Sara OppenheimIn 1959, with the whole world watching,Cubans committed themselves to a new socialorder. A population made up of peasants, laborers,and millionaires underwent a violent shifttoward a system that promised to eliminate hunger,educate the masses, and distribute resourcesequitably. From the beginning, the revolution wasmainly about agriculture. Dating back to Columbus’discovery of the island, Cubans had neverbeen truly autonomous in controlling their land.Spaniards, Americans, the British, and others imposedtheir vision on Cuba and created great richesfor themselves from Cuban sugarcane and Cubanlabor. When Castro rose to power, it was with thefull support of the farming classes, and his promiseto return the land to the people who worked itgalvanized the nation.Sadly, a great leader is not necessarily a greatmanager, and the early post-revolutionary yearswere characterized by chaotic efforts to do goodon all fronts. Although education, health care,and living conditions improved markedly, attemptsto restructure Cuba’s agricultural practices were afailure. Faced with an embargo that cut off sugarsales almost overnight, Castro searched for waysto feed the country. Perhaps inevitably, Cuba eventuallyentered into a reciprocal trade agreement withthe Soviet Union, and Soviet largesse made up forCastro’s excesses and failures in agricultural planning.Cuba supplied sugar to the Soviet bloc andin return received staple foods, petroleum, andmanufactured goods (Castellanos and Alvarez1996).To maintain high levels of sugar production forexport, Cuba adopted a developmentalist agriculturalstrategy similar to that seen in the UnitedStates. In this, Cuba had the full support of theUSSR where high-input, heavily mechanized growingpractices were well advanced. Using Russiangoods and expertise, Cuba’s agriculture was transformed:poor soils were rectified with inorganicfertilizers, crop pests controlled through intensepesticide application, and human laborers replacedby machines. Central-planning techniques wereadopted, and agricultural management movedfrom being a single farmer’s decision about whatto grow, to a huge, state-run enterprise. Increasedyields followed swiftly, and Cuba, although dependentupon the USSR, became the most prosperouscountry in Latin America. In fact, the World HealthOrganization’s Quality of Life Index for 1989(WHO 1989), based on such factors as literacy, lifeexpectancy, and caloric intake, placed Cuba eleventhin the world, with the United States in fifteenthplace.The perils of Cuba’s dependent position wererevealed in 1989 as the Soviet bloc fell into ruin.Within three years, imports of food from the USSRdropped by almost 50%, and the importation ofother goods was affected similarly (Castellanos andAlvarez 1996). The materials necessary to maintainCuba’s input-dependent agricultural systemwere no longer available, and the danger of starvationbecame a reality. Cuba was, therefore, facedwith a unique challenge: to simultaneously increasefood production and reduce or eliminate inputs.To do so, Cuba adopted as national policy an alternativeagricultural model, referred to in theUnited States as Low Input Sustainable Agriculture(USDA 1980).The goals of this system were both philosophicaland practical in nature. In an immediate sense,such a strategy was dictated by the scarcity of fertilizers,pesticides, and farm implements. Yet adeeper issue also was involved: along with many ofthe world’s scientists, Cuban researchers increasinglyhad become disillusioned with classical farmingmethods as practiced by the United States andthe USSR (Levins 1993). The use of inorganic inputswas perceived as poisoning the environment,and the continual escalation of chemical requirementswas regarded as both costly and dangerous.In response, Cuban scientists had for many yearsbeen exploring alternative agricultural techniques.They thus were well prepared to face the challengeof nationwide conversion to sustainable agricul-216 AMERICAN ENTOMOLOGIST • Winter 2001

ture. Since 1989, their commitment to making thealternative model a reality has yielded importantresults in five major areas:With the loss of almost allimported pesticides, Cuba’s crop protection strategynow depends on the effective production andemployment of natural enemies. Although sometechnologies have been imported, others are theresult of goal-oriented Cuban research. Extensivemonitoring of crop pests is now conducted, andtechniques for the evaluation, production, and deploymentof natural enemies have become standardpractice (Dlott et al. 1993).Monitoring forplant disease is carried out on more than 90% ofCuba’s crops, and disease diagnosis involves advancedtechnologies such as electron microscopyand immunofluorescence. In converting fromchemical methods of disease control, Cuba hasdeveloped a large research base in the utilization ofmicrobial antagonists against plant diseases andcurrently produces and employs several agents(Shishkoff 1993). Disease-tolerant cultivars arealso utilized.Weed growth is a commonstumbling block in alternative agricultural practices.In more developed countries, selective herbicideapplications are used, but in Cuba this optionis limited. Instead, Cuba has developed sophisticatedmodeling programs, allowing prediction oflikely weed species and densities. Based on modelpredictions, crop mixtures and crop rotations thatwill reduce the impact of these weeds are designed(Dlott et al. 1993). The development of new tillageimplements has integrated weed destruction withlow till farming practices.Cuba’s soils generally are ofpoor quality, and years of intensive, high inputfarming have further reduced their organic mattercontent. Salinization and erosion are widespreadand limit productivity. To address this, Cuba hascarried out an exhaustive survey of soil contentand quality in all regions, and measures to rectifysoil quality are underway (Gersper et al. 1993).These involve the increase of soil organic mattercontent through vermiculture and other biologicalfertilizers and the reduction of erosion by means oftillage reduction and reforestation. Although one of the mostimportant aspects of alternative agriculture is thereduction of mechanization, the input of humanlabor that this involves can be prohibitive. Toaddress the need for rural laborers, Cuba hasimplemented several strategies aimed at the reruralizationof its populace (Rosset and Benjamin1994). Short-term and long-term volunteer laborprograms are supported through financial andmoral incentives, and the reorganization of laboron state farms has helped form a stronger linkbetween workers and the land.HistoryThe history of Cuban agriculture prior to 1989in many ways mirrors worldwide trends. Followingthe developmentalist ideology described byLevins (1993), agricultural development has proceededalong a steady line from less developed tomore advanced production techniques. As in NorthAmerica and Europe, such progress has followedseveral major trends, including (1) the replacement oflabor-intensive production by capital-intensive production;(2) a change from small, multiple crop productionunits under individual management