MANAGEMENT OF P REHAR VEST I NSECTS 25 l
Barfield and Stimac (1980) critically reviewed IPM from an entomological Chaprer 9 perspective by ( 1) foc using on characrerisrics of agriculture conducive ro creat ing insect pests, (2) identifying characteristics of insects which enable them to become pests, (3) retracing the hisrorical roure of insect control up co IPM, (4) MANAGEMENT OF PREHARVEST INSECTS elucidating discrepancies between theory and practice of IPM, and (5) identi fying relevant problems which must be overcome in dealing with insects as JAMES W. SMITH , JR. , AND CARLS. BARflELD pests and IPM as a philosophical commitment co combatting pests. Our purpose here is to present the utility of IPM fo r the peanut agroecosys Worldwide, some 10,000 species of insecrs are pesrs of man, domesric rem. This presentation can best be accomplished in 4 seeps. First, we will iden animals, food and fiber. A subsrancial indusrry has developed co produce syn tify some bas ic concepts which characterize IPM, then use these concepts as thetic insecticides and other pesticides to combat this myriad of pests (Bor milestones co judge where peanut agriculturists are in relation to the realiza rrell, 1979). Jnsecricides have been of tremendous benefit co man bur have nor tion of IPM programs. Second, we will identify various approaches to com bat been used without deleterious side effects (Luck et al. , 1977; Bottrell, 1979; ting pests and show which, if any, of these approaches is currently utilized in Metcalf, 1980). Boraiko ( 1980) cites specific aspens of environmental and hu peanuts and how such approaches may change as a function of variables such as man health hazards attributed co synthetic pesticide usage. A comprehensive crop mix and geographical location. Third, we will provide a conceptual mod review of the hisrory of insecticide usage and subsequent problems is contained el of the peanut system to serve as a reference for identifying existing and miss in Mercalf( l980). ing information. Fourth, we will place some priority structure on the missing Before the late l930's agriculturists did nor have access co many pesticides; info rmation and justify that structure as relevant ro the development of IPM thus, they were fo rced co rely on culturally inherited farming practices for pest schemes in peanuts. conrrol. Such methods (e.g., crop rorarion) often unknowingly cook advantage At least 5 principles of IPM have been identified (Bottrell, 1979). The first of basic ecological principles co reduce pest arrack. Today, many agriculrurisrs and fo remost principle is that potentially harmful species will continue co exist are directing resea rch efforts toward gaining an understanding of how an at rolerable levels of abundance (Smith and van den Bosch, 1967). Thus, under agroecosysrem functions (i.e., how its components interrelate) so that pest virtuall y all situations, pest eradication is nor consistent with an IMP pro control strategies which arc less c:cologicall y disruptive than blanker usage of gram. Second, the ecosystem is the management unit (Smith and van den insecticides can be developed. Efforts co rekindle studies on agroecosystem Bosch, 1967). We shall see later how the focus on management at the individ form and function have necessitated philosophical, as well as scientific, altera ual peanut field level has resulted in uncertainty, particularly in management tions in the way agricultural scientists approach rhe problems of pest control. of mobile, polyphagous insect pests. Third, IPM encourages maximum utility from naturally occurring mortality agents (parasites, predarors, pathogens) THE IPM PHILOSOPHY (Stern et al. , 1959). Fourth, any applied control procedure may produce unex pected and undesirable effects (Smith and van den Bosch, 1967). Last, an in The latest in a seri es of philosophies on how co combat pest organisms is terdisciplinary systems approach is essential to the development of IPM. Ex called integrated pest management (JPM). Numerous auchors (e.g., Smith amples will be provided later of ongoing efforts which are aimed at using mod and van den Bosch, 1967; Huffaker, 1972; Bottrell, 1979; Barfield and Stim els as cools to understand the peanut agroecosystem prior to managing ir. In ac, L980) define IPM more or less identically as the use of various tactics short, these are efforts co avoid violation of principles 4 and 5 of Bottrell (chemical, cultural, biological, physical) in an integrated fashion so as co yield ( 1979). These 5 principles will be used throughout this discourse in reference predictable economical, ecological and sociological consequences. We shall re to why specific problems (and potential solutions) seem to exist in develop turn co this definitio n of JPM later to provide specific examples of where rhe ment of IPM for specific insects or pest complexes within the peanut agroecos development ofIPM in worldwide peanut, Arachis hypogaerl L. , production sys ysrem. Having reviewed these principles, various approaches to combatting tems is relative co this definition. pests are summarized. Afterward, conclusions will be drawn to determine the Integrated pest management is synonymous with pest management, and status of development of IPM programs fo r insects or pest complexes. both terms evolved from integrated control which was orig inally used co de Barfield and Stimac (1980) reviewed 4 d istinct approaches co combatting scribe the use of biological and chemical controls synchronously (Stern er al. , insects. A brief review of these approaches is necessary for identify ing how spe L95 9). Theoreticall y, IPM represents a combination of actions (tactics) which cific peanut insect pests are being dealt with today. The first approach is no ac can be blended into an overall, balansed arrack (a strategy). Real ization of the tion and involves a lack of action in 2 distinctly different situations: ( 1) in che optimum combination of tactics inro a strategy fo r a given crop, pest, or crop absence of relevant data and (2) as a decision following analysis of relevant data. pesr complex is nor a trivial task. Actual examples show char current IPM pro Secondly, prevention can be uci lized. This approach involves at least 6 catego g rams are in various stages ofdeve lopment (Bottrell, 1979; Barfield and Stim ries of racrics: ( 1) use of resistant plant varieties; (2) manipulacion of crop plan ac, 1980). ting date, tillage and row spacing; (3) conservation or introduccion of pest nat ural enemies; (4) crop roracion schemes; (5) use of attractancs or repellants; and
250 252 PEANUT SclENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 253
(6) preplant application of insecticides. The third approach is suppressibn, and damage. Fourth, evaluate the compatibility of tactics to avoid creating some this approach involves a broad spectrum of actions which may be taken after an pest problems while alleviating others. Lastly, identify what knowledge is insect pest has reached (or is expected to reach) densities considered to be eco missing relative to our ability to design viable IPM strategies for peanuts. nomically important. There are 3 generic categories of suppressive agents: Peanuts are attacked by plant intracellular feeders, foliage consumers, in chemicals, parasites and predators, and microbials. The final approach to com sect-transmitted diseases, and insects feeding on roots, pegs and pods. Each batting insects (or other pests) is directed management and involves the use of type of pest has 1 or more naturally occurring enemies (predators, parasites compatible tactics such that specific consequences, within specified ranges, are and/or pathogens) which theoretically can be manipulated for pest suppres understood prior to action. Thus, directed management involves insects as sion. Besides these natural biological controls (or imported ones), at least 3 well as other pests and complexes of beneficial organisms. The level of knowl other categories of management tactics appear to be in use today against peanut edge about the structure and function of a particular agroecosystem needed to insect pests: resistant plant varieties, various combinations of cultural prac achieve directed management appears far superior to the level of knowledge tices, and various insecticides applied at some economic threshold pest density needed for prevention and/or suppression. or in a preventative manner. How these 4 general categories of tactics are used The basic concepts of IPM have been outlined and 5 principles identified depend upon geographical location, particular pests in question, age of crop, which must be considered in the development of IPM programs. Further, a and philosophy of the managers. summary of 4 distinctive approaches to combatting insect (and other) pests Now that the general IPM philosophy and field level components (plants, have been provided. Integrated pest management provides the theoretical pests, natural enemies, environments and management tactics) in a peanut sys foundation necessary to deal with pests over sustained intervals of time; howev tem have been reviewed, a focus on state of the art for IPM in peanuts world er, it is recognized that the multitude of ongoing programs designed to deal wide becomes pertinent. Insect pests are divided into 2 major categories based with insect pests are in various stages of development. Focus must now be di on habitats: foliage inhabitants (consumers, intracellular and insect transmit rected toward peanuts as a particular crop plant with numerous pests, many of ted diseases) and subterranean inhabitants. The format will include the pest which are cosmopolitan in distribution. Further, concentration will be on the status of each type pest, current management practices used against each, and insect components of chat pest complex. Questions relating to which species current information (biological and ecological) about each. This approach will attack peanuts, where (geographically and in relation to habitat) they attack, accomplish 2 goals critially important in preparing this book chapter. First, when (seasonally and in relation to plant phenology) they attack, and what can focus is directed on both the similarities and differences in the way IPM strate be done to lessen the impact of these attacks on a worldwide basis will be ad gies for specific pests have materialized around the world. A given insect may dressed. This systematic approach focuses on some basic features of the peanut be a key pest in ! location and only an occasional pest elsewhere.· Knowledge of agroecosystem which contributes to insect pest problems and identifies neces why this is true is of central importance in constructing robust management sary information for making progress toward the development of IPM pro strategies (Barfield and Stimac, 1979). Second, a critically sparse amount of grams for peanuts. Lastly, we hope to suggest how features of these IPM pro information exists on the ecology of many peanut insect pests, and this paucity grams might vary geographically. Perhaps the initial step should be a concep has hampered development of better IPM programs for peanuts in many in tual model of the basic features of the peanut agroecosystem. stances. The in-depth discussions of a few well-known species in each of the aforementioned pest groups will provide the evidence that vital ecological and CONCEPTUALIZATION OF THE PEANUT SYSTEM biological information is missing. A final summary will address methodology for overcoming these deficiencies. Toward this end, crop-pest relationships St.ima~ and B~rfield ( 1979) pr~enced a concept_ual model of spatial and pest and pest status categories are explored next to provide a conceptual framework species hierarchies for soybean which may be applicable to peanuts. Using this for discussing specific arthropod pests and pest groups. · conceptual model, we can visualize an analogous spatial hierarchy of how in sect pests may arrive in a peanut field. We can then separate pests into pest CROP-PEST RELATIONSHIPS hierarchies and focus on how insects interact with both the peanut plant and other pests. Within a field, various pests (some ofwhich are insects) attack dif The central issue in the design of crop protection strategies focuses on 3 crit ferent peanut plant parts; further, these pares may be attacked at various times ical questions: (1) when is the plant really susceptible to irrecoverable dam in a particular growing season. To design management strategies for economi age?; (2) how much damage does it take to cause true economic loss?; and (3) cally and environmentally sound production of peanuts, we must focus on 5 as which organisms (singly or in combination) are capable of inflicting true eco pects of a given peanut field. First, identify what parts of the peanut plant are nomic damage? Sufficient data exist for us to explore these questions within available for attack, and what magnitude and timing of attack is needed to re the peanut agroecosystem. To accomplish thi!f exploration, we must move duce yields significantly. Second, focus on the characteristics (behavior and bi beyond general definitions of the economics ofcrop-pest interactions to the de tails of specific experimentation on peanuts. ology) of select pests capab~e of inflicting such damage. Third, identify exactly how these select pests mfhct damage, and what can be done to alleviate such By definition, an insect is considered a pest when ics feeding either directly or indirectly causes economic loss. Such loss resulcs from the ecological syn- 254 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 255 chrony in time and space between specific insect pest populations ansl suscepti same damage will result in significant yield reduction. Several relationships ble crop plants. A more or less static pest density may inflict varying degrees of are depicted in Figure lA (tolerance). Generally, the plant beeomes less toler damage, dependent upon the age of the plant when attacked. The ability of a ant in time to nut feeders. The inverse relationship (more tolerant in time) is plant to withstand injury is related to physiological mechanisms (and resulting also possible (Figure IA-seedling feeder). An insect that injures young peanuts morphologies) dictated by plant age. The relationship between plant age and but cannot injure older, mature plants is an example of a case where the plant is the degree of reaction to injury is of paramount importance in understanding more tolerant in time. A third generalized relationship involves pests which how peanuts and various pests interrelate. This age-injury relationship is consume foliage (Figure IA-foliage feeder). The plant is less tolerant to defoli termed temporal tolerance. ation toward mid-season; it is more tolerant of such damage in early and late Temporal tolerance can be expressed by both gradual and abrupt changes in season. plant phenology. A gradual change (e.g., seed maturity) may cause exposure Another aspect of tolerance is related to the plant part subjected to pest to damage over relatively long intervals of time. A more abrupt change (e.g., damage. Pest injury to harvestable plant parts (pods) usually causes a more se pod appearance) occurs over a much shorter time interval, and insects feeding vere reduction in yield than injury by a defoliator. Pests feeding on pods have a exclusively on pods cannot inflict damage until after this change occurs. In much more direct relationship between damage and yield loss than do defolia gradually maturing plant parts, the amount of real damage inflicted by insects tors whose damage is filtered through plant photosynthetic and partitioning is related to the plant's capability of producing those parts. At certain plant mechanisms. Simply, the plant has a greater propensity for recovering from fo ages, what appears to be significant damage can, in actuality, be replaced by liage loss than from pod loss. The ability of the peanut plant to withstand dam the plant with nonsignificant or no yield reduction. At other plant ages, the age, while not reducing yield significantly, is thus related to the distance be tween the inflicted damage sites and the harvestable sites. That distance is 0 A. TOLERANCE when pods are damaged; thus, maximum loss occurs (eaten pods cannot be harvested). This injury site-recovery ability relationship is termed spatial toler ance. In instances where secondary microbial infection or disease transmission occur as a result of insect damage, the concept of sparial tolerance is modified because effects ofsuch infections are realized through internal plant physiolog ical processes. However, when damage is a direct result of insect feeding, spa tial tolerance is a valid general concept. Various published investigations substantiate the general concepts of tem B. FRUITING ~l.00 poral and spatial tolerance. Williams et al. (1976) reported differential effects ~ .75 of foliage and pod removal in both yield and growth rate of specific plant parts. .50 Plant response to 50 and 75% leaf removal was dependent upon the age at ~ which defoliation was imposed. Pod removal did not change total growth rate; l2: however, yield was obviously reduced. In an effort to quantify the effects of de foliation and disease (Cercospora sp.) on peanut plant canopy photosynthesis, Boote et al. ( 1980) demonstrated that the zone (upper, middle, lower) of can opy damage was important in understanding the peanut plant's reaction to specific injury. The ability of peanuts to recover from various amounts of foliar damage, depending on the plant age at which damage was imposed, was quan tified by Jones et al. (1982). These investigators provided quantification of the temporal tolerance of peanuts. Further, they measured how the plant re sponded in growth, photosynthesis, respiration and yield to impositions of damage. Their approach differs markedly from other studies which imposed damage and measured only yield. This deviation is critically important for gaining greater insight into crop protection schemes aimed at providing pro tection only when needed. Quantitative descriptions of concepts like temporal and spatial tolerance are not merely useful; they are critical to understanding a crop-pest system, and such understanding is essential to economically efficient protection of the crop. 0 30 ~ QO 120 At least 2 commercial types of peanuts are identifiable: runner (prostrate) Pt.ANTING INITIAL IMMATURE MATURING MAI~ HAIMST FLOWER FRUIT FRUIT FRUIT (90'r. MATln!TV) and spanish (bunch). Evidence on phenological events and specifics of growth, photosynthesis and response to damage is available for both types. Obviously, ·Fig. 1. Spanish peanut phenology and pest tolerance. \ . 256 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 257
