....... mm W WW 3 00646 4204 LIBRARY Michigan State | University This is to certify that the thesis entitled TEMPORAL AND SPATIAL DISTRIBUTION AND DISPERSAL PATTERNS OF PARAPHLEPSIUS IRRORATUS (SAY) (HOMOPTERA: CICADELLIDAE), A VECTOR OF X—DISEASE IN MICHIGAN presented by KIRK JON LARSEN has been accepted towards fulfillment of the requirements for M. 5. degree in ENTOMOLOGY %( Pétv’ Major professor [Mark E. Whalon Date g/f/g % 0-7639 MS U is an Affirmative Action/Equal Opportunity Institution MSU RETURNING MATERIALS: Piace in book drop to LIBRARIES remove this checkout from —:I!—— your record. FINES will be charged if book is returned after the date stamped below. OQESQSI‘figfiti TEMPORAL AND SPATIAL DISTRIBUTION AND DISPERSAL PATTERNS OF PZRMPWIETSIUS IRRORATUS (SAY)(ROHOPTERA: CICADELLIDAE), A VECTOR OF X-DISEASE IN MICHIGAN by Kirk Jon Larsen A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Entomology 1987 ABSTRACT TEMPORAL AND SPATIAL DISTRIBUTION AND DISPERSAL PATTERNS OF PMRMPWZETSIUS IRRORATUS (SAY)(HOMOPTERA: CICADELLIDAE), A VECTOR OF X-DISEASE IN MICHIGAN by Kirk Jon Larsen Populations of leafhopper vectors of X-disease, a major disease problem of the Michigan peach industry, were monitored by yellow sticky board traps and sweepnet samples during 1985 and 1986. Parapblepsius jrroratus represented over 70% of all known vectors found. The appearance of symptomatic chokecherry indicated X-disease transmission was occurring throughout the state. Daily activity of P. jrroratus was monitored by light- trap and sweepnet sampling orchard sub-habitats. P. irroratus is found in the groundcover during the day, has a crepuscular flight into cherry trees at night, and returns to the groundcover in the morning. Rate and extent of P. jrroratus dispersal within peach and cherry orchards was studied by a mark, release and recapture experiment. The overall recapture rate was 2.35%, with an average dispersal rate of 3.42 m/day. The major factor influencing leafhopper dispersal was wind, with temperature influencing activity. DEDICATION To Dr. Harvey Blankespoor, who is responsible for introducing ne to a beautiful and complex part of God’s creation, the insects. ii ACKNOWLEDGEMENTS I would like to thank Dr. Mark Whalon, my graduate advisor, for his direction, prayers and support throughout my master’s program, and for offering me the opportunity to learn and work with him. In addition, I thank the other members of my graduate committee, Dr. Alan Jones of the Department of Botany and Plant Pathology, and Dr. James Miller and Dr. David Smitley of the Department of Entomology. Their guidance, encouragement, and helpful suggestions during the preparation of this thesis were most appreciated. The cooperation of the Michigan stone fruit growers whose sites we used in this research is gratefully acknowledged. The assistance of Dr. Robin Taylor of the Department of Entomology, the Ohio State University, in developing the dispersal equations is sincerely appreciated. I sincerely thank field workers Jay Matthes and Martha Zemper for their assistance in the field and enduring the many long miles of travel. Finally, a special thanks.to my wife, Shirley, for her continual support, interest and patience. Spending many of our first evenings together assisting ”sucking bugs" and fighting off the swarms of mosquitoes has surely been an extraordinary expression of her love. iii TABLE OF CONTENTS LIST OF TABLES . . . . . . . . . . . . . . LIST OF FIGURES . . . . . . . . . . . . . . . . . . . GENERAL INTRODUCTION AND LITERATURE REVIEW . . . Review of X-disease . . . . . . . . . . . . Biology of Paraphlepsius irroratus (Say). 0 O Leafhopper movement behavior. . . . . . . . . . . Conclusion. References Cited. . . . . . . . . . . . . . . . CHAPTER I. FIELD MONITORING OF X-DISEASE LEAFROPPER VECTORS AND SYMPTOMATIC CHOKECHERRY Introduction. . . . . . . . Materials and Methods . . . . . . Field Season and Research Sites. Symptomatic Chokecherry Survey . X-disease Vector Leafhopper Survey Results . . . . . . . . . . . . . . . Field Season . . . . . . . . Symptomatic Chokecherry Survey . Vector Leafhopper Survey . . . . . . . . . . Discussion. . . . . . . . . . . . . . . . . . . . References Cited. . . . . . . . . . . . . . . . . CHAPTER II. CREPUSCULAR MOVEMENT OF PMRMPWZEPSIUS IRRORATUS (SAY), BETWEEN THE GROUNDCOVER AND CHERRY TREES . . . . . . . . . . . . . Introduction. . . . . Materials and Methods . . . . . . . . . . . . . . iv vi .viii p—o 0310(0me H 19 20 21 21 22 22 25 25 27 27 34 41 44 44 TABLE OF CONTENTS, continued Results . . . . . . . . . . . . Discussion. . . . . . . . . References Cited. . . . . . . . . CHAPTER III. DISPERSAL OF PMRMPWZEPSIUS IRRORATUS (SAY) IN PEACE AND CHERRY ORCHARDS. . Introduction. . . . . Materials and Methods . . . . . . Leafhopper Capture and Marking Effects of Marking on Survival O 0 Effects of Marking on Flight Activity. Field Release and Recapture. Results . . . . . . . . . . . Effects of Marking on Survival and Flight Activity. . . . . . . Temporal Patterns of Trap Catches. Spacial Patterns of Trap Catches Discussion. . . . . . . References Cited. . . . GENERAL CONCLUSION . . . . . . . . . APPENDIX A. LEAFHOPPER VOUCHER SPECIMENS THE MICHIGAN STATE UNIVERSITY ENTOMOLOGICAL MUSEUM . . . APPENDIX B. FIELD MONITORING, CREPUSCULAR MOVEMENT, AND DISPERSAL DATA TABLES. PLACED IN 46 52 56 57 58 58 58 60 60 61 67 67 67 69 79 86 88 92 94 Table 1. Table I-l. Table I-2. Table I-3. Table 11-1. Table 11-2. Table III-1. Table III-2. Table III-3. Table A. LIST OF TABLES Enown leafhopper vectors (Homoptera: Cicadellidae) of Peach X-disease in North America. . . . . . . . List of X-disease sites for the 1985 and 1986 field seasons with location data. Total number of X-disease vector leafhoppers and relative abundance found in Michigan for both 1985 and 1986 field seasons. . . . Number of X-disease vector leafhoppers found at each field site during 1985 and/or 1986. . . . . . Mean number of P. jrroratus leafhoppers captured by 25 sweeps each 0.5 hr over 24 hrs, three replications in two generations. . . . Circular-linear rank correlation of the number of leafhoppers captured over a 24 hr period in 4 sub—habitats Number of marked leafhoppers released and recaptured and the recapture rate and for both sites, release locations, generations. . . . . . Mean number of P. irroratus leafhoppers recaptured at different trapping distances with the count following transformation by the from release location, interference factor. . Total number of marked P. generations. . Voucher specimen data. vi 0 irraratus leafhoppers recaptured each direction from the release location for both orchard and edge locations at each site during the two 4 23 33 36 51 53 68 74 76 93 LIST OF TABLES, Table Table Table Table Table Table Table continued List of X-disease vector leafhopper species and number captured by site and date during the 1985 and 1986 field seasons. . . . . . . . . . . . . . . . . . 94 Number of X-disease symptomatic choke- cherry observed/5 miles by site and date during the 1986 field season . . . . . . . 98 Number of P. Irroratus leafhoppers captured by subhabitat and time. . . . . .100 Leafhopper dispersal survival data . . . .106 Leafhopper dispersal flight to yellow sticky board trap data . . . . . . . . .107 Leafhopper field recapture data. . . . . .108 Number of male and female P. irroratus captured by method, site and date during 1985 and 1986. . . . . ... . . . . . . . .115 vii Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure I-2. I-4. I-9. II-l. LIST OF FIGURES The field sites monitored for leafhopper vectors of X-disease during 1985 and 1986, and monitored for the appearance of symptomatic chokecherry during 1986. . . . 24 Accumulation of degree day heat units over time at the sites during both 1985 and 1986 field seasons. . . . . . . . . . . . . . . 26 Mean degree day accumulation for the southwest and northwest areas of lower Michigan in 1986 . . . . . . . . . . . . . 28 Mean number of symptomatic chokecherry visually observed/km of two lane roadway in the southern and northern regions of the western Michigan stone fruit belt during 1986. . . . . . . . . . . . . . . . 29 Mean number of X-disease vector leafhoppers captured over time in 1985 and 1986 at all sites. . . . . . . . . . . . . . . . . 30 Mean number of X-disease vector leafhoppers captured, based on average degree day accumulations in 1985 and 1986 at all sites . . . . . . . . . . . . . . . 31 Number of X-disease vector leafhoppers captured at each site during the 1985 and 1986 field seasons . . . . . . . . . . . . 32 Total number of common X-disease vector leafhoppers captured at each site during the 1985 and 1986 field seasons. . . . . . 35 Percent of male and female P. jrroratus leafhoppers captured by monitoring method for both 1985 and 1986 field seasons . . . 37 Mean number of P. jrroratus leafhoppers sampled by light trapping over a 24 hr period for both generations. . . . . . . . 47 viii LIST OF FIGURES, continued Figure Figure Figure Figure Figure Figure Figure Figure Figure Figure II"2. 11-3. III-1. III-2. Ill-3. III-4. III-5. III~6. III-7. 111,8. Mean number of P. irroratus leafhoppers collected in sweep nets every 0.5 hr over a 24 hr period in each subhabitat type for both generations . . . . . . . . . . . . . 49 Mean number (+SE) of P. irraratus leaf- hoppers collected over 24 hrs in sweep nets in the four sub-habitats during first and second generations . . . . . . . . . . . . 50 Location of the two field research sites used in this leafhopper dispersal study. . 62 Release locations, recapture trap layout and the surrounding area at the East Lansing, MI research site . . . . . . 63 Release locations, recapture trap layout and surrounding area at the Lawrence, MI research site . . . . . . . . . . . . . . 64 Arrangement of the yellow sticky board traps used to recapture marked P. irroratus leafhoppers around each release location . . . . . . . . . . . . . 65 Number of P. irroratus leafhoppers recaptured each day for the 21 days following their release. . . . . . . . . . 70 Number of marked and unmarked P. jrroratus leafhoppers captured per yellow sticky board trap per day over the mean temperature during the daily two hour crepuscular flight period following sunset . . . . . . . . . . . . . 71 Actual and transformed mean number of P. irroratus'leafhoppers recaptured at different trapping distances, with the dispersal equations and expected lines . . 73 Number of P. jrroratus leafhoppers recaptured each direction with the mean dispersal and mean wind direction during flight times for recaptured leafhoppers. . 77 ix LIST OF FIGURES, continued Figure III—9. Figure III-10. Mean rate of movement of marked P. irroratus leafhoppers recaptured each direction from release for both sites, release locations, and generations . . . . 78 Mean rate of movement of marked P. jrroratus leafhoppers recaptured each distance from release for both sites, release locations, and generations . . . 80 GENERAL INTRODUCTION AND LITERATURE REVIEW Review of x—disease In the United States in 1982, Michigan ranked first in sour cherry production, third in sweet cherry production, and sixth in peach production (Fedewa & Psocodna 1982). In a 1981 report from Michigan cooperative extension agents, X~ disease was considered a major peach disease problem in southwestern Michigan. An annual loss to X-disease of $1.5- $3.0 million in peach production (M. Whalon, personal communication) is estimated from Michigan Department of Agriculture (MDA) survey data from 1977-82 and 1985-86. The MBA annual survey indicates the incidence of X-disease has increased in peach orchards of southwest Michigan during the past several years with 83% of the peach orchards inspected during 1985 showing the presence of X-disease (Robinson 1985). The causal agent of X-disease is a mycoplasmalike organism (MLO) (Granett & Gilmer 1971, Jones et a1. 1974) that can be transmitted by several species of leafhoppers. MLO’s are microscopic, single-celled prokaryotic organisms similiar to mycoplasmas (Agrios 1978). Two hypotheses have been proposed as to how MLO’s cause disease (Razin 1978). The MLO’s either clog up the phloem tubes, thus inhibiting nutrient translocation through the plant, and/or produce toxins which kill the plant. Leafhopper vectors of X-disease (Ho-optera: Cicadellidae) are primarily of the subfamily Deltocepbaljnae (Gilmer & Blodgett 1976). At least 18 species are known to be capable of transmitting X-disease either naturally or experimentally (Table 1). The species of greatest importance in the spread of X—disease in peach and cherry varies in different regions of the U.S. and Canada (Elliott & Dirks ND). The seasonal distribution and abundance of different leafhopper vectors within the same geographical region also varies considerably (McClure 1980b). The most important vector of X-disease in the eastern U.S. is Scaphytopius acutus (Say) (Palmiter et a1. 1960), while Collodonus montanus (VanD.) is the major vector in the western U.S. (Gilmer & Blodgett 1976). In Michigan, the most important vector of X-disease is presumed to be Parapblepsius jrroratus (Say) (Taboada et a1. 1975). There are thought to be at least two separate strains of X~disease MLO’s, eastern and western, because of variation in symptom development between eastern and western orchards. More recent DNA hybridization research (Kirkpatrick 1986, M. Whalon, personal communication) indicates a lack of homology between California and Michigan isolates of X-disease MLO’s (Whalon, unpublished data). Research is ongoing in the genomic DNA hybridization approach for differentiating the X-disease isolates in host plants and insect vectors. Peach X-disease symptom development begins in mid-June. Healthy and sick trees are most easily distinguished by symptoms during August (Palmiter & Hildebrand 1943). There Table 1. Known leafhopper vectors (Romoptera: Cicadellidae) of Peach X-diseaae in North America (after Nielson 1979 and Chiykowski 1981). Strain Transmission Species Field Greenhouse Author a Date 8883:3888::33:38:33:832883833833833333338888383333388888338338838338:38:33: Eastern collodbnus clitellsrius (Say) FTeberieIle {20:11 (Sta1.) Gyponsns lamina DeLong NbrveIIIns semlnuds (Say) Orientus Isbjdbe (Mat.) Parapblepsius irrorstus (Say). Scspboideus melanotus Osb. S. titans: Ball Scspbytopjus scutus (Say)b Restern chnopterus sngulstus Lawson Collodonus (eminatus (Van Dusee) C. mantsnus (Van D.)° luscelidius vsriegstus Eirsh. FTeberieJIa {Terri (Stal.) leonolla confluens (Uhler) Osbornellus borealis DeL. s Mohr Scspbytopius scutus (Say) 5. delongi Young S. nitridus (DeLong) X 88 Thornberry 1954, Oilmer 1954 Oilmer s Mclwen 1958 Gilmer & McEwen 1958 Cilmer et a1. 1966 Rosenberger & Jones 1978 Gilmer et a1. 1966 Rosenberger & Jones 1978 Rosenberger s Jones 1978 Hildebrand 1953 Purcell 1979 Rolfe et a1. 1950 Rolfe 1955 Jensen 1969 Rolfe et a1. 1951 Anthon & Rolfe 1951 Jensen 1957 Anthon a Rolfe 1951 Swanson 1971 Purcell 1979 'P; Irrorstus is the most important vector of X-disease in Michigan (Taboada et a1. 1975). I'S. scutus is the most important vector of X-disesse in Eastern North America (Palmiter et a1. 1960). cc: montanus is the most important vector of X-disease in Restern North‘ America (Gilmer & Dlodgett 1976). is a slight delay in the foliation of diseased trees which is often missed (Gilmer et a]. 1954). Peach X-disease has similiar yet distinct symptoms to nitrogen deficiency, bacterial spot (Xantbomonas prunj), and Leucostoma canker (Dhanvantari & Kappel 1978). Peach X—disease symptom expression begins with rolling and yellowing of the leaves on infected branches. Blotchy, irregular, and water-soaked spots then develop across the leaf veins which become brittle and fall out, giving the leaf 8 shot-holed and tattered appearance. The older leaves on infected branches fall off, leaving the branches with a small rosetted tuft of leaves at the end. The peach fruit either aborts and drops early or is smaller and more pointed than usual. Fruit that remain on the tree ripen prematurely, and are bitter and unpalatable. Peach trees rarely survive three years after symptoms are noted unless treated with tetracycline. Dieback begins with the diseased branches and spreads branch by branch to the entire tree (Dhanvantari & Kappel 1978, Gilmer et a1. 1966, Palmiter & Hildebrand 1943). X-disease agent has a wide variety of woody and herbaceous host plants. The economically important hosts include Prunus persica Batsch (peach), P. cerasus L. (sour cherry) and P. avjum L. (sweet cherry) (Gilmer et a1. 