tolarge-scale monocultures under centralized or corporatecontrol; and (3) the replacement of local, growerbasedagricultural knowledge with the morespecialized, widely applied results of scientific research.In Cuba, this trend began some centuries ago.Pre-Colombian Cuba was populated by the agriculturalArawak Indians, whose techniques forgrowing maize, cassava, and other crops were,of necessity, low input. Their resource managementstrategies were quite sophisticated, utilizinglocally adapted crop assemblies to minimizecompetition for soil resources and water andplanting of peanuts and other nitrogen-fixinglegumes to enhance soil fertility (Rosset and Benjamin1994).As the Spanish colonized Cuba in the wake ofColumbus’ 1492 discovery of the island, they initiallyrelied on Arawak techniques and labor tosupply needed resources. Soon, however, the Spaniardsbegan to convert to plantation agriculturalpractices. Large plantations, relying on indigenousand imported slave labor, produced sugar and tobaccofor export to Europe. Forests were clearedto free more land for agriculture and animal production,with felled wood contributing to the shipbuildingand hardwood export trades. By the late1700s, sugar was firmly established as the dominantcrop in Cuba, a situation further intensifiedby the slave revolt and subsequent demise of sugarproduction in Haiti, at that time the world’s foremostsupplier of sugar (Baker 1997).Although the political situation changed radicallyin Cuba from 1800 to 1959, reliance on asugar monoculture did not. The vast profits availablefrom exported sugar encouraged an increas-The history of Cubanagriculture prior to 1989in many ways mirrorsworldwide trends.Following thedevelopmentalistideology describedby Levins (1993),agricultural developmenthas proceeded along asteady line from lessdeveloped to moreadvanced productiontechniques.Fields like this one in the Pinar del Rio region of Cuba produce some of theworld’s finest tobacco.AMERICAN ENTOMOLOGIST • Volume 47 Number 4 217

The diversification ofagriculture away fromreliance on sugar wasone of Castro’s firstgoals. By 1962,however, the effects ofthe U.S. embargo hadresulted in widespreadhunger, and rationingwas introduced.Faced with a starvingnation, Castroabandoned attempts atdiversification and turnedwith a new passion tosugar production.ingly narrow crop base, with food crops for internalconsumption taking a distant second place tosugar for exportation. U.S. holdings in Cuba werealmost entirely sugar-based, consisting of both theland on which sugarcane was grown and the facilitieswhere sugar was processed. By 1958, this intensiveproduction was yielding more than 6 milliontons of sugar per year.Along with large scale plantation-style cultivationcame great social inequality. 9% of farmersheld 73% of all agricultural lands, and the sugarcanemonoculture relied on peasant labor. Two particulartraits of the sugarcane system contributed tothe existence of a massive underclass. First, the productionof sugarcane relies on a large number oflaborers, especially at harvest time (prior to mechanization,some 350,000 workers were required toharvest Cuba’s sugarcane crop each year). And second,the long growing period between harvests leftthese laborers unemployed for 7 months of the year.Without a diverse array of crops to harvest, Cuba’sagricultural laborers were left without food, employment,and other necessary resources.The Cuban RevolutionThe existence of this vast and often hungry peasantclass, together with institutionalized political corruption,set the stage for Castro’s 1959 overthrowof the U.S.-supported Batista regime. Advocatingbottom-up social reforms, Castro won widespreadsupport among the lower classes, who supplied himwith food, arms, and volunteers during the 2 yearsof armed struggle that culminated in Batista’s expulsionfrom Cuba on 1 January 1959.Agrarian reform was the first item on Castro’sagenda. All land holdings greater than 67 hectareswere repatriated as state property, driving hugenumbers of upper-class landholders into exile.Further reforms allowed the state to seize controlof foreign-owned oil and sugar processing plantsand resulted in a retaliatory U.S. embargo on Cubansugar. In 1961, the Trade Embargo Act waspassed into law, banning all Cuban exports to theUnited States.The diversification of agriculture away fromreliance on sugar was one of Castro’s first goals.By 1962, however, the effects of the U.S. embargohad resulted in widespread hunger, and rationingwas introduced. Faced with a starving nation,Castro abandoned attempts at diversification andturned with a new passion to sugar production. Awarlike effort to increase the sugar harvest to itshighest level ever was begun. Tens of thousands ofworkers were diverted from their normal jobs andsent to the cane fields, and virtually all arable landwas turned over to sugarcane production. Althougha record harvest was achieved, the entirecountry was left in chaos (Castellanos and Alvarez1996). Clearly, sugar exports alone were not sufficientto feed the nation.Soviet AllianceThe reign of monoculture was strengthened in1968 when Castro, unable to provide food securityfor Cuba, joined the Council for Mutual EconomicAssistance (COMECON), the Soviet bloc’seconomic community. Cuba’s role in COMECONwas to supply sugar to European socialist nationsin exchange for which, the island’s import needswould be met. Thus, the revolution, whose goalhad been Cuban independence, simply paved theway for a new outside power to prop up Cuba’sunstable agricultural situation.The formalization of trade relations with Sovietbloc countries had far reaching implications forCuban agriculture. For all its ideological rejectionof capitalist culture, the Soviet government wascommitted to technological progress following aWestern model. Progress was measured by output,and methods that increased productivity wereadopted widely. The result, for both Cuba and theUSSR, was a highly industrialized farming system.In Cuba, some 75% of all agricultural lands weredevoted to state-run farms, whose main task wasthe production of sugar for export (Castellanosand Alvarez 1996).A Classical ModelThese large, state-run enterprises exhibited manyof the hallmarks of modern agriculture. Whereverpossible, human labor was replaced by mechanization,resulting in an increasingly urbanized populace.Fertilizer inputs were considerable, with thesoil viewed as an expendable resource. Monocultures,which are extremely vulnerable to insect pests,were protected by high levels of pesticide application,and required large-scale irrigation.Such procedures are part of what may be termedthe classical model (Vandermeer et al. 1993) of agriculturalproduction employed by most developedcountries. In the classical model, dependence onexternal sources of food is high. In Cuba’s case,this dependence extended to external sources ofalmost all agricultural inputs. Food consumed inone region generally was grown elsewhere, so efficienttransportation and refrigeration were essential.Reliance on imported technology was high,and involved both machinery and chemicals.Employing this classical model of production,Cuba was far from independent in its food supply.Imports of food, machinery, chemicals and animalfood were extensive. The results were good. Productivityclimbed as tractors replaced animal traction,mechanization replaced human labor, and useof chemical pesticides insulated growers from theeffects of insect damage and soil depletion. Thesetechnologies became fundamental to the Cubanagricultural system and were considered a sign ofits increasing sophistication (Castro himself viewedescalating pesticide use as a sign of Cuba’s progress[Vandermeer et al 1993]). By the late 1980s, 82%of agricultural pesticides used in Cuba were importedfrom the Soviet bloc, and 42% of its fertilizerswere from the same source (Deere 1992).Such a situation, although productive, was extremelyprecarious. Through its dependence onSoviet bloc imports and exports, Cuba’s productivitywas tied inextricably to the fate of the Soviet218 AMERICAN ENTOMOLOGIST • Winter 2001

