




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220 LAMBreported that can be attributed to the introduction of the newcultivars. Instudies of individual pests, no consistent differences in the feeding responsesor damage evels have been detected for the low- and high-glucosinolatecultivars (1,43, 64, 71, 133). Althoughglucosinolate levels are nowat verylow levels in the seeds of somecultivars, levels in the foliage differ (102,131), whichmaypartly explain the continuedattraction of crucifer specialiststo the crop. Nevertheless, the low-glucosinolate seeds are no moresuitable asfood for polyphagous insects that attack seeds than seeds of the high-glucosinolate cultivars (74). One observed effect of the transition to low-glucosinolate cultivars has been an apparent increase in mortality of deer andhares that feed on rape foliage in Europe (103). The increased mortalitydeer results from increased palatability and hence increased consumptionoflow-glucosinolate rape foliage, which still has an unidentified toxic factor(131b).THE INSECT COMMUNITY IN OILSEED BRASSICACROPSThe insects living in oilseed Brassica crops represent a community hatinteracts with other agricultural and nonagricultural habitats. The most strik-ing feature of the rape habitat, particularly in Europe nd Canada, s the recentenormous ncrease in its size. In England n 1971 about 5000ha were plantedin the south-central part of the country, but by 1984 he crop covered 260,000ha spread over muchof the country (69, 134). In Canada n 1950 rape wasgrownon a few hundred hectares in Saskatchewan,but by 1979 a peak of 3.4million hectares was reached, spreading across the three prairie provinces(65). The explosion in the numberof Brassica plants has had a major impacton the insects associated with them. Before the oilseed Brassica crops becamethe dominant rucifers in these areas, the crucifer specialists fed on vegetablecrops, weeds, or scattered native plants. The density of weed and wildcrucifers in and aroundagricultural land is difficult to estimate, but the areadevoted to cruciferous vegetables is known. In England the area devoted torape is at least five times greater than that planted with Brassica vegetables(134), and in Canada he difference is muchgreater. Initially the oilseedBrassica crops probably formeda sink for cruciferous insects (134), but thedensities of pests have responded o the increasing food resource; thus pestproblems have becomemore and more serious on the oilseed Brassica cropsand have overflowedonto other cruciferous crops as well (69, 134). Further-more, the introduction of the oilseed Brassicacrops has changed he temporalavailability of cruciferous hosts (134), particularly in areas whereboth sum-mer and winter forms are grown n adjacent fields (89, 122).The spatial relationships between oilseed Brassica crops and adjacentcommunities ffects the pests of rape, but this subject has received little




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OILSEED BRASSICA ENTOMOLOGY 221attention. Some ests use host plants in habitats other than rape fields (21, 22,38) and overwinter in other habitats (20, 128), but the importanceof thesealternative habitats in the pest dynamics s not known.Rape ields are usuallymuch arger than those of vegetable crops, particularly in Canada, whereCanola fields of 100 ha are common. n small-scale experimental studies,crucifer monocultures upport higher densities of insect herbivores and suffermoredamage er unit area than small patches of crucifers or polycultures (6,100, 114). On he scale of commercial roduction, however, arger rape fieldslose less yield per unit area to flea beetles than smaller ones (65).Little is known bout the diversity and structure of the arthropod com-munities in rape, although many f the individual pests have been studied andmany f the predators and parasites of these insects have been identified. Thebiomassof arthropods was found to be lower in a winter rape field than in ameadow83). The densities of general predators such as spiders are lowrape fields because they must reinvade the large fields each season (131a).Some pecialized predators use the sameolfactory cues as their hosts to locatethe host habitat (96), and thus may e less affected by spatial characteristicsthe crop or temporalvariation in its availability. Interactions between erbi-vores are beginning to be identified. For example, as mentionedabove, D.brassicaeis dependenton pod damage y C. assimilis for oviposition sites (3,112).PEST MANAGEMENTWith he recent increase in the area devoted to the oilseed Brassicacrops inEurope and North America and the increasing importance of pests to cropproduction, much esearch has been conducted on methodsof reducing lossesin yield due to pests. Insecticides are applied to control the most importantpests: aphids (24, 67), M. configurata (129), Phyllotreta spp. (65, 118),chrysocephala4), Meligethes aeneus(90, 122), and C. assirnilis (121). Peststhat attack the early growth stages of the crop are usually controlled withsystemic insecticides applied as seed dressings or in-furrow granules (4, 65,67, 118). Pests that attack at the time of stem elongation or later require one(121, 129) or more 122) foliar sprays. In somecases pesticide applicationsfor one pest also reduce the damage aused by secondary pests (7). Pesticideresistance, pest resurgence, and outbreak~ of secondary pests have not yetbeen reported to result from insecticide applications for control in rape.