1966). Wild hosts can serve as an outside source of X-disease inoculum. Chokecherry (P. Virginians L.) is the most important wild woody host (Gilmer et al. 1954). Other plants known as experimental hosts include over twenty herbaceous species in eleven families (Chiykowski & Sinha 1982), many of which are common within and near most orchards. Insect transmitted plant diseases such as X-disease are difficult to control due to the interactions between plant, pathogen, and insect vector. Historically, control of X- disease has involved eradication of alternate hosts such as chokecherry, application of tetracycline antibiotics, removal of diseased trees, and vector control (Lacy et a1. 1979, Rosenberger 1977). Control of MLO diseases by killing insect vectors with insecticides after they have arrived at the crop has seldom proved effective. Even with good insect control, enough insects survive for sufficiently long periods to spread the pathogen (Agrios 1978). The presence of orchard groundcover often reduces the effectiveness of insecticides used in leafhopper control efforts, and the continuous presence of vectors from June until November makes insecticide use economically impractical (Palmiter & Adams 1957). Current management methods of X-disease in Michigan such as alternate host eradication and unilateral insecticide control of leafhoppers have been ineffective means of X-disease control (Robinson 1985). Under most insecticide programs, peach orchards are not sprayed when vector populations peak in the fall (Taboada et a1. 1975). Integrated pest management (IPM) is the best approach to disease control, yet is often difficult to implement due to the complexity of the disease transmission cycle and the interaction of the various disciplines involved (Rhalon & Croft 1984). IPM requires a holistic approach to the problem, integrating the knowledge of MLO’s by plant pathologists, insect vectors by entomologists, and the host plants by horticulturalists. Current distributions of the disease problem must be known to determine the best management strategy. Information is needed on the occurence of X-disease in the host plants and alternate hosts, vector presence, vector biology and behavior as it relates to disease transmission, and the effect any control tactics may have on X—disease transmission. This information is obtained by monitoring the incidence of infective vectors, X-disease in host plants and in alternate hosts such as chokecherry. From this information, appropriate control tactics can be defined. In order to implement an IPM program for X—disease, several tools are still needed. Probes for detection of MLO’s are necessary to evaluate the relationship of MLO’s to both the host plants and insect vectors. A better understanding of the incidence of X-disease in host plants and insect vectors, the biology, distribution and abundance of vector leafhopper populations, and the short-term and long-term movement behavior of these leafhopper vectors of X-disease is therefore necessary. Biology of Parapblepsjus irroratus (Say) P. jrroratus is a major vector of X-disease in Michigan peach and cherry orchards (Taboada et al. 1975). Little is known about the daily movement and distribution or biology of P. jrroratus, as the first paper on the biology of P. jrroratus was not published until 1985 (Chiykowski). Previous surveys have shown that P. irroratus was the most common X-disease vector leafhopper in Michigan peach and cherry orchards (Taboada et a1. 1975) and the most efficient vector of X-disease in greenhouse tests (Rosenberger & Jones 1978). P. jrroratus was thought to be primarily an herbaceous species, although it has been observed and collected on a wide variety of woody hosts (Hamilton 1975, Chiykowski 1985). Nymphs of P. irroratus have been observed on grasses in cherry orchards (Phillips 1951), and raised experimentally on a combination of barley and clover (Chiykowski 1985). P. irroratus was more common in sour cherry than in peach orchards (Rosenberger 1977) and was bivoltine in Michigan, with the two periods of adult activity being late-June to July and late-September to October (Taboada et a1. 1975). .P. irroratus is thought to overwinter in Michigan in the egg stage. Rosenberger and Jones (1978) suspected that adult P. irroratus were most active in the early evening when they routinely collected leafhoppers at twilight around yellow lights. Increased activity during twilight has been documented in many insects, including leafhoppers (Barker 1961). Yellow is known to attract certain leafhopper species (Alverson et a1. 1977) and has proven to be an excellent means of capturing large numbers of P. jrroratus during their crepuscular active periods (Larsen & Rhalon 1987). Indirect methods of leafhopper sampling have been compared and contrasted with direct censusing of leafhoppers in fruit trees and orchard groundcover (Mowry 1982). Although indirect sampling methods have distinct drawbacks (DeLong 1932, Southwood 1978), relative methods such as light trapping and sweep net sampling were the best methods of obtaining frequent relative density estimates of mobile leafhopper populations (Mowry & Rhalon 1984). Leafhopper Movement Behavior Movement behavior is an important aspect of the distribution and abundance of insect populations. Insect movement is often described as migratory or dispersive. Mfigratjon can be defined as an adaptive departure from a breeding area or other habitat which is no longer fit to support a population. It is often a persistent, uni- directional and long distance movement, during which all activities but flight are ceased, necessary to ensure the survival of a species (Kennedy 1961). Dispersal is an accidental, continuous movement within a habitat, during 10 which insects become scattered over a wider area than originally occupied (Johnson 1969). Since X-disease MLO’s are transmitted only by leafhoppers, understanding the movement of these vectors is necessary to learn more about the epidemiology of X-disease (Purcell 1985). Leafhopper movement may be local, as between plants in a field, dispersive, as from area to area within a habitat, or migratory, in which the leafhoppers may move considerable distances (Chiykowski 1981). Factors that may influence these movements are many and involve biotic factors such as the normal life history of the insect, its host range and preferences, the availability of these hosts and their status as disease reservoirs, and physical factors of the environment (Carter 1961). P. irroratus is not known to be involved in long distance, migration type movements, although P. jrroratus adults have been trapped in low numbers at altitudes of 137.2 m (450 ft) (Osborn 1932) and more than 14.5 km (9 mi) from land (Sterns & MacCreary 1938). Examples of leafhoppers which do migrate over long distances include the aster or six-spotted leafhopper,.MbcrosteIes fascjfrons (Stal), a vector of aster yellows (Chiykowski & Chapman 1965, Drake & Chapman 1965, Nichiporick 1965), the beet leafhopper, Circuljfer tenellus (Baker), the principle vector of curly top virus (Severin 1933, Dorst & Davis 1937, ll Lawson et a1. 1951), and the potato leafhopper, Empoasca fsbae (Harris), a major pest of such crops as potatoes, soybeans and alfalfa (Glick 1960, Pienkowski & Medler 1964). Local insect movement, which usually includes dispersal, often involves a systematic daily or seasonal oscillation between areas, not unlike the human activity of commuting (Taylor 1985). Studies on adult feeding and ovipostion of S. acutus, the major vector of X-disease in Connecticut, indicate the adults mature on wild hosts, fly to peach trees to feed, and then return to wild herbaceous hosts to oviposit (McClure 19808). This "commute", if occurring daily, would significantly affect transmission of any leafhopper—borne pathogen. Dispersal of insects is known to be influenced by meteorological conditions (Taylor 1985). Dispersal behavior of leafhoppers has been studied using mark and recapture techniques for both the western X—disease vector Collodonus montanus Van Duzee (Purcell & Suslow 1982) and the blueberry stunt disease vector Scspbytopjus magdalensjs (Provancher) (Whitney & Meyer submitted). Mark and recapture movement data allow application to an insect dispersal model (Taylor 1978) and can also giwe absolute population estimates (Southwood 1978). Conclusion Knowledge of the temporal and spacial distribution and dispersal patterns of P. irroratus, the suspected major 12 vector of X-disease in Michigan, is needed. This research attempts to quantify local movement, such as daily distributions and host preferences in the orchard, and dispersive movement of P. irroratus. With this information, evaluation of established control procedures will help in the development of new X-disease management strategies. References Cited Agrios, G.N. 1978. Plant diseases caused by mycoplasmalike organisms. Pp. 511-536. In: Plant Pathology. 2nd ed. Academic Press, N.Y. 703 pp. Alverson, D.R., J.N. All and R.R. Matthews. 1977. Response of leafhoppers and aphids to variously colored sticky traps. J. Georgia Entomol. Soc. 12:336-341. Anthon, E.R. and H.R. Rolfe. 1951. Additional insect vectors of western X-disease. Plt. Dis. Rep. 35:345-346. Carter, R. 1961. Ecological aspects of plant virus transmission. Ann. Rev. Entomol. 6:347-370. Chiykowski, L.N. 1981. Epidemiology of diseases caused by leafhopper—borne pathogens. Pp. 105—159. In: Plant Diseases and Vectors: Ecology and Epidemiology. eds. K. Maramorosch and K.F. Harris. Academic Press, N.V. 368 pp. Chiykowski, L.N. 1985. Biology and rearing of Parapblepsjus irroratus (Homoptera: Cicadellidae), a vector of Peach X—disease. Can. Entomol. 117:717-726. Chiykowski, L.N. and R.K. Chapman. 1965. Migration of the six—spotted leafhopper Macrosteles fbscjfrans (Stal). Part 2. Migration of the six-spotted leafhopper in central North America. Univ. Wisconsin Res. Bull. 261:21-45. Chiykowski, L.N. and R.C. Sinha. 1982. Herbaceous host plants of peach eastern X-disease agent. Can. J. Plt. Pathology 4: 8—15. DeLong, D.M. 1932. Some problems encountered in the estimation of insect populations by the sweeping method. Ann. Entomol. Soc. Am. 25:13-17. Dhanvantari, B.N. and F. Eappel. 1978. Peach X-disease in southwestern Ontario. Can. Plt. Dis. Survey. 58:65-68. Dorst, R.E. and E.R. Davis. 1937. Tracing long-distance movements of the beet leafhopper in the desert. J. Econ. Entomol. 30:948-954. 13 14 Drake, D.C. and R.E. Chapman. 1965. Migration of the six- spotted leafhopper Macrosteles fbscjfrons (Stal). Part 1. Evidence for long distance migration of the six- spotted leafhopper into Wisconsin. Univ. Wisconsin Res. Bull. 261:1-20. Elliott, R.M. and V.A.Dirks. ND. Flight periods of two leafhopper vectors of peach X-disease, Scaphoideus titanus Ball and S. melanotus Osb. (Homoptera: Cicadellidae), and related species in Ontario. Unpublished mimeo. Fedewa, D.J. and S.J. Psocodna. 1982. Michigan Orchard and Vineyard Survey. 1982. Michigan Depart. Agric. 49pp. Gilmer, R.M. 1954. Insect transmission of X-disease virus in New York. Plt. Dis. Rep. 38:628-629. Gilmer, R.M. and E.C. Blodgett. 1976. X—Disease. Pp. 145— 155. In: Virus diseases and noninfectious disorders of stone fruits in North America. U.S. Dept. Agric. Handbook 437. 433pp. Gilmer, R.M. and F.L. McEwen. 1958. Insect transmissions of X-disease virus. Phytopath. 48:262. Gilmer, R.M. J.D. Moore and G.R. Heitt. 1954. X—disease virus: Host range and pathogenesis in chokecherry. Phytopath. 44:180—185. Gilmer, R.M., D.H. Palmiter, G.A. Schaefers, and F.L. McEwen. 1966. Leafhopper transmission of X—disease virus of stone fruits in New York. N.Y. State Agric. Exp. Sta. (Geneva) Bull. 813. 22 pp. Click, P.A. 1960. Collecting insects by airplane, with special reference to the dispersal of the potato leafhopper. USDA Tech. Bull. 1222. 16 pp. Granett, A.L. and R.M. Gilmer. 1971. Mycoplasmas associated with X-disease in various Prunus species. Phytopath. 61:1036-1037. Hamilton, E.C.A. 1975. Pbrapblepsius jrroratus (Say). Pp. 30. In: Revision of the genera Paraphlepsjus Baker and Pendarus Ball (Rhynchota: Homoptera: Cicadellidae). Memoirs of the Entomol. Soc. of Canada, No. 96. 129 pp. Harker, J.E. 1961. Diurnal rhythms. Ann. Rev. Entomol. 6:131-146. 15 Hildebrand, R.M. 1953. Yellow-red or X—disease of peach. Cornell Univ. Agric. Exp. Stn. Memoir No. 323. 54 pp. Jensen, D.D. 1957. Transmission of peach yellow leaf roll virus by PieberieIIa fYorji (Sta1.) and a new vector, Osbornellus borealjs Del & M. J. Econ. Entomol. 50:668- 672. Jensen, D.D. 1969. Comparative transmission of western X— disease virus by Colladonus montanus, C. gaminatus, and a new leafhopper vector, Euscelidius variegatus. J. Econ. Entomol. 62:1147-1150. Johnson, 0.9. 1969. Migration and dispersal of insects by flight. Methuen & Co. Ltd., London. 763 pp. Jones, A.L., G.R. Hooper and D.A. Rosenberger. 1974. Association of mycoplasmalike bodies with Little Peach and X-disease. Phytopath. 64:755-756. Kennedy, J.S. 1961. A turning point in the study of insect migration. Nature 189:785—791. Kirkpatrick, E.C. 1986. Characterization of western X- disease mycoplasmalike organisms. Ph.D. Dissertation. University of California, Berkley, CA. 196 pp. Lacey, G.H., M.S. McClure and T.G. Andreadis. 1979. Reducing populations of vector leafhoppers is a new approach to X-disease control. Frontiers of Plant Science 32:2—4. Larsen K.J. and M.E. Rhalon. 1987. Crepuscular movement of Parapblepsjus irroratus (Say)(Homoptera: Cicadellidae), between groundcover and cherry trees. Env. Entomol. (accepted for publication). Lawson, F.R., J.C. Chamberlin and G.T. York. 1951. Dissemination of the beet leafhopper in California. USDA Tech. Bull. 1030. 59 pp. McClure, M.S. 1980a. Role of wild host plants in the feeding, oviposition, and dispersal of Scapbytqpius acutus (Romptera: Cicadellidae), a vector of peach X— disease. Env. Entomol. 9:283-292. McClure, M.S. 1980b. Spatial and seasonal distributions of leafhopper vectors of peach X-disease in Connecticut. Env. Entomol. 9:668-672. l6 Mowry, T.M. 1982. Leafhopper sampling in Michigan peach orchards and serological detection of a spiroplasma associated with X-disease in plant and insect tissue. M.S. Thesis. Michigan State University, East Lansing, MI. 134 pp. Mowry, T.M. and M.E. Rhalon. 1984. Comparison of leafhopper species complexes in the groundcover of sprayed and unsprayed peach orchards in Michigan. Great Lakes Entomologist 17:205-209. Nichiporick, R. 1965. The aerial migration of the six- spotted leafhopper and the spread of the virus disease aster yellows. Int. J. Biomet. 9:219-227. Nielson, M.R. 1979. Taxonomic relationships of leafhopper vectors of plant pathogens. Pp. 3-27. In: Leafhopper vectors and plant disease agents. eds. K. Maramorosch and K.F. Harris. Academic Press, N.Y. 654 pp. Osborn, H. 1932. Supplemental records and notes on Ohio leafhoppers. Ohio J. Sci. 32:513-517. Palmiter, D.H. and J.A. Adams. 1957. Seasonal occurrence of leafhopper vectors of X-disease virus in sprayed and unsprayed peach blocks. Phytopath. 47:531. Palmiter, D.H. and R.M. Hildebrand. 1943. The yellow-red virosis of peach: its identification and control. N.Y. State Agric. Exp. Sta. Bull. 704. 17 pp. Palmiter, D.H., R.J. Coxeter and J.A. Adams. 1960. Seasonal history and rearing of Scapbytopjus acutus (Say) (Homoptera: Cicadellidae). Ann. Ent. Soc. Am. 53:843- 846. Phillips, J.H.H. 1951. An annotated list of Hemiptera inhabiting sour cherry orchards in the Niagara Peninsula, Ontario. Canad. Entomol. 83:194-205. Pienkowski, R.L. and J.T. Medler. 1964. Synoptic weather conditions associated with long—range movement of the potato leafhopper, Empoasca fsbae, into Wisconsin. Ann. Entomol. Soc. Am. 57:588-591. Purcell, A.H. 1979. Transmission of X-disease agent by the leafhoppers Scapbytopjus nitrjdus and Acinqpterus angulatus. Plt. Dis. Rep. 63:549-552. l7 Purcell, A.H. 1985. The ecology of bacterial and mycoplasma plant diseases spread by leafhoppers and planthoppers. Pp. 351—380. In: The Leafhoppers and Planthoppers. eds. L.R. Nault and J.C. Rodriguez. John Wiley a Sons, N.Y. 500 pp. Purcell, A.H. and E.C. Suslow. 1982. Dispersal behavior of Colladonus montanus (Homoptera: Cicadellidae) in cherry orchards. Env. Entomol. 11:1178-1182. Basin, 8. 1978. The mycoplasmas. Microbiol. Rev. 42:414-470. Robinson, 6. 1985. 1985 Peach virus annual report. Michigan Dept. Agric. 4 pp. Rosenberger, D.A. 1977. Leafhopper vectors, epidemiology and control of peach x-disease. Ph.D. Dissertation. Michigan State University, East Lansing, MI. 95 pp. Rosenberger, D.A. and A.L. Jones. 