specific growth patterns and responses to insect (and other) damage are condi mum rate of oil synthesis requires an additional 32 days, then the calculated tioned by local agronomic practices and physical environment. Yer/published peak of oil synthesis is ca. 87 days. The production of lipids represents an information (e.g., McCloud, 1974; Williams et al., 1975; Williams, 1979) energy sink whereby more energy is necessary from photosynthesis to produce reveals few real differences in the phenological sequence between runner and lipids for the seed (ca. 60% oil) than to produce carbohydrates for general spanish type peanuts. We will attempt to describe generally the phenology of plant and pod growth. This high energy requirement for oil synthesis is re peanuts so as to understand when plant pares are available for insect attack and flected in the foliage growth curve and experimentally verified by the excess fo when damage to these respective plant parts is meaningful, rather than concen liage curve (Figure IC). This compilation of certain gross physiological and trate on minor differences between spanish and runner peanuts. morphological processes, coupled with the period of plant growth and devel Certain physiological and morphological events inherent to development of opment when these processes occur, aids in identifying critical damage win peanuts provide the template for the relationship between insect damage, dows for deployment of crop protection tactics. plant growth, and yield. Progressive changes in some of these events are de Requisites for arthropod pest status include synchrony with a susceptible picted qualitatively (Figure lD; adapted from Schenk, 1961) for a spanish type plant growth stage, and population density or feeding voracity sufficient to in peanut. Seedlings emerge in ca. 7 days, and the onset of flowering begins at ca. flict injury for which the plant cannot compensate; thus, yield is reduced. De 30 days. Pod formation and seed genesis occur at ca. 45 days, with a maximum fining an economically damaging population density is difficult, but is ap % mature seed at ca. 120 days (maturity). A quantitative description of leaf proached by weighing the monetary crop loss due co insect damage against the area growth (Figure lC; Smith & Barfield, unpublished) and fruit production cost of control. For an economic benefit to be obtained, the predicted monetary (Figure lB; adapted from Gilman, 1975) is shown. Data on flowering and ma loss must exceed the cost of control (Smith and Holloway, 1979; Berberet et turity (Gilman, 1975) and peg penetration (Smith, 1950) help define and ex al., 1979a). The economic threshold (Stern et al., 1959) is the pest density at. plain pest relationships with regard to plant phenology. which control measures should be applied to maintain an economic advantage. Flower production begins ca. 30 days from planting with aerial peg penetra This is an extremely variable value, subject to changes in commodity value, tion of the soil occurring ca. 10 days after flowering (Smith, 1950). Since the control cost, pest density, local climatic condition, etc. As we have pointed peanut is an indeterminate fruiting plant (i.e., fruits continuously until cli out here, the economic threshold is a function of plant age. The concept, how mate terminates growth), it is imperative to determine the plant age where ever, is important in pest management as it requires knowledge of numerous most harvestable fruits arise. Data (Figure lB) show that pods mature to har facets regarding the particular pest and its relation to the plant. vestable yield after 120 days. This time appears to be an asymptote for maturi Economic thresholds usually are established on a regional basis and fre ty. Mature, 120-day old seed would have had to penetrate the soil by day 70 ., quently revised. Frequent revision of economic thresholds appears co be the since ca. 50 days are required for seed maturity after penetration into the soil state of the art as insufficient data are available to either evaluate local thresh (Schenk, 1961). Correspondingly, precursor flowers were produced prior to olds or develop tools (e.g., systems models) to predict dynamic thresholds as day 60. In summary, mature pods harvested at 120 days arose from flowers functions ofa vector of input variables such as local weather, market value, pest produced during days 30-60 and pegs which penetrated the soil from days 40- density and crop age. One of the major problems facing agriculturists, includ 70 (Figure lB). The plant is most sensitive to pests feeding on pod precursors ing those working on peanuts, is precisely how to arrive at dynamic, realistic during the 40-70 day period, since any pod formation occurring after this date damage thresholds. Barfield and Stimac ( 1980) argue that systems modeling does not contribute to the harvested product (assuming the crop is harvested at appears currently to be the most viable tool for accomplishing this goal. 120 days). Leaves provide energy to the plant through photosynthate production. Leaf PEST STATUS area peaks at 80-85 days (Smith and Barfield, unpublished) which corresponds to the time when the plant is most sensitive to defoliation (Figure IC, Boote et Pest status is an important concept relative to understanding and develop al., 1980; Jones et al., 1982; Smith and Barfield, unpublished). Sensitivity of ing pest management strategies. A phytophagous arthropod may be classified the plant to defoliation has been derived experimentally from defoliation as a key, occasional, secondary or non-pest species. Properly conceived man experiments and expressed as proportion excess foliage. These 2 curves show agement strategies are focused on the key pest (Smith and van den Bosch, that the plant has produced peak foliage area at the same time that foliage is 1967; Bottrell, 1979). A key pest usually causes copsistent economic damage needed most. Results in Boote et al. ( 1980) and) ones et al. ( 1982) show sim annually; whereas, an occasional pest causes economic damage at irregular and ilar results under different environmental settings. Similar experimental de unpredictable intervals, usually not annually. A secondary pest is a species signs have yielded analogous information from other crop systems (e.g., In which originally was an occasional or non-pest species whose status has gram et al., 1981). The important physiological event of seed maturity (filling changed, for some time interval, to that of a key pest. This change in status us with oil) is thus coincident with peak foliage production (Figure lC,D). Oil ually results from major man-induced changes in the agroecosystem such as synthesis begins in spanish peanuts when the seed is 14 days old and continues crop varietal changes, pesticide use, establishment of pest alternate host until the seed is ca. 50 days old (Schenk, 1961). If the midpoint for peg pene plants, changes in planting date, etc. Non-pest species, historically, have nev tration is considered as 55 days from planting, and the midpoint for the maxi- er reached economically damaging levels; however, many of these are poten tially economically injurious (Smith and Jackson, 1975). 258 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OP PREHARVEST INSECTS 259
Management strategies usually are directed at the key pest(s), attempt~ng to in combination with fungicides, created conditions conducive to spider mite maintain the population density below the economic threshold. It is impera outbreaks resulting in a change in pest status from non-pest to secondary pest tive that management tactics directed at the key pest do not disturb the extant (Smith and Hoelscher, 1975a; Smith and Jackson, 1975; Campbell, 1978). natural balance that maintains occasional and non-pest species below economi The specific mechanism(s) involved with spider mite outbreaks cannot be cally damaging levels. Thus, tactics aimed at key pests also must consider oth elucidated individually; however, population outbreaks are due most probably er phytophagous arthropods in the agroecosystem (Barfield and Stimac, 1980). to a combination of events that, when occurring simultaneously or in close Pest status can be tempered by regional environmental conditions. The less temporal proximity, release the spider mite population restraining mecha er cornstalk borer, Elasmopalpus /ignose//11s (Zeller) and southern corn root nisms. Several factors have been identified which contribute to spider mite worm, Diabrotica 11ndecimp11nctata howardi Barber, are key pests of peanuts in population outbreaks. Fungicides can destroy the mite parasitic fungi, Ento the United States, whose pest status in the 3 major peanut production regions mophthora sp., that help regulate mite densities and thus contribute to out (southwest, southeast and Virginia-Carolina) is regulated by climatic and breaks (Campbell, 1978). Insecticides can reduce arthropod natural enemies of edaphic conditions. mites, as well as possibly create physiological changes in the mites themselves. The severity of damage by the E. /ignose//us is related directly to (although Hot, dry climatic conditions also contribute to parameters which cause popu not restricted to) a combination of deep sandy soils and low rainfall (Luginbill lation change. Mite developmental time is much shorter at high temperatures, and Ainslie, 1917; King et al., 1961; Walton et al., 1964; Smith, 1981.). In resulting in an increased net production of new individuals. Dry microclimates the southwest where the growing seasons are characteristically hot and dry, prevent fungal spores from germinating. Regardless, heavy pesticide use must soils are deep sands and irrigation is limited; E. /ignose//m is the key pest and have contributed drastically to the change in pest status ofspider mites on pea annually causes severe economic damage (Berberet et al., 1979a; Smith and nuts, because relaxation of these disruptive practices has resulted in reverting Holloway, 1979). In contrast, in the soucheast and Virginia-Carolina where the pests' status back to the original classification in Texas (Smith and the rainfall is normally more abundant and soils are heavier, the severity of E. Hoelscher, 1975a). Peanut grower acceptance in Texas of a pest management lignose//11s is related to a combination oflight soils and length ofdroughts (Lun program which has as one tactic a selective insecticidal application technique ginbill and Ainslie, 1917; Leuck, 1966; French, 1971). In normal rainfall which reduces the number of applications and the exposure of nontarget spe years, E. /ignose//11s is an occasional pest since rainfall tends to suppress popula cies to insecticides (Smith and Hoelscher, 1975b; Smith and Jackson, 1975), tion outbreaks; however, with prolonged droughts, the status may change to has resulted in spider mites presently being reclassified as non-pests. key pest. ,l Climatic and edaphic factors favoring population growth of D. 11ndecimp11nc ·~ ARTHROPOD PESTS tata howardi are antithetical to factors favoring E. /ignose//us. Although D. un decimpunctata howardi damage is not restricted to certain soil types, it is more Phytophagous arthropods reported to attack peanuts worldwide fall into 3 likely to be severe where peanuts are grown in heavier, poorly drained soils classes: Arachnida, Diplopoda, and Insecta. Further, these arthropods occupy (Grayson and Poos, 1947; Fronk, 1950). Diabrotica 11ndecimp11nctata howardi at least 2 distinctly different habitats (foliage and soil) which are of paramount oviposition, egg eclosion, larval survival and adult longevity are enhanced by importance in the design of management strategies. Identification of pest hab relative humidities in excess of 75% (Arant, 1929; Campbell and Emery, itat provides ecological insight crucial to management activities such as (1) as 1967). High rainfall and medium textured soils, which result in the moist certaining relationships between particular pests and complexes of natural ene soils necessary for enhancing population growth, are characteristic of the Vir mies within distinct habitats (Johnson and Smith, 1981), (2} directing partic ginia-Carolina area and certain areas of the southeast where D. 11ndecimp11nctata ular management tactics (e.g., an insecticide) at a particular habitat so as to howardi is a key pest (Miller, 1943; Hays and Morgan, 1965; Campbell and minimize deleterious side effects within the entire system (Smith and Jackson, Emery, 1967; Chalfant and Mitchell, 1967). In contrast, it is only an occasion 1975), and (3) developing relevant sampling plans (allocation, unit size, al pest in the southwest (King er al., 1961; Smith and Jackson, 1975). numbers) for ascertaining pest densities (Southwood, 1978; Jones and Bass, The vast majority of phytophagous arthropods inhabiting peanut fields are 1979). Such ecological knowledge is consistent with the methodologies pre occasional and non-pest species (Smith and Jackson, 1975). Populations ofar sented by Barfield and Stimac ( 1980) toward design of reliable 'management thropods in these classifications usually are maintained bel2w damaging levels strategies for a myriacl, of pests. by climatic factors and/or natural enemies. Biologically perturbing factors, This section has been designed to illustrate the diversity of pests attacking such as unnecessary or non-selective insecticide applications, can disturb the peanuts. Sufficient space and knowledge of pest bionomics are not available balance between the natural regulating agent and pest, and create conditions here to develop the biology, natural history,· damage caused, and tactics usable conducive for outbreaks of occasional or non-pest species. The twospotted against every pest known (or reported) to attack peanuts. This problem has spider mite, Tetranychus urticae Koch, and carmine spider mite, T. cinnebari been addressed herein by presenting a tremendous volume of information in nus(Boisduval), provide excellent examples of peanut non-pests changing sta tabular form with a relevant literature citation(s) to guide the reader to infor tus. Prior to 1970, these mites were considered non-pest species on peanuts mation sources. Further, specific examples of pests have been selected for con throughout the southern United States peanut belt (King et al., 1961; Smith centration on varied biologies and natural histories while illustrating manage and Jackson, 1975; Campbell, 1978). The heavy use of insecticides alone and ment practices from around the world. This should result in an appreciation for 260 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 261 the diversity of pests which attack peanuts and the various ways with which Table I (Continued) each may be managed or controlled. An in-depth look at where IPM is relevant Haplothysanus 011bang11im1is Africa s Pierrard 1968 Pierrard to peanut insects worldwide will be summarized in rhe last section of this chap- Ptri4ontopygt 'onani Senegal s Gillier 1976 ter. Ptrithntopyge pwplitata Africa s R.oubaud 1916 A list of arthropods attacking peanuts worldwide has been compiled (Table Ptrid4ntopygt rubtscms Senegal s Gillier 1976 1). Classes, orders, families, genera and species (unless unavailable) of peanut Ptri"4ntopygt 1'ho111etkni Attem Africa s Pierrard 1968 s Raheja 19n pests are provided. Geographical distribution is listed as well as habitat occu- Pwid4111opygt sp. Africa Ptrid4111opygt spinosissirna Nigeria s Misari 1975 pied within the peanut field. References provided are to the earliest or best (Silvestri) available source on the biology of each particular pest and are intended to pro- Symks~gmus mimuwi Senegal s Gillier 1976 vide a checklist to facilitate entry into the massive literature on peanut pests. Tibi011J1'J ambit111 (Am:ms) Africa s Pierrard 1969 Many of these references provide more localized distribution maps and, to T ibiom11s gossypii Pierrard Africa s Pierrard 1969 some extent, may deal with management tactics available for that particular Oass: lnsecta pest. Order: Orthoptera Table 1. A world list of arthropods attacking preharvest peanuts, Ar«his hypogtUa L. Family: Tetrigidae Paratttix tarinat11s Kirby Nigeria F Yayock 1976 Arthropod Distribution Feeding Site' Reference Family: Acrididae Amistylus pairudis H. S. Nigeria F Yayock 1976 Al/Jlrams g11tt11/osa Walker Asia F Cotrercll-Dormer 1941 Class: Arachnida Chondracris roJM DcGeer Japan F Sonon 1940 Order: Acarina Chrotogonm hanipttrtJS Schaurn India F Kevan 1954 Family: Astigmacidae Chralogo111a rotandat11s Kirby Africa F Jepson 1948 Santauania sp. South Africa, USA s Aucarnp 1969, Chralogo1111s smegalnuis Nigeria F Yayock 1976 ( = Calog/yphas) Shew & Beute 1979 Krauss Tyrophag11s sp. South Africa s Aucamp 1969 Chrotogonus trd(hypUrlls India F Srivastava et al. 1965 Family: Eupodidae (Blanchard) Pmthalt11s major(Dugcs) Queensland F Smith 1946 Colemania sphmarioides, Boliver India F Scshagiri Rao 1943 Family: Tetranychidae Conipoda calcarata Saussure Senegal F R.oubaud 1916 Mononych11s planki (McGregor) Brazil F Flechtmann 1968 Kraussaria ang11/ftra (Krauss) Nigeria F Oyidi 1975 Oligonychm prattn1is USA F Smith Meyer 1974 ltx11Jtd migratoria migratorioides Africa, Asia, F Vriiash 1932, (Banks) Reiche and Fairmaire Formosa Jeps Table I (Continued) Table I (Continued) Gryllotalpa eryllotalpa Calio1hrip• f umipmnis Sudan F Clinton 1962 var. cafta Egypt f Serry 1976 (Bagnall and Cameron) G ryllotalpa gryllotalpa L. North Caucasus S/F Shchegolev &. Weroneb 1930 Cali01hrip1 indims Bagnall India, Africa F Corbett 1920, Gryllotalpa htxadactyla Percy Florida S/F Metcalf er al. 1962 Panchabhavi &. SrapttriJCuJ abbnviatm USA S/F Wisecup&. Hayslip 1943 Thimmaiah 1973 Scudder Ca/io1hrip1 suda11tSis Sudan F Schmutrerer 1971 Scapuriscus a>fetus USA S/f Metcalfet al. 1962 (Bagnall and Cameron) Rehn &. Hebard EnnttJlhrip•flawm Moulton B=il F Almeidaetal. 1977 Scapttrisrus vicinus Scudder USA S/F Metcalf et al. 1962 ·l Franklinitlla bispinosa Morgan USA F Morganetal. 1970 Family: Blattidac \',.I Franlelinitlla /usca (Hinds) USA, Brazil f Morganctal. 1970, Blattdla sp. Nigeria f Yayock 1976 (;' Almeida et al. 1965 Order: Dermapcera Frank/initlla 1Kcitltn1ali1 USA F Smith &.Jackson 1975 Family: Labiduridae (Pergande) Anisolabis ( = Eu6ortllia) Israel s Melamcd-Madjar 1971 Franlelinitlla schu/ui (Trybom) Australia F Hill 1975 anm1/ipes (Lucas) Frankliniella tritici (Fitch) USA F Morgan et al. 1970 Eubonllia Jtali Dohrn S. India s Hill 1975 ·.• Haplothrips gal/arum Pricsncr Africa F Nonveiller 1973 Order: lsoptera '· Stirlothrips dorsalis Hood India, Sri Lanka, Kenya F Hill 1975 Family: Termiridae Stlenothrips YNbrocinam Giard Cosmopolitan F Anon. 1977 Allodanterma morogoremiJ Tanzania s Jepson 1948 Seri(Olbrips o«ipitalis Hood Africa F Hill 1975 Harris T amiothrips JistaJis Karny India F Hill 1975 Amitmnes t111mcifer Silvestri Nigeria s Feakin 1973 T amiothrips intonsutumJ Uiel USA F Watson 1923 Ancis1ro1ermes mt T able I (Concinucd) Table I (Continued) C) d1m1 sp. T:inzan1a F Jepson 1948 Apbi1 glyrintJ Matsumura lndoncsin F Rocchan er al. 1978 1\l(11ukz lorn r111r1J Germ:ir Nigeria F Yayock 1976 tlpbi1 go11ypi i Glover Cosmopoliran F Hill 1975 Ntz11 ra pallttlo Table l (Continued) Table I (Continued) Podalgw ( = Crator) amiculm Africa s Roubaud 1916 Apophylia nigrico/liJ Allard Nigeria I.' Yayock 1976 Burmeister Apophyllia murina Gc:ntaecker Rhodesia I.' Jack 1922 Popillia japonira Newman China, Japan, N. America l.'/S Anon. 1952 Barombia htm11ralis Lab. Nigeria F Yayock 1976 Rlxipata magniroriJ Blackburn Australia s Hill 1975 Buphontlla nigrrwio/aua Jacoby Nigeria F Misari 1975 Schizonytha a/ritana Cast. Africa, Sudan, Egypt FIS Roubaud 1916 var. mtlalica SchizonydJa sp. Sudan s Hill 1975 ColaJpis janssmi ( = M""olaspis) Brazil F Almcidactal. 