Union. Some 84% of all Cuban trade was with theSoviet Union and Eastern Europe, and other formsof aid from these sources totaled more than US $3billion (Baker 1997). This dependence was intensifiedby the favorable terms of trade the Sovietsoffered Cuba. Sugar, Cuba’s main export, waspurchased at a fixed price, regardless of worldmarket fluctuations. By trading this sugar, whichcomprised 75% of all Cuban exports in 1988 (Pastor1992), Cuba was able to obtain the food, chemicals,and manufactured goods needed to sustainits populace.Fall of the USSRThe collapse of the Soviet bloc in 1989 dealt acatastrophic blow to Cuba’s economy. Importsfrom Eastern Europe fell by half from 1989 to1990, and by 1992 plunged to a third of theirformer value (Castellanos and Alvarez 1996).During the same time, the U.S. embargo againstCuba was strengthened through the Mack Amendment,passed in 1992, which made it illegal forforeign subsidiaries of U.S. companies to trade withCuba; shipment of any medicinal or food items toCuba was forbidden.These events precipitated a crisis that affectedalmost all levels of society. Cuba’s sophisticated,high-input agricultural system faced an 80% dropin pesticide and fertilizer availability, and access tofuels and irrigation also fell sharply. Machine partsfor transportation and agricultural devices becameunavailable, rendering many mechanized processesuseless. Imports of food for human consumptiondecreased quickly and extensively. Average caloricintake per individual declined by 30% from 1989levels, and nutritional deficiencies led to the outbreakof new diseases such as optical neuritis(Eckstein 1997). Cuba, the only country in LatinAmerica to have eradicated hunger, was faced withwidespread malnutrition and food shortage.The focal point of these trade losses was agriculture.“ The food question,” said Castro, “ hasnumber one priority “ (as quoted in Rosset andBenjamin 1994). Long dependent on outsidesources for the majority of its production and produceneeds, Cuba appeared ill equipped to dealwith this new crisis.A New ModelWithin Cuba, however, as in many countries, aquiet agricultural revolution had been taking place.Even as the classical model of agriculture was beingimplemented, a vocal minority was decrying itsdangers. The 1962 publication of Rachel Carson’sSilent Spring brought worldwide attention to thecosts of the classical model. The intensive use ofpesticides for pest control led to escalating cycles ofpest resistance, demanding new chemicals and largerdoses to achieve control. Pesticides were demonstratedto have significant and long-term healthrisks for humans and other organisms, causingwidespread fear about their presence in foods.Huge monocultures, designed to exploit economiesof scale, required constant inputs of fertilizers tocompensate for declining soil fertility and erosion,and the mechanized growing techniques used insuch monocultures put an entire class of rural laborersout of work.As such trends became widely acknowledged,an alternative model of agricultural productionemerged. Attempting to address the health, environmental,and social costs of the classical model,the alternative model called for a change in inputs,a reduction in scale, and new systems of production.By using organic and biological fertilizers,chemical fertilizer input would be reduced or eliminated;the introduction of natural enemies and biologicalpesticides would decrease reliance on chemicalpesticides; strategic crop mixes would help in weedcontrol and soil rectification; and seasonal rainfallpatterns would be exploited to reduce the need forirrigation. Additionally, the substitution of humanlabor for mechanization, and of animal traction fortractors, would significantly reduce fuel requirements,and help reestablish viable rural communities.This alternative model was given a boost in 1989when the National Academy of Sciences publishedits report on alternative agriculture, concluding thatavailable data gave no evidence that the classicalmodel offered higher production levels than thealternative model. Coupled with the noneconomicbenefits of the alternative model, such conclusionsgenerated increasing support for the alternativemodel. Its lower cost to growers and decreasedreliance on outside sources of inputs made the modelparticularly well suited to developing countries(Altieri and Hecht 1990), and the reduction inhealth-threatening chemical inputs led the UnitedStates Department of Agriculture to endorse lowinput sustainable agriculture (USDA 1980).The philosophical basis of the alternative modelgenerally is accepted throughout the developedworld, yet adoption of this production system isstill rare. Full conversion from conventional toalternative production methods can involve a profitlagof up to 5 years during which productivity andprofitability may suffer. This financial sacrifice,combined with the deeply entrenched interests ofpesticide and fertilizer producers, has hamperedthe comprehensive adoption of the alternativemodel, and conversion generally has been limitedto individual farms (Thrupp 1996).The Alternative Model in CubaAs the full implications of the Soviet bloc dissolutionwere realized, Cuba’s adoption of the alternativemodel became a matter of necessity ratherthan choice. A “special period in a time of peace”was declared during which Cuba’s primary goalwas to be the achievement of food security andeconomic independence. Castro called on agriculturalscientists to “produce more food without feedstock,fertilizers or fuel” (as quoted in Rosset andBenjamin 1994). The alternative model was declaredofficial government policy, and Cuba’s scientistswere expected to provide the neededbiological control, biological fertilizer, soil-rectification,and production technologies.AMERICAN ENTOMOLOGIST • Volume 47 Number 4 219