1978. Leafhopper vectors of the peach X-disease pathogen and its seasonal transmission from chokecherry. Phytopath. 68:782-790. Severin, H.H.P. 1933. Field observations on the beet leafhopper Eutettix tenellus, in California. Hilgardia 7:281—360. Southwood, T.R.E. 1978. Ecological Methods. 2nd ed. Chapman and Hall, N.Y. 524pp. Sterns, L.A. and D. MacCreary. 1938. Leafhopper migration across Delaware Bay. J. Econ. Entomol. 31:226-229. Swanson, E.C. 1971. Environmental biology of the leafhopper Scaphytopius delongj. Ann. Entomol. Soc. Am. 64:809- 812. Taboada, 0., D.A. Rosenberger and A.L. Jones. 1975. Leafhopper fauna of X-diseased peach and cherry orchards in southwest Michigan. J. Econ. Entomol. 68:255-257. Taylor, R.A.J. 1978. The relationship between density and distance of dispersing insects. Ecol. Entomol. 3:63-70. Taylor, R.A.J. 1985. Migratory behavior in the Auchenorrhyncha. Pp. 259-288. In: The Leafhoppers and Planthoppers. eds. L.R. Nault and J.C. Rodriguez. John Wiley & Sons, N.Y. 500 pp. 18 Thornberry, H.H. 1954. Preliminary report on insect transmission of eastern X~disease virus in Illinois. Plt. Dis. Rep. 38:412-413. Rhalon. M.E. and D.A. Croft. 1984. Apple IPM implementation in North America. Ann. Rev. Entomol. 29:435-470. Whitney, S.P. and J.R. Meyer. 1987. Movement between wild and cultivated blueberry by two species of sharpnosed leafhoppers in North Carolina. J. Entomol. (submitted). Rolfe, H.H. 1955. Transmission of the western X—disease virus by the leafhopper, Colladonus.montsnus (Van D.). Plt. Dis. Rep. 39:298-299. Rolfe, H.H., E.R. Anthon and L.S. Jones. 1950. Transmission of western X—disease of peaches by the leafhopper Colladonus geminatus (Van D.). Phytopath. 40:971. CHAPTER I FIELD MONITORING OF X-DISEASE LEAFHOPPER VECTORS AND SYMPTOMATIC CHOEECHERRY 19 Introduction The X—disease research effort of the current stone fruit decline project (USDA grant no. 85-CRSR-2-2551) requires up-to—date field monitoring of the abundance of X- disease vector leafhoppers and chokecherry. These data are needed to aid in assessing year to year variation in X- disease and leafhopper incidence, evaluating established control procedures and in developing new X-disease management strategies. Past research (Taboada et a1. 1975, Rosenberger 1977, Rosenberger & Jones 1978) has demonstrated that at least nine species of leafhoppers (Homoptera: Cicadellidae) that occur in Michigan are vectors of X-disease. Parapblepsius irroratus (Say) is the most common known vector of X—disease in Michigan peach and cherry orchards (Taboada et a1. 1975, Rosenberger 1977). It is also the most efficient vector in greenhouse tests (Rosenberger & Jones 1978). Both P. jrroratus and Scapbytopjus acutus (Say) are bivoltine in Michigan, with the two periods of adult activity being late June to July and late September to October (Taboada et a1. 1975). Michigan Department of Agriculture (MDA) annual peach surveys indicate the incidence of X-disease has increased in peach orchards of southwest Michigan during the past several years. Chokecherry as an alternate host of X-disease (Gilmer et a1. 1954) is considered the major source of X— 20 21 disease inoculum outside the orchards. For this reason, MDA X-disease regulation No. 612 requires the removal of all chokecherry within 500 ft of peach and cherry orchards. X-disease is a major peach disease problem in southwestern lower Michigan, but has not been a severe problem north of Kent County. Many factors may be limiting the distribution of X-disease. Past monitoring of X-disease in Michigan (Taboada et a1. 1975, Rosenberger 1977, Mowry 1982) has not been done north of the Peach Ridge area on a regular basis. About 58% of Michigan’s peach acreage is located in Berrien and Van Buren Counties (Fedewa & Pscodna 1982), and these are the counties hardest hit by X-disease (Robinson 1985). The leafhopper monitoring reported here was a survey of the entire southern Michigan stone fruit belt. The objectives of this survey were to determine how the abundance and distribution of X-disease vector leafhoppers and symptomatic chokecherry differ temporally and spatially throughout the west coast of Michigan. Materials and Methods Field Season and Research Sites During the 1985 and 1986 field seasons, traps were placed in the field during the first week of May. Monitoring occurred weekly in 1985 and biweekly in 1986 and ended ca. November 15 after several hard frosts and the first snow. 22 Five sites were monitored in 1985 and six sites in 1986 (Table 1). All sites were located in Michigan’s lower penninsula (Fig. 1). Weather data such as temperature and the resulting degree day accumulations for each site were obtained from the M.S.U. Cooperative Crop Monitoring Service (CCMS) using agricultural weather observation stations located at or near each field site (Table 1). Symptomatic Chokecherry Survey The abundance of wild sources of X-disease inoculum in Michigan was surveyed by biweekly monitoring of chokecherry. In 1986, a 8 km route leaving each field site along two lane roadways was selected and all chokecherry clumps or individual bushes visually observed exhibiting X—disease symptoms were counted. The average number of symptomatic chokecherry/km was then calculated for each site. X—disease Vector Leafhopper Survey The abundance and distribution of known X-disease vector leathppers were monitored. In 1985, monitoring was performed weekly at the Lawrence, Hartford, Fennville, Clarksville, and East Lansing sites. In 1986, monitoring was performed biweekly at the Lawrence, Bainbridge Center, Fennville, Walkerville, Manistee, and Northport sites. Monitoring was performed with yellow sticky board traps and by sweep net sampling. Six yellow sticky board traps were hung at each site ca. 1.5 m above the orchard groundcover. The traps were 12.5 x 25 cm made of 0.25 in 23 .:0muoun accuse: can cums cassava Aaxv oocouswaa lIIIIIIIIIIIIIIIIII'ItI‘l-llll'l'l'sll'lv|IIIEIEIIIIIE'ElIIIIIIIIIIIIIll-l'l"l-l“"l'l"|IE" .om gem .me one m.n~ acme: accuse acoooo mmmmsaoxmo: mm .mm com .m one o.m sanamoog .ss gooom accesses scoasecoz em .em com .m~ ave m.v exam Leon accuse oneness: assumes: em .4 see .NH owe e.e 3am sou zooms sauna =o> oonoszaa mm.mm .os gem .v~ owe m.m Loassccsa: some; sues: nus - success: mm .m can .mm cue o.o omfia>asms suave =amo-< a m-a>anoa em.em .om gem .me one m.m atom one: am: stucco accuse manages Luau mm .mH one .Nm one o.o oama>axuoao :oaom omcoH ofimw>aguamo mm .s~ 0mm .s one o.m smam>cosaz scams sanctum sesame sweetness: mm .1122212......:o;c mouecmvuooo guesses“: IélllIII'IIIIIIIIIIIIIIIII.IlllllIIIIIIIIII'IIIIIIIII'II'I[Ill‘ll'Il'I'IIllIIIIII'I'I'iIIIIII .nuov :ofiumOOH same can .cOwunum sesame: .oass ensue sea: snowman enema mean can swam was com modem omaemae-x co sues .H «Haas 24 Northport . l Manistee O Walkerville :1: Clarksville $Fennville *East Lansing *Hartford ®Lawrence Bainbridge Center * 1985 01986 L--_--_--_ Fig. l. The field sites monitored for leafhopper vectors of X-disease during 1985 and 1986, and monitored for the appearance of symptomatic chokecherry during 1986. 25 plywood and painted with sun yellow enamel (Benjamin Moore & Co., Montvale, NJ) and coated with Tree TanglefootT" (The Tanglefoot Company; Grand Rapids, MI). These traps were replaced each visit to the site and returned to the lab for examination and removal of captured leafhoppers. Sweep net samples were taken from different areas in and around each orchard site. Four sweep samples were taken, each consisting of 25 sweeps with a 37.5 cm dia net. Each sweep was ca. 8 1.5 m pass through the groundcover foliage. The sweep samples were deposited in plastic bags, placed in a cooler for transport back to the laboratory, and then frozen at ~20° C in the lab to kill all insects. Sorting, leafhopper identification to species, and counts of abundance and sex took place in the laboratory. Results Field Season During 1985, temperature effects as measured by degree day accumulations (Baskerville & Emin 1969) was similiar at all sites (Fig. 2). The 1986 total accumulations are similiar to the 1985 total accumulations for both the Lawrence and Fennville sites. Generally higher temperatures were experienced in both mid—July and early-October of 1986. The difference in total degree day accumulation between the Northport (1820 DD) and Lawrence (2585 DD) sites was dramatic, where an average accumulated difference of 765 DD was realized. Average accumulated degree days showed a 478 26 Hartford Lawrence / Fennville " Clarksville East Lansing 11m A CD U0 a) 8 .o 1985 V (D O 'I'flfit'l‘fii'lI't'llj'VII'Y'IIT—rrlufy' 5‘ 4 o—o Bainbridge Center 0 2400— H Lawrence CD - +—+ Fennville 8 2100-1 e—e Walkerville / 8 « H Manistee / , . // 'afl“ _ A a D 18004 H Northport / / .. .. 1200-1 // 900- // soo— ./ 300‘ /'/ 1986 O 7 I I T T r I I I I I fiT l I I Tfi [’jiTfi I fl Tfi If , j . 1 . . . . T 125 150 175 200 225 250 275 300 325 May Jun Jul Aug Sep Oct Nov JUIIOI’l Date Fig. 2. Accumulation of degree day (base 50) heat units (Baskerville & Emin 1969) over time at the sites during both 1985 and 1986 field seasons. 27 DD difference between the average of northwestern (1980 DD) and southwestern (2458 DD) weather stations (Fig. 3). Symptomatic Chokecherry Survey During 1986, chokecherry exhibiting symptoms of X- disease were first observed in southwestern lower Michigan in late-June and in northwestern lower Michigan in mid-July (Fig. 4). By early September, up to six symptomatic chokecherry/km were visually evident. This delay in symptom expression between southwest and northwest is similiar to the mean degree day accumulation for those areas. Vector Leafhopper Survey Leafhopper populations were about five times greater in 1985 than in 1986 (Fig. 5). Although the generations peaked at different dates in 1985 and 1986, the peaks did occur at approximately the same number of accumulated degree days (Fig. 6). Differences in X-disease vector leafhopper density occurred both between field sites (P50.05, LSD test of data)(Fig. 7) and between 1985 and 1986 field seasons (P)0.05, ANOVA). Representatives of all leafhopper species known to vector X-disease in Michigan were found during both the 1985 and 1986 field seasons. Only four of these, P. irroratus, S. acutus, C. clitellarius, and N1 seminuda were present in numbers greater than 1% of all the known vector leafhoppers captured (Table 2). The relative abundance of these 28 2700 2400- 8 1 Southwest Lower Michigan 0 2100- 3’: 1eoo~ O 1 .0 V 1500- m .. g 1200- Northwest Lower Michigan g 900- L .. 8’ 600- C) . 300- 1 986 1 O 5".'5'IIIIUEUIUf'rIUINI]Ij'Ul'IIr'['5' 125 150 175 200 225 250 275 300 325 May Jun Jul Aug Sep Oct Nov Julian Date Fig. 3. Mean degree day (base 50) accumulation for the southwest (Bainbridge Center, Lawrence & Fennville sites) and northwest (Walkerville, Manistee & Northport sites) areas of lower Michigan in 1986. 29 51 Southwest Lower Michigan Northwest Lower Michigan Mean Number Chokecherry/km .p I . 1985 O 1 Y 1 1 :1 1 7 1 1 r 1 1 1 1 I 1 1 1 1 I 1 1 1 1 I 1 1 1 1 1 1 1 1 r1 1 1 1 1 i 25 150 175 200 225 250 275 300 325 May Jun Jul Aug Sep Oct Nov Julian Date . Fig. 4. Mean number of symptomatic chokecherry visually observed/km of two lane roadway in the southern and northern regions of the western Michigan stone fruit belt during 1986. 30 1501 13 . x—x 1985 8 1 H 1985 3 125- 4.; . CL . 8 -l 100« (I) a. Q) 3- l o 751 .C .. H. cl 0 a 33 50- :n= 1 l 8 25- 0 I 5 l 0" F1l1111111111111fl1111'1111'11 125 150 175 200 225 250 275 300 325 May Jun Jul Aug Sep Oct Nov Julian Date Fig. 5. Mean number of X-disease vector leafhoppers captured by yellow board traps and in sweep nets over time in 1985 and 1986 at all sites. 31 150— : x—x 1985 I o—o 1986 1251 100$ Mean # Leafhoppers Captured 01 \l O U" I. l ' I i I l I C IV. I '1' I v 5 5 fi' 5 U 0 300 600 900 1200 1500 who 21'00 {4100 27'00 Degree Days (base 50) Fig. 6. Mean number of X-disease vector leafhoppers captured by yellow sticky board traps and in sweep nets, based on average degree day accumulations in 1985 and 1986 at all sites. 32 200 l [3'—El Hartford 175; 0—0 Lawrence ; 4—+ Fennville Z 9—0 Clarksville ° 150—- H East Lansing 100—: 75—: '0 E > 93 50‘. 1 :5 .1 “a I o 25‘. £3 : e 0-' .1 . <1) ‘ B—El 8: : H Lawrence E - +—+ Fennville ~— 40: e—e Walkerville 8 - H Manistee —' ~ x—x Northport =§= ' 30- 20- 10‘: 1986 A ,, J ~- 0 300 600 900 1200 1500 1800 2100 2400 2700 Degree Days (base 50) Fig. 7. Total number of X-disease vector leafhoppers captured by yellow sticky board traps and in sweep nets at each site during the 1985 and 1986 field seasons. Note: The y-axis is expanded in 1986 to accentuate lower leafhopper counts. 33 Table 2. Total number of X-disease vector leafhoppers captured by yellow sticky board traps and sweep nets, and percent relative abundance of each found in Michigan for both 1985 and 1986 field seasons. 1985 1986 Species Total X of total Total X of total P. irroratus 1790 72.47 278 60.57 S. acutus 529 21.42 144 31.37 C. clitellarius 20 0.81 17 3.70 N; seminuda 92 3.72 5 1.09 Scapboideus app. 23 0.93 4 0.87 F. florii 2 0.08 3 0.65 0. isbidae 1 0.04 4 0.87 G. lamina 13 0.53 4 0.87 Totals 2470 459 34 leafhoppers in the field during 1985 and 1986 was P. irroratus: 73.1%, S. acutus: 22.0%, C. clitellarius: 1.5%, and N; seminuda: 3.4%. Some sites supported larger populations of these vectors than others (Fig. 8, Table 3). P. irroratus was very common in the East Lansing, Lawrence, Hartford, and Fennville sites. S. acutus was found easily at the Hartford site and in good numbers in Lawrence and Fennville. C. clitellarius was found most commonly at the Manistee site, while N. seminuda was found easily in East Lansing and often in Fennville, but was not found at or north of Walkerville. Yellow sticky board traps captured 90.3% of all known X-disease vector leafhoppers captured during 1985 and 1986. There was no significant difference in this monitoring method capture rate between the two generations (P}0.05, ANOVA). The sex ratio of P. irroratus leafhoppers did not significantly differ between the yellow board trap and sweep net monitoring methods (Fig. 9), with male leafhoppers accounting for 65% of the captures on yellow sticky board traps, and 42% of the captures in sweep nets. There was no significant difference in this captured leafhopper sex ratio Discussion The similarity of the degree day accumulations during 1985 was due to the concentration of all 1985 field sites in 35 800- ‘ 1 9 85 700- - V " l 500- g l U m L D H 8 0 U I l I e . f 8 f f a. d 4? o \I .C u— U G) "J 1 gar §t= 100... g - coed . ozozoze 0990 75 ‘ om: ' :.:....' r """ ;' ll ' " ‘0 ‘ ¢ ‘i 'l' l .. / I. ¢ i l1] 1 'l / V 50: Z 4. g . so»: 4 . ’o’o’o‘ we“: '4 ’o’o’o’ 'o’o’o’ 1 'o’o°o°~ ’o’o°o’ 2 5 - ’o’e’o’ ’o’e’o’ . J :.:,:,: :,:,:,: E: N. semunuda . 533:0: 63:93 % mm C. clitellorius - 1:33:32 33;: ¢ 2 s. acutus O ' {02020: 510,3:- ‘Y/ m P. irroratus ' 1 1 o o. a :9 P 39.: 6" a” 5 3 8 \. g Q ' '. ¢ -°.c . . - s s r 6% .3 «P s a 13 Fig. 8. Total number of common X-disease vector leafhoppers captured by yellow sticky board traps and in sweep nets at each site during the 1985 and 1986 field seasons. Note: The y-axis is expanded in 1986 to accentuate lower leafhopper counts. 36 Table 3. Number of X-disease vector leafhoppers captured at each field site during 1985 and/or 1986. Species Site P. irroratus S. acutus c. clitellarius N2 seminuda Bainbridge Centerc 40 21 2 Clarksvilleb 101 45 3 3 East Lansingb 641 19 4 45 Fennville' 217.5 94.5 3.5 13.5 Hartford° 276 145 5 3 Lawrence. 244 96 1 5 8.5 Manisteec 39 17 12 Northportc 2 23 Walkervillec 46 22 2 Totals 1606.5 482.5 32.0 75.0 Means 178.5 53.6 3.6 8.3 % of total 73.1 22.0 1.5 3.4 ‘average of 1985 and 1986 data. b1985 data. c1986 data. 37 60— EZ] Females ‘QQQIMMS ‘0 50- 8 .3 _ 8-40— (J 8 d / <1) 30— CL r//// 8- ~ é E 20— / C3 3 1 Z 88 10— éjj / ‘ /7 O / // Sweep Net YeHow Board Fig. 9. Percent of male and female P. irroratus leafhoppers captured by monitoring method (sweep net or yellow sticky board trap) for both 1985 and 1986 field seasons. 