1977, Strigoderma arboricola F. USA s Hill 1975 (Bcchyne) Fcakin 1973 Trinodon( = /s/Xkn)punctiro//is Queensland s Smith 1946 Diabrotira bal1ta1a I..c:Conre Americas SIP Wolfenbarger 1963, Maclure Fcakin 1973 X ylotrufJ lt.111ographa ( =Pby1011U1ra) Bulgaria, USSR F Popov et al. 1972 Israel s Plaur 1975 Sphrigodu globulw Marshal gammd (L.) Tanzania F Jepson 1948 S:;st111es altit'Ollis Marshal EllXlld temtra (Hubner) Bulgaria F Popov et al. 1972 Tanzania F Jepson 1948 Sy11111es D1ap111s Marshal Feltias11btm'11Rt11(F.) USA F Hill 19n Rhodesia s Broad 1966 Systalts sp. HdiuJbi.r annigera (Hubner) Cosmopolitan F Anon. 1952 Africa F Hill 197'.i TridM1UU1JkK111s tkmi111 Hulst Hdiotbb Jipsacea (L.) Bulgaria, USSR F Shchegolev & Weroneb Argentina FIS Brewer&Varas 1973 Order: Lepidoptcra 1928, Popover al. 1972 Family: Limacodidae Hdiotbb jlflligera USSR F Shchegolev & Weroneb Parasa i-inda(Walker) 1928 E. & W. Africa F Hill 197'.i Family: Pyromorphidae Hdio1bi.r p11naigera Walker Asia, Cocos-Keling F Anon. 1977 AtrRflomorpha mnu/a/a ( F.) Is., Australia, Pacific Is., India F Srivastava et al. 1965 Family: Pyralidae N., S. & Central America Elasmopalpus lignosellus (Zeller) HdiorbiJ llimtmJ (F.) USA F Hill 19n N., S. & Cenrral America s Hill 1975, Smith 1980 Hedy/epta ( = Lamprosema) Cov" e c.. '"' ·• .. Htliotbi.r :ua (Boddie) N., S. & Central America F Hill 1975 Mauritius F Dove&Williams 1971 imlica1a (F.) llania «111 (Guenee) China F Wu 1977 Loxosttge s1rit1i,alis L. Mamutra ( = Baratbra) braJsi Table I (Continued) among peanuts, other cultured plants and native vegetation is axiomatic to the Family: Lepridae Senegal s Roubaud 1916 problem. Such interplant movement undoubtedly effects pest dynamics and Family: Empididae Senegal s Roubaud 1916 Family: Plarysmmafid~ subsequent pest status (Stimac and Barfield, 1979; Barfield and Stimac, 1980, Ri111//iasp. Africa s Seeger & Maldague 1960 1981); however, the details of these intricate ecologicaUbiological relation Family: l.auxaniidae ! ' ships are noc dealt with in this section. The lase section of this paper will pro Homonmra sp. Nigeria Yayock 1976 vide the details of what is and is not known about such relationships and will Family; Chloropidae chan a course toward improved management of peanut pests worldwide. First, Hippe/;:w pusio Loew USA s Snoddy er al. 1975 we must provide adequate details about select peanut pests to set the stage for a Pachylophus sp. Nigeria Yayock 1976 Family: Muscid~ compare and contrast approach to IPM worldwide in the peanut agroecosys A1herigona sp. Nigeria Yayock 1976 tem. Order: Hymenoptera Family: Formicidae FOLIAGE INHABITING PESTS Atta (apiguara Goneal ves S. & Cenrral America F Amante 1967 Camponotus ma,11/a111s F. Nigeria Yayock 1976 Dory/111/11/vus Westwood Senegal s Roubaud 1916 Foliage inhabiting phytophagous anhropods may be divided into 2 groups Dory/111 orimtalis Westwood India s Roonwal 1976 according to method offeeding and characteristic injury inflicted to the peanut E.11ponera smnaarensis Mayr Africa s Roubaud 1916 plant. These are ( 1) foliage consumers which remove foliage with mandibulate Musorspp. E. Africa F Hill 1975 mouthpans (orders Orthoptera, Coleoptera, Lepidoptera, Hymenoptera) and Mononwri11m bkolor Emery Africa s Roubaud 1916 So/mops is f11gax 1.atreille USSR s Shchegolev & Weroneb (2) intracellular feeders which extract plant cell contents by aspiration with 1930 piercing-sucking mouthparts (orders Acarina, Thysanoptera, Hemiptera, Ho TelrarilQrimn lation attributes such as developmental time, reproductive rate, consµmption 23 days (3 7. SC). Studies were conducted using artificial diet. Development rate, longevity, propensity co move, and natural enemy induced morcality. and oviposirion studies on excised peanut foliage were reported by Nickle The following examples will serve to highlight the similarities and differences (1976). among foliage feeding lepidopteran pests of peanuts worldwide. Initial infor Feltia subterranea (F.) is a New World pest whose larvae are nocturnal feed mation on the individual species mentioned is referenced in Table 1. ers. The eggs are laid singly on the peanut plant and are often confused with H. Biology. Species in the lepidopteran families Arctiidae, Noctuidae, Py zea eggs. Feltia s11bterranea eggs have 36'-40 longitudinal ribs (Crumb, 1929). ralidae and Gelechiidae constitute the major defoliating pests. Most species are whereas H. zea eggs have fewer. Newly hatched larvae feed on the foliage sim polyphagous and host on a wide range of grasses, legumes and/or ocher planes. ilar to Heliothis spp. Larvae soon become nocturnal feeders and hide in the soil The adults characteristically are highly vagile and may move great distances or trash beneath the plant during the day (Snow and Callahan, 1968). Larval from their pupation sites. feeding damage by the later larval instars of F. subterranea is easily distin The genus Spodoptera contains 5 economically important pest species on pea guished from other defoliators because larvae cut the leaflets off at the petiole nuts: S. frugiperda J. E. Smith, S. exigua (Hubner), S. ornithogalli (Guenee), S. and feed on the excised leaflets on the soil. Leaf stems appear to be the leaflets littoralis (Boisduval), and S. litura (F .)(Brown and Dewhurst, 1975; Smith and snipped off rather than the ragging appearance left by other foliage consumers. Jackson, 1975). The eggs of all 5 species are laid in scale-covered masses either The larval stage develops in ca. 24 days (Snow and Callahan, 1968) with pupa on the peanut foliage and stems, or the vegetation of hose plants adjacent to or tion in the soil. The pupal stage lasts for ca. 16 days at 25 C (Lee and Bass, within peanut fields. Upon hatching, larvae initially are gregarious and skelec 1969). A complete life cycle should rake 33-89 days dependent on ambient onize the leaf surface. Later insrars disperse and become solitary. Larval devel field temperatures (Lee and Bass, 1969). opment requires 2-3 weeks with pupation occurring in the soil. Moths emerge The arctiids-Amsacta moorei (Butler), A. albistriga (Walker}, A. lactinea ca. 1 week after pupation with the number of generations per year changing (Cremer), and Diacrisia obliqua (Walker)--a.re major defoliators in India (Rai, with latitude. In the tropics and subtropics, continuous breeding occurs. De 197 6). Moths of A. moorei begin emerging after the first heavy monsoon show tailed biologies are available for S. frugiperda (Vickery, 1929; Luginbill, er, mate, and oviposit in groups of small rows on the lower surface of the leaves 1928), S. ornithogalli (Crumb, 1929), S. exigua (Wilson, 1932, 1934), S. litto of peanuts and weeds (Ramaswamy et al., 1968). Newly hatched larvae feed ralis (Hill, 1975) and S. lit11ra (Hill, 1975). gregariously during the early instars (Mathur, 1966). Dispersal to solitary Developmental biology of S. frugiperda and S. exigua fed peanut foliage did feeding occurs in approximately the 3rd instar. Pupation occurs both in culti not differ drastically from the general format already given. Larval develop vated fields and land adjacent to cultivated fields (Patel and Patel, 1965). A ment of S. exigua in laboratory experiments was 15 days, and pupal develop portion of the adults emerge after 6-34 days while the remaining complement ment was 7 days. Eighty-three percent of the larvae pupated and 88% of the delays emergence until the onset of the next monsoon (Rai, 1976). The pupae emerged as adults (Verma et al., 1974). Spodoptera frugiperda develop number of generations vary from 1 to 3 dependent upon geographic location mental time (egg to adult) was ca. 25 days when fed peanut leaves (cv. Florun (Singh and Singh, 1956; Bindra and Kittur, 1961; Yadava et al., 1966). The ner) from 45-92 day old plants. However, developmental time increased to 28 ·~ . biologies ofA. albistriga, A. lactinea and D. obliqua are very similar to A . moorei days when leaves from 92-120 day old plants were used (Barfield et al., 1980). with a few minor exceptions (Sen and Makherjee, 1955; Nagarajan et al., Heliothis armigera (Hubner), H. zea (Boddie), and to a lesser extent H. vires 1957;Pandeyetal., 1968;RamaswamyandKuppuswamy, 1973;Rai, 1976). cens (F.) cause severe, but sporadic defoliation. Heliothir armigera is present in Stomopteryx sRbsecivelta (Zeller) eggs are laid 1-2 per leaf and seldom on the the Old World, while H. zea and H. virescens are New World species. Eggs are stem (Rai, 197 6). Newly hatched larvae mine into the seem for 10-15 days be laid singly on the foliage, stems and inflorescences with the newly hatched lar fore pupating inside the leaf or in leaves folded together by the larvae (Krishna vae preferably feeding on leaves in the terminal buds. Larvae are extremely nanda and Kaiwar, 1965). Multiple generations (4-5) occur each year as the variable in color. Larval development on peanut leaves requires 3 5 and 30 days generation time is less than 1 month (Rao er al., 1962; Yang and Liu, 1966; for H. zea at 26 and 30C, respectively (Huffman and Smith, 1979); and 25 Gujrati et al., 197 3). days for H. armigera (Pretorius, 1976). Pupation occurs in the soil with the Other gelechiids, Stegasta bosqueella (Chambers) and S. capitetla (F.), have adult emerging in 8-12 days (Isley, 1935; Prerorius, 1976). The entire life cy been considered pests of peanuts in the New World (Bondar, 1928; Walton cle lasts about 4-6 weeks on peanuts. Mortality and larval developmental time and Matlock, 1959; Briceno, 1971; Wall and Berberet, 1980). Larval feeding increased when larvae feed on peanut foliage as compared to other cultivated is restricted co the unopened leaf buds, which causes the unfolded mature crops (Pretorius, 1976; Huffman and Smith, 1979). leaves to have symmetrical damage on either side of the midrib (Arthur et al., Anticarsia gemmatalis Hubner is a New World pest whose immature stages 1959; Wall and Berberet, 1979). Although this damage may be readily appar feed predominately on legumes (Watson, 1916). A fairly complete literature ent, yield losses resulting from defoliation are questionable (Wall and Ber compilation on this insect can be found in Ford et al. (1975) and Moscardi beret, 1979). ( 1979). Mean developmental periods for most life stages ofA. gemmatalis across Development ofS. bosqReelta eggs requires 66.5 C degree days above 12.2 C a broad range of constant and variable temperatures were derived by Johnson (Wall and Berberet, 1980). Females laid an average of 16 eggs in the labora ( 1980). Mean egg-to-adult development time ranged from 90 days (15.6C) to tory, a figure probably much below that for field populations. Three genera- 274 PEANUT 5CJEN E AND TECI INOLOGY MANAGEMENT OF PREHARVEST INSECTS 275 tions occur annually wirh a generation completed in as few as 23 days when plants were 35 days pose emergence and continuing until 10 da~s prior t? har remperatures are hig h (\' (ca. 4 leaflets/individual). ~ ~ ~ ~ ro oo ~ ~ ~ Smith and Barfield (unpublished) hand defoliated 0, 25, 50, 75 and 100% AGE IN DAYS of rhe leaflets from peanuts (cv. Scarr) at weekly i nrervals beg inning when rhe Fig. 2. Response surface depicting rhe relat.ionship of plane age, % defoliarion and % yield reduction for a spanish peanut vancry. 276 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 277 Consumption of flowers by foliage consumers should have little effect on ent, species-specific parame~ers .. Youn~ larvae consume small 8?1ounts of fo pod production and final yield. Dehiscence and pollination occur almost si liage with foliage consumption increasing as age progi:sses until the popula multaneously with petal expansion in the early morning hours. Breaking the tion begins to pupate. Since pupae do not. feed, the feeding rate then d~cr~es hypanthium near its base 4 hours after anchesis does not prevent fruit develop as an increasing proportion of the population reach the ~upal stage. Variab1hty ment (Smith, 1950). Thus, flower consumption, except during the first 4 in the duration oflarval development (Huffman and ~m1th, 1979) preven~s the hours after anthesis, should not seriously reduce yield. However, inflorescence abrupt cessation of foliage consumption by a population that would be evident consumption by insects prior to hypanthium elongation would prevent polli with individuals. . nation and could be decrimencal to pod set. If insect feeding on inflorescences The phenological stage of the plant available whe_n foliage co?5umpt1?n oc- did prevent normal pod setting, the peanut plant could compensate by produc curs can conversely affect the population dyn~1cs. of cercam d~fohators. ing more flowers (Smith, 1954) as the flowering rate is determined by previous Changes in plant phenology are reflective .o~ physiological p~esses 1~ternal to fruit (pod) setting and pollination. Thus, the peanut plant possesses a mecha the peanut which, in turn, alter the nutrmve value of the ~ohage. D1ffere?ces nism to restore lost pods (or pod precursers) by producing more inflorescences; in fall armyworm population parameters (developmental time, consumption, i.e., physiological tolerance. The damage window is limited to4 hours per day oviposition and longevity) as a function of peanut .plant age consume~ empha and the plant is temporally most sensitive when inflorescences destined to be size the impact of plant phenology on pest dynamics. Spodoptera frugtperda lar come fruit are most prevalent. vae fed peanut leaves (cv. Florunner) from planes 67-92 days old (peak pod set The relevance of insect consumption of peanut flowers can be further devel and onset of pod fill) had higher consumption rates than larvae fed leaves from oped by examining data resulting from cross-pollination by native bees and ho 45-70 (vegetative growth and initial flowering) or 92-120 day old plants (peak neybees (Hammons et al., 1963). Observations revealed bees are most abun pod fill and onset of leaf senescence) (Barfield et al., 1980). However, larvae dant in peanut fields in the early morning when the most efficient natural pol fed leaves from 92-120 day old plants took significantly longer (o: = .05) to lination occurs (Hammons et al., 1963). When plants were caged co prevent complete development to adults. Adult females result~ng from ~he larvae with bee visitation, genetic markers revealed cross-pollination was negligible; the highest consumption rates had a shorter longevity but laid more eggs. while plants available for bee visitation resulted in levels of cross-pollination Leuck and Hammons ( 197 4a) obsecved S. frugiperda foliage feeding was great ranging from 0-2.37% with most cases less than 0. 5% (Leuck and Hammons, est on the most vigorously growing peanut plants, which could have represent- 1965a, b; Hammons and Leuck, 1966; Girardeau et al., 1975). These data ed preference for high nutritive value fol~age_. . . . support the concept chat pollination occurs rapidly with petal expansion Peanut defoliation by insects is dynamic with a multitude of interactive pro (Smith, 1950), as evidenced by the extremely low level of cross-pollination by cesses occurring simultaneously. Practical utility of a model capturing the es bees, and that insect activity, such as flower consumption, would cause only a sence of this process was developed by Smith and Barfield (unpublished data) small yield reduction. Girardeau and Leuck ( 1967), however, found some evi in a Peanut Insect Defoliation Model used in Texas IPM programs for peanuts. dence chat bee tripping of flowers may increase self-pollination and yield in Data from the plant defoliation response surface (Figure 2) were coupled wit~ some varieties. This evidence was not substantial enough for changing the cur age specific feeding rates forlarvae ofS. /rugiperda (Figure 3), S. exigua, S. orm- rent concept that flower consumption by insects does not drastically curb 100 yield. ,...., Insect populations consume foliage at a rate dependent upon species diversi N~ ty, population density and population age distribution. Different pest species ~ 80 have unique foliage consumption rates and resulting coral foliage consump z Q tion. Age distribution of a pest population also governs consumption rate. For ,_ a. FALL ARMYWORM example, S. frugiperda consumes an average of ca. 1.67 cm2/day during the me ~ 60 dium larval stages and ca. 9. 99 cm1/day during the large larval stages (Barfield et al., 1980) with total consumption ca. 100 cm2 (Smith and Barfield, unpub lished). Heliothis zea larvae consume a total of 176 and 195 cm2 foliage at 26 ~ 40 and 30 C, respectively (Huffman and Smith, 1979). while F. subterranea aver ~ age 168 cm2 (Snow and Callahan, 1968). The last 2 instars of F. subterranea and ~_, :::> H. zea consume 73-97% of the total foliage consumed. Daily consumption of ~ 20 peanut (cv. Florunner) foliage by A. gemmatalis across 6 constant temperatures u::> demonstrated that most consumption occurred in the penultimate and ulti mate instar. Consumption by the last instar ranged from 0.46g/larva/day (21 0 C) to 0.04g/larva/day (35 C) (Nickle, 1976). 1 5 10 15 20 25 Average cumulative foliage consumption rares by populations oflepidoprer LARVAL AGE (DAYS) ous larvae shouid follow the general shape of the fall armyworm (S. frugiperda) Fig. 3, Spanish peanut foliage consumption by a larval population of Spodapttra fr11giptrda consumption rate (Figure 3; Smith and Barfield, unpub. data), but with differ- J. E. Smith at 26 C. 279 278 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS thoga/li and H. zea to create a computerized model for predicting yield loss. In sive generations. A nuclear polyhedrosis virus was responsible f9r high levels of puts into the model (from field scouts) include planting date, date of insect larval mortality, killing in excess of 90% in some generations. Total genera sampling, age distribution and density of defoliating larvae by species and de tion real mortality ranged from 88.13 - 99.97%. The impact of natural mor sired prediction interval. The model converts sampling dare to peanut pheno tality is easily visualized when 666, 707 eggs per hectare yield only 3 ,564 large logical time (as chronological time based on planting date), allows the larvae to larvae per hectare (Sears and Smith, 1975). If no mortality had occurred_, t~e grow and defoliate the plant, and predicts estimated yield loss (as% yield loss) large larval population would have excee~ed 5. 5 larvae per row meter, whic~ is at desired time intervals following the sampling date. A maximum 7-day pre damaging at mid-season. Natural mortality, however, reduced the population diction interval from sampling is recommended to reduce error due to changes to less than O. 3 large larvae per row meter which is decidedly not economically in pest density parameters. important. . Management. Natural Enemies. Numerous listings of natural enemies Parasitism of A. gemmatalis larvae on peanuts appears to be extremely low m of lepidopterous defoliators are available. Compiled records are given in Table all reports. Nickle (1976) found only 1. 5% ofA. gemmatali~ ~arvaecollected i_n 2 as a guide to a source; thus, no attempt is made to rewrite a complete list. Al peanuts parasitized, which represented the lowest % pa~1~ism ~f the 6 Lepi though the parasite fauna of foliage consumers is rich, few efforts have been doptera species studied. Berberet (1978a) also reported similar differences be made co evaluate the impact of natural enemies on suppressing foliage consu tween A. gemmatalis and H. zea parasitism in peanuts. Heliothis zea larval para mer populations. Luna ( 1979) and Collins ( 1980) made significant progress in sitism by Microplitis croceipes (Cresson) and Euce~toria "."!'igera (C~quille~) w~ understanding natural mortality due to predators of Lepidoptera in soybeans. 61 and 24%, respectively, whereas A. gemmatalts parasm~m w_as ml._ ~ntzcarst~ The techniques appear usable in peanuts, where many of the same pests (e.g., gemmatalis parasitism was again the lowest of the 10 foliage mhabmng Lepi- A. gemmatalis and H. zea) occur. doptera investigated by Wall and Berberet (1975). . Two studies in the southwestern United Stares (Sears and Smith, 1975; Conservation of extant natural enemies is an important aspect m manage Wall and Berberet, 1975) evaluated the impact of extant natural enemies of ment of foliage consumers (Smith and Hoelscher, 1975a, b; Smith and Jack certain lepidopcerous pests. Mean seasonal lepidopteran larval para son 1975· Wall and Berberec, 1975; Mangold, 1979). The use of biologically· sitism on peanuts in Oklahoma ranged from 1.5-63.3% with the species and sele~tive i~seccicides (e.g., Bacillus thuringiensis Berliner; Sams and Smith, incidence of parasitism(%) as follows: S. bosquee//a (21.4), S.frugiperda (41.8), 1980), ecologically selective application techniques (Smith and Jackson, S. ornithoga//i (63.3), S.exigua (42.9), Strymon me/inus (Hubner) (62. 5), H. zea 1975) and strict adherence to economic thresholds (Smith and Hoelscher, (57.5), Estigmene acrea (Drury) (15.6), F. subterranea (53.8), A. gemmatalis 1975a) helps minimize: ( 1) nontarget pest resurgence by conserving natural (1. 5), Trichoplusia ni (Hubner) (38. 7) and Platynota nigrocervina Walsingham enemies, (2) target and nontarget pest resistance to insecticides by reducing (25.0) (Wall and Berberer, 1975). This 3-yearstudy concluded chat larval par exposure to selection pressures, and (3) changes in pest status. asitism had a definite impact on suppressing foliage consumer populations and Insecticides. Numerous modern insecticides are very effective in con that conservation of natural enemies was necessary in preventing these pests trolling foliage consuming Lepidoptera larvae on peanuts when applied as from achieving key pest status. sprays, baits or dusts (van der Laan and Ankers~it, 1951; Ai:hur et al., 1959; Table 2. Resources for information on natural enemies oflepidopterous foliage consu King et al., 1961; Bastos Cruz et al., 1962; Vmal and SaroJa, 1965; Stoeva, mers on peanuts. 