This daunting task was not as impossible as itseemed, for Cuba’s scientists have a long traditionof exploring alternative technologies (Vandermeeret al. 1993). The National Institute for Research inCitrus and Other Fruits, for example, had embarkedupon a 5-year plan to reduce pesticide inputswell before the advent of the special period.Biological control of citrus pests dates back to the1930s in Cuba. In 1930, for example, the introductionof Eretmocerus serius Silvestri, a parasitoidof Chinese origin, achieved control of the citrusblackfly, Aleurocanthus woglumi Ashby (DeBach1974). Further successes in the same year weregained through the release of Lixophaga diatraeae(Townsend), an endemic parasitoid fly, for the controlof the sugarcane borer, Diatraea saccharalis(F). (Scaramuzza 1930). Although these biologicalcontrol efforts were carried out under the auspicesof U.S. scientists, the 1960s witnessed the implementationof Cuban biological control researchprograms. Since 1968, such programs have sponsoredthe release of L. diatraeae over 100% of thearea under sugarcane seed production (Dlott et al.1993).Research on alternative agricultural practicesgained momentum in the 1970s and 1980s as agroup of Cuban scientists in various disciplinesincreasingly became alarmed about the long-termnegative effects of classical agriculture. In the mid-1970s, scientists at the Institute of Botany refusedto carry out the Forestry Department’s plans forterracing mountainsides with monocultures of teakand then clear-cutting the trees, stating that theplan’s risks of pest outbreak and soil erosion wereunacceptable (Levins 1993). In the mid-1980s,this vocal minority of scientists in favor of the alternativemodel began to gain a hearing. Theirsuggestions were valued primarily as a means ofsaving money on petroleum products purchasedfrom other countries and were, therefore, swiftlyadopted (Vandermeer et al. 1993). By the late1980s, the Ministry of Agriculture adopted biologicalcontrol as national policy (Levins 1993).Reforms After 1989Cuba’s agricultural scientists thus were well preparedto face the economic catastrophes of 1989by rapidly implementing alternative agriculturalpractices. Preliminary data on alternative farmingtechniques already existed, and this knowledge baseallowed for the rapid expansion and redirection ofCuba’s agricultural research program. The eventsof 1989 simply triggered and formalized a philosophythat long had been gaining popularity withCuban scientists.Yet conversion from classical to alternative agriculturalprocesses does not come easily, no matterhow much popular support is available. InCuba, the difficulties of such a conversion wereincreased by the triple challenge the country faced—to simultaneously increase food production, reduceor eliminate inputs, and maintain a steadycrop of export sugarcane to exchange for hardcurrency. To meet these challenges, Cuba not onlyhas undertaken the largest shift to organic farmingtechniques the world has ever known but also hasrestructured its social and economic systems fromthe ground up (Eckstein 1997).This realignment of society along ecological linesrequired a general strategy reaching into every areaof production. In accordance with the philosophyof Che Guevara, who emphasized the inseparabilityof social and economic development (Levins1993), Cuba is attempting to achieve food selfsufficiencyby involving the entire populace in theeffort. Central planning helps to determine researchand education programs, disseminating the knowledgeneeded to employ alternative agricultural techniques.Economies of scale are exploited and scarceresources directed to where they are needed most.At the same time, the units of production are keptas small as possible, empowering individual growersand drawing upon their specialized knowledgeof local agronomic conditions. Additionally, suchsmall-scale units of management have created numeroussmall research centers, allowing farmersand scientists to interact more frequently and effectively.National ConversionIntroducing alternative farming techniques inan entire country involves complexities far beyondthose encountered in a single farm conversion.Cuba’s conversion has focused on the development,implementation, and coordination of new techniquesin several key areas: insect management,plant disease management, weed management, soilmanagement, and labor mobilization.Insect ManagementThe massive decline in pesticide imports thatfollowed the dissolution of the Soviet bloc in 1989left Cuba’s agriculture system extremely vulnerableto insect pests. Since the introduction of agrochemicalsinto Cuba in the 1940s, these inputs hadbecome increasingly vital to Cuba’s food production.Although research on alternatives was beingcarried out well before 1989, wide-scale implementationof alternative techniques had not been attempted.In recent years, a countrywide shift to biologicalcontrol methods has been set in motion (Table1). This effort is overseen on a national level byCuba’s Ministry of Agriculture, which includes cropprotection organizations such as the National Serviceof Plant Protection, the Central Research Laboratory,14 regional research laboratories, 60regional plant protection stations, 27 diagnosticlaboratories, and 218 Centers for the Reproductionof Entomophages and Entomopathogens.Important crops such as tobacco and sugarcanehave their own specialized agencies.The Ministry of Agriculture’s work since 1989has focused on the development and applicationof biological control against insect pests. Traditionalcontrol methods based on peasant farmingpractices, such as the use of ants for the control ofbanana and sweet potato pests, have been reintro-220 AMERICAN ENTOMOLOGIST • Winter 2001

Table 1. Biological control agents employed in CubaBacillus thuringiensis (Berliner)Heliothis spp. (Noctuidae) on tomato and tobacco; Erinnyis ello (L.);(Sphingidae) on papaya and cassava;Spodoptera spp. (Noctuidae) on tomato, pepper, sweet potato, and watercress;Plutella xylostella (L.) (Yponomeutidae) on cabbageBeauvaria bassiana (Balsamo)Lissorhoptrus brevirostris Kuschel (Curculionidae) on rice;Diatrea saccharalis (F.) (Pyralidae) on sugarcane;Cosmopolites sordidus (Germar) (Curculionidae) on plantain;Pachnaeus spp. (Curculionidae) on citrus;Cylas formicarius elegantulus (Summers) (Curculionidae) on sweet potatoMetarhizium anisopliae(Metschnikoff)Lissorhoptrus brevirostris Kuschel (Curculionidae) on rice;Galleria mellonella (L.) (Pyralidae) in beehives;Cosmopolites sordidus (Germar) (Curculionidae) on plantain;Mocis spp. (Noctuidae) on forage crops;Diatrea saccharalis (F.) (Pyralidae) on sugarcanePaecilomyces lilacinus (Thom)Meloidogyne spp. (Nematoda) on coffee and guava;Radopholus similis (Cobb) (Nematoda) on plantainVerticillium lecanii (Zimmermann)Bemisia tabaci Gennadius (Aleyrodidae) on sweet potato, tomato, cucumber, beans,and peppersTrichoderma spp.Phytophthora nicotianae Breda de Haan (fungus) on tobacco;Rhizoctonia solani Kuhn (fungus) on tobacco;Fusarium spp. (fungus) on tobacco;Sclerotium rolfsii Saccardo (fungus) on peanuts and beansLixophaga diatraea (Townsend)(Tachinidae)Diatrea saccharalis (F.) (Pyralidae) on sugarcaneTrichogramma spp.(Trichogrammatidae)Mocis spp. (Noctuidae) on forage crops;Erinnyis ello (L.) (Sphingidae) on yucca;Diatrea saccharalis (F.) (Pyralidae) on sugarcaneTelenomus minimus Ashmead(Scelionidae)Spodoptera spp. (Noctuidae) on legumes(Formicidae)Pheidole megacephala (F.)Cylas formicarius elegantulus (Summers) (Curculionidae) on sweet potato;Cosmopolites sordidus (Germar) (Curculionidae) on plantainTetramorium guineense BernardCosmopolites sordidus (Germar) (Curculionidae) on plantainHeterorhabditis heliothidis (Khan, Cylas formicarius eleganutulus (Summers) (Curculionidae) on sweet potato;Brooks & Hirschmann)Plutella xylostella (L.) (Yponomeutidae) on cabbageSources: Perfecto (1994), Thrupp and Perez (1989).AMERICAN ENTOMOLOGIST • Volume 47 Number 4 221