38 the southwestern and central lower penninsula. The 1986 sites had greater latitude differences from south to north and a corresponding decrease in degree day accumulation with distance north. The two week lag in degree day accumulation probably explains the delay in chokecherry development and X-disease symptom expression. The presence of chokecherry along roadways indicates that many bushes are not being eradicated per MDA regulations and therefore may once again be serving as a major alternate host of X-disease pathogen. Of all the known species of X-disease vector leafhoppers found present in 1985 and 1986, only four seem to be common enough to warrant our attention unless one of the rare species is found to have a very high MLO infection rate or its feeding behavior predisposes it to transmit more frequently. P. irroratus is still the most common vector leafhopper in Michigan, representing 73% of the total number caught, with S. acutus second most common at 22%. This confirms the earlier work by Taboada et a1. (1975) and Rosenberger (1977) that P. irroratus is the most numerous X- disease vector in Michigan. The graphical evidence (Figs. 5, 6 & 7) that X-disease vectors are bivoltine is largely influenced by the two generations of P. irroratus, which constitutes the largest portion of the vector population. Further work on the number of generations of other vector species found in Michigan would help to clarify this observation. 39 Distributions of leafhopper populations was influenced by sample location in the state. P. irroratus was commonly found in the southwest and central sites. Since the second generation of P. irroratus occurs at degree day accumulations greater than 2200 DD (Fig. 7), areas that do not reach this degree day accumulation probably do not have a second generation. This is most likely the reason why P. irroratus is rare in Leelanau County, where less than 1900 DD (base 50) were accumulated in 1986, and only in exceptional years are more than 2000 DD accumulated (CCMS data). C. clitellarius was found in significant numbers only at the Manistee site, and thus may be an important vector in that area. Since the most common vector leafhopper found in Leelanau County was S. acutus, but at a low density when compared with other sites, the chance of X-disease transmission by leathppers there seems low. Sampling methods used in future X—disease monitoring efforts should reflect the effectiveness of the yellow sticky board traps, with over 90% of all vector leafhoppers captured by this method. Although sweep net sampling is the best method of detecting leafhoppers moving into and out of orchards in a short period of time (Mowry & Whalon 1984), sweep net sampling alone is of minor importance and an inefficient, labor-intensive, and incomplete sampling method for long-term Xedisease vector monitoring. 4o Presence in the northwest area of symptomatic chokecherry indicates that a wild source of X-disease inoculum is present, and that transmission among chokecherry does occur. However, the limited distribution of populations of vector leafhoppers in this region may be preventing the vector transmission of X-disease to peach and cherry in the northwest part of the state. References Cited Baskerville, G.L. and P. Emin. 1969. Rapid estimation of heat accumulation from maximum and minimum temperatures. Ecology 50:514-517. Fedewa, D.J. and S.J. Pscodna. 1982. Michigan orchard and vineyard survey 1982. Michigan Dept. Agric. 49 pp. Gilmer, R.M. 1954. Insect transmission of X-disease virus in New York. Plant Dis. Rep. 38:628—629. Gilmer, R.M., J.D. Moore, and G.R. Keitt. 1954. X-disease virus: 1. Host range and pathogenesis in chokecherry. Phytopath. 44:180-185. Gilmer, R.M., D.H. Palmiter, G.A. Schaefers, and F.L. McEwen. 1966. Leafhopper transmission of X-disease virus of stone fruits in New York. N.Y. State Agric. Exp. Sta. (Geneva) Bull. 813. 22 pp. Hildebrand, R.M. 1953. Yellow-red or X-disease of peach. Cornell Univ. Agric. Exp. Sta. Memoir No. 323. 54 pp. Jensen, D.D. 1957. Transmission of peach yellow leaf roll virus by FTeberiella florii (Sta1.) and a new vector, Osbornellus borealis DeL. & M. J. Econ. Entomol. 50:668-672. McClure, M.S. 1980. Spatial and seasonal distributions of leafhopper vectors of peach X-disease in Connecticut. Env. Entomol. 9:668-672. Mowry, T.M. 1982. Leafhopper sampling in Michigan peach orchards and serological detection of a spiroplasma associated with X-disease in plant and insect tissue. M.S. Thesis. Michigan State University, East Lansing, MI. 134 pp. Mowry, T.M. and M.E. Rhalon. 1984. Comparison of leafhopper species complexes in the groundcover of sprayed and unsprayed peach orchards in Michigan. Great Lakes Entomologist 17:205-209. Robinson, G. 1985. 1985 Peach virus annual report. Mich. Dept. Agric. Memorandum of October 4, 1985. 4 pp. 41 42 Rosenberger, D.A. 1977. Leafhopper vectors, epidemiology and control of peach X-disease. Ph.D. Dissertation. Michigan State University, East Lansing, MI. 95 pp. Rosenberger, D.A. and A.L. Jones. 1978. Leafhopper vectors of the peach X—disease pathogen and its seasonal transmission from chokecherry. Phytopath. 68:782-790. Taboada, O., D.A. Rosenberger, and A.L. Jones. 1975. Leafhopper fauna of x-diseased peach and cherry orchards in southwest Michigan. J. Econ. Entomol. 68:255-257. CHAPTER II CREPUSCULAR MOVEMENT OF PARAPHIEPSIUS IRRORATUS (SAY) BETWEEN THE GROUNDCOVER AND CHERRY TREES 43 Introduction Little is known about the daily movement of P. irroratus and its distribution within orchards. Rosenberger and Jones (1978) suspected that adult P. irroratus were most active in the early evening when, and previous studies have shown increased activity in many insects during twilight (Barker 1961). Based on field observations, we hypothesized that P. irroratus moves daily from orchard groundcover to fruit tree foliage and back again. The purpose of this study was to determine how populations of P. irroratus fluctuate temporally and spatially between the different orchard sub-habitats of groundcover and the foliage of sweet cherry, sour cherry and apple trees. Materials and Methods Light trapping and sweep sampling were performed on six separate occasions in a 1 ha unsprayed orchard located on the Michigan State University campus in East Lansing, Michigan. The orchard was composed of blocks of apple, sour cherry, and sweet cherry trees ca. 7 yr old, with a mixed groundcover of rye fescue, red clover, and broadleaf weeds. The experiment was replicated three times during each of the two population peaks of P. irroratus as determined by weekly monitoring with yellow sticky board traps. The traps were 12.5 x 25 cm made of 0.25 in plywood and painted with sun 44 45 yellow enamel (Benjamin Moore & Co.; Montvale, NJ) and coated with tree tanglefoot (The Tanglefoot Company; Grand Rapids, MI). Light trapping samples were taken at 0.5 hr intervals beginning 1.5 hr before sunset until 1.5 hr after sunrise and blocked by three separate dates during each generation. Sample locations were randomly selected from a grid layed out on the orchard. The light trap was a 30.5 x 66 x 91 on wooden box painted flat black with the exception of the interior back which was a glossy white and illuminated by two yellow 60 watt "Bug Lites" (General Electric; Cleveland, Ohio) mounted on the white surface. The trap was situated ca. 1.8 m above the orchard floor and located between tree rows. The lights were turned on or off at 15 min intervals throughout the sampling period. All attracted P. irroratus were immediately aspirated by hand, counted, and retained to eliminate recaptures. Monitoring for leafhoppers in the sub-habitats was accomplished by sweep sampling, organized as a split plot design. The sub-habitat was the whole plot factor and time was the split plot within each sub-habitat. Randomization was by sample location selection within each sub-habitat, and the experiment was blocked by the three sampling dates in each generation. The four orchard sub-habitats were groundcover, sour cherry foliage, sweet cherry foliage, and apple tree foliage. Sweep samples consisted of 25 sweeps with a 37.5 cm dia net. Each sweep was a 1.5 I pass through 46 the groundcover or fruit tree foliage. Each sub-habitat was sampled at 0.5 hr intervals over the entire 24 hr sampling period and numbers of P. irroratus were counted. The data were subjected to analysis of variance and multiple comparison of the means by Scheffe’s test of data (SAS Institute 1985). All data were analyzed by individual generations before any combining of generations occurred. Periodicity of the data was analyzed by non-parametric circular-linear rank correlation (Batschelet 1981) which gives a correlation coefficient Dn, and its test-statistic Un. Results Periodic sampling of P. irroratus adults with light traps indicated that the daily flight period is crepuscular. P. irroratus fly for ca. 2 hr after sunset with peak flight time ca. 45 min after sunset (Fig. 1). Sunset was at ca. 21:15 first generation and ca. 19:45 second generation. The mean time of flight first generation was 22:05 and 20:34 second generation. During the peak 0.5 hr flight period, well over 100 P. irroratus adults were captured per light trap. Low numbers of leafhoppers were captured during other times of the night. No leafhoppers were captured during morning twilight. The number of leafhoppers captured 0.5 and 1 hr after sunset were statistically different (Pg0.05, ANOVA) from numbers captured during all other trapping times both first generation (Dn=0.938, n=24, Un=18.76, P<0.01, rank correlation) and second generation (Dn=0.973, n=24, 47 Flight Periods 300 .0 8 .... 1st Generation 3 ‘ X—K 2nd Generation ‘ 4.) o. 4 a J o Q) y d ‘3 a) 200- _ 8t 0 Es g d d “a ., a) '1 .J q :' 1 W- 100- 1‘ _ o : | 0' ‘ . : l . Z . lst generation sunset at 2115 , 1‘ 1 - ' l S . 2nd generation sunset at 1945 lb ‘1’ h K 2 o 0 300 600 900 1200 1500 1800 2100 2400 Time (24 hours) Fig. 1. Mean number of P. irroratus leafhoppers sampled by light trapping over a 24 hr period for both generations. Means followed by the same letter are not significantly different (Pg0.05; Scheffe’s test of data). 48 Un=19.46, P<0.0l, rank correlation). The mean number of P. irroratus leafhoppers captured each 0.5 hr sampling period for 24 hrs by light trapping was 4.2 and 6.2 for the first and second generations respectively. These leans were not significantly different (P?0.05, ANOVA). There were no significant differences between sample dates (P30.05, ANOVA). Sweep net samples of the groundcover indicated leafhopper density was highest during the day and began rapidly declining until 2 hr after sunset. During the night, the highest leafhopper densities were detected on sour and sweet cherry. At sunrise the groundcover leafhopper density began to increase, returning to its highest density by 15:00 (Fig. 2). Very few P. jrroratus were detected on apple at any time. There was no significant difference in number of leafhoppers between the sample dates within a generation (PZ0.05, ANOVA). Sweep sampling indicated that the leafhopper densities between the four sub-habitats were significantly different (P30.05, ANOVA). The density was highest in sour cherry first generation and in sweet cherry second generation (Fig. 3). The mean density in the groundcover was the same both generations, lower than that of either sweet or sour cherries but greater than the apple (Table 1). Circular- linear rank correlation indicated the number of leafhoppers captured in the groundcover (P<0.05), sour cherry (P<0.01) and sweet cherry (P<0.01) correlated significantly with the 49 30 L 1 st Generation 25- y‘x A, sunrise suuslex \ \ I I I ff] rIil I I I I I I I I l I I l ' 2nd Generation 25- « SUN1RISE SUNSET 20~ 1 .50. Mean Number of Leathppers Captured h“ [’4' ." M 10 ' ‘a 5.... 0 I I l I I I I T I I I I I T I I I ' I r O 300 600 900 1200 1500 1800 2100 240 Time (24- hours) 1 H Sour Cherry o-o Sweet Cherry Heroundaovsr Fig. 2. Mean number of P. irroratus leafhoppers collected in sweep nets every 0.5 hr over a 24 hr period in each sub- habitat type for both generations. 50 15 30391 0...... VV .9 V 3’3 F-—4 e a ‘%& ‘V 0 ° 0 O o «3 A 0;. ‘5 9% IIIApme :32: Sweet Cherry E Sour Cherry Graundcaver , T 1- / / 02 ___ 4 1 st Generation 2nd Generation 0 AGL 1% 0 fi’ '0 .9 O O o <£ O ‘3 Mean No. of Leafhoppers Captured o Fig. 3. Mean number (:SE) of P. jrroratus leafhoppers collected over 24 hrs in sweep nets in the four sub-habitats during first and second generations. 51 Table 1.. Mean number of P. jrroratus leafhoppers captured by 25 sweeps each 0.5 hr over 24 hrs, three replications in two generations lst 2nd Combined Generation Generation Generations $372123};—"m—"m’QfIé; ----- EELS-"”2538."- Sweet cherry 2.76b 7.56c 5.16e Groundcover 2.84ab 2.68d 2.76ef Apple 0 07b 0 03d 0 05f Means followed by the same letter are not significantly different (P<0.05; Scheffe’s test of data). 52 time of day sampled for both generations, but was not significantly correlated with time of day sampled in the apple (Table 2). Discussion Earlier work by Mowry and Whalon (1984) on absolute and relative sampling methods, indicated that relative sampling methods were the best way to detect leafhoppers moving into and out of orchards within a short period of time. Although sweep samples have distinct drawbacks (DeLong 1932), they were an effective means of getting frequent relative density estimates of a highly mobile leafhopper population. After the leafhoppers had dropped off the trees down to the ground during the early morning hours, they were unable to be collected efficiently by sweeping. Sweep sampling appears to be increasingly effective until ca. 15:00, when the leafhopper density in the groundcover is greatest. The current study was limited to a single site for several reasons, including a known high population of P. jrroratus and our ability to control pesticide spray applications. A systematic bias in the spacial position caused by the three blocks of fruit trees was unavoidable and created factors unable to be addressed by our experimental design. Sweep samples taken from groundcover indicated the groundcover leafhopper density was uniform throughout the orchard. 53 Table 2. Circular-linear rank correlation (Batschelet 1981) of the number of leafhoppers captured over a 24 hr period in 4 sub-habitats. Significance indicates the number of leafhoppers captured correlated with the time of day sampled. Sour Cherry lst generation 48 0.983 10.66 (0.01 2nd generation 48 0.970 10.53 (0.01 Sweet Cherry lst generation 48 0.985 10.66 (0.01 2nd generation 48 0.963 10.45 (0.01 Groundcover lst generation 48 0.885 7.86 (0.05 2nd generation 48 0.872 7.75 (0.05 Apple lst generation 48 6.4x10‘3 4.2x10'2 n.s. 2nd generation 48 6.4x10‘3 4.3)(10‘2 n.s. Dn=correlation coefficient, Un=test statistic. 54 At this site, P. jrroratus had an evening crepuscular activity period both generations during which they were responsive and attracted to the light traps. Movement out of the trees back into the groundcover the following morning occurred when leafhoppers were visually observed dropping out or gliding down from the orchard trees. Leafhoppers were generally inactive until the sun came up and temperatures rose. Leafhoppers were then active during the day within the orchard groundcover. Observations at other sites during the 1986 field season support these conclusions. Differences in host suitability may be why apple trees were not occupied at any time, and why first generation P. jrroratus seem to prefer sour cherry during the night while second generation leafhoppers seem to prefer sweet cherry. Circular—linear correlation coefficients support the conclusion that the leafhopper density in the groundcover, sour cherry and sweet cherry sub-habitats is correlated with the time of day. The lower mean leafhopper density in the groundcover is due to possible movement by P. jrroratus into and out of the orchard from outside habitats, and the large volume of groundcover foliage Compared with the volume of the young fruit tree foliage present in this orchard. Groundcover was the major orchard sub-habitat occupied by P. irroratus during the day. This observation supports 55 McClure’s (1982) assertion that groundcover type may have an important influence on the distribution and abundance of leafhoppers within the orchard. Groundcover manipulation and other management techniques may indirectly influence the transmission of X- disease by the night feeding leafhoppers. Knowing that P. irroratus visits cherry trees will aid in evaluating established control procedures and developing new X-disease management strategies. Recommendations to spray orchards after sunset with a quick-knockdown insecticide to reduce leafhopper populations have already been made (Whalon & Larsen 1985). Crepuscular movement by these leafhoppers helps them avoid being hit directly by daytime spray applications and could therefore be providing a mechanism of resistance to insecticides. References Cited Batschelet, E. 1981. Circular statistics in Biology. Academic Press, N.Y. 371 pp. DeLong, D.M. 1932. Some problems encountered in the estimation of insect populations by the sweeping method. Ann. Entomol. Soc. Am. 25:13-17. Barker, J.R. 1961. Diurnal rhythms. Ann. Rev. Entomol. 