1968; Venkataraman et al., 1970; Morgan and French, 1971; Castro et al., · 1972; Bass and Arant, 1973; Feakin, 1973; Kareem et al., 197~; Hill, 1975; Pest Species References Morgan and Todd, 1975; Sinha et al., 1975; Harvey, 1976; Ra1, 1976; Sada Spodoptera frugiperda Bass& Arant 1973, Wall & Berberec 1975, Ashley 1979 S. txigua Wall& Bcrberet 1975, Rai 1976 kathulla et al., 1978; Urs and Kothai, 1977; Berberet, 1978a, b, 1979; Bass, S. ornithogalli Bomell 1969, Wall&Berberet 1974, Wall&Berberet 1975 1979· Berberet and Guilavogui, 1980; Sams and Smith, 1980). Within the S. litura Rai 1976 New 'world, S. exigua (Brown, 1961; Cobb and Bass, 1975), S. frugiperda Heliothis zu Sears&Smith 1975, Wall&Berberer 1975 (Young, 1979), H. zea (Brown, 1968) and H. virescens (Nemec and Adkisson, H.armigtra Rai 1976, Singh er al. 1978b, Sroeva 1973, Bhatnagar& Davies 1978 1969) have shown high levels of resistance to certain insecticides. Insecticidal Ftltia suhterranu Wall & Berberct 1975 efficacy against certain species may be dependen~ upon the status of re~ional A msaaa 1111/0rti Rai 1976 insecticidal resistance; thus, management strategies dependent upon this tac A. alhimiga Rai 1976, Sundaramurthy er al. 1976 tic must be reviewed frequently. Diacrisia ohliq11a Ramaseshiah 1973, Rai 1976 Several organic and inorganic compounds have adverse effects on foliage S1011111pttryx substt:ivtlla Rai 1976 S1tgasta bosquttlia Badar 19'12, Wall&Berberet 1975 consumer feeding and biology. Inorganic nutrients applied as foliar sprays r~ Anti south of the Sahara, although it has been reported from India, Australia and In- Bottomley, 1928; Evans, 1954; Kousalya et al., 1971), no sexual forms have donesia. _+ been reported (Real, 1955). Aphis craccivora overwinters on numerous unculti Ten aphid species are reported from peanuts (Table 1, Table 3). The role vated host plants (mainly the Leguminosae) with the alate migrants from these each species plays in virus transmission in peanuts is not always known. Aphis sources invading cultivated peanuts (Evans, 1954; Booker, 1963; Adams, craccivora, important in the transmission ofseveral plant viruses, is a key pest of 1967; Davies, 1972). Local overwintering and alternate hosts of A. craccivora, peanuts in several geographical areas and illustrates management primarily however, are not sources ofGR V virus (Adams, 1967; Davies, 1972; Gibbons, based on pest ecology; thus, this section will concentrate on that particular 1977). This does not preclude the possibility that wild, uncultivated hosts of aphid. the virus do exist (Rossel, 1977). Table 3. Peanut viruses and arthropod vectors. Pastures planted with Sty/osanthes could be perennial reservoirs of GRV (Okusanya and Watson, 1966; Gibbons, 1977). Dry season maintenance of Disease Vecmr(s) Reference(s) GRV could be volunteer peanuts in East Africa but not in Sudan, Nigeria and Peanut Mortie Aphis cra«iJ10ra Koch Herold& Munz 1969, Malawi (Gibbons, 1977). Viruliferous alates from outside the field provide the MyzNJ pmifat Sulzce Bock 1973, Kuhn & initial inoculum of GRV and are responsible for primary infection (or primary Demski 1975, Paquio &Kuhn 1976 spread) of GRV within the field. These immigrant, alate aphids are transport Aphis gossypii Glovers Behncken 1970 ed on weather fronts (Davies, 1972). Amphorophora lamJtat (L.) Behncken 1970 A higher incidence of primary GRV infection is found in fields with wide Rhopalosiphum padi (Fitch) Behncken 1970 spacings between plants as opposed to closely spaced plantings (Hull, 1964). PcanutStunt Aphis crtl«iwra Tolin er al. 1970, Myz11s pmkm Herberr 1967 Hull (1964) concluded that immigrant alaces were more attracted to sparsely Aphis crilkola ( = spira«ola) Herbert 1967, Feakin 1973 spaced plants due to greater exposure and proliferation of apical buds and Franklin idla /NJra Porrereral. 1975 young leaves. Young and senescent leaves ofsparse plantings increase the expo TomaroSpotced Wilr Sdrtoshrips ursalis Amin et al. 197 8 sure and incidence of the color yellow which in turn increases the alighting re Thrips sobaci Bald&Samuel 1931, Ghanekaret al. 1979 sponse of A. cracdvora alates. The plant parts appearing yellow are fewer and Franklinitlla sch11/ui Bald&Samuel 1931 camouflaged by green mature leaves in close plantings. A'Brook (1968) ex Franklin it/la /111ca Bald&Samuel 1931 panded this working hypothesis to include an additional increased alighting Franklinitlla ocridtnralis Bald & Samuel 1931 response from the contrast ofplants and soil, the optomotor response, as well as Groundnut Rosetrc Apbis t:ra«iwra Davies 1972 color attractiveness for sparsely spaced plants. Thus, high density plantings are Aphis grmypii Adams 1967 Peanut Green Aphis go1sypii Reddy' 1980 not attractive to flying aphids searching for a host plant, as they do not receive Mosaic Myzuspmkm Reddy 1980 the correct visual cues (yellow color and contrast between bare earth and Groundnut Eyespor Aphis cra«iwra Dubern & Dollet 1978 plants). Rugose Leaf Curl A111troagallia torrida Evans Grylls 1954 Secondary spread of GRV is attributed to both alate and apterous aphid (a leafhopper) forms produced from initial colonization within the field (Storey and Ryland, 1955). Widely spaced plantings result in higher aphid populations (Farrell, Biology. Mose aphids overwinter in the egg stage which gives cise to an 1976b) which in turn enhances the very sensitive crowding stimulus for alate immature female in the spring. The immature female moles several times and production in A. craccivora (Johnson, 1965) with 50% alates produced in pop develops into an adult, winged, parthenogenetic, ovoviviparous female known ulations of ca. 200 aphids/m2 (Farrell, 1976b). Crowding could increase secon as the fundatrix (Eastop, 1977). The fundatrix may give birth to several dary spread through the increased production of alates which emigrate to re hundred females which also are parthogenetic and may be either alate (winged) duce the population densiry. However, apterous forms infected 4 times as or apterous. The progeny of the fundatrix are the spring migrants' (alates) · many plants adjacent to the GRV inoculum source as alates (Storey and Ry which disperse from the original host plant. In most aphids, a winged genera land, 1955). Gibbons ( 1977) reported alates in Nigeria and apterae in Uganda tion is followed by several wingless generations. Overcrowding or unsatisfac and Malawi most responsible for secondary spread of GRV. cory condition of the host plant may cause production of winged forms. Shorter The rate of increase of A. craccivora is lower in closely spaced peanut plants daylengchs in the autumn induce the production of sexuparae which, in turn, (Farrell;- 1976b). This phenomenon has been attributed to a lower level of nu produce a single generation of sexuals (males and sexually reproducing fe trition in the densely planted fields (Real, 1955; Farrell, 1976b). Waghray males). After mating, the female lays eggs which overwinter (Eastop, 1977). and Singh ( 1965) showed decreased fecundiry in A. cracdvora with low levels of Variation in polymorphism is common in aphids and results from external sti nitrogen. Therefore, close plantings not only reduceprimaryGRV spread, but muli (temperature, host plant condition, photoperiod, crowding, etc.). These also lessen secondary spread by reducing aphid buildup and decreasing emigra stimuli dictate aphid hormonal balance which, in turn, regulates polymor tion. phism. The GRV is transmitted in a persistent manner by A. cracciV01'a and requires In Africa, where A. craccivora is important as the veccor of GRV (Storey and an acquisition period of ca. 2-3 days (Watson and Okusanya, 1967). Once A. 284 P EANUT SCIENCE AND TECH NOLOGY MANAGEMENT OF PREHARVEST I NSECTS 285 craccivora acquires GRV, rhe virus can remain viable for ar leasr 10 day~ (Srorey .spreading cultivars. Several culrivars showed promise as providing resistant and Ryland, 1955). Borh immarure and adulr srages can vecror GRV (Srorey germplasm. Amin and Mohammad (1980) have also shown greatly reduce? and Ryland, 1955). . . . progeny production by A. a-accivora when i.solared on deta~hed shoots of cultt Damage. AphiJ cracci11ora is considered a key pesr 111 ".'-f'.1ca (Hill, 1?75) vars of ArachiJ chacoeme and A . hypogaea . High levels of resistance to GRV have and certain areas of l ndia (Rai, 1976). In Africa rhe economic 1mporrance is re been reported from a group of cultivars from the Ivory Coast and Upper Volta lated ro transmission ofGRV, whereas in India direct feeding damage as well areas of West Africa (Gibbons and Mercer, 1972; Rossel, 1977). as disease transmission is serious. Cultural. The major management tactics available for use against A . In India, aphids feed on the succulenr vegerarive rips prior ro rhe iniriation craccivora and transmission ofGRV involve sanitation, early planting and uni of fl owering causing leaf curl and srunred growth. Larer rhey migrare ro rhe form, dense plantings of peanuts within a field. Sanitation includes removal of floral shoots, seriously reducing pod formarion (Rangaswany and Rao, 1964). volunteer peanut plants to reduce the GRV inoculum (Gibbons, 1977) and Yield reducrions of 40% are reported (Khan and Husain, 1965) from A. cram- removal of weed hoses (Kousalya et al., 1971). Early planting allows the plant 11ora feeding damage alone. ro partially develop either prior co the arrival of viruliferous alates or prior to Kousalya er al. ( 1967) described primary and secondary spread of a roserre the buildup of aphids locally (temporal colerance) (Booker, 1963; Farrell, disease of groundnurs vecrored by A . craccivora in India. When rhe crop was 15 1976a). This approach also allows the plant co reach a more mature, unattrac days old, 39% of rhe planrs were infested with A . craccivorc1. By 60 ~ays, 88% tive stage and obtain maximum groundcover which tends co reduce primary of rhe planrs were infesred. Maximum disease incidence (0.25% w1~h symp spread of GRV . Plants 20-30 days old may be infested with 3-5 times as many roms) was manifested when the plants were 75 days old. The propom on ofap alate aphids as plants 50-60 days old (Feakin, 1973). Winged aphids invade rerae was 0. 7 2 at 15 days and 0. 94 ar 90 days afrer planring with a posi rive cor the crop in large numbers beginning 50 days after 2. 54 cm of rain has fallen in relarion between % planrs infested ·wirh aphids and disease incidence. the growing season (Feakin, 1973). ln Africa, A. crnccivora is imporranr as rhe vector of GRV (Hill, 1975). · A uniform, close spaced planring drastically reduced the alate A. craccivora Groundnut rosette virus is mani fes ted by 2 main rypes of symptoms: chlororic alighting in the field (A'Brook, 1964; Farrell, 1976c). Close spacing also re and g reen rose tte (Gibbons, 1977; Mercer, 1977): Both cause s.runring ofrhe duces the rate of increase of the colonizing aphids (Farrell, 1976b). Early plan planr with severity of rhe disease related ro the earl iness of rnfecrion. Plan.rs in ting of uniform, closely spaced plants reduces the damage from GRV if the fected early by GRV produce few, if any, pods resulting 1.n v1rrual crop fail ure, plane is infested, reduces primary spread by deterring aphids from alighting whereas aphid conrrol in later infections may increase yield ~y 50% (Eastop, and further reduces secondary spread by reducing aphid reproduction. Densi 1977). Crop loss is related ro rhe incidence of the disease (seventy) and the phe ties of 197 ,600 planes/ha for the late crop and 98,800 planes/ha for the early nological age of the plant when i nfecrion occurs (temporal tolerance). The lacer crop are sacisfacrory in minimizing GRV infection (Feakin, 1973). rhe infection (or rhe closer ro harves t), the less severe the crop loss. Thrips. Seventeen species of Management. Natural Enemies. Numerous general predators are re chrips (Thysanoptera:Thripidae) have ported ro arrack A. rraccivora in India (Patel er al., 1976; Rai, 1976) and USSR been reported feeding on peanuts (Kesten, 1975), mostly from rhe families Coccinellidae (Coleoprera) and Syr worldwide (Table 1). The economics phidae (Diprera). The aphid parasite, Lysif1hleb11J teJtacei/m (Cresson)_. was in of thrips control with insecticides has troduced inro India from the U nired Srares (1966-67) for suppression of A. been a controversial issue for over 30 craccivora (Ramaseshiah et al. , 1968). The parasite reproduced readily during years and still remains unresolved on rhe cooler seasons but was successful at temperatures above 32 Conly when A. a worldwide basis. The role of thrips rraccivora was hosting on OolichoJ lab-lab. A related aphid species, L. /abamm as veccors of peanut diseases (Table 3) (Marshall), parasirizes 85% of the /\. craccivora in June in th~ USSR (~esren , will receive more attention in the 1975). For general information on the impact of narural enemies on aphids, see next decade, but such information, at Hagen and van den Bosch ( 1968). . . . present, is scarce. Insecticides. Sprays, granules and dusts are effecnve against A. cram Biology. Thrips found on peanuts are minute (0. 5-2.0 mm long), incon vora in India (Sarup er al. , 1960; Vasanrharaj er al., 1965; Dorge er al., 1966; spicuous insects. Adult females oviposit in the plane tissue and all life stages Gangadharan et al. , 1972) and the U nire~ St a re~ ~S mi~h and. Culp, 1968) .. Da (egg, larva, pupa and adult) are found on the hose plane. Hatching gives rise to vies ( 197 5) reviewed the currenr starus of 1nsccnodcs 111 Africa for A. cramvora 2 successive larval stages which feed on the plant tissue, followed by 2 pupal control and reported on large scale trials with menazon and conventional trials stages which are active but do not feed. All immature stages are wingless but with numerous ocher compounds. - resemble adults otherwise. The metamorphosis of thrips is typical of the pau Resistant Cultivars. Brar and Sandhu ( 1975) evaluated 50 culrivars rometabolous insects whose juvenile stages are called nymphs. The terms larva ( 19 bunch, 16 spreading and 15 semi-spreading) for resistance ro J\. craccivora and pupa usually are used co describe the juveniles of the insects having com in India. Dara on d ifferences in aphid reproduction were significanr wirh the plete metamorphosis; however, the terms larva and pupa are used here to avoid bunch culrivars having lower aphid reproducrion than spreading or semi- confusion with the thrips literature (Lewis, 1973). 286 . PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST INSECTS 287 Developmental time from egg to adult varies with temperature ang species. thrips population decline in the buds (Leuck et al. , 196?; _Tappan an? G~rbet, Adult tobacco thrips, Frankliniella fusca (Hinds), live ca. 30 days and lay 50- 1979). This lag represents the time delay between the mJury occumng m the 60 eggs (Watts, 1934), which require ca. 16 days to hatch during summer unfolded leaflets and its appearance on the exposed leaf surface. . (Watts, 1936). Numerous generations occur on cultivated and wild hosts Information on the role of the thrips as vectors of peanut pathogens is s~e throughout the warm seasons, while in temperature zones thrips generally en (Table 3). Frankliniella fmca has been implicat~d as a v~cto~ of pe~nut stu~t vi ter diapause or quiesence during cold climatic conditions as fully grown larvae rus in Virginia (Porter et al., 1975 ). Several thnps species, mcl_udmg. Thnps ta. or adults (Lewis, 1973). In warmer zones, there is a period of inactivity during baci Lindeman Frankliniella schulzei (Trybom}, F. fmca, Sartothrips dorsa/1s cool periods (Watson, 1918), but breeding is essentially continuous. Hood and F. o;cidentalis (Pergande) vector tomato spotted wilt virus (Bald and Thrips initially move into fields from wild or cultivated hosts. This move Samuel, 1931; Ghanekar et al., 1979; Amin and Mohammed, 1~80; Anan ment is most pronounced in the spring when peanuts are in the seedling stage. thakrishnan, 1980). Thrips are unusual in comparison to most virus vectors Peanuts planted downwind of small grain fields readily obtain high densities of because the larvae must feed on infected plants before either the larva or adult thrips (Smith and Sams, 1977). Thrips emigration from the maturing grain can transmit a virus (Bald and Samuel, 1931; Lewis, 1973). crop in the late spring coincides with the planting time for peanuts. Detrimental effects of direct thrips feeding on plant growth, anthesis, pod Migrant adults oviposit on peanuts and successive generations occur on the yield and seed quality have been a subject of controversy for ~everal decade~. plant throughout the growing season (Tappan and Gorbet, 1979). Populations ' ' The initial reports by Poos (1945) and Poo~ et al. ~1947) provided the.genesis are highest in the terminal buds during the first 30 days after planting (Smith for the thrips control controversy by repomng thrt~s reduced peanut yields up and Sams, 1977; Tappan and Gorbet, 1979) and decline to a lesser density for to 37%. Successive reports by numerous other United States authors (Arant, the remainder of the season. Phenologically, the plant begins to flower at ca. 1954; Arthur and Arant, 1954; Howe and Miller, 1954; Dagger, 1956; Ar 30 days after emergence, which coincides with the thrips population decline in thur and Hyche, 1959; Harding, 1959; King et al., 1961; Morgan et :11·• the terminal buds (Sams and Smith, 1978; Tappan and Gorbet, 1979). Ham 1970· Smith 197 lb 1972a; Minton and Morgan, 1974; Sams and· Smith, mons and Leuck ( 1966) reported immature thrips to be predominant in flowers 1978'. Smith ~nd Sam~. 