The reintroductionof traditionalpeasant practicesfor management ofinsect pestsalso has provenfruitful in Cuba.duced and more widely applied. Additionally,modern technologies have been utilized for the massrearing and release of natural enemies and for thedevelopment, mass production, and application ofentomopathogenic organisms.The first step in applying any biological controlmethod is to monitor for the presence of crop pests.The system now in place in Cuba is among theworld’s most comprehensive. Monitoring was firstdeveloped in the 1970s, when it was used as ameans of tracking and delaying insect resistance topesticides. Today, each of the 60 research stationsoverseen by the Ministry of Agriculture includes amonitor plot. These plots are planted with theimportant crops and varieties of their regions, andon-site weather stations constantly record climateconditions. Scientists regularly evaluate crop phenology,pest population levels, disease onset anddevelopment, and crop yield data. Additionally,several fields in each region are selected as test plots,and data on crop threats are collected. When potentiallydangerous climatic conditions, diseases,or pests are discovered, local farmers are advisedof the threat. Control measures employed in responseto such a situation depend upon local productionand availability of biological control agents.The mass production of biological controlagents was an early goal of Cuba’s shift to alternativeagricultural practices. Lack of currency meantthat such agents could not be purchased from othercountries, and, in fact, large-scale production ofbiological control organisms is rare. In Cuba, theCenters for the Rearing of Entomophages andEntomopathogens (CREEs) are responsible forproducing the needed numbers of organisms.These centers function as artisanal (i.e., staffed byskilled laborers rather than scientists) productioncenters, which are located on collective farms andoperated by members of the collective. Operationof these centers by local residents, whose educationlevels range from Ph.D. scientists to high schoolgraduates, has helped destroy the myth that biotechnologyrequires high-tech, sophisticated laboratoriesand personnel to achieve success. Theentomopathogens and entomophages producedare available free of charge to cooperative membersand are sold for profit to others.Biological control agents produced at the CREEsare varied, and are targeted to meet local pest controlneeds. Entomophage production has focusedprimarily on Trichogramma, a hymenopteran eggparasitoid. Trichogramma colony stocks are collectedfrom the region’s target pests, ensuring ahigh level of host preference and adaptation by theparasitoid. These field-collected Trichogramma arereared in the laboratory and allowed to oviposit inthe eggs of either the rice moth, Corcyra cephalonica(Stainton); or the Angoumois grain moth, Sitotrogacerealella (Olivier); which are stored product pests.Parasitized eggs are collected in vials and transportedto pest-infested fields when 50% emergencehas occurred. Release rates are determined by theintensity of pest infestation and range from 8,000to 30,000 Trichogramma per hectare of agriculturalland. To date, Trichogramma releases havebeen used successfully to control the sugarcaneborer; the cassava hornworm, 1 Erinnyis ello (L.);the tobacco budworm, Heliothis virescens (F.); andseveral Mocis spp.; which are lepidopteran foragepests (Dlott et al. 1993).Entomopathogenic fungi and bacteria are alsomass-produced at local CREEs. Cuba’s productionand deployment of these biological controlagents are among the most highly developed in theworld and have allowed for control of a great varietyof pests. Agents employed include Bacillusthuringiensis (Berliner), Pasteuria penetrans(Thorne), Beauvaria bassiana (Balsamo),Metarhizium anisopliae (Metschnikoff),Paecilomyces lilacinus (Thom), and Verticilliumlecanii (Zimmerman). Organism efficacy is ensuredby constant quality control and monitoring; foreach formulation of bacteria or fungi, virulence isevaluated in the laboratory prior to deployment.Application rates are determined by evaluating pestsexposed to standardized doses of entomopathogens;spores per pest organism per application volumeare evaluated and adjusted for maximumbenefit. After wide application of a formulation,treated crops are scouted frequently to determineincidence and severity of entomopathogen infection.In 1995, some 500,000 hectares of agriculturalland were treated with biological controlagents, up from 77,000 hectares in 1990 (F. F.Monzote, personal communication). Good controlof nematodes, lepidopterans, and weevils hasbeen achieved through use of locally manufacturedbiological pesticides.The reintroduction of traditional peasant practicesfor management of insect pests also has provenfruitful in Cuba. Based on input from farmers,Cuba has pioneered the use of generalist antsagainst the sweet potato weevil, Cylas formicariuselegantulus (Summers). Previous research in othercountries (Carroll and Risch 1990) had indicatedthat the bigheaded ant, Pheidole megacephala (F.),could not provide effective pest control in annualcrops, but Cuban farmers, observing the voraciousnessof this ant against the sweet potato weevil, feltthat the potential for control existed. Preliminarydata from the 1980s were promising, and todaythis system is employed in the entire Pinar del Rioprovince, a prime growing region.Directed research on the use of the bigheadedant has generated an effective management system.Reservoir colonies of this ant are created in areaswith high natural populations, and all pesticideapplications are prohibited. At critical infestationperiods, ants are transported to sweet potato fieldsvia a labor intensive method of inducing them tocolonize banana or plantain pseudostems liberallydosed with sugar. After colonization, thepseudostems are transported to target field wheresubsequent desiccation induces the ants to move1Common name not currently among common namesof insects and related organisms approved for use bythe ESA Committee on Common Names of Insects.222 AMERICAN ENTOMOLOGIST • Winter 2001