6:131-146. McClure, M.S. 1982. Factors affecting colonization of an orchard by leafhopper vectors of peach X-disease. Env. Entomol. 11:695-700. Mowry, T.M. and M.E. Whalon. 1984. Comparison of leafhopper species complexes in the groundcover of sprayed and unsprayed peach orchards in Michigan. Great Lakes Entomol. 17:205-209. Rosenberger, D.A. and A.L. Jones. 1978. Leafhopper vectors of the peach X-disease pathogen and its seasonal transmission from chokecherry. Phytopath. 68:782-790. SAS Institute. 1985. SAS User’s Guide: Statistics, Version 5 Edition. SAS Institute Inc., Cary NC. 956 pp. Whalon, M.E. and K.J. Larsen. 1985. X-disease leafhopper vector alert. Pest Alerts 24:85. 56 CHAPTER III DISPERSAL 0F PARMPHIEPSIUS IRRORATUS (SAY) IN PEACH AND CHERRY ORCHARDS 57 Introduction Understanding vector leafhopper movement is necessary to learn more about X-disease epidemiology. Until now, dispersal of P. jrroratus, a major vector of peach X-disease in Michigan, has not been examined. P. irroratus is known to spend the day within the orchard groundcover, and at twilight it moves into fruit tree foliage (Larsen & Whalon 1987). The purpose of this research was to estimate population density and evaluate the temporal and spatial distribution and dispersal patterns of local populations of P. irroratus around peach and cherry orchards. Mark, release and recapture techniques can be used to estimate population densities (Southwood 1978), and have also been used to study dispersal behavior of leafhoppers (Ito & Miyashita 1961, Purcell & Suslow 1982, Whitney & Meyer submitted). Using fluorescent dye dusts to mark the leafhoppers, we evaluated their dispersal rate, distance and direction of movement \ within, into, and out of the orchards. Materials and Methods Leafhopper Capture and.Mbrking Leafhoppers to be marked and released were captured during the evenings of the two generation peaks of P. jrroratus. First generation adult activity peaks between late-June and July, while second generation activity occurs 58 59 from late-September to October. Leafhoppers were captured from natural populations near East Lansing and Lawrence, MI by aspirating leathppers from box light traps. Each light trap was a 30.5 x 66 x 91 cm wooden box painted flat black with the exception of the interior back which was glossy white and illuminated by two yellow 60 watt "Bug Lites" (General Electric; Cleveland, Ohio) mounted on the white surface. The traps were situated ca. 1.8 m above the ground and powered by a 12 volt automotive battery connected to a 12v DC to 120v AC inverter. The light traps were set up at capture sites located at least 1 km away from the release sites. Leafhopper capture began at 0.5 hrs before sunset and continued until 2 hrs after sunset to sample during the peak flight period of P. jrroratus which occurred ca. 0.75 hr after sunset (Larsen & Whalon 1987). Leafhoppers were aspirated into holding vials in groups of 100 for marking. The leafhoppers were marked using six Day-GloTH fluorescent powder dyes (Day-Glo Color Corporation, Cleveland, Ohio). Different colors were used to indicate a release date and location. Rocket Red, Signal Green, Arc Yellow, Horizon Blue, Creme, and Corona Magenta were the colors used. The groups of 100 aspirated leafhoppers were placed in a dry 1 qt mason jar with 1/8 teaspoon of dye. The jar was then gently agitated for one minute. The undusted controls used in some of the tests were handled in the same manner, except that the control insects were not dusted. The marked 60 leafhoppers were immediately removed from the jar by dumping the contents onto an open petri dish cover placed on the ground at the release point or in the middle of the laboratory flight cage. Only those leafhoppers that flew away were counted as released. .Effects of4Marking on Survival To test the effect of the marking method on leafhopper survival, seven sets of 3 male and 3 female leafhoppers were treated with one of the six colors of flourescent dye or were an undusted control. The experimental design was randomized block, replicated in six cages each generation. Test leafhoppers were released into small 30.5 cm tall x 20.25 cm dia cages on clover and barley host plants housed in a walk—in growth chamber. The experiment was conducted in a light-dark regime of 15:9 LD for the first generation or 12:12 LD for the second generation. Daily counts of leafhopper survival were made for 21 days by searching the cages and removing dead leafhoppers, then identifying and recording leafhopper treatment and sex in the laboratory. Effects of.Marking on Flight Activity To test the effect of the'marking method on flight activity, seven sets of 25 male and 25 female leafhoppers were marked with a flourescent dye color or were an undusted control. These leafhoppers were released together on clover and barley host plants in a 0.6 x 1.0 x 2.0 m cage in the greenhouse under a light—dark regime of 15:9 LD for the 61 first generation and 12:12 LD for the second generation. Four yellow sticky board traps were hung in the cage and trapped leafhoppers were monitored for 21 days following release. Field Release and Pecapture The leafhopper field release and recapture study was performed at research sites in East Lansing and Lawrence, MI (Fig. 1). The first research site was a 1 ha unsprayed mixed orchard of apple, sour cherry and sweet cherry blocks ca. 7 yrs old located on the Entomology research farm at Collins Road on the Michigan State University campus in East Lansing, Michigan (location: 42° 41’ N. Lat. 84° 30’ W. Long.) (Fig. 2). The second research site was a commercial orchard of peach, tart cherry and apple near Lawrence, Michigan in VanBuren County (location: 42° 12’ N. Lat. 86° 4’ W. Lang.) (Fig. 3). Each orchard research area contained two release sites, one within a peach or tart cherry block and the other 10 m outside the edge of the same orchard block. At the Lawrence site, the edge release location and recapture traps were set up in what we considered a large field, recently planted with widely spaced young apple trees. At each release site, yellow sticky board traps were hung ca. 1.5 m above the groundcover at distances of 5, 10, 20, 40, and 60 m in six directions, radiating from the release point (fig. 4). The yellow sticky board traps were 12.5 x 25 cm made of 0.25 in plywood and painted with sun 62 >1: East Lansing * Lawrence --—---—---—-- —---—-1_-- __-.. —-—-- Fig. 1. Location of the two field research sites used in this leafhopper dispersal study. Beans 63 Corn oc>oo oaoo<>000 00 o . ODOQWSQSOW cnocadbooooobooooooooo <3om>oooooooooo:8<::::::)E53 00 00 00 OO 00 00 00 OO 00 00 Sweet Corn 5' . 0 e N181“ [n mum Tm Corn EJ ......... 3...... «e * Release Location 0 Recapture Traps Fig. 2. Release locations, surrounding area at the East Lansing, MI research site (location: Woods recapture trap layout and T4N RZW Sec 36 W 1/2). 64 00 000000 ooooooooooooooooooooooooooooo ooooooooaooooooooooqoooooooooo ooooooooooooooooooooooooooooooo oooooooooooooooooooooooooooooooo ooooooooooooooooooooooooooooooooo ooooooooooooooooooooooooooooooooo Woods oooooooooooooooooooqoooooopoooooo oooooooooooooéooooooooooooooooooo o o oooooooooooooooooopqooooooooooooo o o o o oooooooooooooooooogdldoooooooooooo o o o b ooooooooooooooooooodooooooooooooo o O O o oooooooooooooooooooooooooopoooooo o o o o ooooooooooooooooooooooooooooooooo 0 o 0 .0 ooodooooodooooooooooooooooooobooo ' oooooooooooooooaoooooooooooooooo : :‘o'q ' ooooooooooooooooooooooooooooooo 0 O 0 0,0 0.0 C200000000§0é§00000000000000m0 .0 O O O 0 000000000 000000000000000 0 o 0 0 o 00(b000000000000000000000 + 0000 0 0 O 0 ° ° 0' 0000000000 0 o o o o- oooooooooooooooo 0 0 ° ° ° 0 o o‘o' oooooooooooooooooooooooo g ooooooooooooooooooooooo 0 ° ° oooooooo oooooooooooooo ° ° 'oooooooooooooooooooooooo o 59 100 fi- ; =1 1 = sisters 96 Release Location 0 Recapture Traps Fig. 3. Release locations, recapture trap layout and surrounding area at the Lawrence, MI research site (location: T38 R15W Sec 20 SE 1/4). 65 0C) '9” ’o z e 300 60 e 240 120 * Release Location 0 Recapture Traps e 180 Fig. 4. Arrangement of the yellow sticky board traps used to recapture marked P. irroratus leafhoppers around each release location. 66 yellow enamel (Benjamin Moore & Co.; Montvale, NJ) and coated with Tree TanglefootT" (The Tanglefoot Company; Grand Rapids, MI). Leafhoppers captured on yellow sticky boards were monitored daily for 21 days following a release. Leafhoppers were removed from the yellow sticky boards carefully with a small spatula and placed in a coded petri dish. A black light was used to determine marked and unmarked leafhoppers. Weather data were obtained from nearby agricultural weather observation stations at Paw Paw and the M.S.U. Horticultural farm in East Lansing. The mark/recapture field experimental design was a randomized block, 4 factor factorial combined over 2 sites. Three dates of release (the blocks or replicates) occurred during each generation. The four factors were release location, direction, distance, and time following release. Randomization occurred by random assignment of the marked leafhoppers to release site and location. Field data were entered into R:Base System V (Microrim 1986) for data management and analysis. Data were subjected to further analysis of variance and multiple comparison of means using SAS (SAS Institute 1985). Directionality of the data was analyzed by circular correlation and circular-linear rank correlation tests (Batschelet 1981). Dispersal equations describing the distance data were fitted and analyzed using GLIM (Baker & Nelder 1978). 67 Results Effects of Marking on Survival and Flight Activity Survival of the leafhoppers treated with the six dyes was not significantly different (P10.05, ANOVA) than that of the undusted control group for either generation. For first generation, the average days of survival following treatment was 12.9-b1ue, 12.1-green, 11.9—magenta, 11.6-yellow, 11.4— creme, 10.7-unmarked control, and 9.3-red. For the second generation survival was 16.5-red, 15.4-yellow, 15.4-green, l5.0~unmarked control, 13.7—creme, 13.5-magenta, and 12.7- blue. Capture on yellow sticky board traps, as a measure of flight activity was also not significantly different (P?0.05, ANOVA) between marked and unmarked leafhoppers for either generation. The average days until capture for first generation was 4.2-yellow, 3.4-green, 2.8-redn 2.4-magenta, 2.4wblue, 2.2—unmarked control, and 2.1-creme. For second generation, average days until capture was 3.6-unmarked control, 3.2—creme, 3.1-magenta, 2.7-yellow, 2.5—blue, 2.5" green, and 2.3—red. Temporal Patterns of Trap Catches The recapture rates for released leafhoppers (Table l) ranged from 1.47% first generation at the Lawrence orchard release location, to 3.68%, also first generation, at the East Lansing orchard release location. Over both sites, release locations, and generations, 16,237 marked 68 Table 1. Number of marked leafhoppers released and re- captured and the recapture rate for both sites, release locations, and generations. Release Location East Lansing Lawrence Totals Generation 1 , - Orchard 1440/53 (3.68) 1975/29 (1.47) 3415/82 (2.40) Edge 1347/47 (2.46) 1975/35 (1.77) 3322/82 (2.47) 2787/100 (3.59) 3950/64 (1.62) 6737/164 (2.43) Generation 2 ‘ Orchard 2400/64 (2.67) 2350/52 (2.21) 4750/116 (2.44) Edge 2400/59 (2.46) 2350/43 (1.83) 4750/102 (2.15) 4800/123 (2.56) 4700/95 (2.02) 9500/218 (2.29) Totals ‘ . Orchard 3840/117 (3.05) 4325/81 (1.87) 8165/198 (2.42) Edge 3747/106 (2.85) 4325/78 (1.80) 8072/184 (2.28) 7587/223 (2.94) 8650/159 (1.84) 16237/382 (2.35) # released/t recaptured (X recapture rate) 69 leafhoppers were released and 382 recaptured for an overall recapture rate of 2.35%. There were significant differences in leafhopper recaptures over time between release dates at each site during each generation (P50.05, ANOVA). There was an exponential decrease in the number of leafhoppers recaptured over the 21 days following their release (Fig. 5), with the largest number of leafhoppers recaptured in the first several days following release. Temperature influenced the total number of leafhoppers (both marked and unmarked) captured each day. The average temperature for similiar leafhopper capture between the two generations was lower in the second generation than in the first (Fig. 6). For example, to trap 1 leafhopper/trap/day first generation required temperatures near 85°F (29°C), while the same results second generation required temperatures ca. 20°F (11°C) colder near 65°F (18°C). Spacial Patterns of Trap Catches There were significant differences in leafhopper recaptures over time between research sites both generations (P(0.05, ANOVA), but not between the two release locations within a site for either generation. No leafhoppers released at the orchard location were captured in the edge location trapping network, and neither were any leafhoppers released at the edge location recaptured in the orchard trap network. Trap catches of marked leafhoppers was highest at traps nearest the release location and decreased linearly with 70 25 .4 - x 20- 13 . U . L. 3 4 x a ‘ o 15"“ o < I q) .. Q: - L .. m 10- I _O - E - 3 .1 23 5‘ * q I X X " X I X 0 T I l I l I I l l I I l l l ' I . T T ' 1O 12 14 '16 18 20 22 C N .s 05 @— Doys Following Release Fig. 5. Number of P. irroratus leafhoppers recaptured each day for the 21 days following their release. 71 2 - lst Generation 1 1* xx > d o 1 CD . \ 0- cl c) . L 1.— - \ . U E 'l :3 O ""l "'FI""'IF*"‘I';" l""l "r'l" "r' "r o. ‘ . 8 i 2nd Generation L- 4 Q) .. x .D x E .- ID 2: O IIIIIIIIIITIII'IIIIIIIrrlefIlIITIIIIITIIrI 4O 45 50 55 60 65 70 75 80 85 Temperature (Fahrenheit) Fig. 6. Number of marked and unmarked P. irraratus leafhoppers captured per yellow sticky board trap per day over the mean temperature during the daily two hour crepuscular flight period following sunset. 72 distance (Fig. 7). Equations which were a special form of Taylor’s general dispersal model n=exp(a+bxc) were fitted to the data. The dispersal model (Taylor 1980) which best fit the actual data was: n = exp (3.503 + -0.133 * ln(X) + -0.28*10‘5 x X3 7) where ‘n’ is the number of insects at distance X, 'exp’ is exponent, and '1n’ is natural logarithm. Interference by other recapture traps close to the release location was assumed to occur because of the recapture trap layout and the increasing numbers of unmarked, endemic leafhoppers captured with increasing distance from the release point (Table 2). An interference factor (It) was computed for each set of traps at different distances from the release location. This factor incorporated both the number of and distance to nearby traps, with traps closer influencing the interference factor more than traps further away. The interference factor was determined by the equation: It: 1+ 2(1/d) 1-1 Where ‘It’ is the interference factor for all recapture traps at distance 't’ from the release point, 'd’ is the distance (m) to each interfering trap within 30 m of the recapture trap, and 'i’ is the number of interfering traps. The '1’ was included in the It equation as traps with no interference from other recapture traps could not transform capture counts. 73 . x Actual Data 70- 0 o Transformed N=exp(4.3+(—O.012)*X1'3) Number Leafhoppers Captured .p ‘3 l N=exp(3.5+(-.13)*ln(X)+(—.28*10‘5)*X3-7) O T r I T I T I T l T O 10 20 3O 4O 50 60 7O I T T Capture Distance from Release (meters) Fig. 7. Actual and transformed mean number of P. irraratus leafhoppers recaptured at different trapping distances, for both sites, generations and release locations, with the dispersal equation and expected line. 74 Table 2. Mean number of marked and unmarked P. irroratus leafhoppers recaptured at different trapping distances from release location, with the count following transformation by the interference factor. Trap Count Interference Transformed Distance=d Marked Unmarkeda Factor=Itb Count ""3 """"" £3?“—“iiilm'mZEEEé """"""" $33"-— 10 25.0 20.3bc 2.3498 58.7 20 21.5 24.8b 1.7759 38.2 40 16 0 32 6a 1 1333 18 l 60 6 5 32 9a 1.0513 6.8 aMeans within column followed by the same letter are not significantly different (P30.05; Scheffe’s test of data). bInterference factor was determined from the equation: It: 1 4' :3 (l/d) 1-1 Where 'It’ is the interference factor for all recapture traps at distance 't’ from the release point, ‘d’ is the distance (m) to each interfering trap within 30 m of the recapture trap, and ‘i’ is the number of interfering traps. 