1977; Tappan and Gorbet, 1979, 1981) failed to iden and postulated thrips changed microhabitats and food source, moving from tify y'ield increases by controlling. thrips with .insecticides, even with thrips the terminal buds to flowers with anthesis. Comprehensive data from Tappan populations as high as 50 per cermmal bud (Smith and ~ams, 1977) or 92% of and Gorbet ( 1979) discounted movement of immatures to flowers, since 90% the leaflets damaged (Minton and Morgan, 1974). Thnps damage also has not of the thrips population in terminal buds were immatures throughout the sea been correlated with seed maturity (Sams and Smith, 197 8), rate of flowering son. Correspondingly, 92% of the population in the flowers were adults. The or plant growth (Morgan et al., 1970). Close examination of i:he reports by decline of thrips density in terminal buds at the onset of flowering could result Poos (1945) and Poos et al. (1947) reveal multiple insecticide applications from a dilution of che population from a concentrated few terminal buds early were used over a long period of plant growth which could have suppressed a in the season to the numerous flowers and terminal buds produced later in the nontarget pest species and resulted in spurious concl':15ions rega~ding the_ be season as the planes grow. nefits of thrips control. Existing data clearly show thnps control m the ~ mted Thrips found on peanuts are not all phycophagous; some are predaceous and States is not a sound economic investment (Bass and Arant, 1973; Smith and mycophagous. The most commonly encountered predaceous thrips is Scolo ;.. ~. .'; Sams, 1977; Tappan and Gorbet, 1979). . . . thrips sexmtKulatui (Pergande), which preys on several species of spider mites. Enneothrips flavens is considered a key peanut pest m Brazil: Control of ~his Scolothrips sexmaculatus rarely is abundant on peanuts, although it is quite com thrips on peanuts up to 60 days after plant emergence result~ m ab~olute yield monly found on mice infested plants (observation of senior author). The bio increases of 790 kg/ha (Almeida et al., 1977) and proportional mcreases of nomics and predaceous habits of S. sexmacrdatus are reviewed by Gilstrap and 39% (Cakagnolo et al., 1974), 50% (AlI?eida et al., 1965J and 35% (Lara ~t Oatman (1976). Euphysothrips minozii Bagnall is a mycophagous species that al., 1975). In several instances, pest species other than thnps were ~resent m feeds on spores of peanut rust,Puccinia arachidis Spegazzini, in India (Shanmu these evaluations, but their contribution to yield loss was not readily deter gam et al., 1975). mined. Damage. Adult and larval chrips rasp che leaf tissue and extract the plant In central Africa (Cameroon and Sudan), several species of thrips, Taenio juices, preferring the unfolded, developing leaflets in the terminal buds. Feed I thrips sjostedti (Trybom); Haplothrips gal/arum Friesner; Serkothrips occifital~s ing is manifested as scarring, chlorosis and deformation of the leaflets. Damage i' Hood; Caliothrips sp.; C. s111/anensis (Bagnall and Cameron); and C. fumipenms is visually apparent as the leaves unfold. The plant is considered most suscepti (Bagnall and Cameron}, attack the unfolded leaf resulting in chlorosis and d~ ble to feeding injury by F .fusca in the United States from emergence to 30 days formation on the leaflets (Clinton, 1962; Nonveiller, 1973). The economic old and by Enneothrips flavens Moulton in Brazil up to 60 days old (Batista et importance of thrips on African peanuts is questionable (Hill, 1975). al., 1973). Severity of damage can range from minimum chlorosis and scarring Scirothrips dorsalis Hood and Caliothrips indicm (Bagnall). heavily dama~e to leaflet abscission. Approximately 1 thrips per bud can result in injury to 33- peanut plants in India (Rai, 197 6; Sapathy et al., 1977), causmg !eaf c~loro~is 80% of the leaflets (Tappan and Gorbet, 1979). The number of damaged leaf and abscission. Control of S. dorsali.s and Empoasca sp. resulted ma yield m- lets declines within 1-2 weeks after the onset of flowering, lagging behind the crease of 1536 kg/ha (Saboo and Puri, 1978). . Management. Natural Enemies. Thrips are attacked by both parasttes MANAGEMENT OF PREHARVEST INSECTS 289 288 PEANU:f SCIENCE AND TECHNOLOGY migration into peanut fields which occurs in August and January. Bud necrosis and pr~dacor~. General arthropod predators include anthocorid bugs in the ge incidenc~ is related to. i~migran.t thrips (i.e., secondary spread is unimpor nus Orms w~1ch are cosmopolitan enemies; Geocoris sp. (Lygaeidae), Chrysopa sp. (Chrysop1dae), Hemerobius sp. (Hemerobiidae), in the United States· Cheilo tant)'. High plant den.s1t1es result ma lower percentage ofinfested plants. Early menes vicina Mulsant (Coccinelidae), Ischiodon aegypticus Wied in the planting dates and high plant densities coupled with properly timed insecti (Syrphida~) cide applications and using less susceptible cultivars are effective in manageing Su~an,. and Psallus sp ..<~irid~e) in India (Lewis, 1973). Insects parasitic on bud necrosis disease in India (Amin and Mohammad, 1980). thr~ps mclu~e. the fa?11hes Tnchogrammatidae, Scelionidae and Mymaridae which parasmze thnps eggs, and Eulophidae which attack larvae (Lewis Spider Mites. Several species· of 1973). , soil and foliage inhabiting acarines Insecticides. Thrips are controlled easily with most modern insecti are associated with the peanut plant ~ cides. Granular systemic, spray and dust formulations give highly effective (Table 1). The soil inhabiting astig control (Arthur and Hyche, 1959; Castro et al., 1972; Smith, 1972a; Bass and matids have been implicated in the spread and increase of soil borne fun M Arant, 1973; Nonveiller, 1973; Minton and Morgan, 1974; Almeida et al., 1977; Mateus and Gravena, 1977; Rensi et al., 1977· Saboo and Puri 1978· gal diseases of peanut pods (Aucamp, Sams and Smith, 1978; Rohlfs and Bass, 1980). Cam~bell et al. (1976). how~ 1969; Shew and Beute, 1979), whereas foliage feeding by the tetra ever, ha~e shown insecticidal performance is not independent of peanut vari ety. Their research produced results which allude co unquantified interactions nychids causes leaf chlorosis and defo liation. The soil inhabiting astigmat ber~een soil applied sysce~ic insecticides and cultivars with different growth habits (bunch, runner and intermediate growth types). Insecticides that gave ids are presented in this section rather 90% thrips control on one cultivar (or growth type) gave ca. 50% control on than the soil pest section to maintain another. biological continuity of pest groups. Resistant Cultivars. Thrips resistant peanut cultivars have been iden Biology. Tetranychid mites develop through the metamorphic stages of tified in the United States (reviewed by Smith, 1980) and in India(Panchabha egg, larva, protonymph, deutonymph and adult (van de Vrie ec al., 1972). vi and.Thimmaiah, 1973). Resistance to F. fusca in the virginia, spanish and The larval stage has 6 legs, and the other stages have 8 legs. Variations in the valenc1a type peanuts has been verified by both laboratory and field evalua dev~lopmental stages may occur among mite families. For example, the eu tions. Collectively, the spanish types have the most resistant germplasm. The pod1d, Penthaleus major (Duges), has a deutovum (prelarval) stage. Length of spanish culrivars Starr and Argentine are immediate sources of agronomically the life cycle is correlated with temperature, humidity and host plant quality suitable resistant germplasm (Leuck et al., 1967; Young et al., 1972; Kinzer (Watson, 1964; van de Vrie et al., 1972; Jeppson et al., 1975). Copulation of adults occurs immediately after hatching of the female with et al., 1973). Plant Introduction 280688, a valencia type cultivar, is non-pre ferred for larval and adult feeding, has low larval survival (antibiosis) and low diploid eggs giving rise to female progeny and haploid eggs yielding male pro geny (van de Vrie et al., 1972). Tetranychus urticae (Koch) females produce le~el~ ~foviposition (Kinzer et al., 1972; Kinzer et al., 1973). Cultivars NC 6, from 42-204 eggs per female depending upon host plant (van de Vrie et al., ~1rgin1a Bunch 67 and Pl 290599 are sources of resistant germplasm for virgi 1972) and moisture (Boudreaux, 1958). nia type peanuts (Leuck et al., 1967; Young et al., 1972; Campbell et al., . The total life cycle for T. urticae females is 8-12 days at 30-32 C. Females 1977; Campbell and Wynne, 1980). Resistance co Ca/iothrips indicus(Bagnall) live ca. 30 days and lay 90-110 eggs Qeppson et al., 1975). Thus spider mites was identified in several cultivars: 21008, 21016, HG-10, 21018 and EC- have a tremendous capacity for increase and can rapidly produce enormous 20979 (Panchabhavi and Thimmaiah, 197 3). Cultivars 21008 and 21016 population densities if conditions are optimal. were considered the most resistant, with ca. 23% of the leaves showing no The dormant stage which passes through adverse environmental conditions damage as compared to 1.39% for the cultivar commonly under cultivation. is the mated, diapausing female which does not feed until the adverse condi Fecundity of F'. schultzei (expressed as eggs/female/24 hours) was greatly re tions .ease (van de Vrie et al., 19_7~). Factors inducing diapause include pho duced on Arachu chacoeme(0.0), A. glabrata (0.0)and A. duranensis(0.4) com topenod, temperature and nutrmon (Parr and Hussey, 1966). Tetranychid pared to A. hypogaea (4.4) (Amin and Mohammad, 1980). ~ttes are less host specific than other mite families, as most species have a rela C~tU:al· Several cultural practices are conducive to thrips manage tively narrow host range. Telranychus cinnebarinus (Boisduval), T. urticae and T. men~. S~01tat1on of ear~y v~lunceer see~lings in the spring is important in pre 1urkes1ani (Ugarov and Nikolski) have a wide host range including many plant venting infield population mcreases prior to planting the crop (Bass and Arant, genera Qeppson et al., 1975). 19~3?· Peanut fiel?s planted a~jacenr to winter small grains, especially when 1:he astigmati~ biologies are poorly known with the exception of species in posmoned downwmd, are available for migrant adults when the grain begins festing ~cored grains (Hu~hes, 19~ 1). Caloglyphus life cycle requires 8-9 days at to mature (Smith and Sams, 1977). 22 C with adequate moisture with females laying ca. 200 eggs in 24 hours In India •. ea~ly planted peanuts usually escape heavy losses from thrips-born (Hughes, 1961). The biology of P. major, Eupodidae, is summarized by Jep bud necrosis disease caused by tomato spotted wilt virus. Infection levels are pson et al. ( 1975) and Smith ( 1946). lowest in peanuts planted at least 6 weeks prior to peak thrips (F. schultzei) im- 290 PEANUT SCIENCE AND T ECHNOLOGY MAN AGEMENT O F PREHJ\RVEST I NSECTS 29 1 D a mage. Tcrranychid mites ft:ed by penetrating the plant tissues and are cons idered important predarors of spider mires. The coccinellid genus, Ste removing the cell contents. Feeding causes the chloroplast ro disappear with rhorm, is restricted ro mire predation and has been found in peanut fields in the small amount of remaining cellular material coagulating ro fo rm an am ber T exas (observation of senior author). Ocher insect predarors and their impacr mass (Jeppson er al., 1975). Continued feeding results in rhe fo rmation of ir on spider mires are reviewed by H uffaker er al. ( 1970), McMumy er al. ( 1970) regular ch lororic spots on the leaf ri ssue typical of spider mire damage. 1njec and van de Vrie er al. (1972). rion of roxins or g rowth regularors by spider mires was questioned by J eppson I nsecticides. Spray and g ranular formulations of insecticides and er al. (1975), although the reactions of specific plant species ro feeding by rhe acaricides effective against Tetranyrhm on peanuts have been reported from same spider mite species may drastically differ. The severity of plant damage Egypt (Arriah and Rizk, 197 3; Osman and Rasmy, 1976), 1ndia (Gupta er al., resulting from tecranych id feeding is related ro crop species, mire density, lo 1969), Pakistan (Moiz and Qureshi, 1969). Bulgaria (Atanasov, 197 1) and the cal environmental conditions, plant nutrition, moisture stress and phenolog i United Stares (Campbell er al. , 1974; Smith, 1976a; Smith and Mozingo, ca l growth srage (Watson, 1964; van de Vrie et al. , 1972). Spider mires on 1976, 1977). peanuts are especially destructive during hoc, dry weather (Campbell et al., A preponderance of evidence exists ro implicate pesticides (insecticides, 1974; Osman and Abdcl-Faccah , 1975). Significant mite densities may kill acaricides and fungicides) as agents inducing spider mire outbreaks (Teich , large areas of plants (Smith and J ackson, 1975). resulting in considerable yield 1969; van de Vrie et al., 1972; Smith and Jackson, 1975; Campbell, 1978). loss. The change in spider m ire pest status on peanuts in the United Stares (from The asrig marid genera, Tyrophc1gm and CaloglyphllS, were isolated frequent nonpest ro secondary pest) was discussed earlier under Pest Status. This change ly from rhe subterranean parts of the peanut plants in South Africa (Aucam p, in pest status has nor been resrricred reg ionally co the U nired Stares (Smith and 1969). Caloglyph11S micheali and ocher Ca/oglyph11S species were isolated from Jackson, 1975; Campbell, 1978) bur geographicall y encompasses worldwide field soil , decaying pods and rhe root and pod zone of healthy plants in rhe peanut production as evidenced by reports from Pakistan (Mioz and Q ureshi, Uni red Stares (Shew and Beute, 1979). CaloglyjJhm are capable of acting as a 1969), 1ndia (Gupta and Sandhu, 1969; Gupta er al., 1969), Israel (Teich, disseminating agent for the AspergillllS ffav11s (Link) and Pythi11111 111yrioty/11111 1969) and Bulgaria (Aranasov, 197 1). The application of certain insecticides Drechsler fungi by internal contamination of gur contents wirh viable spores in and fung icides in combination fu rther exacerbates the spider mire outbreak defecated fecal pellets (Aucamp, 1969; Shew and Beute, 1979). Caloglyplms l . phenomenon. Campbell ( 1978) reported multiple applications of carbaryl + spp. in South Africa were reported as feeding on the peanut pods and seed, as benom yl ro peanuts resulted in a mire density increase of 344X over the densi well as on fungi (Aucamp, 1969); whereas C. mirhecdi was reported as rotally ty of the uncreated control. T he increased use and dependence upon agricultu m~1co phagous (Shew and Beute, 1979), acting wholly as a fungal disseminat ra l chemicals as the sole management racric fo r peanut pest suppression in ing and nor a wounding agent. These mires can spread fung i, bur fo r infections many production areas should continue ro exacerbate the spider mire problem. ro occur, entrance ro rhe seed musr be gained. Caloglyph11S can nor penerrare in Spider mire populations have a high propensity fo r developing resistance ro tacr pods and environmental conditions muse be in accordance with rhe ecolog insecticides and acari cides. Hisroricall y rhe genus Tetra11ychm has been capable ical requirements of the fungi for germination and fungus growth (Aucamp, of rapidly developing resistance ro a wide variety of roxicants, when repeared 1969). Soil applications of acaricides significantly reduced both peanut pod roxicant applicarions provide selection pressure (Jeppson er al. , 1975). T he ror, caused by P. myrioty/11111 , and m ire populations in borh field and green problem of a high incidence of pesticide resistance in mites is magnified fur house studies (Shew and Beute, 1979). T hi s report enhances the implication of ther by cross-resistance ro chemically related and ro some unrelated com the role of certain asrigmarid m ites as vecrors of soil fu ngal pathogens. Ocher pounds (S mith, 1960). Populations resistant ro one organophosphorous (OP) soil pests, i\leloidogy11e are11aria (Neal) and Diabrorira 1111deri111p1111cta1a ho1vardi insecticide often show resistance ro ocher OP and carbamare compounds while Barber, also have been reported ro disseminate and enhance P. 111yriotyl11111 in remaining susceptible ro organochlorine compounds (Jeppson er al., 1975). fection of peanut pods (Porter and Smith, 1974; Garcia and Mitchell , 1975 ). On peanuts, T . 11rticcte and T. ci1111ebari1111s have developed resistance co some Manage ment. Natural Enem ies. Spider mires are attacked by fungi, O P compounds as a result of prophylactic OP use (Smith and Jackson, 1975). predaceous mires and insects (Huffaker er al. , 1970; McMurrry er al., 1970). Resistant Culrivars. Resistance and susceptibility co spider mires The fu ngi, E11to111oph1hora spp. and E. frese11ii Nowakowski, have been reported have been identified in both commercial culrivars and wild peanur species. l n infecting T. 11rticae and T. ri11nebrll'i11m (Ca rnerand Canerday, l968)wirh E1110- 1ndia, T. 11r1icae infestations were higher on semi-spreading culrivars (e.g., 111oph1hora spp. mycosis being most effective in hot, humid weather (Campbell , Exotic 5, C50 l, Asirya M wirunde and HG lO) than bunch-type varieties 1978). The desrrucrion of Ento111oph1hora spp. fung i by fung icides applied for (Gupta er al. , 1969); rhus, erect g rowing fo liage may offer some prorecrion. control of plant pathogenic fung i (leafspors, rusts, ere.) on peanuts may be a Campbell er al. ( 1974) reported the culrivars Va 72R, NC-Fla 14 and NC 17 major factor in spider mire outbreaks (Campbell, 1978). had the lowest leaf damage while NC 2 had the hig hest leaf damage resulting Predaceous phyroseiid mi res are of maj or importance in suppression of rerra from T. 11rticae feeding. Ranges in suscepribiliry were from 23.8 - 76.9% for nychids of many crops (McMurrry er al. , 1970). Alrhough predaceous phyro rhe 11 culrivars evaluated in the greenhouse. sci ids have nor been reported from peanurs, ca. 10 species have been reported Germplasm possessing rhe highesr levels of resistance co spider mires is acracking T. 11rricae and/or T. ci1111ebari11 m on various ocher crops in the United within rhe wi ld species of Arachis. Mose species in the section Rhizomarosae Srates, Egypt, J apan and Canada (McMurtry er al. , 1970). Several insects also 292 P EANUT SCIENCE AND TECHNOLOGY MAN AGEMENT O F PREHARVEST INSECTS 293 are hig hl y resistant as wild species Pf 338296, 3383 17 , 262840 and 262827 onmental conditions to express their bioric potential. The deep, sandy soils of remained vi rrua ll}' free ofT. 11rticc1eUohnson er al., 1977). Mires feeding on Pl Texas and Oklahoma share rhe same fol iage consuming lepidoprerous pest 262286 and 262840 had reduced fecundiry when compared ro the commercial complex as rhe heavier soils of N orth Carolina but the soil pests are different. 1ine NC 5 U ohnson et al. , 1980). Adu! r females also exhibi red a hig h degree of E/as111opalp11s lignosellm (Zeller) is an annual , key pest in the southwestern non-preforcnce for feeding on both plant introductions. Only a few plant intro United States (Berberer et al. , 1979a; Smith and Holl oway, 1979), whereas ir ductions from the sections Arachis, Erccroidcs, Ex trancrvosae, and Caulori zae occurs infrequently in North Carolina (W. V. Campbell, pers. comm.). Psm exhibited res istance Uohnson ct al., 1977). Resistance ro T. 111111irlellm Pri rlococms sp. (Smith, 1946), Diabrotica spp. (Fronk, 1950; Feakin, 1973) and chard and Baker has also been reported (Leuck and H ammons, 1968). Dysmicoccm brevipes (Cockerell) (Feakin, 1973) are pests of poorly drained soils. Cultural. The microclimatc produced by intcrplanring between rows White grubs are pests of friable, volcanic, red soils in Queensland (Smith, of citrus and the cultural measures assoc iated wit h cirrus production were pre 1946). sumed as the cause of an our break ofT. ci1111ebt1ri1111s and T. 11rticaeon peanuts in Management strategies fo r many soil pests must be developed on rhe indi Egypt (Osman and Rasmy, 1976). Sprinkler irrigation was effe([ive in sup vidual species basis and cannot be classed collectively by plant injury as has pressing T. 11rt icae populations on peanuts in Egypt (Osman and Abdel-Farrah, been done w ith fo liage inhabitants. The major reasons mandating species di ( 1975). Mire populations were 3 ro lOX hig her under overflood irrigation as visions in management may include: ( 1) most soil pests are key pests, thus they compared ro sprinkler irrigation. Osman and Abdel-fatrah ( 1975) suggested are rhe initial rarget of management; (2) a good quantitative description of the sprinkler irrigation washed the mires from the leaves. This observation is sup relationship berween pest injury and peanut growth and frui ting is nor availa ported by research on mites infesting orher crops (van de Vrie er al. , 1972). ble, (3) edaphic facrors which provide the template for pest population dynam ics are more mosaic than the environmental facrors which drive foliage inhabi SOIL INHABITING PESTS tants, and (4) information on soil pest bionomics is limited, restricting the use of biological common denominarors. Many soil inhabit ing pests characteristically feed upon the fruit (pods) and/ or fruit precursors. Secondary infections of fung i and other plane d isease orga Lepidoptera nisms may gain entry into rhe seems, roots and pods as a res ult of soil arthropod feed ing and may further reduce pod production. Information on interactions ElasmopaljJ11s lignose/Lm (Zeller) is between soi l borne d iseases and soil arthropod feeding are limited, bur will be rhe only lepidopteran considered a presented when available (also see section on spicier mires; rhe soil inhabiting rrue pest as all life stages, except as rigmatids). Planes exhibit temporal tolerance ro borh primary and secondary adult, are soil inhabiting . Several injury, bur spat ial tolerance is nil. noctuids (e.g., Agrotis and Feltia) Soil inhabi ring arrhropods are rcpresenrecl in rhe orders Acarina, Jul ida, Or have diurnal soil inhabiting larvae, rhoprera, Dermaprera, lsoprera, H emiprera, Homoprera, Coleoprera, Lepi bur most of rhe damage is from noc doprera, Diprera and Hymenoptera (Table l). Of the 2 management groups turnal foliage consumpt ion. (soil and foliage), soil pests probably ;-epresent the key pests of peanuts; howev Recently, Stylopalpia costalimai Almeida was reported attacking peanuts in er, t he ecology, biology, damage and plane phenological relationships are the Brazil (Almeida, 1960, 1961; Almeida and Pigatti, 1961; Bastos Cruz et al., least understood. Special problems are inherent with soil arthropod research 1962). T ranslation of these reports revealed S. costalimai feeds on the fo liage and management which may explain the absence of biological knowledge fo r and may spend part of the larval stage in or on the soil bur is not considered a understanding pest bionomics and fo rmulating management strategies. true soil pest. S1ylopalpic1 sp. near costali111ai has been reported as a key pest of Soil a rthropods occupy a cryptic habitat which res rricrs undisturbed, direct peanuts in Paraguay (Unruh, 1981). This species feeds on pegs and pods and observation of pesr li fe srages, associated behavior and damage inflicted upon spins feeding webs in the soil which is similar to E. lignosellm feeding behavior. the plane. Curbed biological observations hinder hypothesis formulation and Elasmopalpus lignosellus (Zeller). This pest is restricted in distribution dara collecting. Dara gathering must rely on labor intensive sampling tech ro rhe New W orld where it attacks peanuts and numerous other legume crops niques for censusing population density, age disrriburion and inflicted plant as well as numerous grasses. An ecolog ical equivalent of E. lignosellm from pea damage. Visual observations of damage ro t he roors and main stem are delayed nuts in the Old World has not been reported. The genus Elam1opalpm was con temporall y unril rhe injury is manifested in vegerarive (aerial) plane parrs. Pod sidered monospecific by H einrich ( 1956); however, more recently Gares damage may go rorall y unnoticed , as secondary infections of soil microorga Clarke ( 196 5) removed E. angmtellm Blanchard from Heinrich's ( 1956) synon nisms camouflage rhe arrhropod damage at harvest or decompose rhe pod in omy. The larval habitat and hosr plants of the closely related genera, Adelphia, rhe soil. These cond itions tend either ro mask the importance of soil arthro Tota and Ufa are unknown (H einrich, 1956) except fo r U. mbedinella (Zeller) pods or result in the arbitrary assignment of the damage of soil microorga- which attacks t he pods of pigeon pea in the Lesser Antilles (Fennah, 1947). 