Phyllotreta spp. in Canadaand M. aeneus in parts of Europe can causesevere damage nnually over wide areas, and insecticides are applied routine-ly to control these pests. Other pests are more sporadic (66), and economicthresholds in conjunction with pest monitoring have been used to minimizethe use of insecticides (105). For example, n the control of M. configurata,an economichreshold for larvae was developed n caged field trials (14, 15)




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222 LAMBand then verified in commercial fields (16). An area-wide pheromone-monitoring system was devised to warn of pest outbreaks (125), and larvalsamplingmethodswere tested that mayenable farmers to assess pest densitiesefficiently in individual fields to determine if control is necessary (127).Economichresholds have also been defined for M. aeneus (90), C. assimilis(36, 121), and flea beetles in Finland (7). Pest monitoring and economicthresholds are not effective tools for flea beetle control in Canadabecause heinvasion of the newly emerged crops is sudden and weather dependent.Seedlingmortality can reach high levels before farmers are able to assess pestderisities or apply insecticides (61).

Well-planned nsecticide applications are economically ustified, but never-theless costly. A number of alternative control measures have been in-vestigated for the important pests of oilseed Brassicacrops, Trap cropping isan effective control for M. aeneusin Finland (52). In Canada, all cultivationcontributes to the control of E. americana(42) and M. configurata (126).Biological control of Phyllotreta spp. has been attempted in Canada(139).Host-plant resistance is being investigated for aphids in India (9, 76) and forPhyllotreta spp. in Canada(60, 63).CONCLUSIONSThe entomology f the oilseed Brassica crops is in an early stage of develop-ment (70). The recent enormousexpansion of the crop area has led to theestablishment of a new agricultural ecosystem with a novel pest complex.Although many of the pests of this crop are also well-known pests ofcruciferous vegetables, their status in rape was previously unknown. Aconsensus is only nowbeing reached on which insects are the primary pests.Furthermore, he drastic difference in scale and in value per unit area betweenrape and the cruciferous vegetable crops assures that the ecology of the pestsand the strategies adopted to control themwill be substantially different. Theoilseed Brassica crop has been qualitatively dynamicas it has becomees-tablished and has expanded. The three Brassica species vary spatially andtemporally in their dominance, s do the annual and biennial forms of the cropin certain parts of Europe. As the crop has become stablished there has beena major transformation in the composition of the secondary plant compoundsof the crop, which is still under way. This dynamiccrop has provided manychallenges and opportunities for entomological research.The future is sure to bring newentomological challenges as plant breedersand agronomists continue to transform the crop. At the moment, esearch isunder way to develop hybrid Canola (106). If a suitable commercialmethodof producing hybrid seed becomesavailable, the yield per unit area willincrease substantially and the economicsof pest control will be transformedaccordingly. Furthermore, the hybrid plants maygrow more vigorously and




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OILSEED BRASSICA ENTOMOLOGY 223may compensate better for insect damage. Research is also under way todevelop B. juncea as a Canola crop for North America. Production of B.juncea could extend the crop into the hotter and drier areas of the NorthAmerican prairies, a new habitat for many crucifer pests. The chemicalcharacteristics of the crop will continue to change as plant breeders tailorcultivars to the oilseed markets. The ability to modify the glucosinolatecontent of foliage through plant breeding opens up the possibility of develop-ing a rape crop lacking the attractants and feeding stimuli used by thecruciferous pests. However, muchmore needs to be learned about the orienta-tion cues and feeding stimuli used by these insects before this goal can beachieved. Mixed cropping systems of field peas and Canola are being tested inCanada because they are thought to have agronomic advantages and becausethe seed of the two crops can be readily separated. This practice is likely tohave a substantial effect on the pest complex n Canola if experinaental resultson pest densities in polycultures (6, 100, 114) are borne out.The insect pests of oilseed Brassica crops are sufficiently important thatcontinued research will be demanded to minimize the damage they cause.Because of the cropping area involved and the annual requirements for largeamountsof insecticides to control the pests, there will be a concerted effort todiscover alternative methods of control. Given the intimate relationship be-tween many of the pests and the host plants, and given our increasingknowledge of the secondary compounds of the Brassicas plants, the chemicalaspect of the insect-plant relationship will undoubtedly be the starting pointfor develbping alternative control measures.ACKNOWLEDGMENTSI thank C. E. Shearer for helping prepare a bibliography on the crop and itspests and for checking the manuscript. L. Burgess provided many of thereferences and advice as the project developed. R. Bodnaryk, P. Pachagoun-der, and W. J. Turnock provided useful comments on the manuscript.

This paper is Contribution No. 1317, Agriculture Canada Research Station,Winnipeg.Literature Cited

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OILSEED BRASSICA ENTOMOLOGY 229flower and other crucifers. Phytochernis-try 15:763-66133. Weber, G., Oswald, S., Z611ner, U.1986.Suitability of rape cultivars with adifferent glucosinolate content for Brevi-coryne brassicae (L.) and Myzus per-sicae (Sulzer) (Hemiptera, Aphididae).Z. pIlanzenkr. Pflanzenschutz 93:113-24 (In German)134. Wheatley, G. A., Finch, S. 1984.Effects of oilseed rape on the status ofinsect pests of vegetable brassicas. InPests and Diseases, Br. Crop Prot.Conf. Brighton, UK, 2:807-14.Croydon, UK: Br. Crop. Prot. Counc.135. Williams, I. H., Free, J. B. 1979. Com-pensation of oil-seed rape (Brassicanapus L.) plants after damage o theirbuds and pods. J. Agric. Sci. 92:53-59

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