to the ground and begin feeding on the sweet potatoweevil. Efficacy rates of up to 99% have beenachieved in some regions, and use of the bigheadedant has replaced chemical pesticides in sweet potatocrops, producing higher yields at a lower cost(Dlott et al. 1993). Currently, biological controlusing ants is being extended to plantains for thecontrol of the banana root borer, Cosmopolitessordidus (Germar).Plant Disease ManagementThe biological control of plant pathogens haslagged behind the development of biological controlof insects. Cuban research on plant diseasetraditionally has focused on the epidemiology ofpoorly understood diseases, and fungicides andother chemicals have been relied upon for diseasecontrol. In recent years, the emphasis of plant diseaseresearch in Cuba has shifted to methods ofdisease diagnosis and reduction, both fairly newresearch disciplines. Until the early 1980s, diagnostictests did not exist at Cuba’s research stations,and development of biological agents fordisease control did not begin in earnest until 1990.Disease detection has been enhanced by the incorporationof techniques from veterinary andhuman health research. The Cuban National Animaland Plant Health Center has developed diagnosticprotocols employing electron microscopy,dot blot tests, immunofluorescence, and serologicaltests (ELISA). The production of ELISA kitshas proven so successful that Cuba now exportsthese tests worldwide for the detection of importantplant diseases. Additionally, the local pestmonitoring stations maintained by the Ministry ofAgriculture are used to track the development andspread of plant diseases, and local farmers arealerted to any worrisome trends.As fungicide availability has declined, Cubanresearchers have begun to seek alternative meansfor coping with plant diseases. On banana plantations,the fungus Mycosphaerella musicola (Leach)causes yellow sigatoka disease and is responsiblefor large reductions in yield. This fungus kills bananaleaves, and a linear relationship exists betweenthe number of leaves killed and the degree ofyield reduction. Monitor plots to track the incidenceof yellow sigatoka infection have been establishedat all major growing sites and until 1989were used to determine the optimum timing of pesticideapplication. More recently, however, sigatokatolerantcultivars have been discovered, and thesenow are widely planted (Shishkoff 1993). Wheninfections do arise, treatment consists of leaf strippingand the application of mineral oils, which appearto inhibit infection.The discovery of microbial antagonists for thecontrol of plant disease has become a major aim ofCuban research since 1989. Prior to that time, theexhaustive screening and evaluation proceduresrequired for such research were not regarded ascost effective. In 1990, however, the Institute ofPlant Health began to investigate microbial antagonistsfor the control of tobacco root diseases. Soilsamples were collected from both infected and disease-freefields, and 25 colonies of Trichodermaspp. fungi were isolated for evaluation as antagonistsagainst a variety of root-disease pathogens.Paired tests of antagonists and targets were carriedout to evaluate antagonist efficacy, and four highlyeffective Trichoderma spp. isolates were discovered.These were then tested in greenhouse conditionsagainst heavily infested tobacco seedlings. Theseisolates have proven so effective that fungicide usein tobacco seedbeds has been eliminated almostentirely. Trichoderma now is mass-reared at severalCREEs, and streamlined regulatory processeshave facilitated its widespread use. Although biologicalcontrol agents for agricultural use must meetregulatory guidelines modeled on those of the U.S.Federal Drug Administration and the EuropeanCommunity, this process is reported to be extremelyrapid in comparison to U.S. procedures (Rossetand Benjamin 1994).Weed ManagementThe control of weeds within an alternative farmingsystem can be extremely challenging. A majorcomponent of the alternative model of agricultureis the reduction of tillage to improve soil health,but this reduced tillage allows weeds to flourish. Indeveloped countries employing low-till or zero-tilltechniques, selective herbicide applications are usedto combat weeds. In Cuba, the reduced availabilityof herbicides following the collapse of the Sovietbloc made the development of alternative weedmanagement strategies a necessity. The resultingfocus of Cuban weed control science is twofold:(1) to reintroduce the traditional methods of weedcontrol employed by farmers prior to the modernizationof Cuban agriculture and (2) to scientificallyevaluate the basis for the success of thesemethods and implement these methods for widespreadapplication.Weeds are evaluated in the context of the complexecological systems they inhabit. Monitoringof test sites is used to generate data on weed speciesand densities from year to year, the compositionand viability of seed banks, the type of crop to beplanted and its competitive relationship to theCrop rotation for weed control: beans planted after a year of corn cultivation.AMERICAN ENTOMOLOGIST • Volume 47 Number 4 223

weeds present, the effectiveness of herbicides againstthose weeds, and the degree of similarity betweencrop and weed growth habits (Dlott et al. 1993).These data are then incorporated into a mathematicalmodel to predict what weed problems will arisein each year and location and to devise plantingschemes to alleviate the effects of those problems.Crop rotation is one of the most important traditionaltechniques now being applied based onmodel predictions. Crops with high competitiveability against a given weed community are employed,and a single year of planting a highly competitivecrop can provide nearly weed-free growingconditions in the following year. Corn, for example,can shade out low growing weeds, allowingbeans to be planted the following year. Regionswith particularly severe weed problems may beplanted with a dense cover crop such as sweet potato,which smothers out any competing weeds.Further, selective rotation can allow the applicationof herbicides at a time when herbicide-tolerantcrops are present, paving the way for weed-freegrowing of herbicide-susceptible crops in subsequentgrowing seasons.Soil cultivation techniques also have a strongimpact on weed control. Incorporation of tillagefor weed control with a minimum-tillage philosophyhas called for creative solutions on the part ofCuban growers. One such solution is the multiplowdesigned by Cuban agricultural engineers. Unlikeroto-tillers and discs, this device lifts and opens thesoil without turning it. Thus, subsoil cultivationoccurs without exposing weed seeds for germination;more importantly, any weeds present are essentiallysliced off at the root.Soil ManagementSustainable agriculture under the alternativemodel is dependent completely upon high qualitysoil resources. Prior to the crisis of 1989, Cuba’ssoil management practices were similar to thoseemployed in most developed countries, relyingCuba produces over 90,000 metric tons of earthworm humus per year in plotslike this one.heavily on inorganic inputs of fertilizer to maintainhigh productivity levels. Production-oriented practicescontributed to significant fertility loss, erosion,and salinization of Cuba’s already poor soils,and addressing these problems has been an importantpart of Cuba’s agricultural conversion.The first task carried out by Cuba’s soil scientistswas the classification of all the island’s soils.Mapping on a 1:250,000 scale was carried out, andsoil productivity potential evaluated. A mere 8% ofCuba’s soils were rated as having very high potentialproductivity (Gersper et al. 1993), whereas mostsoils fell into the low or very low categories. Existingproductivity levels could