111 sms. Biology. Adult females oviposit 33-420 eggs (Luginbill and Ainslie, Edaphic conditions are paramount ro rhe population ecology of soil arthro 1917; W alton et al., 1964; Leuck, 1966; Srone, 1968a; Razuri, 1975). Mosr pods. Different pest species require certai n soil types as well as optimal envir- eggs are laid sing ly in the soil (less than 2 mm deep) under the drip line of pea- 294 PE1\ NUT IE 1CE AND TECHNOLOGY MANAGEi\IENT OF PREHARVEST I NSECTS 295 nut plants. Two % of rhe eggs may be deposited on rhe foliage (Smith er al., absence of the classical diapause evident in most temperate Lepidoprera with 1981 ). Eggs are white rhe first day, rum red by the second day and harch rhe similar host ranges and geographical distributions. Cooler temperatures sig third day (Leuck, 1966). Hatch may require up to 5 days during the cooler fall nificantly increased larval and pupal developmental time; thus, these stages periods (Luginbill and Ainslie, 1917). negotiate overwintering by a gradual, but prolonged, larval-pupal develop . The small, red, first insrar larvae crawl across rhe soil from rhe oviposirion ment period . sire to the plant or ro other edible organic matter. Larvae feed slightly below Elasmopalpm lignosel/11J is mobile and highly polyphagous, feeding on 62 the soil surface and construct a si lken tunnel covered with soil particles which plant species representing 14 families (Stone, 1968c). It is a pesr of many culti is attached to the feedi ng site (Luginbill and Ainslie, 1917 ; King et al., 1961; vated legumes (Luginbill and Ainslie, 1917; Isley and Miner, 1944; King er Basros Cruz et al. , 1962; Leuck, 1966). Larvae feed read ily on dead organic al., 1961; Leuck, 1966; Razuri, 197 5 ), rice (Sauer, 1979) and sugarcane matter (Cheshire and All, 1979) as well as numerous plant species (Scone, (Plank, 1928; Bennett, 1962). The role of wild or cultivated host planes on l 968c). Larval development in the field has been found to require 13-24 days population outbreaks is unknown but may fit the pattern proposed by Stimac (Leuck, 1966), 11-39 days (King et al., 1961), 24-46 days (Dupree, 1965), or and Barfield ( 1979). 14-42 days (Luginbill and Ainslie, 19 17). Laboratory studies on larval and pu Population outbreaks of E. lignosellus occur during periods of hot, dry wea pal development by Holloway and Smith ( I 976b) showed developmental rare ther (Luginbill and Ainslie, 1917; King et al., 1961; Walron et al., 1964; dependent upon temperature, with larva l-pupal development predicted by rhe Bertels, 1970; French and Morgan, 1972; Smith, 1981) with high: soil mois following equation: developmental days= 19 1.245 days - 5.202 (C tempera ture inhibiting population outbreaks (Bertels, 1970; All and Gallaher, 1977). ture). Berberet er al. (l 979b) reported 5 30 degree days above 13 C to be re Thus, deep, sandy soils with good water percolation favor E. lig11osell11J popula quired for development from egg deposirion co adulr emergence. tion increases (Luginbill and Ainslie, 1917; Walton et al., 1964; Dupree, The number of larval ins tars is va riable. Leuck ( I 966) and Dupree ( 1965) re 1965; Leuck, 1966). Edaphic facrors undoubtedly are extremely important in port 6 insrars; Razuri (1975), 5-6 insrars; and Lug inbill and Ainslie (1917) 4- E. lignosell11J outbreaks. Elucidation of the interactions among soil type, cli 7 instars. The laner author seated that the number of insrars increased \~ir h mate and f.. lignosell11J population biology is paramount for temporally and temperature. Larvae reared at constant temperatures on artificia l diet in these spatially predicting epidemic popular ions. Analysis of a study on the popula nior author's laborarory have completed 5-9 insrars, wirh 6 instars being most tion dynamics of E. /ig11osel!11S immature stages in peanuts revealed mortality common. was nor density-dependent; thus, abioric facrors were again implicated as dic Pupation occurs in the soil, usually at a greater depth rhan larval feeding (se tating population levels (Johnson, 1978). nior author's observation). The pupal chamber is consrrucred of material sim D amage. Elas111opalp11S Lig11ose//11J is a.key pest of peanuts in the New ~lar ro the larval feeding tunnel bur is of much stronger construction. Pupation World, especially when hot, dry climatic conditions prevail (Bastos Cruz et 1n the field lasts ca. 10 days (Luginbill and Ainslie, 1917; Leuck, 1966). al., 1962; Walron et al., 1964; Leuck, 1966; French and Morgan, 1972; The life cycle from oviposirion to adult emergence spans 33-65 days under SmirhandJackson, 1975;Berbereteral., 1979a;SmirhandHolloway, 1979). field conditions. The number of generations per year in rhe United States is Yield losses in excess of 70% can occur under severe attack (Smith et al., v~riab l e with 4 in South Carolina (Luginbill and Ainslie, 1917) and Mississip 1975). pi (Lyle, 1927), 3 in Arizona (Vorhies and Wehrle, 1946), 3 plus a partial Larvae arrack all phenological stages of plane growth, feeding on subterra fourth in Georgia ~ Leuck, 1966) and 3 distinct generations in Texas (Johnson, nean plant parts. Leuck ( 1967) described 2 broad types oflarval damage to pea 1978) and Oklahoma (Berberet et al., l 979b). nut plants. The early instars (1st and 2nd) feed on vegetative buds, flower ax Adults are nocturnal (H olloway and Smith, 1975, l976a), mate the first ils, ground level stems and leaves. These larvae do not consume much plant day after emergence and begin ovipositing the second day (Stone, 1968b). Fe material during these early instars with damage resulting from scarification of male moths release a sex pheromone from 0-96 hrs. post emergence which at plant parts. Older larvae feed on pegs (gynophores) and pods. Damage co the tracts males (Payne and Smith, 1975). Ganyard and Brady ( 1972) also reported pegs and pods is considered the most damaging on runner and spanish type E. /1g11osell11J males attracted to virgin female Plodia i11terpu11ctella (Hubner), peanuts (Leuck, 1967; Berberet et al., 1979a). Smith and Holloway ( 1979), Cadra ca11tel~a (Walker) (~JOth Pyralidae), Spodoptera fmgiperda and S. exigua however, reported spanish type peanuts may be damaged more heavily by lar (both Noctu1dae). The acnve compound for attractiveness was considered co be vae scarifying tissue destined co become inflorescences and consuming the (Z,E)-9, 12-tecradecadien- l-ol acetate. Fi eld experiments by Mitchell er al. minute flower buds concentrated in the plane crown area prior ro gynophore ( 1976) revealed male E. lig11ose!!11J are not attracted to this compound, bur a and pod formation. They argue chat spanish type plants at a chronological age field permeated with the compound may d isrupt mating. of 28-58 days post planting are phenologically more susceptible to damage A co~siderab l e di.v~rgence in the overwinteri ng stage may occur, depending from a given larval population than at later stages. During this susceptible upon cl1manc conditions. All reports agree that f.. lig11osell11J overwinter as a plant age, larvae can cause a greater yield loss from a smaller amount of tissue mature larva and/or pupa (Luginbill and Ainslie, 1917; Lyle, 1927; Vorhies consumed. Leuck (1967) and Berberer er al. ( 1979a) report E. lignosell11S popu and Wehrle, 1946; King er al., I 96 I; Leuck, I 966). Holloway and Smith lations in Oklahoma and Georgia are very low until most planes begin co form ( 1976b) showed all life srages unresponsive ro changes in photoperiod with an pods, whereas in Texas high larval populations occur much earlier in plant phenology (Smith and Holloway, 1979). 296 P EANUT SCIENCE AN D TECI INOLOGY M ANAGEMENT OF P REl-I ARVEST I NSECTS 297 Yield losses from larval feeding have been descri bed q uanrirarively for span T able 4. Parasites of Elam1opalpw lig11oullm (Zeller). ish-rype peanuts during 2 phenologica l periods: 28-58 days and 60- l 10 days posr planting. Smith and Holloway ( 1979) reporred rhar larval popularions up Para.sire Reference Braconidac ro 14,448/ha (10% infesred plants) are rolerared by plants 28-58 days posr Che/011u1 imulari1 ( = 1txa11111) (Cresson) Wall&Berberer 1975 planting wi rhour a yield loss. Yield losses from larval densiries exceeding Che/011111 elaJmopalpi McComb Wall & Berberer 1975,Johnson&Smith 198 1 14,448/ha were described in a 3 paramerer nonlinear function. Berberer er al. Orgilm tlrumopalpi Muesebeck Wall & Berbcret 1975, Falloon 1978,Johnson &Smi th 198 1 ( l 979a) descri bed yield reducrions 60-110 days posr planting as a function of Orgi/11111i1id111 Mucscbeck Johnson &Smith 198 1 % infested plants using a linear equarion. Borh scudies provide the essential in Orgilmsp. W all & Bcrbcrct 1975 1-ltibrobraron gtlerhiae(t\shmead) Joh nson &Smith 198 1 sect density yield loss input dara for calculating economic rhresholds. Brt1ro11111ellitor Say Leuck & Dupree 1965 Secondary invasion of larval damaged p lants or pods by microorganisms has Aga1h11 rubriri11r1111 Ashmead Beg & Bennet! 197 4 nor been reporred. Scarificari on of pods by larvae (Leuck, 1967) should assisr Aga1h11 sp. J ohnson & Smith 1981 secondary invasion as reported for orher soil pests (Porter and Smith, 1974 · Apa111elt1 sp. Wall & Berbercr 1975,Johnson &Smith 1981 Garcia and Mitchell , 1975). ' /\!irropli1i1 rroceipeI (Cresson) Wall & Bcrbcrct 1975 Marrocm1m1 sp. Beg & Bennen 1974, Johnson &Smith 198 1 Management. Natural Enemies. A lisr of egg-larval, larval and p upal parasi res of E. lignosel/11s arc presented in Table 4. Levels of E. /ig11osel!m para lchneumonid ac sirism reported from peanu rs vary; 2.5-8% Oohnson and Smith, 1981), 5% PriJ10111er1111pi11a1or(F.) Wall &Berberet 1975,J ohnson&Smith 1981 (Wall a nd Berberer, 197 5 ), 8% (Schuster er al. , 197 5) and 13% (Berberet er Eiphosomf/ de111a1or(F.) Falloon 1978 Dit1deg111a sp. Frank & Bennet! 1970 al. , 1979b). H owever, parasitism of E. lignosel/11s on other crops is subsrantial Ny1hobia sp. Metcalfe 1965 ly hig her; 35-6 1% on cowpeas and soybeans (Leuck and Dupree, 19 65) and 12-1 4 % on sugarcane (Beg a nd Bennerr , 1974; Falloon , 1978). This disparity Chalcididae in parasirism between peanuts and other crops was d iscussed by J ohnson and /,m·cia tit f{l/orGrissell & Schauf Johnson&Smith 1981 lnvreia 1brea G rissel! & Schauf J ohnson & Smith 198 1 Smirh (198 1) They posrulared rhe diffe rences were d ue ro E. !ig11osel/11s larval /n vrtia 11110 G rissel! & Schauf J ohnson & Smirh 198 I b e ha~ ior on certain plants and resulting rachinid paras itism, as rachinids only parasmze exposed larvae. Larvae arracking seedling plants frequently abandon Bombyliidac the cryptic microhabitar of rhe feeding runnel in search of new hos ts as seed Gtro11 arit/111 Painter Johnson & Smith 1981 lings perish . This behavior is d icrared by plant growrh stage, as older plants T nchinidac are less likely ro peri sh . £. lig11ose!!11s larvae usua ll y arrack peanurs ar a g rowrh S10111f//0111yit1 floridemiJ (Townsend) W all & Berbercr 1975,J ohnson &Smiih 198 1 m tge where plant clearh is unlikely and larval exposure ro rachinid parasitism is S10111a10111yi" 1ri11i1a1i1 (T hompson) Beg & Bennett 1974 limited. Parasitism by 1-l ymenoprera , however , is approximately rhe same on S10111a10111yra parvipalpis (Wulp) Leuck & Dupree 1965 all crops. T he cryp ric soil m icrohabirat of rhe larvae may further resrricr para sirism by concealing rhe larvae from rhc numerous general hymenoprerous erage maximal remperarures were 36. 5 C (Johnson and Smith, unpublished). parasites of orher lepidoprerous larvae inhabiting rhe fo liage Oohnson and Extremely high soil temperatures could inhibit both functional and numerical Smirh, 198 1). responses of 0 . elasmo/1a!pi Oohnson and Smith, 1980), as well as or her E. ligno The mosr im portant hymenopterous larva l and pupal parasites are Orgi!m se!!us parasi res . ~!C1s111opalpi Muesebeck and I 11vreia sp., respectively. Orgilm e!asmopa!pi parasit Predators atracking E. lignosellm larvae include: lygaeids, Geocoris sp; a ca ism accounted fo r 13-37% (Leuck and Dupree, 196 5), 33.3% (W all and Ber rabid, Philophuga vfridico!is LeConte; and 2 rherevids, Psilocephala amta Adams berer, 1975), 24-100% (Johnson , 1978) and 4-20 % (Berberer er al., 1979b) and F11rcifera mfiventris (L W.) (Johnson, 1978). Although no rares of predation Jf roral parasitism . lnvreia sp. accounted for 9-77% (Bcrberer et al., l979b) could be assig ned quantitatively, these predarors were abundant in peanut tnd 16% parasitism (W all and Berberer, 1975) (erroneously reporred as lnv1·eia fi elds and were often observed feeding on E. !ignosel!m larvae. uimbi!is (Boucek), a vali d Old W orld species (Grissell and Schauff, 198 l.) In a J ohnson ( 1978) also reporred a virus infecting E. !ignosellus larvae in peanut )-year T exas study of E. lig11osel!m natural morraliry, J ohnson and Smith fields. Two rypes of infection were noted: an acute infection which results in 198 l) reported lnvreia spp. parasitism ro be errari c, rang ing from 0 .2-7.2% rapid death, cuticle disinteg ration and black cadaver; and a chronic form ind only occurring in 2 of the 5 generations studied. which results in prolonged larval li fe, a color change from greenish-blue ro pal Parasitism by 0 . elasmo/NtfjJi may be curbed by hig h rem perarurcs (J ohnson l id red , and the co ntinued srrucrural i nreg ri ty of the cadaver cut icle. Small ( 1- nd Smirh, 1980). Parasite survival from egg ro larval emergence from the hosr 2 insrar) and medium sized (3-4 insrar) larvae succumbed ro the acure infec vas 34% (32.2 C) and 23% (35 C) as com pared ro a favorable 85% (26.7 C). tion, whereas the chronic infection was primarily in large (5-6 insrar) larvae. 1 ig h levels of parasi re morral iry also occurred after emergence from rhe hosr ar Viral infection of larvae and the subsequent morraliry ranged from 0- 19. 8% he upper rempcrarure exrremes; 59% and 0 survival at 32. 2 and 35 C, respec- bur played a minor role in E. !ig11ose!!11s regulation. 1vely. The levels of fi eld parasi tism by 0 . elm111opalpi averaged 3-4% when av- Further studies by Mitchell (1980) identified rhe £. lig11ose!!11s virus as an Entomopoxvims with a hig h level of virulence, arracking rhe la rval and pupal fa r 298 P EANUT SCIENCE AND TECHNOLOGY MANAGEMENT O F PREHARVEST INSECTS 299 bodies and haemocyres. lnfecred larvae are active and ear normally unti! just tion values and risk. Smith and H olloway (1979) concend chat economic prior ro death. All infected larvae die as larvae, prepupae or pupae. Chronically threshold for E. lig11oseff11S cannot be fu rther refined until population densities infecced larvae live an average of 48 days beyond the normal developmental pe can be predicted accurately in advance of actual losses, thus allowing the calcu riod at 27 C. larion of reliable estimates of probable yield loss. Esrimares of probable loss are Partial ecological life rabies (Southwood, 1978) construcr~d on E. li~110Jell11J mandacory for balancing probable losses against coses and optimizing rhe net egg, larval and pupal srages in peanuts showed meal generanon morcal1cy (real return (Berberec er al., 1979a; Smith and Holloway, 1979). morcali cy) ro vary between 87. 0 and 99. 9% Qohnson, 1978). T he greacesr age Field sampling co estimate pesr population densities is inherent co IPM pro specific morcal icy occurred in rhe egg and newly harche~ (< 1 da~ old) ~arva l g rams in which economic thresholds diccare timing of insecticide applications. scages (49.7-86.8%) with an average of60% + morcal1 ry occurring pri.or ro Two main sampling plans for estimating E. Lig110Jefl11S larval population densi che medium larval scage (3-4 inscar). Morcal iry facrors were found ro ace inde ties currently are in use in rhe United Scares. In rhe souchwesr, individual pendencly of E. fig110Jell11S density; ch us, biori.c factors (natural enemies~ were planes are examined co determ ine t he proportion of planes infested with larvae nor regulating pest density. Johnson and. Smith ( !981) suggesce? that ~ntro (Smith and H oelscher, 1975b; Berberec and Pinkscon, 1978); and in rhe duccion of an exoric natural enemy rhar 1s ecologically synchronized w1rh £ . southeast, thresholds are based on the proportion of 0 . 91 m sections of row in /ignoJe/lm could help regulate populations. Since E. lig11osel/11S b~ l o n gs t~ a ?10- fested wich larvae (French and Weeks, 1978). Boch sampling plans involve a nospecific genus and its distribution is limired ro r~ e ~mencas (H einrich, time consuming manual search of the subterranean plant pares and adjacent 1956), candidare exotic natural enemies for suppression in rhe U n1ted Scares soil fo r larvae. Early inscar larvae are normally difficult co find; wee soils in would need robe obtained from South and Central America or from an ecologi- crease the difficulty oflocaring all inscars (Smith and Hoelscher, l 975b). Large ca l equivalent in rhe Old World. . . . . (< l. 3 cm) E. Lig11ose!l11S larvae caught in pitfall craps provide as accurate an es l nsecticides. Sprayable and g ranular formu lanons of insecc1c1des have timate of field density as sampling 0.91 m sections of row Qones and Bass, shown varying degrees of efficacy against£. lignose!lllS on peanurs (Arthur and 1979). Pitfall sampling reduced sampling time 33-50% and enabled sampling Arant, 1956; Cunningham er al., 1959; Harding, 1960; King er al., 1961; under wet conditions. All currently used sampling programs fo r estimating E. Basros Cruz er al., 1962; Walron er al. , 1964; French, 197 1; Lee, 1971; lig11oseff11J larval densities in peanuts underestimate rhe actual density because French and Morgan, 1972; Corseuil and Terhorsr, 1975; Smith er al. , 1975; rhe small, earl y inscars are difficult co census. Sampling also requires destruc Berberec and Wall, 1976; Sams and Smich, 1979). Inseccic idal efficacy is en tion of the planes , is rime consuming and laborious and is sensitive co soil hanced when eicher sprays or granules are appli ed co che soil surface (Cun mo1srure. ningham er al. , 1959; Smith and H oelscher, 1975b; Smith er al., 1975). In The use of ecologically selective insecticide formulations (i.e., granules) and che souchwesrern Unired States, sprays are recommended fo r dryland peanut application techniques which direct rhe insecticide co the soil habitat of the production and granules fo r irrigated production (Smirh e.c al. •. 