75 Transformation of the actual data by multiplying by the interference factor resulted in the expected exponential decrease with increasing distance from the release location (Table 2). The dispersal equation (Taylor 1978) which best fit the transformed data was: n = exp (4.297 + -0.0117 8 X1 3) where ‘n’ is the number of insects at distance X and 'exp’ is exponent. Trap captures of marked leafhoppers were highest to the southeast (120° from north) for all releases and lowest to the northwest (300° from north)(Table 3). The mean dispersal direction and mean wind direction were significantly correlated (circular correlation coefficient r=0.87) for both sites and both generations (Fig. 8). Rate of movement of the dispersing leafhoppers was greater at the Lawrence site than at the East Lansing site, and also greater second generation than first generation. At the Lawrence site, marked leafhoppers were moving 3.35 and 4.12 m/day for the first and second generations respectively, and at the East Lansing site they were moving 2.66 and 3.57 m/day for the first and second generations. Rate of movement also varied with both recapture direction and distance. The mean rate of movement (m/day) for each capture direction was fastest to the southeast (3.96m/day at 120° from north) and was slowest to the northwest (2.59m/day at 300° from north) (Fig. 9). The mean Table 3. Total number of marked P. 76 irroratus leafhoppers recaptured each direction from the release location for both orchard and edge locations at each site during the two generations. Dispersal Direction Orchard Edge 60° Orchard Edge 120° Orchard Edge 180° Orchard Edge 240° Orchard Edge 300° Orchard Edge 41 73 117 85 40 10. 18. 29. 21. 10 25 25 25 .00 77 East Lansing Ist Generation Lawrence T 80 No. Leafhoppers Receptor“ 1O 20 30 Lao-u" ‘ufl- A v .0 ................... In 4 b 4030 5““ Mean Dispersal Direction Int-mu. Mean Wind Direction Fig. 8. Number of P. irroratus leafhoppers recaptured each direction with the mean dispersal and mean wind direction during flight times for recaptured leafhoppers (circular correlation coefficient r=0.87). 300 240 120 Fig. 9. Mean rate of movement (m/day) of marked P. irroratus leafhoppers recaptured each direction from release for both sites, release locations, and generations. 79 rate of movement (m/day) for each capture distance increased with distance from release (Fig. 10) from 2.86m/day at 5m to 5.09m/day at 60m. Population estimates were made using the Lincoln Index method (Lincoln 1930) based on the numbers of recaptured marked leafhoppers and numbers of unmarked leafhoppers captured while monitoring dispersal. The index equation used was: N = a * n / r where ‘N’ is the estimated number of individuals in a population, 'a’ is the total number of individuals marked and released, 'n’ is the number of marked and unmarked individuals captured, and ‘r’ is the number of marked individuals recaptured. The estimated leafhopper density at the Lawrence site was 0.292/m2 and 0.395/m2 for the first and second generations respectively, and similarly 0.251/m2 and 0.328/m2 at the East Lansing site. Discussion The marking method had no significant affect on leafhopper survival or attraction to yellow sticky board traps. Concern that the marking method would disturb the leafhoppers (Southwood 1978) was diminished by the release of the marked leafhoppers as soon after capture as possible during the post crepuscular inactive period. Marked leafhoppers were easily distinguished as the dye was neither 80 O) L 0‘ J .s L Rate of Movement (meters/day) \ \\\\\\\\\\ C) \\\\\\\\\\\\\\\\\\\\\ \\\\\\\\\\\\\\\\\\ \ \\\\\\\\\\\ \ \\\\\\\\\\ I I I I I I I I 1 I I I 10 20 30 40 50 50 70 0 Capture Distance from Release (meters) Fig. 10. Mean rate of movement (m/day) of marked P. irroratus leafhoppers recaptured each distance from release for both sites, release locations, and generations. 81 washed or groomed off from the neck area between the head and thorax, or from the thoracic sterna between the bases of the legs on the thorax. The recapture rate was low (1.47-3.682) compared to some past leafhopper mark/release/recapture studies (26.0- 36.4%; Ito & Miyashita 1961, 17.3-43.78; Whitney & Meyer submitted), but similiar to the study by Purcell & Suslow (l-QX; 1982). Differences in dispersal activity between release dates at each site were due to temperature effects. When combined, the different temperatures cancel out the release date differences. This effect gives an expected smooth decrease in leafhopper recaptures over time. Temperature plays an important role in insect development and behavior (Tschinkel 1985). Ambient temperature influences activity of P. irroratus, and this influence differs significantly between first and second generations. The results indicate that second generation adult P. irroratus are active in temperatures as low as 40°F (4°C). These temperatures are not uncommon in Michigan during late-September and October and would indicate that cold nights will not necessarily reduce P. irroratus activity. Differences between the Lawrence and East Lansing research sites may be due to several factors. The Lawrence site is a commercial orchard which is regularly treated with pesticides, while the East Lansing site is an unsprayed 82 research orchard. Other differences included peach orchard vs. cherry orchard, and the configuration of the surrounding habitat. Inspite of these differences, the dispersal patterns were remarkably similiar. The position of the yellow sticky board traps used for recapture (Fig. 4) simplified our data analysis, but generated problems in that the effective trapping space of each trap interfered with that of other traps, especially those close to the release location. For this reason, distance capture data were transformed using the interference factor (It) which was specific for each set of traps at different distances from the release location. The increasing number of randomly distributed, unmarked, endemic leafhoppers captured with increasing distance and decreasing interference support our hypothesis of trap interference. The dispersal models derived from these data do not represent functional forms of dispersal, but rather are empirical descriptions of the dispersal by P. irroratus in this study. Differences in sites or release dates did not influence the direction the leafhoppers were moving. Wind seems to be the major factor influencing the leafhopper dispersal direction, not the presence nearby of P. irroratus habitats such as clover fields, meadows, fence rows, or wood lots. The rate of movement may be influenced by site differences. The suspected presence of insecticide residues in the Lawrence orchard may possibly encourage the 83 leafhoppers to move faster than in the unsprayed East Lansing orchard. It also appears that rate of movement increases with increasing recapture distance from the release point. Wind also influences the rate of movement, as it increases with downwind direction of dispersal. Average rate of dispersal does not imply that the leafhoppers are incapable of moving further and faster than this speed, but that under the conditions present in the field, the leafhopper average movement is a certain straight line distance per unit of time. Visual observations during light capture for marking indicate P. irroratus is a strong flyer and easily capable of quick flights over 10 m in length. The population estimates using the Lincoln Index method gave a rough idea of the leafhopper density and allowed comparisons between generations and sites. Problems with using this method (Southwood 1978) include ignoring the mortality rate, addition of emerging adults, and immigration and emmigration of unmarked individuals from the population. The predicted densities ranged from 0.25 to 0.395 leafhoppers/m2. A leafhopper density of 0.25/m2 would yield a population of 2500 leafhoppers/ha, but if 5% of those leafhoppers are infective vectors of X-disease, 125 leafhoppers would be present inoculating their host plants in the area. 84 P. irroratus outside the orchard are not making significant contributions to endemic orchard populations because of the lack of any cross captures between orchard or edge released leafhoppers. The lack of a daily movement by P. irroratus into and out of the orchard contrasts with predicted movement patterns (Mowry & Whalon 1984) and past research on other eastern X-disease vectors which invade orchards often from outside sources (McClure et a1. 1982). Further research is needed to quantify the flight distance, duration, and speed capabilities of P. irroratus to complete the picture of the movement and dispersal of this leafhopper. This could include both flight mill and wind tunnel studies to determine flight characteristics such as speed and length of flight, physiological flight capabilities and dispersal behavior tendencies of adult leafhoppers from both generations. One significant concern is the possible non-random selection of P. irroratus adults attracted to yellow. This selection could account for behavioral differences not evident in this study. Research into the effects of age, sex and the vector or non-vector status on the behavior of P. irroratus is needed. As the differences in rate of movement between the two sites indicate, current control methods, such as applications of insecticides may actually be increasing X- disease transmission by encouraging leafhopper movement. In 85 Michigan, cultural practices such as clean-till or short cut, grass—only groundcover, as suggested by Rosenberger (1977) may create unfavorable habitats for P. irroratus and may lower populations to a level which would significantly reduce X-disease transmission. References Cited Baker, R.J. and J.A. Nelder. 1978. Generalised Linear Interactive Modelling (GLIM) System, Release 3, User’s Manual. Royal Statistical Society, Rothamsted Experimental Station, Rerts, England. 257 pp. Batschelet, E. 1981. Circular Statistics in Biology. Academic Press, N.Y. 371 pp. Ito, Y. and E. Miyashita. 1961. Studies on the dispersal of leaf- and planthoppers. I. Dispersal of Nephotettix cincticeps Uhler on paddy fields at the flowering stage. Jap. J. of Ecol. 11:181-186. Larsen, K.J. and M.E. Whalon. 1987. Crepuscular movement of Parapblepsius irroratus (Say)(Romoptera: Cicadellidae), between the groundcover and cherry trees. Env. Entomol. (accepted for publication). Lincoln, E.C. 1930. Calculating waterfowl abundance on the basis of banding returns. USDA Circ. 118:1-4. McClure, M.S., T.G. Andreadis, and G.H. Lacy. 1982. Manipultaing orchard groundcover to reduce invasion by leafhopper vectors of peach X-disease. J. of Econ. Entomol. 75:64-68. Microrim. 1986. R:Base System V User’s Manual, Version 1.0. Microrim Inc., Redmond, WA. 413 pp. Mowry, T.M. and M.E. Whalon. 1984. Comparison of leafhopper species complexes in the groundcover of sprayed and unsprayed peach orchards in Michigan. Great Lakes Entomol. 17:205-209. Purcell, A.H. and E.C. Suslow. 1982. Dispersal behavior of Collodonus montanus (Homoptera: Cicadellidae) in cherry orchards. Env. Entomol. 11:1178-1182. Rosenberger, D.A. 1977. Leafhopper vectors, epidemiology and control of peach X-disease. Ph.D. Dissertation. Michigan State University, East Lansing, MI. 95 pp. SAS Institute. 1985. SAS User’s Guide: Statistics, Version 5 Edition. SAS Institute Inc., Cary NC. 956 pp. 86 87 Southwood, T.R.E. 1978. Ecological Methods. 2nd ed. Chapman and Hall, N.Y. 524 pp. Taylor, R.A.J. 1978. The relationship between density and distance of dispersing insects. Ecol. Entomol. 3:63—70. Taylor, R.A.J. 1980. A family of regression equations describing the density distribution of dispersing organisms. Nature 286:53-55. Tschinkel, W.R. 1985. Behavior and physiology. Pp. 391-435. In: Fundamentals of Insect Physiology. ed. M.S. Blum. John Wiley & Sons, N.Y. 598 pp. Whitney, S.P. and J.R. Meyer. submitted. Movement between wild and cultivated blueberry by two species of sharpnosed leafhoppers in North Carolina. J. Entomol. GENERAL CONCLUSION 88 This research has supplied information regarding the temporal and spatial distribution and dispersal patterns of leafhopper vectors of X-disease. This information is essential in the continuing development of a management plan for X—disease of stone fruits in Michigan. The monitoring of leafhopper populations has confirmed earlier work that P. irroratus is the most common known vector present in Michigan, and that vector population density can change considerably from year to year. Populations of vector leafhoppers vary between different regions, with P5 irroratus being present in high numbers in the southern portion of the state. Other leafhopper species, such as S. acutus and C. clitellarius, may be important vectors in other areas of Michigan. The observations of symptomatic chokecherry throughout the state indicate X-disease is present and transmission does occur. The daily movement cycle of the most common vector species in Michigan, P. irroratus, is now understood. The leafhopper is found during the day in the orchard groundcover and has a crepuscular flight into the fruit trees, where it spends the night. Certain fruit trees, such as sweet or sour cherry, are preferred woody hosts of P. irrora t as . The dispersal of P. irroratus within, into and out of the orchards is now known. The leafhoppers are not moving 89 90 as fast, or moving into and out of the orchards from outside habitats daily as we had previously hypothesized. Thus, P. irroratus populations outside the orchard are not making significant contributions to endemic orchard populations. Factors within the orchard, such as the presence of insecticides or cultural practices seem to influence dispersal. Wind is the major environmental factor in leafhopper dispersal direction, while temperature has a major influence on P. irroratus activity. The presence of symptomatic chokecherry may indicate a need for renewed enforcement of the MDA chokecherry eradication regulations, as chokecherry could be serving as the major source of X-disease inoculum outside the orchards. With the known dispersal behavior of P. irroratus, some other vector may be much more effective in importing X~ disease inoculum into orchards from outside sources. Cultural practices, such as clean tilling or short mowing of a groundcover of unfavorable leafhopper host plants, as suggested by Rosenberger (1977), may significantly reduce endemic orchard leafhopper populations. Biological and behavioral timing of insecticide applications is very important. Application during the two adult populations, and use of a quick knock—down insecticide after dark during the post-crepuscular inactive period when the leafhoppers are exposed on the trees could possibly reduce present and future leafhopper populations. 91 Further information is needed regarding the biology and behavior of the Michigan leafhopper vectors of X—disease. Leafhopper species should be closely examined to determine the most efficient and economically important vector. Effects X-disease MLO’s may have on the physiology and behavior of the Michigan vectors following ingestion is unknown. Of particular importance is whether some leafhopper vector species other than P. irroratus may be importing X—disease inoculum into orchards from outside sources. The natural habitat of immature stages of leafhopper vectors, particularly P. irroratus must be discovered. This will aid in the search for any natural biological control factors which may exist, including natural enemies such as parasitic wasps or nematodes, and any fungal pathogens. Further work into the flight capabilities and quantification of the effect of color on the behavior of P. irroratus is needed to complete our understanding of the movement behavior of this leafhopper. APPENDIX A LEAFHOPPER VOUCHER SPECIMENS PLACED IN THE MICHIGAN STATE UNIVERSITY ENTOMOLOGICAL MUSEUM APPENDIX A Record of Deposition of Voucher Specimens* The specimens listed on the following sheet(s) have been deposited in the named museum(s) as samples of those species or other taxa which were used in this research. Voucher recognition labels bearing the Voucher No. have been attached or included in fluid-preserved specimens. Voucher No.: 1987-02 Title of thesis or dissertation (or other research projects): Temporal and Spacial Distribution and Dispersal Patterns of Paraphlepsius irroratus (Say)(Homoptera: Cicadellidae), a major vector of X-disease in Michigan Museum(s) where deposited and abbreviations for table on following sheets: Entomology Museum, Michigan State University (MSU) Other Museums: none Investigator's Name (3) (typed) Kirk J. Larsen Date 23 April 1987 *Reference: Yoshimoto, C. M. 1978. Voucher Specimens for Entomology in North America. Bull. Entomol. Soc. Amer. 24:141-42. Deposit as follows: Original: Include as Appendix 1 in ribbon copy of thesis or dissertation. Copies: Included as Appendix 1 in copies of thesis or dissertation. Museum(s) files. Research project files. This form is available from and the Voucher No. is assigned by the Curator, Michigan State University Entomology Museum. 