not be expected to meetCuba’s food needs, and an ambitious program ofsoil management involving minimum tillage, recycledand biological fertilizers, green manure, reforestation,and vermiculture was undertaken.The low organic matter content of Cuba’s soilsis being addressed through the addition of organicand biological fertilizers. The introduction of nitrogen-fixingmicroorganisms into agricultural soilshelps increase the amount of nitrogen available tocrops. Rhizobium bacteria are mass-produced foruse on leguminous crops, and inoculation with thesebacteria is estimated to supply 80% of the crops’nitrogen needs (Gersper et al. 1993). Azotobacter,free-living nitrogen-fixing bacteria, are used onnonlegumes. The large-scale production and useof Azotobacter is a technique unique to Cuba, andis estimated to supply more than half of the nitrogenneeded by nonlegumes (Perfecto 1994).Another uncommon technology employed inCuban agriculture is the widespread use of Bacillusbacteria. Many Cuban soils are high in aluminumand iron oxides, which bind with phosphatesinto insoluble complexes. Phosphorous, althoughpresent, then becomes unavailable for plant uptake.In the past, this situation was rectified throughartificial inputs of phosphates. Bacillus organisms,however, function as phosphosolubilizing bacteria,breaking down these complexes and freeingphosphorous for absorption by plants.Finally, investigations of vesicular arbuscularmicorrhizae (VAM), a group of symbiotic fungithat function as auxiliary roots, currently are underway.Fifty-three species of VAM native to Cubahave been identified, and arrangements for massproductionare underway (Perfecto 1994).Green manure is another important source ofsoil nutrients. By growing and then plowing underplants with high nutrient-uptake capabilities, soilfertility can be increased. One particularly usefulgreen manure is Sesbania. This saline-tolerant legumecan produce up to 60 tons of green manurein 45 days (Gersper et al. 1993), providing some75% of the nitrogen required for crops plantedsubsequently. The ability of Sesbania’s to grow insaline soils is especially important in Cuba, becausethe long, narrow geography of the island subjectsalmost all soils to saline stress.Cultural techniques also are used to increasethe health of Cuba’s soils. Exhaustive surveys ofcrop/soil productivity have been carried out, iden-224 AMERICAN ENTOMOLOGIST • Winter 2001

tifying the crops best suited to a region’s soils. Croprotations are designed to maximize the effects ofgreen manure, and crop residues commonly are plowedunder to contribute nutrients to the soil. Intercroppingpractices also have been studied and employed.Although traditional peasant farming madeextensive use of intercropping, the practice fell awaywith the introduction of large-scale monocultureproduction. Today, the value of intercropping istwofold: field productivity is increased, as in theintercropping of soybeans for animal food withsugarcane for export; and soil erosion is prevented,particularly through the use of leguminous intercrops.Additionally, the replacement of tractorpower by animal traction has reduced erosionthreats to soil significantly.Many of the plant nutrition needs not met bythese techniques are addressed by the use ofvermiculture. Vermiculture, the large-scale use andproduction of earthworm humus, has grown rapidlyin Cuba, from 2,000 metric tons in 1987 to96,000 metric tons in 1992 (Gersper et al. 1993).Earthworm humus stimulates plant growth, increasesnutrient uptake, and helps protect plantsfrom soil-borne diseases (Gersper et al. 1993).Earthworm humus is an efficient fertilizer, and 4tons of humus are reported to provide as muchnutrition as 40 tons of cow manure (Werner 1994).Humus production in Cuba is state of the art.Fresh animal manure is fermented for 30 days,then mixed with soil and seeded with earthworms.The worms feed on the top layer of manure anddeposit castings into the lower layers. Compost isadded continuously for 90 days, then the wormsare removed and the humus is ready for agriculturaluse. Several regional research stations havemass-production facilities for earthworm humus,and these stations are responsible for educatingpeople in their region on vermiculture techniques.Annual vermiculture conferences further spreadknowledge, and vermiculture is encouraged forhousehold and community use as well as on farms.Organic materials from livestock manure, foodwaste, human waste, and sugarcane productionare recycled as vermiculture substrates.Waste recycling also is common for uses otherthan vermiculture. Sugarcane is the best exampleof effective waste recycling in Cuba, and some sugarmills are run entirely on energy from their ownwaste (Kaufman 1993). Sugarcane productioninvolves extremely high levels of waste byproductsincluding bagasse, wastewater, and filter press cake.Bagasse, a dry pulp, is used both for animal feedand as biomass for energy production; wastewateris diverted to local farms for irrigation; and filterpress cake, a processing byproduct high in calcium,phosphorous, and potassium, is used as a cropfertilizer. The extraction of steam energy from bagassehas been so successful that excess energy oftenis available for the local municipal power grid.Cuba’s efforts at soil rectification have producedone of the world’s most impressive reforestationprograms. After the 1959 revolution, Cuba beganreplanting thousands of hectares of forestland felledby Spanish and American sugarcane growers.Cuban revolutionaries have a long-standing appreciationof trees, dating back to Jose Marti, whosaid “A region without trees is poor. A city withouttrees is sickly, land without trees is parchedand bears wretched fruit.” Even before the eventsof 1989, Cuba had increased tree cover from 14 to20%. In the years since, the reforestation programhas been intensified, and under the Manati Planseeds are provided to all interested people for thereforestation of degraded lands. As a result, Cubahas added 697,000 hectares of forest in the last 8Additionally, thereplacement oftractor power byanimal tractionhas reducederosion threats tosoil significantly.Plows drawn by oxen such as these have largely replaced gasoline-powered tractors in Cuba.AMERICAN ENTOMOLOGIST • Volume 47 Number 4 225