1975; target organism has been shown co have biological as well as economical bene Hoelscher, 1977; Berberet and Pinksron, 1978). Adequate sod moisture is fits. Granular insecticide fo rmulations applied in irrigated peanut cultures for fundamencal co efficacious results wirh granular insecticides. suppress ion of£. lignoseffm conserved natural enemies of the numerous fol iage The economics of insecticide use for E. lignoseflus control on peanuts was inhabiting arthropods (Smith and Jackson, 1975) and increased insecticidal ef original ly questioned (Cunningham er al. , 1959; Harding, 1960; King er a.I. , ficacy against the target species (Smith et al., 1975). Basal directed sprays in 1961). However, larval density-yield studies (Berberec et al. , 1979a; Sm1ch creased insecticidal effi cacy when compared co broadcast sprays (Cunningham and H oll oway, 1979) and recent insecticidal efficacy-yield studies (French a.nd et al., 1959; Smith and Hoelscher, 197 5 b; Smith et al., 1975); however, basal Morgan, 1972; Smith er al., 1975; Berberer and Wall, 1976) have shown in directed sprays are nor selective enough co conserve rhe extant, noncargec, fo secticidal control co be economicall y efficacious, dependent upon larval popu liage inhabiting arthropods (Smith and Jackson, 1975). Basal directed sprays lation density and plant age. Yield increases of up co 250% (Smith et al., do decrease the coca! insecticide load on the peanut ecosystem by reducing the 1975) and 29% (Berberer and Wall, 1976) have been reported in the south number of applications necessary co maintain E. /ignosellm below established western United Scares on dryland peanuts where sprayable insecticides have economic thresholds. Reducing the roral number of applications aids in con been used. Under irrigation or adequate soil moisture, g ranular insecticide fo r servation of natural enemies during rhe entire growing season. Ecologically se mulae ions resulted in yield increases of 78% in the southwest (Smith et al., lective pesticide application techniques, used in concert with economic 1975) and 45% in the southeast (French and Morgan, 1972). thresholds, help circumve nt nomarger pesr upset. Problems of conservation of Economic thresholds fo r insecticide applications in the southwest are 5% in natural enemies, changes in pest status, selection of pesticide resistant popula fested planes prior co the initiation of Regging and 10% after che initiacio.n ~f t ions and economic losses (Smith and Jackson, 1975; Smith et al., 1975) are pegging (Hoelscher, 1977; Berberetand Pinkscon •. 1978). ~ hres h o l ds for 1m. minimized. gared peanuts are higher- IO and 15 % , respecr1vely-since adequate so.ii The economic benefits of ecological ly selective insecticidal application rech moisture allows fo r a g reater probability for plane damage recovery. Economic niques for suppression of E.lignosef/m in Texas are d ramatic. The major thrust thres holds are based primaril y upon darn from insecricide crials and larval d~n of an IPM program introduced co peanut producers in 1971 utilized ecologi siry-yield studies. To dace, no efforts have been published which refine exist cally selective insecticide applications and economic thresholds for the key ing economic thresholds by including the variables of control coses, produc- pesr, E. lignose/Lm. Prior ro 197 l, insecticidal application techniques were 300 P EANUT SCIENCE AND TECH 1 O LOGY MA AGEMENTOF PREHARVEST I NSECTS 301 mainly broadcast SJ rays applied by fi xed wing aircraft (nonsele rive). During Cultural. Culturally oriented management tactics include planting 197 I, aerial application decreased 5 3% on irrigated fields and 3 l % on non irri dares, sanirarion, winter plowing, irrigation and crop rotation. Many cultural gated (Smith and H oelscher, 1975a). Insecticides applied by g round driven practices have resulted from observations on peanuts and other agronomic equipment (granules and d irected sprays) were unchanged. The effectiveness of crops in earlier reports and lack a sound data base. the p rogram in managing£. lignosellm is further illustrated by changes in the Removal of crop residues, fa ll and winter plowing and weed removal report proportion of producers using multiple applications. In 1970, 30 % of the pro edly help prevent severe infestations (Lug inbill and Ainslie, 1917; Watson, ducers used 5 + applications as opposed ro 6% in 1971. Also in 1970, 69% of 19 17; Guyton, 1918; Box, 1929; Stahl, 1930; Hayward, 1943; Isley and the producers used 3 or more applications while 72% used I co 2 applications Miner, 1944). In crops such as corn, sorg hum and peas, E. lignosell11s is mosr in 197 1 (Smith and Hoelscher, 1975a). Thus, within I year 82% of the g row destructive when attacking the seedling stage. Many authors (e.g., Isley and ers had adopted the selective application method. The drasticall y reduced in Miner, 1944; Wilson and Kelsheimer, 1955; Reynolds er al., 1959) felt crop secticide load on the peanut ecosystem was attributed ro adherence ro econom res idues and alternate hosrs (weeds) in the fields prior to planting were the ic thresholds and increased effectiveness of the insecticides on the target soil sources of damaging larvae on seedling susceptible crops, since seedlings pesr (Smith and J ohnson, 1977). would grow out of this susceptible srage before eggs oviposited in the planted Resistant Cultivars. Resistant germ plasm has been identified in crop could develop into the damaging, older larval stages. Cultivating all in commercial cultivars, noncommercial cul ti vars and wi ld species. Levels of re fested hosts after planting could force the entire larval infestation to concen sisrance identified have been low, but even those low levels have utility in an trate on the crop seedlings; therefore, sanitation should be practiced several lPM program (Smith, 1980; Smith et al., 1980b). weeks in advance of planting (Reynolds et al., 1959). Dupree (1964) reported A wide variation in seed li ng survival among 108 peanut lines artificiall y in that tillage and mai ntenance of weed free fields 8-10 weeks prior to planting fested with £. lignosellm eggs provided the first evidence of resistant germ cow peas gave E. lig11osel!11S control equal to insecticides. The level of larval in plasm (Leuck and H arvey, 1968). Field evaluations of 14 adv;.nced peanut f~srat ion was shown to be different in rill (clean field) versus no-till (crop re lines, infested in a similar manner but at a more advanced stage of plant sidue) corn (All and Gallaher, 1977; All er al., 1979). The feeding behavior of growth, fa iled ro show any differences in plant response between lines or types larvae apparently is modified when crop residues are present (Cheshire et al. , (e.g. , spanish, runner and vi rg inia) (Leuck et al. , 1967). Several of the lines in 1_977 ), as larval. damage to seedling corn was reduced g reatly when plant re vestigated by Leuck et al. (1967) also were included in the Leuck and H arvey sidues were available as a food source (Cheshire and All, 1979). ( 1968) report. In t he latter report, several of the Iin es reevaluated as seed Iin gs Flood irrigation readily controlled E.lig11ose/111s larvae (Reynolds et al. , showed a more variable response. Smith et al. (1980 a, b) screened 490 peanut 1959). Irrigation also increased soil moisture in corn and was an important fac cultivars and identified 81 cultivars which scored significantly less damage tor in prohibiting£. lig11ose/111s infestations (All and Gallaher , 1977). Increased than Starr, a commonly grown cultivar in the southwestern U nited States plant vigor (by eliminating drought stress) and increased soil moisture are both identified as being the most susceptible of t he 490 cul t ivars. Among these pu associated with irrigation and should help reduce damage from E. lig11osel/11s tative resistant cultivars were the commercial cul ti vars Fl orunner (tolerance), larvae. Early Runner (antibiosis), and Virginia Bunch 67 (tolerance). Further trials Peanuts p lanted so that the most susceptible phenological scage(s) will not with Florunner, Early Runner and Virg inia Bunch 67 carried to maturity un synchronize temporally wirh peak£. /ig11osellm populations would minimize der field conditions substantiated rhe g reenhouse results. Yield reductions, damage. In the soutl:eastern U ·~ir ed States, peanuts planted prior to mid-April due ro E. lignose/lm infestations, measured by comparing the same variety with escape peak p_opulat1?ns of E. /1g11osellus (Leuck, 1967). Corresponding ly, pea and without an insecticide umbrella, were: Earl y Runner, 8 .4 % ; Florunner, nuts planted 10 May 10 Texa.s s hou l ~ obtain sufficient maturity to escape heavy 19% ; Virg inia Bunch 67, 19% ; Florig iant, 15 % ; Dixie Spanish , 26% ; and damage by t he pest populanons which peak near mid-Aug ust (Johnson 1978· Comet, 45% (Schuster et al. , 1975). Comet is a commercial cultivar, suscepti Smith and Holloway, 1979). ' ' ble ro E. lig11ose/111s (Kamal, 197 3). Florig iant, Florunner and Early Runner al so reduced larval survival better than Comet, offering a level of antibiosis. Coleoptern These varieties, Florunner, Florigiant, Early Runner and Virg inia Bunch 67, are t hought ro possess onl y low but useful levels of resistance co E. lig11osell11S The Coleoprera are a large and diverse g roup of phytophagous insects on (Smith, 1980). peanuts (Table l ). Many species are fo liage consumers as adults and subterra Schuster et al. ( 197 5) observed parasitism off.. lignose/lm larvae by hymen nean feeders as larvae. Subterranean feeding larval stages constirute the g reat opterous parasites to be hig her ( 12. 5- 18. 7% ) in prostrate g rowing culrivars est peanut pest problem , as spatial tolerance is nil. The scarabaeids (white (cv. Florunner) when compared ro erect g rowing spanish cultivars (0.0- 1.6%) g rubs), elarerids (wireworms), tenebrionids (false wireworms), certain curculi and suggested that combining intermed iate levels of resistance wi th enhanced onids and chrysomelids (Diabrotica spp.) are representative of major coleopter parasitism from g rowth habit could be important in E. lignosellm manage ous pests where adults and larvae occur in separate habitats. The subterranean ment. Berberet er al. ( 19 79b), however, fo und rhar parasitism was not en coleopterous pes ts as a g roup are poorly known, with published information hanced by prosrrare plant g rowrh habit. existing as a potpourri of miscellaneous notes. 302 PEANUT SCIENCE AND TECHNOLOGY MANAGEMENT OP PREHARVEST INSECTS 303 Diabrotica spp. Diabrotica baltea tral Missouri. Adults are mobile and highly polyphagous, feeding on 208 ta LeConte, D. speciosa Germar, and plant species (Sell, 1916). The larvae attack corn, cucurbits and most agro- D. 11ndecimp11nctata howardi Barber nomic legumes. . . have been reported attacking pea Damage. Adult D. 11ndecimp11nctata howard1 feed on pean?t _fo~1age_. pre- nuts. These beetles have a biology ferring the terminal buds (Hays and Morgan, 1965). Economic miury 1s due different from the other chrysomelid principally to larval feeding on ~ubter~~ean pegs and p?<2s (Grayson ~nd Poos, genera listed in Table 1 as adult fe 1947). Pods are highly susceptible to miury from the ume they begtn to form males oviposit on the soil and the lar until they approach maturity (Grayson and Poos, 1947). Larvae prefer young, vae are subterranean feeders. In the soft pods to the older, more mature pods (Fink, 1916). Pegs are also heavily United States, D. 11ndecimp11nctata damaged when they first penetrate the soil, prior to enlarging to form pods howardi has been recognized as a pest (Grayson and Poos, 1947). Losses from larval feeding can reduce the yield of of peanuts since the beginning of the whole seed by as much as 80% (Feakin, 1973). 20th century (Fink, 1916) and has re- Larval injury predisposes the pod to attack by soil microorg~nisms (Grayso~ ceived considerable attention by en- -·•0~;;!}:?!1:}'·~~(fJ~·'IZl~[iJ·~ and Poos, 1947). Pod damage facilitates entry of the pathogemc fungus, ~yth1- tomologists. 11m myriotylum, which has caused increasing damage to the peanut crop m the The preponderance of literature on D. 11ndecimp11nctata howardi dictates con Virginia-Carolina area of the United States (Porter and Smith, 1974). centration on this species. However, in certain geographical areas (e.g., Management. Natural Enemies. Information on specific natural ene United States Gulf Coast area), D. balteata may be the dominant species (Fea mies of Diabrotica spp. is quite limited. Tachinids parasitic on D. 11ndecimp11nc_ , kin, 1973). The difficulty in separating Diabrotica spp. larvae taxonomically . tata adults include Pseudomyothyria anci//a (Walker) (Arnaud, 1978), Celatoria (Bass and Arant, 1973) may have led co an incomplete understanding of the to ( = Chaetoph/eps) setosa (Coquillett) (Bussart, 1937) and C. diabroticae (Shime~) tal species involved throughout the New World. (Fronk, 1950); the latter are also parasitic on larvae (Arant, 1929). Celatorza Biology. Overwintered adult female D. 11ndecimpunctata howardi oviposit bosqi Blanch attacks D. speciosa Germar in Argentina (Christensen, 1944). in the soil preferring a shady, moist substratum (Howard, 1926; Campbell and Parasitic nematodes include Diplogaster spp., Neoaplectana sp. and Howardu Emery, 1967). Females oviposit over an average of 48 days (Isley, 1929). Field la benigna Cobb (Fronk, 1950). Entomogenous fung~ have ~en reported fro~ collected females laid from 116-895 eggs (Sweetman, 1926). Oviposition is South America (Christensen, 1944). Apiomerus crampes crasszpes (F.) (Reduvu related to both temperature and relative humidity (Campbell and Emery, dae) (Morrill, 197 5), and species of Xantho/inus, Anisodactylus, Agonum, '!-mara 1967). Average eggs laidperfemalewere0.3, 20.3, 110.6, 106.0, 284.5 and and Pterostrichus, (all Coleoptera) (Fronk, 1950) have been observed preymg on 168.7 at 7, 13, 18.5, 24, 29.5 and 35 C, respectively. Total eggs laid in 7 D. undecimpunctata howardi. Only the parasitic fly, C. diabroticae, and nema days (24 C) also increased from 19. 7 to 29.0 with relative humidities (RH) of tode, H. benigna, occurred with regularity, parasitizing 3.7% and 23.6%, re- 55 to 97%. Females did not oviposit at a RH of 25%. spectively, of adult D. undecimpunctata howardi ~Fronk, 1950): . . . Eggs hatch in 6-13 days depending upon temperature (Sweetman, 1926). Insecticides. Prior to the early 1960 s, the cyclod1ene msect1c1des Eggs will not hatch below 75% RH with peak% hatch occurring above 85% (e.g., aldrin, heptachlor) were used extensively as soil broadcast and banded RH (Campbell and Emery, 1967). Larvae feed on the subterranean plant parts treatments for D. 11ndecimp11nctata howardi. This method of control was so effec for ca. 21 days and then form a pupal cell in the upper 7. 6 cm of soil (Sweet tive and in such widespread, general use as a preventative concrol tactic that D. man, 1926). The prepupal period requires 6. 3 days and the pupal 8. S, with a undecimpunctata howardi was seldom considered a pest (Boush et al., 1963). total of 46. 3 days (average) needed ro complete a generation (Sweetman, Aldrin and heptachlor gave season long control and were effective as spray, 1926). Under laboratory conditions of 27 C and 60-70% RH, egg incubation dust and granular formulations (Boush et al., 1963). These formulations were was 7 days, larval stage 10 days and prepupal and pupal stages 10 days (Hays available in convenient form for preventative treatments (e.g., mixed with fer and Morgan, 1965). Under laboratory conditions, the life cycle is completed tilizers) (Ritcher, 1953) and were used heavily over a wide geographical area. in ca. 27 days (27 C constant), whereas Sweetman ( 1926) recorded in excess of The result of this pesticide load was D. 11ndecimp11nctata howardi resistance to 2 months for development at a northern latitude. cyclodiene insecticides in ca. 10 years (Boush ec al., 1963). Cyclodiene insecti Three to 4 generations occur each year in the southern United States (Hays cides persist in the soil for the entire growing season with high residue concen and Morgan, 1965); 1 generation occurs in the north (Sweetman, 1926). In the trated in the pods and seed (Beck et al., 1962; Dorough and Randolph, 1967; southern states, D. 11ndecimp11nc1ata howardi does not enter a completely dor Morgan et al., 1967). mant state during the winter; rather', it overwinters in organic debris near Many organochlorine, organophosphate and carbamate insecticides give fields as an adult which is active when temperature permits (Hays and Morgan, adequate control of D. undecimpunctata howardi larvae (Howe and Miller, 1954; 1965). According to the observations of Smith and Allen (1932), D. 11ndecim Arthur and Arant, 1956; Boush et al., 1963; Boush and Alexander, 1964; p11nctata howardi migrates northward during the spring and early summer. Pro Hays and Morgan, 1965; Smith, 197la, 1972b, 1976b, 1977a, b). However, geny migrate southward during the fall, with no winter survival north of cen- the organochlorines, DDT and chlordane, leave undesirable residues in the 304 PEANUT S CIENCE AND T ECHNOLOGY MANAGEMENT OF P REHARVEST INSECTS 305 peanut pods (Doroug h and Randolph , 1967; Morgan cc al., 1967). Historical 100 larvae (Smith and Porter, 1971). Differences between% infested fruir oc ly, all insecticides have been used in a prevenrarive manner , applied eirher ar curred ar rhe low levels of infestation (25 and 50 larvae), but no differences planring or when the planrs begin ro peg. were discernible at the high infestation level. The authors concluded that high Cerra in peanur varieries (a nd resul ring growth habit) may influence the effi levels of resistance were nor available in the culrivars evaluared, and char dem cacy of granular insecricides (Campbell er al. , 197 6). Diazinon and erhoprop ons trated levels of resistance would not g ive commercially acceptable conrrol. were ineffective in controlling D. 1111deci111p111l(fafa howardi on NC Ac 15 7 5 3 It is interesring to note that NC 6, a commercial virginia type culrivar re (bunch growrh habit and 0. 1111deci111p1111ctara howardi susceptible), gave moder sistant ro D. 1mdecimp1111ctata howardi, was developed from a cross ofNC 343 x ate conrrol on cv. Florigiant (runner growrh habir and suscepri ble) and gave Va 61R (Cam pbell er al., 1971; Wynne er al., 1977). Both parents of NC 6 excellenr concrol on NC 343 (inrermediare growth habit and resisranr). Cul were considered previously susceptible to larval feeding in greenhouse studies rivars resisranr to 0. 1111deci111p1111ctata howardi (e.g. , NC 6, NC 343) require less (Smith , 1970; Smirh and Porter, 1971). The absence of the expression of resis insecri cide (as much as less) ro achieve adequate conrrol (Campbell er al. , 75 % tance by NC 6 parenrs in the greenhouse tends co indicate resistance is related 1976; Wynne er al., 1977). ro multiple facrors expressed under field conditions and nor in rhe laborarory. Resis ta nt Culrivars. After screening 2,500 peanut culrivars for resis Also, Boush and Alexander (1965) and Hays and Morgan (1965) stated that tance ro D. 1111deci111/J1mctalct howai·di, Boush and Alexander (1965) concluded spanish type cul ti vars are sources of resisranr germ plasm. T~is .could be related rhar spanish culrivars were more resisranr rhan valencia, wirh vi rginia culrivars ro rates of change in plant phenology and growth charactensncs as speculated being rhe mosr susceptible. A highly resisranr spanish culrivar (Pl 262048) by Fronk ( 1950). NC 6 is especially adapted ro heavier soil typ.e~ where D .. 