92 IIIIJllIII I \ 5! \.man vbwmwvo swam kua< mm some . a: I (aim. . Qtléfl . - “we 9.3.th \ < QNWIWN sans: muoHoEOuam zuwmwo>waa aumum :mwflsofiz ecu aw uwmoaap . paw msoEwooam poumwa e>opm azu po>waoom smoked n x»«& Noumwofi .oz ua£o=o> Apaahuv Amvaeaz n.sounwwuwo>cu Aawmmmmooc ma nuoocm Hmcowufippm omav S a e t co m n... n A m 1— m a nu e F. . . . mwo am: a 33.322 m r _ 33(2th "Hz .00 comma?» B 1 h 0 0 O u s m z 1 £2 2% m % .s.m.z a mess sax NN Page .038 macaHBaA "Hz .00 cawamam> .28.: m e 32 32. S wcwmcaq ummm “Hz .00 EmswaH Azamv nauwuouwfi maflnmaanmmumm 93 mace poufiwoaop pom can: no pouooHHoo :oxmu wmcuo so mawowam m _ s s s e mcoE comm no a e a a u e s r t t e .n a H m u p H a A e r o.d e .1 .1 a p. w s s e 0.8 .n u u D. m co u.n e t t .d .d u v. a 25 deiOAAPNLE "mo wapfiaz APPENDIX B FIELD MONITORING, CREPUSCULAR MOVEMENT, AND LEAFHOPPER DISPERSAL DATA Table 1. List of X-disease vector leafhopper species and number captured by site and date during the 1985 and 1986 field seasons. Leafhopper species coded as follows: Pi = Paraphlepsius irroratus, Sa = Scaphytopius acutus, Cc = Collodonus clitellarius, Ns =.N0rvellina seminuda, Oi = Orientus isbidae, Ff = Fieberiella florii, Ss = Scapboideus spp., 61 = Gyponana lamina. Page 1 of 4 Site Species - Number Captured Total Date JDate DD50 Pi Se Cc Ne 01 Ff Se 61 Bainbridge Center Site 06/24/86 175 736 l 1 07/07/86 188 955 3 3 6 08/04/86 216 1565 1 1 09/03/86 246 1986 1 1 09/16/86 259 2083 1 1 2 10/01/86 274 2349 8 12 2 22 10/17/86 290 2391 15 1 16 10/31/86 304 2441 5 4 l 10 1986 totals 40 21 2 1 64 1 Clarksville Site 06/13/85 162 588 l 6 7 06/26/85 176 750 7 13 l 21 07/10/85 190 1012 4 3 1 1 9 08/07/85 217 1539 2 2 4 08/20/85 230 1757 2 3 1 6 09/04/85 245 1996 3 7 1 11 09/18/85 259 2186 12 7 1 1 21 10/03/85 274 2308 17 72 19 10/17/85 288 2353 33 3 36 10/31/85 302 2370 17 1 1 19 11/14/85 316 2379 3 l 4 1985 totals 101 45 3 3 4 1 157 East Lansing Site 07/23/85 203 1326 4 3 7 08/07/85 217 1598 18 4 1 2 2 27 08/20/85 230 1823 22 3 6 2 33 09/04/85 245 2090 35 2 l4 1 l 53 09/18/85 259 2288 57 6 8 71 10/03/85 274 2414 124 1 l 126 10/17/85 288 2453 266 2 3 2 273 10/31/85 302 2470 113 1 14 128 11/14/85 316 2475 2 2 1985 totals 641 19 4 45 1 3 7 720 94 95 Table l. (cont’d.). Page 2 of 4 m. 34.12:. uni;E'E;;Z;;I§"""';ZEII Date JDete DD50 Pi Se Cc ls Oi F! 6s 61 8:2:3:8883388888823883828:8:38:8:888‘888888888888388328888883388;;E==888=88 'Fennville Site 05/29/85 148 385 6 1 7 06/13/85 163 534 22 4 26 06/26/85 176 700 16 17 33 07/10/85 190 947 6 4 l 11 07/23/85 203 1192 7 4 3 1 15 08/07/85 217 1451 6 8 4 2 20 08/20/85 230 1665 11 10 3 1 4 29 09/04/85 245 1911 17 12 2 3 34 09/18/85 259 2114 46 41 1 2 90 10/03/85 274 2247 59 26 1 86 10/17/85 288 2314 116 18 13 147 10/31/85 302 2336 59 2 2 3 66 11/14/85 316 2345 1 1 1985 totals 373 147 6 26 1 11 3 567 05/28/86 148 318 2 2 06/09/86 160 427 1 3 4 06/24/86 :175 708 6 1 7 07/07/86 188 918 5 2 7 07/21/86 202 1256 l 2 3 08/04/86 216 1530 3 1 1 5 09/03/86 246 1968 5 1 6 09/16/86 259 2061 4 5 9 10/01/86 274 2301 22 23 1 46 10/17/86 290 2317 13 3 16 10/31/86 304 2343 6 1 1 8 1986 totals 62 42 1 1 3 3 1 113 .Bartford Site 05/29/85 148 475 4 2 1 7 06/12/85 162 651 4 22 26 06/26/85 176 845 6 20 26 07/10/85 190 1111 1 3 l 5 07/23/85 203 1365 l 2 1 4 08/07/85 217 1640 1 1 2 08/20/85 230 1863 4 4 09/04/85 245 2123 7 12 1 20 09/18/85 259 2325 33 41 2 2 1 79 10/03/85 274 2455 42 33 75 10/17/85 288 2518 134 7 1 1 143 10/31/85 302 2550 39 2 1 42 11/14/85 316 2557 1 1 1985 totals 276 145 5 3 3 2 434 96 Table 1. (cont’d.). Page 3 of 4 Site - Species - Number Captured Total Date JDate DD50 P1 SI CC “I 01 If 58 Cl ======================================================::=======:========g== Lawrence Site 05/29/85 148 491 5 1 6 06/12/85 162 669 11 12 23 06/26/85 176 842 58 27 85 07/10/85 190 1120 8 1 l 10 07/23/85 203 1370 4 5 9 08/20/85 230 1866 l 1 09/04/85 245 2133 3 4 1 8 09/18/85 259 2336 56 65 8 1 130 10/03/85 274 2470 60 39 3 102 10/17/85 288 2535 141 15 1 3 160 10/31/85 302 2562 50 4 1 l 56~ 11/14/85 316 2571 2 2 1985 totals 399 173 2 15 l 2 592 06/09/86 160 507 1 1 06/24/86 175 757 5 1 6 07/07/86 188 988 13 5 18 07/21/86 202 1337 2 2 09/03/86 246 2119 1 1 09/16/86 259 2251 3 1 4 10/01/86 274 2500 22 8 1 31 10/17/86 290 2539 25 4 1 30 10/31/86 304 2585 18 1 19 1986 totals 89 19 l 2 1 112 Manistee Site 06/10/86 161 407 1 1 06/26/86 177 576 1 1 07/09/86 190 772 1 2 3 07/22/86 203 1079 3 1 4 08/05/86 217 1350 1 1 2 08/21/86 233 1615 2 1 2 5 09/04/86 247 1785 1 4 5 09/17/86 260 1835 l 2 3 10/03/86 276 1973 18 6 3 27 10/18/86 291 1981 12 5 2 19 11/01/86 305 1989 1 l 1986 totals 39 17 13 2 71 Northport Site 06/10/86 161 327 l 1 06/26/86 177 483 2 2 08/05/86 217 1189 3 3 08/21/86 233 1438 4 4 09/04/86 247 1589 3 3 09/17/86 260 1650 2 2 10/03/86 276 1800 1 8 9 10/18/86 291 1804 3 3 11/01/86 305 1819 1 1 1986 totals 2 23 1 2 28 97 Table l. (cont’d.). Page 4 of 4 Site Species - Number Captured Total Date JDate DD50 Pi Se Cc Ns Oi Ff Se 01 ======================33=======3:==:==============3========2=3==g==2:82:32: Walkerville Site 07/09/86 190 807 1 1 07/22/86 203 1119 l l 2 08/05/86 217 1390 2 l 3 08/21/86 233 1647 1 1 2 09/04/86 247 1820 2 1 3 09/17/86 260 1886 6 2 8 10/03/86 276 2089 20 10 1 31 10/18/86 291 2100 11 7 1 19 11/01/86 305 2133 2 2 1986 totals 46 22 2 1 71 Table 2. Number of X-disease symptomatic chokecherry observed/5 miles by site and date during the 1986 field season. Southwestern Lower Michigan. Page 1 of 2 Site Date Count Bainbridge Center 05/14/86 0 05/28/86 1 06/09/86 1 06/24/86 2 07/07/86 2 07/21/86 13 08/04/86 13 08/20/86 29 09/03/86 27 09/16/86 49 10/01/86 34 10/17/86 35 10/31/86 39 1 Lawrence 05/14/86 0 l 05/28/86 3 06/09/86 1 06/24/86 1 07/07/86 9 07/21/86 18 08/04/86 20 08/20/86 47 09/03/86 52 09/16/86 59 10/01/86 50 10/17/86 46 10/31/86 47 Fennville 05/14/86 0 05/28/86 2 06/09/86 2 06/24/86 1 07/07/86 2 07/21/86 9 08/04/86 7 08/20/86, 35 09/03/86 43 09/16/86 55 10/01/86 49 10/17/86 53 10/31/86 52 98 99 Table 2. (cont’d.). Northwestern Lower Michigan. Page 2 of 2 Site Date Count Walkerville 05/16/86 0 05/30/86 0 06/10/86 0 06/26/86 0 07/09/86 1 07/22/86 1 08/05/86 11 08/21/86 13 09/04/86 24 09/17/86 37 10/03/86 33 10/18/86 36 Manistee 05/16/86 0 05/30/86 1 06/10/86 1 06/26/86 0 07/09/86 0 07/22/86 1 08/05/86 8 08/21/86 17 09/04/86 23 09/17/86 27 10/03/86 28 10/18/86 42 Northport 05/16/86 0 05/30/86 0 06/10/86 0 06/26/86 0 07/09/86 0 07/22/86 1 08/05/86 11 08/21/86 12 09/04/86 32 09/17/86 36 10/03/86 43 10/18/86 49 irroratus Light Trap 1 0000 0 1 0030 0 1 0100 0 1 0130 0 1 0200 0 1 0230 0 1 0300 0 1 0330 0 1 0400 0 1 0430 0 1 0500 0 1 0530 0 Page 1 of 6 1985 Apple 1 0600 0 1 0630 0 1 0700 0 1 0730 0 1 0800 0 1 0830 0 1 0900 0 1 0930 0 1 1000 0 1 1030 0 1 1100 0 1 1130 0 .1 1200 0 1 1230 0 1 1300 0 1 1330 0 l 1400 0 1 1430 0 l 1500 0 1 1530 0 1 1600 0 1 1630 0 l 1700 0 1 1730 0 1 1800 0 1 1830 0 1 1900 0 1 1930 0 1 2000 0 2030 0 1 2100 0 2130 5 1 2200 168’ 1 2230 5 1 2300 2 2330 0 1 l 1 000000000 0 00000000 0 0000000000 00000 0000 mooo 0°00 3m: Imuoam Insozommmmmomm ‘044‘0MM‘0440‘MM444‘.‘4‘0‘04‘0‘444‘4‘4440‘4‘44“ 11111111111111111111else-111111111111111111111111111 Number of P. 111111111111111111111111111111111111111111111111 Replicate: Sour Cherry Sweet Cherry 1120MW34 Table 3. Crepuscular Movement Data. 1 leafhoppers captured by subhabitat and time. Generation: Groundcover 6H01639.87 ““31‘20010200100000000000 00000200000417.1058 000 000 o o o 000 0 wmomm m:mnm mmmo 0mm 25m mmmmmnmmmnmnwnmmmn 333m333333333333m333MH3333333333-03333333 111111111111111111111111111111!1.1.11.1.1.111 13 1 20 2 1 1 3 0330 S 1111111111111111111111111111111111111111 7 0 Nwmmwn Muw m “w 0.0mw0nwW mwMHwMWM mwouw 00000000001003105-n49“l536—/798mph”.hww9wu89w9ww~llflsfilbio 000000000 0 Oahu-000000.100 O mmmm mmmmo swam macs mxnmamnwummwnmmmmmaommmmo m 00000000000 0000 01111111111111.1111”Iwmmmflntmmnl 1111111111111111111111111111151111111111111111 111111111111111111111111111111111111111111111I 1111111111111111111111111111111111111111111111 100 101 Table 3 (cont’d.). 2 = July 8-9, 1985 Page 2 of 6 Replicate: 1 Generation: P Light Tra Sweet Cherry Apple Sour Cherry Groundcover 2 0000 l 1 1 2 0030 0 10000001 00000000 03030303 11223344 00000000 22222222 11111111 000000 wmmmmmm 11 21‘ 000° 1 2 0500 0 1 2 0530 0 1 2 0600 0 OMOOMOIMOMOOOOOIoooooooowooooom 5 7 0000000000000000000000000000001mn~ul3 000000000000000000000000000000 0 0 303030303030303030303030303030m0m0m 67738990011223344556677889900112233 Onunuonun00.1111.1111.1111.1111.11.1.1111.1112n4922n292254 22222222222222222222222222222222222 1.11.1.11.1.1111.1111.11.1.11?11.1.1111.11?1111.11.1.11.1 -0. Mal-0000000 MI.W.U 0 83 mm .23 mm- o 111111mm1u11m11111113222222 ‘44444M4WH4‘44MM44444.4.4.4.4444444444444‘42‘244‘444 01 1 000000000000C1004V036711 ; wmmmmmommwmomomom mmm a 11111111111111125L2 222222222222222222222205222222222222 11111111111111.411111111111111111.1111 7...t,..n,..4l7ur)t‘.3n£54674658678453854 7J716fln11510 ”2.11.000 00 000.0 000 0 0 m0 000000 677889 011 3:4 5667788 may 0112 000000311111111111111111111 2222 l1111111111111111111111111111111111 22222222222222222222222222222222222 1111111111111111111111111111111111'; 2 102 1985 Page 3 of 6 July 17-18, 3: l Replicate: Table 3 (cont’d.). Generation: Light Trap Sweet Cherry Apple Sour Cherry Groundcover 9“ 10 100100000100000000000000000000000000000000021232 000000000000000000000000000000000000000000000000 0303030303030a0303030303030303030.603030303030303 001122334455667738990011223344556677389900112233 nuOnunuonunuonunuonunvonunuonunu01L1Tl1l1.l1.1.L1.1w11ll.l1.1.L1.1n£952n49:Zo‘2 333333333333333333333333333333333333333333333333 .11.1.l1.l.11.1.l1.1.11.1.l1.1.11.1.1131.11.1.l1.1.Lfi.1.l1.1.L1.l.l1.l.11.1.¢1.1 0000000000000000000000000000000OOOOOOOIOIOOOOOCK» 303 m 3030 3 3 3 3 3 3 3 MOMOM mmmmw& 3M0mmuumummuufifiMMfiflmmmlm 22 $38 ‘4‘Q‘4uumm44444”44‘44444“‘44Q4444‘444Q4‘444‘444 333333333333333333333333333333333.633333333333333 1111111111111111111111111111111111.111111101111111 67740534160311010100000000100.000000000001000153~/ oommmmo mumo oomo 0mm o:mum oo mmmmmmo mmmmmoom mmm 19° 3 30 30 333333333333333333333333333333333333333333333333 333333333333333333333333333333333333333333333333 i 7 393 GLMMMMWNWAN1947W324OI 010001000IOOOIOCOOOMIOOIIQL4119. 000 0000 o o 000 0 o o 0000 0 M2370 MW677 WW 9001.]. 2 45 40107 233 momoooom 000 W0111111111111111111112m 222 222222222HZ2222222222222222222222222222222222222 333333333333333333333333333333333333333333333333 000010000010103223234479.49...»64557870-42637: CJIUS4CI5322110 o o 0 0000 ooooourvoco 0000000000 0 w: mmsmmommm awn wmmmwumnmnwummmmwnmmwnmmmumo n 111111111111111111111111111111111111111111111111 333333333333333333333333333333333333333333333333 111111111111111111111111111111111111111111111111‘ Light Trap 1985 Page 4 of 6 Apple 16-17, Sept. 103 Sweet Cherry Replicate:l 2 Sour Cherry Table 3 (cont’d.). Generation: Groundcover n... 31 000000000000000000000000000000000000000021160100 0 0 0000000000000000000000000000000000 mmmmmmmmwm0w0m0303030303030303030303030303030303 00112233445566778899001122334.4556677839.900112233 OnunuonunuonunuonunuonunuOnunuonu1.l.i1ll.i1.11i1711i1.i1ll.i1.1.lo‘2nzegZA4952 1.i1.1.i1.1.l1i1.11i1.11.l.11i1.11i1.l1l1.i1.1.11i1.l1.1.i1.1.i1.1.11i1.i1.1.i1i 2n¢952n4952n49‘2nao‘2n49.2fizog2n¢952n¢952n40.2n49‘2nzo52n4952nzoa2“49‘2a4952n4n. 00000000ooooooooooooooooooooooooOZOOIOOHOOOOOOOO 00 000 000 oomoooo .JMWSAuzgmmm 3;vz. W3WvW.MQwM"Mwmm mmmmmm 3wmm :mmmmmmmmm MWOWwoZUWMM ”1122 EOJIO“ 011 000000 “omoow 11111u1‘1111111111mw 222 4‘-“-““‘mm””“.‘4“m“““‘u“4““‘4‘-44“””‘4“ J 5 ml. 0000000 0 4654541 643 .71.. 1 000 000000 0 0 0 00000 0 mmmmm233 o3o om3 oommumnmnm3mmmmm nmmwnwwmmmwmn moooooooooo ooo “callllillilflllllmll1112222 22 300 «MO “00 $00 $0: 11‘111111111111111111111111111111111111111111111 111111111111111111111111.111111111141111.111111114111 222222222222222222222222222222222222222222222229; 104 Table 3 (cont’d.). Page 5 of 6 1985 17-18, Sept. 2 Replicatez2 = Generation: Light Trap Sweet Cherry Apple Sour Cherry Groundcover 9 0472 000000000000000000000000000000000000000012210000 000000000000000000000000000000000000000000000000 0303030303030303030303030303030303030303030.00303 00112233445566778n.990011223344556677889900112233 OaunuonunuonunuonunuOnunuonunuonu1.1.11i1.i1i1.i1i1.i1ll.i1i1.1171042.¢¢g2n¢9.2 222222222222222222222222222229.222222222222222222 222222222222222222222222222222229.222222222222222 oooooooooooooooooooooo ooooooooooooooooooooooo000 mmmmmmmmmwwwmmmmmmmmmmmm mmmmmmm mmmmmmmmmum mmmm ‘4‘0‘““.‘.‘.“‘.04-000‘44“““.“‘u““““‘4‘”““. 22222222222222322227.22222222222222222222222222222 2220.122222222222222222222222222222222222222222222 7MM57HM3m100000 1.1.2 5077 mzzmm 678776 0000 1 o om 0000 o wmm3m w3mwo m m m: 0000 003 om11¢ll II 1 2222 333333333W3H33“”33333333333333333333333333“:3333 222222222222222222222222222222222222222222222222 222222222222222222222222222222222222222222222222 1“»0 1430 0 1500 0 1530 0 1600 0 1630 0 1700 0 1730 1 1830 0 1900 0 1930 0 1230 0 1MNO 54.4323544365552011010000000000100010000M237u47v554 mmwmmom :mm mmm owmmmmmmmM3 omooommmmmmm wwwwwmm 2222222222222222222222222229.22222222222222222222 222222222222222222222222222222222222222222222222 222222222222222222222222222222222222222222222222 0000000000012344‘56555463557‘875‘6654514010150 mfi mm mmmmmmmmmmmxllmoommmmm mm mmmemmm mm o 1111.11181115111111111111111111111111.111111115111311 22Alp-(2222222222222222222222222222222222222222222 229.222222222222an2222222222222222222222222222422 2 1 2330 0 105 6 f 0 6 e g a D... 5 8 9 1 0 2 _ 9 1 t p e S .- 3 e t a C .1 .1 )0. .8 dB t n 02 C (00 n 30 .1 et 18 hr. 88 TD 8 G Groundcover Light Trap Sweet Cherry Apple Sour Cherry “ 3 m 898 0000000000000000000MB0000000000000000000023212000 0 O 0 0 0 000000000000000000000000000000000000 mm0m0m0m0m0m030303030303030303030303030303030303 001122334455667788950011223344556677889900112233 nu0nunuonunuOnunuOnunuonunuOnun30.L1‘l.L1.1.;1.1.;1;1.;1‘1.L1.1.L1.2nz052n49.26¢ 3333333333333333333:3333333333333333333333333333 222222222222222222272222222222222222222222222222 00000000000000000000010000000000000000000000ooon w ”mm mwmowmmmwmmmm MM” moowm M 0M momwm M MM nmwmmmwmmww wmmnumummuummmwnmmm1w2mmm nmm ‘444““.‘4“‘44““‘4‘u4‘44“.““‘4“44“‘44““‘ 33333333333333.33193333333333333333333333333333333 222222222222222122222222222222222222222222222222 0000000000010001 N2fiMRflflNWfl H67120000123flfiflnfiuw mmmmmmmmwmmmwmwmmmw mmmmmmmm mmmmmmmmmmmmmmwmm 0. 39876 5645 ‘1... 11102.“ a... 1111.11 1111111 OOOOOIOrnuoco «win/‘22 19 3 3 3 3 30303 3 omo wmmmmom «mm u77m w onxnuunmum: 9mm2 n22 11111111 _ 00 0 0 o 0 0. o o 00 1 1815 3 ‘- ‘. C3 79 In. 5 6 7 8 67! 8 I07 6 n, .1] S ‘U n] pl. 7 C5 3 ”‘5 1 0 10 O C o 000 000 000000 0000 0000 000 o o 00 m1! 233 ‘55 112233 4 566778899 112233 000000 00 0°1m1111111111111111112 222222 11111111IIIIIIHHIIIIIIIIIIIIIIII11111111111111.1511 333333333333333333333333333333333333333333333333 222222222222222222222222222222222222222222222222 Leafhopper dispersal survival Count is the mean number of days Table 4. data. Page 1 of 1 Generation 2 until captured. Generation 1 uaaoo acaoo mmav :vo assoc heaoo ammo cmo 15.6 4 10.33 2 13.83 5 16.17 6 13.17 .7 16.83, _1_3 3334.4. 73222 230: 6 6 6 2 2 2 a... 2 2. n49.4n32 106 -o-..l_._.= 14.83 7.33 14.33 13.17 1 6 6 14 5 10.17 1,3 6 10.33 13.5 _l_6_l_19433_. 11.5 6 7 1 6 3 14.17 13 1 1 16 _1_6_A__19_..BS_ 16 1111114111 Count Page 1 of 1 Generation 2 Leafhopper dispersal flight 5. to yellow sticky board trap data. Table is the mean number of days until captured. Generation 1 _ _ 46c 4 91. 7 4.8 99L.“ 214—81._37 _ 4. 1.5 76 6 oAu and? 5.1 :36763 4.6 —H::°D O .0 0 O O. 0.9.. 00.... O. n 4 233*“.3 43 12632612422133#92 n w e 1 "uoaou 134 23 56 12 L45471234567 edl "6?; 1 11¢.22393Z43 L33334i44444 flu” " can 2 22622o22722622122122222222 _._... £66.: 8 r o 1 o c .td n..a ebut m..n e t_a..m . numnae _ enua r _ 85 51 1 £71 9. .6 5 =7 4 7. rumcb .55 79727 98 63 21 3797 4.4.5 6 6 t _H. 000400 00400 00 00 O O 0 2:2: _ 22422431122 23 21.. 32 22a. 23.. 2 8 n123 "peace 12 45 71¢ 34.. 67 23 56 12 45 7 i 4 "maps 11 11 12o 22.. 