years, making it one of the very few countries in theworld to have increased the amount of forestedland in recent decades (Baker 1997).Labor MobilizationOver the past 40 years, Cuba’s population hasbecome more urban. As farm work became increasinglymechanized, this flight to the cities wasexacerbated, so that today an estimated 30% ofCuba’s population lives in Havana or its environs.The advent of the special period following the dissolutionof the Soviet bloc, therefore, gave rise to acrisis in labor because petroleum was no longeravailable to carry out mechanized production.Moreover, nationwide conversion to alternativeagricultural techniques demands a huge labor pool.Although 30 years of progress under Soviet guidancewere aimed at reducing human inputs in agriculture,the events of 1989 have made a reversal ofthis trend imperative.Many alternative growing techniques are notamenable to mechanization; the harvesting of intercroppedfields, for example, must be done manually.Although some practices may be carried outby machine eventually, such technology is in itsinfancy. As a result, any large-scale implementationof sustainable agriculture depends upon themustering of a vast agricultural work force.Cuba is attempting to redirect labor in severalways. In the short term, incentive plans have beeninstituted to encourage city dwellers to volunteertheir services as agricultural laborers for periodsranging from 2 weeks to 2 years. These workersare guaranteed a salary at least equal to their realjob’s wages and receive somewhat better food andhousing than is standard. Their city jobs are heldfor them, and, should they choose to remain in thecountry, they are provided with permanent housing.Another form of volunteerism is the PlanTurquina whereby young people can substitute 2years of agricultural work for their required militaryservice. By targeting young people at the thresholdof marriage and childbearing, Cuba hopes toreestablish rural communities.The division of labor on state farms also hasbeen restructured to increase rural satisfaction.Previously, workers were sent to various parts ofthe farm as their specialized tasks needed to beperformed; now, it is standard for each cohort oflaborers to be associated with a particular tract ofland, generally about 80 hectares in size (Rosset1996). The same group carries out all growing operationson this tract, from sowing to harvest. Simultaneously,productivity incentives have beenestablished in which the base pay is augmented accordingto production levels. Management responsibilitieshave been decentralized similarly, and manyland parcels are managed by those who farm them.The call to agricultural productivity has not beenlimited to rural populations. City dwellers increasinglyare encouraged to plant organoponicos (urbangardens) with the aim of decreasing dependenceon food from rural sources. The loss of petroleumimports has resulted in severe problems with therefrigeration and transport of food, and urbangardens are seen as a way of shifting the burden.Unused land parcels are available free of charge toanyone who will grow food on them, and thegrower is entitled to keep all produce. There noware more than 2,000 small gardens in the city ofHavana, and growers range from individual familiesto entire neighborhoods, workplaces, andschools.ConclusionsThe quiet revolution in Cuba is well on its wayto success. For the first time in post-Colombianhistory, Cuba is approaching economic independence.That such gains have been made using sustainableagricultural practices is a lesson we shouldall note. Although many scientists around theworld have lamented the persistent spread of agriculturalpractices based on the classical model ofproduction, nowhere else has a large-scale conversionto alternative techniques been attempted.By making agriculture essentially a small-scaleendeavor on a national level, Cuba has drawn uponlocal and individual knowledge in designing solutionsto problems such as soil fertility and insectmanagement. Revolutionary to their core, Cubanshave adopted alternative agriculture as swiftly andpassionately as they did the Revolution of 1959.Rather than excluding the populace, the new modelin Cuba integrates all areas of society in a commonstruggle for food self-sufficiency. By asking not ifsustainable agriculture is possible, but rather howit can best be carried out, Cuba has paved the wayto a more rational form of food production.AcknowledgmentsI thank G. Kennedy (Department of Entomology,NCSU), F. Gould (Department of Entomology,NCSU), B. Zivkovic (Department of Zoology,NCSU), and two anonymous reviewers for criticalreadings and discussion of the manuscript; R.Oppenheim (Department of Neurobiology, BowmanGray School of Medicine) for the opportunityto visit Cuba; A. Mittal and the folks at Food Firstfor information on the current state of Cuban biotechnology;and Fernando Funes Monzote (Pasturesand Forages Research Institute, CubanAssociation of Organic Agriculture, ApartadoPostal 4029, C.P. 10400, Ciudad de La Habana,Cuba) for information on biological control practicesin Cuba.References Cited Boca Raton, FL.Baker, C. P. 1997. Cuba handbook. Moon TravelHandbooks Series. Moon Publications, Chico, CA. Silent spring. Houghton Mifflin,Boston, MA.An evaluation ofants as possible candidates for biological control intropical annual ecosystems, pp. 30-46. In S. R.Gliessman (ed.), Agroecology. Springer, New York,NY. The transfor -226 AMERICAN ENTOMOLOGIST • Winter 2001

mation of the state extensive growth model inCuba’s sugarcane agriculture. Agric. Hum. Values13: 59-68. Biological control by natural enemies.Cambridge University Press, London. Socialism on one island? Cuba’snational food program and its prospects for foodsecurity. Working papers series number 124. Institutefor Social Studies, The Hague, The Netherlands. Management of insectpests and weeds. Agric. Hum. Values 10: 9-15.The limits of socialism in a capitalistworld economy: Cuba since the collapse of theSoviet bloc, pp. 193–217. In M. A. Centeno and M.Font (eds.), Towards a new Cuba? Legacies of arevolution. Lynne Reiner, London.Soil conservation in Cuba: a key to the new modelfor agriculture. Agric. Hum. Values 10: 16-23. From red to green: Cuba forced toconserve due to economic crisis. Agric. Hum. Values10: 31-34.The ecological transformation ofCuba. Agric. Hum. Values 10: 52-60. Alternativeagriculture: committee on the role of alternativefarming methods in modern production agriculture.National Research Council; National AcademyPress, Washington, DC.. External shocks and adjustment incontemporary Cuba. The International and PublicAffairs Center, Occidental College, Los Angeles, CA. The transformation of Cuban agricultureafter the Cold War. Am. J. Altern. Agric. 9: 98-108. Cuba: Alternative agriculture duringcrisis, pp. 64-74. In L. A. Thrupp (ed.), New partnershipsfor sustainable agriculture. World ResourcesInstitute, Baltimore, MD. The greeningof the revolution: Cuba’s experiment with organicagriculture. Ocean Press, Melbourne.Preliminary report on astudy of the biology of Lixophaga diatraeae TennesseeJ. Econ. Entomol. 23: 999-1104. New approaches to plant pathologyin Cuba. Agric. Hum. Values 10: 24-30. Introduction, pp. 1-38. In L. A.Thrupp (ed.), New partnerships for sustainable agriculture.World Resources Institute, Baltimore, MD. Breaking chemicaldependency in agriculture: the remarkable case ofCuba. Unpublished manuscript as quoted in Perfecto(1993). Energy and Resource Group, Universityof California, Berkeley. Cuba and the dilemma ofmodern agriculture. Agric. Hum. Values 10:3-8. Cuban agriculture looks tovermiculture. The Cultivar (newsletter of UCSCCenter for Agroecology & Sustainable Food Systems)12(2): 2-4.. WorldHealth Organization Yearbook, 1989. Geneva,Switzerland.Sara Jane Oppenheim, Department of Entomology,North Carolina State University,840 MethodRoad, Unit 1, Raleigh, NC 27612 (e-mail:sjoppenh@unity.ncsu.edu).AMERICAN ENTOMOLOGIST • Volume 47 Number 4 227

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