1~11- was crossed with 2 virg inia cul rivars in an arrempr to transfer resistance (Alex decimpunctata hOU1ardi is consistently a problem. As add1t1onal pos1t1ve ander and Smirh, 1966). Resulring F 1 planrs were more resisranr rhan rhesus qualities, it possesses moderate resistance ro rhrips (Fra11kliniella /usca ), E111- ceprible parents (virginia culrivars) bur were more susceptible rhan rhe resis poasca fabae H arris and Heliothis zea (Campbe!I ~ nd Wynne, 1980) . . tant parenrs. Cultural. Moisrure content of rhe sod 1s probably the mosr important Survival of D. 1111decimp1111ctrtlct howarrli larvae was measured on germinating single environmental facror in determining population levels of 0. 1111decim seeds of 172 culrivars under laboratory condi rions (Chalfanr and Mi rchell , p1111ctata howardi (G rayson and Poos, 1947). High soil moisture and a 1967) . Resistance (anribiosis) was identified in rhose culrivars with larval sur roughened soil surface make a preferred oviposition substratum for adult fe vival~ 10% , which comprised ca. 5% ofrhe coral culrivars. Selected putative males (Howard, 1926). Avoidance of heavy soils with high organic matter con res istant a nd susceprible culrivars were further evaluared in field experiments tent in poorly drained areas helps prevent larval population buildup (Fronk, (Chalfanr and Mirchcll , 1970). Resulrs from field evaluarions were nor always 1950). congruenr wirh laboratory dara. For example, Pl 22 1068 was susceprible in Larval injury is g rearer on crops thar follow winter cover crops of green ma rhe laborarory and resistant in the field rrials. The reverse was rrue fo r cv. nure but injury may also be severe in winter fallow fi elds (Bissell, 1936, 1940; Georgia Station Runner. Arremprs to explain variabiliry in pod damage by Feakin, 1973). Injury may be g reater when peanuts are grown annually on the planting according to pod maruriry (pl anr phenology), as opposed to a single same field (Fink, 1916). Fronk (1950) found no differences in 0. 1mdeci111p11nc planring dare, were unsuccessful. T he aurhors did, however, explain that re tata howardi injury between 3 different dares of planting in Virginia. Plant sisrance evaluarions based on % damaged, immarure and undamaged pods spacings of 15, 23 and 30 cm apart and row widths of 0. 61 and 0. 84 m had no could result in spurious concl usions since maruriry and peg formarion rares are effect on plant injury. hi ghly variable among culrivars. The ra re of pod maruriry may also be an ex White Grubs. Numerous species of white grubs (Scarabaeidae), arrack pea tremely imporranr resisrance factor. Fink ( 19 16) reported rhar peanur planrs in nuts throughout the world (Table 1). which rhe pods are marured or are maruring rapid!y were ei rher free from larval Some species are pests as both larvae damage or decidedly less injured. Spanish culrivars marure fasrer rhan mosr and adults, while others are pests as virginia or runner culrivars, which may explain rhe general resulr of spanish either adults or larvae. Many species culrivars possessing resisrance (Boush and Alexander , 1965; Hays and Mor are highly polyphagous and have l or gan, 1965). 2 year life cycles. Long white grub life The original observarion of Fronk ( 1950) rhar plant growrh habir, pod size cycles present special problems in and seasonal insect and planr developmenr were related to resistance were con crop rotation especially when peanuts sidered by Smith ( 1970) in developing a greenhouse screening technique fo r are planted behind new land or weedy idenrifying D. 1mdecimp1111ctata howctrcli resistance ro peanuts. Smirh (1970) fallow. In general, the life hisrories measmed larval damage to both immature and marure fr uit and found no pref are poorly undersrood and informa- erence for eirher. Cultivar NC 10211 sustained the least damaged fruits tion relating damage and plant phe- ,, (9.2%) while cv. Argentine and NC 343 were the highest damaged (28.5 and nology is nil. -··· . . 34.0% , respectively). Nine culriva rs considered possibly resisranr (Smirh, Biology. The biology of Lachnoster11a co11sanguinea (Blanchard) 111 India 1970) were reevaluated by subjecring rhe cul t ivars to 3 ;JCSr levels: 25, 50 and has been studied by several authors (Kalra and Kulshresrha, 1961; Desai and 306 P EANUT SC IENCE AND TECHNOLOGY MANAGEMENT OF PREHARVEST I NSECfS 307 Patel , 1965; Patel ct al., 1967; Rai ct al., 1969; Srivasrava et al., 197 la). The Passlow, 1969; H. brevicoflis, Smith, 1946) or feed on the main plant stem sev following biology is a compilation of these reports. ering the rop of the plant from rhe roots (e.g., T . p1111cticollis, Smith, 1946). The adult beetles are polyphagous, nocturnal feeders of the foliage and inflo Adults of several species in which the larval stages are also pests prefer ro feed rescence of numerous plants. Adults emerge from the soil pupation site, leav on the foliage of trees (e.g., L. consa11g11inea, Rai er al., 1969; Srivastava er al., ing behind a round hole on the soil surface. Maximum adult emergence and 197 la; E. masho11a, Rose, .1962; Broad, 1966). mating coincides with the onset of the rainy season. Beetles emerge from the Larvae feed on the roots, pods and nitrogen nodules on the roots (Miller, soil sharply between 7:30 and 8:00 p.m. and fly ro adult host plants to feed. 1943; Smith, 1946; Rai er al., 1969; Mercer, 1978b). The tap root system of Sharply between 5:00 and 5:30 a. m., the beetles return ro the soil. Caged bee peanuts make it particularly susceptible ro root feeding (Rai er al., 1969; Sri tles also exhibit this distinct periodici ry in ac ri vity. Adu! r beetles prefer to feed vastava er al., 197 la). Dry weather exacerbates the wilting caused by larval on the foliage of numerous trees adjacent ro the cultivated crops and fallow root pruning. Larvae feeding on root systems and pods may facilitate secondary fields where larvae developed rather than cultivated crops. invasion of fungi (Mercer, 1978a). Adult females lay eggs singly in the soil up to a depth of IO cm. Oviposition Management. Natural Enemies. Lach11os1erna spp. grubs are parasit coincides with the onset of the monsoon. Eggs incubate for 7- 12 days. White ized by the scoliid, Scalia a11reipe11nis, a fungus, Metarrhizium anisopliae (Kalra g rubs complete 3 instars feeding on the subterranean plant parts for 8 to 10 and Kulshrestha, 196 l) and a milky disease, Bacillm spp(?) (Patel et al., weeks. The grubs remain in the upper 15 cm of soil when the soil is moist, de 1978). Anthia sexg111tala (F.), a carabid (Rai er al., 1969), 811/0 melanostictm, a scend deeper (up to 0 . 75 m) during dry periods and rerurn close to the surface road and Gecko gecko, a gecko (Kalra and Kulshresrha, 1961) prey on adults. when soi l moisture returns. The g rub population rends to be denser in rhe Birds may be an important larval predaror, especially in newly plowed fields raised portions of infested fields. Full grown g rubs descend deeper in the soil (Yadava et al., 1975). Prasad ( L96 l) reviewed the importance of natural ene (up to 0. 75 m) to pupate. Pu par ion requires 13 days at 27C. The adults hiber mies for regulating white grubs in India and suggested the introduction of B11- nate in the soil until the onset of the next monsoon. Only I generation occurs fo 111ari11m, the Surinam road, ro bolster predation. annually. The natural enemies of the white grubs in rhe genus Leucopho/is were re In Australia, the scarabs Rhopaea magnirornis Blackburn, He1ero11yx brevicollis viewed by Leefmans ( 1915). Numerous scoliid wasps were reported to parasir Blackburn and Trissodon ( = lsodo11) p111wicollis Maclure (Smith, 1946) are pests ize Le11copholis with l species, Die/is thoracica, parasirizing 26% of the larvae. ofpeanurs. Only R. magnicornis and H. brevicollis larvae are damaging with T. Insecticides. Insecticidal control of L. consang11i11ea in India has re puncticollis feeding on peanuts only as adults. ceived a great deal of attention (Desai and Patel, 1965; Kaul et al., 1966; Patel Adu! t R. magnicornis are active between November and J anuary with females et al., 1967; Ravindra and Thobbi, 1967; Rai er al., 1969; Sharma and oviposiring in the soil. Larvae hatch within 2 Ldays and feed for approximately Shinde, 1970; Srivastava er al., 197 lb; Bindra and Singh, 1972; Yadava and 16 months. The last larval insrar is a voracious feeder and occurs in November Yadava, 1975; Prasad, 1977; Yadava er al., 1977; Yadava et al., 1978). Con of the second year. Pupation occurs in an earthen cell deep in the soil. The re trol has been targeted roward both adults and larvae. Grub control with a gran sulting adult emerges wirh the onset of the rains. This scarab has a 2-year life ular insecticide has increased yields by 794 kg/ha when the insecticide created cycle and attacks peanuts planted on previously fallow or pasture land where area had 24% plant mortality and 68% grub control (Prasad, 1977). the adult oviposired the year prior to rhe crop being planted. The adults also Applying an insecticide to the trees where the adult beetles congregate to rend to be active on friable, volcanic, red soils (Smith, 1946). feed has had some success (Yadava et al., 1977). It is imperative that the tim Adults of H. breviroflis oviposir ar the base of peanut plants and weeds. ing of the sprays coincide with peak adult emergence after the monsoon onset. Adults prefer oviposirion sites which offer a hi gh degree of physical cover and Control must be applied on an areawide basis. are arrracred ro organic manure (Passlow, 1969). In Africa, Eulepida mashona Cultural. Species of white grub adults attracted ro organic manure Arrow is artracred ro organic manure for oviposition with outbreaks more (e.g., E. mashona, Rose, 1962; Broad, 1966; Heteronyx spp., Passlow, 1969) prone to occur in areas where Brachystegict spp. trees are dominant (Rose, 1962; for oviposition could be managed by avoiding the practice of deep burial or or Broad, 1966). The adults are abundant for the first few weeks ofrhe rainy sea ganic manure immediately after application. White grubs with 2-year life cy son , feeding on the foliage of Bmchystegia spp. and related trees. cles (e.g. , R. 111ag11icornis), where oviposition originates in weedy fallow or pas Strigoderma arborirola F. is a common, widely distributed, white grub feed ture lands, are manageable by planting peanuts on land previously under culti ing on the pods of peanuts in the eastern Uni reel States (Miller, 194 3; Grayson, vation for 1 or 2 years (Smith, 1946). 1947). This insect has 1 generation per year with rhe third insrar overwinter ing at a soil depth of !8-20 cm (Gray.son, !946). Adult females lay an average OVERALL PERSPECTIVE of26. 3 eggs which require 8-2 1 days to hatch. The first 2 larval insrars last 12- 30 and 10-36 days, respectively, with rhe third and final insrar lasting 238- The body of knowledge in population ecology comprises the theoretical ba 292 days (Grayson, 1946). Peanuts grown in dark soi l, high in organic matter sis for the use of integrated pest management (IPM), and violations in ecologi and where the plants are large, are more prone ro attack (Miller, 1943). cally founded management principles have caused severe constraints in design Damage. Adults are either defoliators of peanuts (e.g., I-Je1ero11yx spp., of pest management strategies (Bottrell, 1979; Barfield and Stimac, 1980). 308 P EANUT SCIENCE A D TECHNOLOGY MANAGEMENT OF PREHARVEST ] NSECTS 309 ~o rl dwide pea~ut crop pr~d u crion and protecrion have nor been immune ro Table 6. Pesr stacus o f selecr pea nut insect pests w o rldwide. rrn srakes m~de in orher ag n c ulr~ra l producrion systems. The present chapter United States South America Africa Asia h_as summ.anzed ~est types, panrcular pests, variant pest biolog ies and protec tron practices for 1nsecc pesrs of peanucs worldwide. Several observations can be Th rips Non-pest Key pest {unknown) (unknown) made concerning th!s summa~ which wi ll allow analysis of where improve m.ents can be made in the desig n of peanut crop protection strategies worl d Aphids Non-pest Non-pest Key pest Key pest wide. Spider Mites Secondar}' pest (unknown) Secondary pest Secondary pest . An overview of tactical approaches to insecr management in peanuts world wide er: able 5A) '.eveals rhar, b~ far, chemical pesricides comprise rhe mainstay E.111poa1w Non-pest {unknown) (unknown) (unknown) pr~ r ec t1on practice; yet, published :esea.rch depicts a rremendous potential (Table 58) .for d ev~ l opment of mulmarnc lPM strategies sufficiently adapt Foliage Occasional Occasional Occasional to Occasional Consumers pest pest Secondary pest pest able to pam cular sites. Furche:, select insect pesrs vary from key pesrs to non p.escs ( T~b l e 6) ~ ve r geographical space. This obviously reflects rhe effects of E.la1111opalpm s 1ce -spe~ 1fi c e n~1ronments (b i ~ r ic and abiocic components) on pesr dynamics. lig11oull111 Key pest Key pest Does not occur Does noc occur Thus, s.1ce specific lPM strategies .m usr be ~crucrured consistent wirh local pest dynamics. Only rhen can we begin ro decipher the general icy of a g iven man Diabrorira spp. Kc)• pest Occasional pest (unknown) (unknown) agement scraregy. (regionally) Tables 5"., B reflecr a basic problem seemingly common ro mosr all current \Xlhite Grubs Occasional pest Occasional pest Occasional pest Occasional pest crop.pro.rccrwn prograi:is: almost unilateral dependence upon pesticides often applied in a prophylacnc mann ~ r. Economic thresholds, sampling merhodolo Termites Non-pest Non-pest Key pest Key pest g1e.s and systems level desrgn , inherent to urilizing pesricides under rhe lPM philosophy, are generaJl.y lacking. Wirhin rhese 3 areas lie rhe keys ro improv- The concept of an economic threshold is not new ro agricul ture and has been 111g peanut crop p rotection srrareg1es based on rhe pesticide tactic. reiterated earl ier. Despite much dialogue on the concept, researchers simply have not addressed the complexity of construction of realistic, dynamic thresh Table S · Cu~r e nc u sage o f select !PM ca.ctic~ .an d sampling for select groups of peanut in olds. Peanut researchers are no exception. The literature is clogged with dis sccc pcscs worldwide (A ) a nd avadabd1ty of cactics and sampling plans (8) whe the r cussions that true economic thresholds are functions of pest density, crop o r n~t accually ~ sed. L~rg e d e nominario n (X) rcpresenrs s ignificanc use; small (x) stage, physical environment, market value, pest combinations, etc. Yee, che reprcsenrs relauvely mmo r use. Blank spaces rc prescnc n o n-availa bilit)r. fact remains rhac, in actual icy, economic thresholds (where used at all) are used as static val ues for sing le pests. Until agriculrural researchers (i n this case, for B1olos1cal peanuts) address che conscruccion of more realistic thresholds, crop protection Control " schemes will continue ro be based more on experience rhan on the dynamics of ,,,~ specific ecological/sociological/economic srages. Barfield and Stimac (1980) .§ ~ e v ~ ;; r: "- ~ argue char systems models offer che most promising cools currently available " " c -0 .\< -0 0::" ~ ~ t :g c ""§ a ."\< "E "" g for derivation of these thresholds. § ;;; " 9 8. ~ v :; -0 c :; ..: 0 ~ 0 0 :t :,,;" E c v " v ~ As a whole, peanut (as ocher) lPM practitioners have a poor perception of the "- u <""" The proper use of pesricides (i. e. , d ynamic economic rhresholds, reliable We also appreciate rhe assistance of Dr. Horace Burke, Dr. J oseph Schaffner sampling merhodologies, rarger pesr selecriviry, ere.) requires more rechnol (Texas A&M U niversity) and Dr. Fred Bennett (Director, Commonwealth In ogy on the user level than orher IPM racrics (e.g . , hosr plant resistance, culru stitute of Biolog ical Control, Curepe, Trinidad, \' Aeanasov. N. 1971. The Adantic mite (Ttlranychus atlantiCJ11 McGregor): A pest of ~roundnut. Ras- Bertels, A. 1970. Esrudos da influencia da umidade sobre a dinamica de populacoes de lcpidopteros pragas teniev"dni Nauki 8: 149-155. · . do milho. Pesquisa Agropecuaria Brasileira 5:67-79. Aetiah, H. H. and R. A. Rizk. 1973. On the: control of the green spider mite Tttranychus arabi&llJ Attiah to· Bhardwaj, S. C. and K. S. Kushwaha. 1977. Effect of host planes on the larval and post-larval development fcsting peanut plants in Egypt. Agric. Res. Rev. 51:109-112. . . of hairy caterpillar, Amsana /into/a Fabricius. Indian J. Entomol. 38:230-232. Aucamp, J. L. 1969. The role ofmite vectors in the development ofaflatoxm m groundnuts. J. Stored Prod. Bhatnager, S. P. 1970. Records of new Cctonides pests in Rajasthan State. Labdev J. Sci. and Tech. 5:245-249. . 85:119-120. Backman, P.A.,J. D. Harper,). M. Hammond, and E. M. Clark. 1977. Antifeeding effects of the fungi Bhatnager, V. S. and J. C. Davies. 1978. 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V., D. A. Emery, and W. C. Gregory. 1971. Registration ofGP-NC343 peanut germ Bergman, B. H. H. 1956b. On three Jassidac of Arachis hypogata in Java. Entomol. Ber. 16:64-71. plasm. Crop Sci. 11:605. 3 14 PEANUT S CIENCE AND TECHNOLOGY MANAGEMENT OF PREHARYEST I NSECTS 315 Campbell. \VI . V., D . A. Emery, J .C. Wynne, and R W . Batts. 1976. lntcrnccion ofpean\I< variety and Dupree, M. 1964. Insecticide and cul rural concrol of 1hc lesser cornsialk borer. Mimeogr. Univ. Ga. Agric. insecundc. Peanu1 ci. 3:75-77. Exp. Sia., N. S. 197. 21 pp. Campbell, \VI . V. , J .C. W ynne, D. A. Emery, and R. \'II. Mozingo. 1977. Regis1rarion of NC 6 peanu1s. Dupree, M. 1965. Observations on 1he life his1ory of 1he lesser cornstalk borer. J. Econ. Emomol. 58: 1156- Crop. Sci. 17:346. 1157 . Canerda)•, T . D. Jnd F. S. Arnn!. 1966. The looper complex in Alabama (Lepidmera, Plusiinac). J . Econ. Duzgunes, Z . 1959. Tt1rar1;rh111111midtllm, the ground nut mite. Plane Pro1. Bull. I: 10-14. 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Al l. 1979. f eeding behavior oflesser cornsialk borer larvae in simulat ions of 197 pp. no-1illagc. mulched conveniional 1illag e, and conwmional 1illagc corn cropping sys1cms. Environ. En- Fennah , R. G . 1947. The insect pem offood crops in the Lesser Antilles. Dept. Ag ric., Antigua, British 1omol. 8:26 1-264. West Indies. 207 pp. Cheshire. J . M .. Jr.. J . Henningson. andj. N. All. 1977. Radiolabcli ng lesser corns1alk borer larvae for Fink, D. E. 19 16. Injury co peanu1s by 1he rwelvc-spo1tcd cucumber beetle (Diabrotira 12-p1111aa111). J. monirnring movcmcm in soil habitats. J. Econ. Enrnmol. 70:578-580. Econ. Enrnmol. 9:366-369. China, \Y/. E. 1928 . Two new species ofCir11tl11/11111, China (Homop1cra, Jassidae) from the G:11nbia, \Xlcs1 Flechrmann, C. H . \XI. 1968. Phyrophagus mires associaicd wi1h forage plants. Pesquisa Ag ropecuaria Bra Africa. Bull. Entomol. Res. 19:6 1-63. sileira. 3: 17 1-172. China, \XI. 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W., Jr., L. Posada, and 0. D. Smich. 1980a. Greenhouse evaluation of 490 peanur lines for re Singh, S. and J. Singh. 1956. On rhe control of kurra, t\111Jfl{ffl moorei Buder(Arcriidae: Lcpidoptera). Indi sisrance ro rhe lesser cornstalk borer. Texas A&M Univ. Syst., Tx. Agric. Exp. Sta. Misc. Publ. 1464. an). Hort. 13: 107. 42 pp. Singh, S. R. , 11. F. van Emden, and T . A. T•ylor, eds. 1978a. Checklist of insect and mite pesrs of grain Smith, J. W., Jr., L. Posada, and 0. D. Smith. 1980b. Greenhouse screening peanut germ plasm forresis legumes, pp. 399-'1 17. / 11 Pesrs of Grain Legumes: Ecology and Control. Academic Press. New York. rance to rhe lesser cornstalk borer. Peanur Sci. 7:68-7 l. 45'1 pp. Smirh, J. W., Jr. and R. Sams. 1977. Economics ofrhrips cont rol on peanu cs in Texas. Southwestern En Singh,S. R., II. F. van Emden. and T .1\ . Taylor, eds. 1978b. Checklisr ofn:u ural comrolagentsofgrain tomol. 2: 149- 154. legume pests. lu Pests of Grain Legumes: Ecology and Control. 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