22 33 33 44 44 4 w “ :mo 11.11.111.11 1.1.11 11 .11 .1141 M 107 Table 6. Leafhopper field recapture data. Page 1 of 7 --------- Release---------- ---Capture----- Distance Site Date Paint Duration Dist Dirac # Per Day East Lansing 06/25/86 3 4 10 60 1 2.5 East LanSing 07/05/86 e 1 10 60 1 10.0 East Lansing 07/05/86 e 1 5 240 1 5.0 East Lansing 07/05/86 e 1 5 0 1 5.0 East Lansing 07/05/86 e 2 5 120 1 2.5 East Lansing 07/05/86 9 2 5 120 1 2.5 East Lansing 07/05/86 9 2 5 60 1 2.5 East Lansing 07/05/86 2 2 10 180 1 5.0 East Lansing 07/05/86 0 2 5 180 1 2.5 East Lansing 07/05/86 9 2 10 240 1 5.0 East Lansing 07/05/86 9 3 5 120 1 1.7 East Lansing 06/23/86 2 15 40 240 1 2.7 East Lansing 07/05/86 2 3 10 120 1 3.3 East Lansing 07/05/86 e 4 10 180 1 2.5 East Lansing 07/05/86 9 4 10 0 1 2.5 East Lansing 07/05/86 e 5 5 180 1 1.0 East Lansing 07/05/86 2 5 40 60 1 8.0 East Lansing 07/05/86 2 7 10 60 1 1.4 East Lansing 07/05/86 a B 20 0 1 2.5 East Lansing 07/05/86 9 8 10 180 1 1.3 East Lansing 07/12/86 9 1 5 120 1 5.0 East Lansing 07/05/86 e 9 20 180 1 2.2 East Lansing 06/23/86 9 21 20 60 1 0.0 East Lansing 07/05/86 2 10 60 120 1 6.0 East Lansing 07/05/86 3 10 20 60 1 2.0 East Lansing 07/05/86 2 11 40 120 1 3.6 East Lansing 07/12/86 e 5 5 180 1 1.0 East Lansing 07/12/86 e 5 5 0 1 1.0 East Lansing 07/05/86 e 12 20 180 1 1.7 East Lansing 07/12/86 9 5 5 300 1. 1.0 East Lansing 07/12/86 e 5 5 60 1 1.0 East Lansing 07/05/86 2 13 20 120 1 1.5 East Lansing 07/05/86 6 14 '60 180 1 4.3 East Lansing 07/12/86 e 7 20 300 l 2.9 East Lansing 07/05/86 e 14 20 300 1 1.4 East Lansing 07/05/86 e 14 40 180 1 2.9 East Lansing 07/05/86 0 17 60 60 1 3.5 East Lansing 07/05/86 9 17 20 60 1 1.2 East Lansing 07/05/86 0 17 40 120 1. 2.4 East Lansing 07/12/86 9 10 5 180 1 0.5 East Lansing 07/12/86 0 11 20 180 1 1.8 East Lansing 07/12/86 0 12 20 180 I 1.7 East Lansing 07/05/86 9 19 5 60 1 0.3 East Lansing 07/05/86 9 21 20 180 1 0.0 East Lansing 07/12/86 a 14 5 180 1 0.4 East Lansing 07/12/86 e 14 5 120 1 0.4 East Lansing 07/12/86 2 14 20 180 1 1.4 East Lansing 06/23/86 0 2 5 180 1 2.5 East Lansing 07/05/86 0 1 5 240 1 5.0 East Lansing 07/05/86 0 1 5 120 1 5.0 East Lansing 07/05/86 a 2 10 180 1 5.0 East Lansing 07/05/86 0 2 10 120 1 5.0 East Lansing 07/05/86 0 2 5 0 1 2.5 East Lansing 07/05/86 0 2 10 60 1 5.0 East Lansing 07/05/86 0 2 5 240 1 2.5 East Lansing 07/05/86 0 2 5 120 1 2.5 108 109 Page 2 of 7 Distance Per Dav 3' U! N H UI H \l \1 Table 6. (cont’d.). --------- Releas —---Capture----- Site Date Faint Duration Dist Direc East Lansing 07/05/86 0 3 5 0 East Lansing 07/05/86 0 ~3 20 180 East Lansing 07/05/86 0 3 10 180 East Lansing 07/05/86 0 4 10 60 East Lansing 07/05/86 0 4 S 180 East Lansing 07/05/86 0 4 5 120 East Lansing 06/23/86 0 17 60 180 East Lansing 07/05/86 0 5 20 300 East Lansing 07/05/86 0 5 10 300 East Lansing 07/05/86 0 5 10 300 East Lansing 07/05/86 0 5 5 60 East Lansing 07/05/86 0 5 5 0 East Lansing 07/05/86 0 6 10 60 East Lansing 07/05/86 6 7 10 240 East Lansing 07/05/86 0 7 20 120 East Lansing 07/12/86 0 1 5 180 East Lansing 07/05/86 0 8 10 120 East Lansing 06/23/86 0 21 60 180 East Lansing 06/23/86 0 21 40 300 East Lansing 06/23/86 0 21 60 240 East Lansing 07/05/86 6 9 20 180 East Lansing 07/05/86 0 10 40 60 East Lansing 07/05/86 0 10 40 120 East Lansing 07/05/86 0 10 20 240 East Lansing 07/12/86 0 3 10 60 East Lansing 07/05/86 0 11 40 180 East Lansing 07/12/86 0 4 10 240 East Lansing 07/12/86 0 5 5 60 East Lansing 07/12/86 0 6 60 120 East Lansing 07/05/86 0 13 20 180 East Lansing 07/05/86 0 13 20 120 East Lansing 07/12/86 0 7 5 60 East Lansing 07/05/86 0 14 40 120 East Lansing 07/12/86 0 8 40 120 East Lansing 07/05/86 0 16 40 120 East Lansing 07/05/86 0 16 5 300 East Lansing 07/05/86 0 17 2O 60 East Lansing 07/05/86 0 17 60 60 East Lansing 07/12/86 0 10 10 0 East Lansing 07/12/86 0 10 10 120 East Lansing 07/12/86 0 11 10 120 East Lansing 07/12/86 0 12 20 60 East Lansing 07/05/86 0 20 20 0 East Lansing 07/12/86 0 14 20 180 Lawrence 07/17/86 e 2 5 120 Lawrence 07/17/86 e 3 10 60 Lawrence 07/17/86 e 3 5 0 Lawrence 07/20/86 e 2 5 60 Lawrence 07/20/86 e 2 5 180 Lawrence 07/20/86 e 2 5 300 Lawrence 07/20/86 e 3 5 0 Lawrence 07/17/86 e 6 20 60 Lawrence 07/20/86 e 3 10 60 Lawrence 07/20/86 e 4 5 120 Lawrence 07/20/86 e 4 10 180 Lawrence 07/20/86 e 4 40 120 HHHHHHHHHHHHHHHHHHHHHHHMHHHHHHHHHHHHHHHHHHHHHHHHI‘HD-‘HHHHH H OI'JHHUHMNN"UNHFFOHO‘UPOMMNOHHOHNMUNbDNNHNHUINHF‘HHNNDMHHMIL-40‘" C-0001(de\JUUILINIUUIbOVOOOUMHLIOQNLRL'IOOUO‘HC'C-ONOOOCJOObVC-C'C-O' p.- 110 Table 6. (cont’d.). Page 3 of 7 --------- Release---------- ----Capture----- Distance Site Date Point Duration Dist Direc # Per Day Lawrence 07/20/86 e 4 5 240 1 1.: Lawrence 07/19/86 e 5 20 0 1 4.0 Lawrence 07/20/86 e 4 20 60 1 5.0 Lawrence 07/20/86 e 4 10 120 1 2.5 Lawrence 07/17/86 I 7 10 60 1 1.4 Lawrence 07/20/86 e 5 10 60 1 2.0 Lawrence 07/20/86 e 5 5 120 1 1.0 Lawrence 07/20/86 e 5 10 120 1 2.0 Lawrence 07/20/86 e 6 20 60 1 3.3 Lawrence 07/20/86 e 7 10 0 1 1.4 Lawrence 07/20/86 e 7 20 120 1 2.9 Lawrence 07/20/86 e 7 40 60 1 5.7 Lawrence 07/17/86 e 11 60 60 1 5.5 Lawrence 07/20/86 e 8 40 60 1 5.0 Lawrence 07/19/86 I 11 40 300 1 3.6 Lawrence 07/19/86 e 11 40 300 1 3.6 Lawrence 07/20/86 e 12 40 120 1 3.3 Lawrence 07/20/86 e 14 20 60 1. 1.4 Lawrence 07/20/86 e 15 60 120 1 4.0 Lawrence 07/20/86 e 15 60 120 1 4.0 Lawrence 07/17/86 0 2 10 180 1 5.0 Lawrence 07/17/86 0 2 10 120 1 5.0 Lawrence 07/19/86 0 1 5 180 1 5.0 Lawrence 07/19/86 0 1 5 60 1 5.0 Lawrence 07/17/86 0 3 10 60 1 3.3 Lawrence 07/19/86 0 1 10 120 1 10.0 Lawrence 07/17/86 0 4 10 120 1 2.5 Lawrence 07/20/86 0 1 5 120 1 5.0 Lawrence 07/20/86 0 2 5 120 1 2.5 Lawrence 07/20/86 0 2 5 0 1 2.5 Lawrence 07/20/86 0 3 5 180 1 1.7 Lawrence 07/20/86 0 3 10 180 1 3.3 Lawrence 07/20/86 0 4 5 120 1 1.3 Lawrence 07/20/86 0 4 20 0 1 5.0 Lawrence 07/20/86 0 4 10 300 1 2.5 Lawrence 07/20/86 0 5 20 60 1 4.0 Lawrence 07/20/86 0 5 10 20 1 2.0 Lawrence 07/20/86 0 7 20 120 1 2.9 Lawrence 07/20/86 0 7 40 180 1 5.7 Lawrence 07/20/86 0 7 5 0 1 0.7 Lawrence 07/20/86 0 9 20 240 1 2.2 Lawrence 07/20/86 0 9 60 120 1 6.7 Lawrence 07/17/86 0 13 40 300 1 3.1 Lawrence 07/19/86 0 11 20 0 1 1.8 Lawrence 07/20/86 0 13 40 120 1 3.1 Lawrence 07/20/86 o 16 40 240 1 2.5 Lawrence 07/20/86 6 17 60 120 1 3.5 east lansing 10/07/86 0 1 20 180 1 20.0 east lansing 10/07/86 I 1 10 120 1 10.0 last lansing 10/07/86 e 1 5 300 1 5.0 east lansing 10/07/86 e 1 5 180 1 5.0 east lansing 10/07/86 e 2 5 120 1 2.5 east lansing 10/07/86 e 2 5 240 1 2.5 east lansing 10/07/86 e 2 10 180 1 5.0 east lansing 10/09/86 e 1 5 180 1 5.0 east lansing 10/09/86 e 1 5 120 1 5.0 lll Table 6. (cont’d.). Page 4 of 7 --------- Release---------- ----Capture----- Distance Site Date Paint Duration Dist Direc # Per Day east lansing 10/09/86 e 1 5 0 1_ east lansing 10/07/86 e 3 20 180 1 east lansing 10/09/86 e 1 5 60 1 east lansing 10/07/86 e 3 5 300 1 east lansing 10/07/86 e 3 10 240 1 east lansing 10/07/86 e 3 10 0 1 east lansing 10/07/86 e 4 20 120 1 east lansing 10/09/86 e 2 10 120 1 east lansing 10/09/86 e 2 10 240 1 east lansing 10/07/86 e 4 20 180 1 east lansing 10/09/86 e 2 5 120 1. east lansing 10/07/86 e 4 20 180 1 east lansing 10/07/86 e 4 5 180 1 east lansing 10/07/86 e 4 20 60 1 east lansing 10/11/86 e 1 5 180 1 east lansing 10/11/86 e 1 5 180 1 east lansing 10/09/86 e 3 20 180 1 east lansing 10/09/86 e 3 5 240 1 east lansing 10/11/86 e 1 5 120 1 east lansing 10/07/86 e 6 20 180 1 east lansing 10/11/86 e 3 5 120 1’ east lansing 10/07/86 e 8 20 180 1 East Lansing 10/11/86 e 5 40 240 1 East Lansing 10/07/86 e 9 5 180 1 East Lansing 10/11/86 e 5 10 0 1 East Lansing 10/07/86 e 10 40 120 1 East Lansing 10/11/86 e 6 20 180 1 East Lansing 10/09/86 e 8 20 0 1 East Lansing 10/07/86 e 11 40 180 1 East Lansing 10/09/86 e 9 20 300 1 East Lansing 10/09/86 e 10 20 120 1. East Lansing 10/11/86 I 10 10 120 1 East Lansing 10/07/86 e 14 20 300 1 east lansing 10/11/86 e 11 10 120 1 east lansing 10/09/86 e 13 20 120 1 east lansing 10/09/86 e 13 40 240 1 east lansing 10/09/86 e 13 40 180 1 east lansing 10/07/86 e 16 20 300 1 east lansing 10/09/86 e 14 40 120 1 east lansing 10/07/86 e 16 40 180 1 east lansing 10/11/86 e 12 10 0 1. east lansing 10/11/86 e 13 5 0 1 east lansing 10/07/86 e 19 40 240 1 east lansing 10/09/86 e 17 20 180 1 east lansing 10/11/86 e 16 40 120 1 east lansing 10/11/86 e 16 40 120 1 east lansing 10/11/86 e 17 60 120 1 east lansing 10/07/86 e 21 40 180 1 east lansing 10/11/86 e 18 40 180 1 east lansing 10/09/86 e 20 60 120 1 east lansing 10/07/86 0 1 5 180 1. east lansing 10/07/86 0 1 5 0 1 east lansing 10/07/86 0 1 10 60 1 1 east lansing 10/07/86 0 2 5 180 1 east lansing 10/07/86 0 2 5 12 1 east lansing 10/07/86 0 2 20 120 1 1 ONNOUUUNHMMNHNOONNPHHHOHHNMHNHbl-JOIDNHHUN-‘0‘UILIUIHUMUUIUILHHMHUIOUI “'UILflOOOOl'-J~OLIUUNHme~OHO-‘HUI~OhOOfJO‘LflMO00GUI\IMOVVC'C-C'MOUIOOOOHH\JOVO 112 Table 6. (cont’d.). Page 5 of 7 --------- Release ----Capture----- Distance Site Date Point Duration Dist Direc * Per Day east lansing 10/07/86 0 3 5 300 1 1.7 east lansing 10/09/86 0 1 5 20 1 5.0 east lansing 10/07/86 0 3 10 240 1 3.: east lansing 10/09/86 o 1 5 0 1 5.0 east lansing 10/09/86 0 1 5 120 1' 5.0 east lansing 10/07/86 0 3 40 120 1 3.3 east lansing 10/09/86 0 1 10 60 1 10.0 east lansing 10/09/86 0 1 5 180 1 5.0 east lansing 10/07/86 0 4 5 180 1 1.3 east lansing 10/09/86 0 2 2O 0 1 10.0 east lansing 10/09/86 0 2 20 180 1 10.0 east lansing 10/09/86 0 2 5 120 1 2.5 east lansing 10/07/86 0 4 10 6O 1 2.5 east lansing 10/07/86 0 4 40 180 1 10.0 east lansing 10/07/86 0 4 20 120 1_ 5.0 east lansing 10/09/86 0 2 5 0 1 2.5 east lansing 10/09/86 0 2 5 300 1 2.5 east lansing 10/09/86 0 3 10 180 1 3.3 east lansing 10/09/86 0 3 10 0 1 3.3 east lansing 10/11/86 0 1 5 120 1 5.0 east lansing 10/11/86 0 1 5 60 1 5.0 east lansing 10/11/86 o 1 5 180 1 5.0 east lansing 10/11/86 0 2 5 120 1 2.5 east lansing 10/11/86 0 3 10 60 1 3.3 East Lansing 10/09/86 0 7 20 120 1‘ 2.9 East Lansing 10/11/86 0 5 5 60 1 1.0 East Lansing 10/09/86 0 8 20 180 - 1 2.5 East Lansing 10/09/86 0 8 20 180 1 2.5 East Lansing 10/07/86 0 10 10 240 1 1.0 East Lansing 10/09/86 0 B 10 0 -1 1.3 East Lansing 10/09/86 0 8 20 60 1 2.5 East Lansing 10/07/86 0 10 40 120 1 4.0 East Lansing 10/09/86 0 9 40 120 1 4.4 East Lansing 10/09/86 0 9 20 180 1 2.2 East Lansing 10/07/86 0 12 10 180 1_ 0.8 East Lansing 10/11/86 0 8 20 120 1 2.5 East Lansing 10/07/86 0 13 20 180 1 1.5 East Lansing 10/11/86 0 10 20 0 1 2.0 East Lansing 10/09/86 0 12 20 240 1 1.7 East Lansing 10/07/86 0 14 40 120 1 2.9 East Lansing 10/07/86 0 14 20 120 1 1. east lansing 10/09/86 0 13 20 60 1 1.5 east lansing 10/09/86 0 13 10 180 1 0.8 last lansing 10/11/86 0 11 40 180 1 3.6 east lansing 10/11/86 0 11 10 O 1. 0.9 east lansing 10/07/86 0 15 40 240 1 2.7 east lansing 10/07/86 0 15 20 120 1 1.3 east lansing 10/09/86 0 14 20 300 1 1.4 east lansing 10/09/86 0 14 40 120 1 2.9 east lansing 10/09/86 0 14 10 120 1 0.7 east lansing 10/11/86 0 12 4O 60 1 3.3 east lansing 10/07/86 0 16 40 180 1 2.5 east lansing 10/09/86 0 17 10 120 1 0.6 east lansing 10/07/86 0 19 20 0 1 1.1 east lansing 10/11/86 0 16 40 60 1, 2.5 east lansing 10/11/86 0 16 20 180 1 1.3 b 113 Table 6. (cont’d.). Page 6 of 7 --------- Release ----Capture----- Distance Site Date Point Duration Dist Direc # Per Day east lansing 10/11/86 0 18 20 60 1 1.1 east lansing 10/09/86 0 21 40 120 1 1.9 east lansing 10/11/86 0 20 60 120 l 3.0 east lansing 10/11/86 0 20 60 120 1 3.0 Lawrence 10/09/86 e 1 5 300 1 5.0 Lawrence 10/09/86 e 1 5 60 1 5.0 Lawrence 10/09/86 e 1 10 180 1 10.0 Lawrence 10/10/86 e 1 20 0 1 20.0 Lawrence 10/09/86 e 2 10 240 1 5.0 Lawrence 10/09/86 e 2 10 300 1 5.0 Lawrence 10/09/86 e 2 10 60 1 5.0 Lawrence 10/09/86 e 2 60 120 1 30.0 Lawrence 10/10/86 e '1 5 120 1 5.0 Lawrence 10/11/86 e 1 5 0 1‘ 5.0 Lawrence 10/09/86 e 3 20 120 1 6.7 Lawrence 10/10/86 e 2 10 60 1 5.0 Lawrence 10/11/86 e 1 5 180 1 5.0 Lawrence 10/10/86 e 3 10 120 1 3.3 Lawrence 10/11/86 e 3 5 240 1 1.7 Lawrence 10/09/86 e 7 20 240 1 2.9 Lawrence 10/09/86 e 7 20 240 1 2.9 LAWRENCE 10/10/86 e 7 10 60 1 1.4 LAWRENCE 10/11/86 e 6 10 300 1 1.7 LAWRENCE 10/09/86 e 8 40 120 1‘ 5.0 LAWRENCE 10/09/86 e 9 40 180 1 4.4 LAWRENCE 10/10/86 e 9 20 60 1 2.2 LAWRENCE 10/09/86 e 10 5 180 1 0.5 LAWRENCE 10/09/86 e 11 20 60 1 1.8 LAWRENCE 10/10/86 e 11 40 120 1 3.6 LAWRENCE 10/10/86 e 11 20 60 1 1.8 LAWRENCE 10/10/86 e 11 10 60 1 0.9 LAWRENCE 10/10/86 e 11 20 60 1 1.8 LAWRENCE 10/10/86 e 2 10 0 1 0.8 LAWRENCE 10/11/86 e 11 40 60 1. 3.6 LAWRENCE 10/11/86 e 11 5 60 1 0.5 LAWRENCE 10/09/86 e 13 10 60 1 0.8 LAWRENCE 10/10/86 e 12 40 120 1 3.3 LAWRENCE 10/09/86 e 13 20 240 1 1.5 LAWRENCE 10/11/86 e 11 20 120 1 1.8 LAWRENCE 10/10/86 e 13 40 180 1 3.1 LAWRENCE 10/09/86 e 14 20 120 1 1.4 LAWRENCE 10/10/86 e 15 20 120 1 1.3 LAWRENCE 10/09/86 e 16 40 120 1 2.5 LAWRENCE 10/09/86 e 17 40 120 1, 2.4 LAWRENCE 10/10/86 e 17 60 120 1 3.5 LAWRENCE 10/10/86 e 18 20 240 1 1.1 LAWRENCE 10/09/86 e 20 60 120 1 3.0 LAWRENCE 10/11/86 e 19 40 180 1 2.1 LAWRENCE 10/11/86 e 21 60 12 1 2.9 Lawrence 10/09/86 0 1 5 60 1 5.0 Lawrence 10/09/86 0 1 5 120 1 5.0 Lawrence 10/09/86 0 1 5 60 1 5.0 Lawrence 10/09/86 0 1 5 240 1 5.0 Lawrence 10/09/86 0 1 5 0 1, 5.0 Lawrence 10/10/86 0 1 20 60 1 20.0 Lawrence 10/10/86 0 1 5 300 1 5.0 114 Table 6. (cont’d.). Page 7 of 7 _________ Release ----Capture----- Distance Site Date Point Duration Dist Direc # Per Day Lawrence 10/09/86 o 2 20 180 1 10.0 Lawrence 10/10/86 o 1 40 120 1 40.0 Lawrence 10/09/86 0 2 10 240 1 5.0 Lawrence 10/09/86 0 2 10 0 1 5.0 Lawrence 10/10/86 0 2 5 120 1 2.5 Lawrence 10/09/86 o 3 10 240 1 3.3 Lawrence 10/11/86 o 1 5 180 1 5.0 Lawrence 10/11/86 0 1 5 300 1, 5.0 Lawrence 10/09/86 0 4 10 180 1 2.5 Lawrence 10/11/86 0 3 10 180 1 3.3 Lawrence 10/09/86 0 6 2 120 1 3.3 Lawrence 10/09/86 0 7 20 120 1 2.9 Lawrence 10/10/86 0 6 5 0 1 0.8 LAWRENCE 10/11/86 0 6 10 300 1 1.7 LAWRENCE 10/10/86 0 7 20 240 1 2.9 LAWRENCE 10/09/86 0 8 40 120 1 5.0 LAWRENCE 10/10/86 0 7 40 180 1 5.7 LAWRENCE 10/09/86 0 9 60 180 1. 6.7 LAWRENCE 10/10/86 0 8 40 120 1 5.0 LAWRENCE 10/09/86 0 10 20 0 1 2.0 LAWRENCE 10/11/86 0 8 10 240 1 1.3 LAWRENCE 10/10/86 0 9 5 120 1 0.6 LAWRENCE 10/09/86 o 11 20 240 1 1.8 LAWRENCE 10/10/86 0 10 1O 0 1 1.0 LAWRENCE 10/09/86 0 12 20 180 1 1.7 LAWRENCE 10/11/86 0 10 20 240 1 2.0 LAWRENCE 10/11/86 0 10 10 60 1 1.0 LAWRENCE 10/11/86 0 10 20 180 1, 2.0 LAWRENCE 10/09/86 o 13 10 240 1 0.8 LAWRENCE 10/11/86 0 11 20 60 1 1.8 LAWRENCE 10/10/86 o 12 20 0 1 1.7 LAWRENCE 10/11/86 0 11 40 60 1 3.6 LAWRENCE 10/10/86 0 12 40 120 1 3.3 LAWRENCE 10/10/86 0 13 40 60 1 3.1 LAWRENCE 10/11/86 0 12 10 180 1 0.8 LAWRENCE 10/09/86 0 14 5 120 1 0.4 LAWRENCE 10/09/86 0 15 20 180 1 1.3 LAWRENCE 10/09/86 0 16 60 60 1_ 3.8 LAWRENCE 10/10/86 0 16 40 120 1 2.5 LAWRENCE 10/09/86 0 17 60 60 1 3.5 LAWRENCE 10/10/86 0 17 20 300 1 1.2 LAWRENCE 10/11/86 0 17 60 120 1 3.5 LAWRENCE 10/09/86 0 20 40 240 1 2.0 LAWRENCE 10/11/86 0 19 20 240 1 1.1 LAWRENCE 10/10/86 0 21 60 120 1 2.9 Table 7. Number of male and female P. irroratus captured by method, site and date during 1985 and 1986. Page 1 of 4 Site Yellow Board Sweep Net Total Date 2 Male Female Male Female Eaiybridge Ceataswfiiis 06/24/86 1 1 07/07/86 1 1 1 3 08/04/86 1 1 09/03/86 1 1 09/16/86 1 1 10/01/86 2 3 1 2 3 10/17/86 3 7 2 3 15 10/31/86 1 4 5 1986 Totals 10 15 4 e 35 Claxksyillem§ite 06/13/85 1 1 06/26/85 3 1 1 2 7 07/10/85 2 1 1 4 08/07/85 2 2 08/20/85 1 1 2 09/04/85 1 2 3 09/18/85 5 3 1 3 12 10/03/85 3 8 3 3 17 10/17/85 10 17 4 2 33 10/31/85 14 2 1 17 11/14/85 2 1 3 1985 Totals 40 39 1o 12 101 Baasmkapaiasmfiita 07/23/85 3 1 4 08/07/85 12 5 1 18 08/20/85 7 7 3 5 22 09/04/85 19 10 2 4 35 09/18/85 27 23 3 4 57 10/03/85 84 27 5 8 124 10/17/85 207 55 4 266 10/31/85 66 47 113 11/14/85 1 1 2 1985 Totals 425 176 13 26 641 115 116 Table 7. (cont’d.). Page 2 of 4 Site Yellow Board Sweep Net Total Date Male Female Male Female Fennville 8139 05/29/85 2 4 6 06/13/85 2 1 ll 8 22 06/26/85 3 6 1 6 16 07/10/85 4 1 l 6 07/23/85 1 4 1 l 7 08/07/85 3 3 6 08/20/85 2 5 2 2 11 09/04/85 9 6 2 17 09/18/85 18 28 46 10/03/85 40 16 3 59 10/17/85 81 34 1 116 10/31/85 31 26 2 59 1985 Totals 194 130 20 27 371 05/28/86 1 l 2 06/24/86 2 1 1 2 6 07/07/86 2 2 1 5 07/21/86 1 1 08/04/86 1 1 l 3 09/16/86 1 1 1 1 4 10/01/86 9 11 1 1 22 10/17/86 5 5 2 1 13 10/31/86 5 1 6 1986 Totals 27 23 6 6 62 Nartford Site 05/29/85 2 1 1 4 06/12/85 1 2 1 4 06/26/85 3 3 6 07/10/85 1 1 07/23/85 1 1 08/20/85 1 l l 1 4 09/04/85 1 3 2 1 7 09/18/85 16 4 6 7 33 10/03/85 30 7 2 3 42 10/17/85 99 35 134 10/31/85 26 12 1 39 11/14/85 1 1 1985 Totals 180 70 12 14 276 —-—-——_———————————_————————_--.——--—————-—-~-————————-——.--- 117 Table 7. (cont’d.). Page 3 of 4 Site Yellow Board Sweep Net Total Date Male Female Male Female Lawrence Site 05/29/85 1 1 3 5 06/12/85 3 4 4 11 06/26/85 24 25 2 5 58 07/10/85 3 4 1 8 07/23/85 1 3 4 08/20/85 1 1 09/04/85 2 1 3 09/18/85 38 15 2 1 55 10/03/85 29 27 1 3 50 10/17/85 97 43 1 141 10/31/85 33 17 50 11/14/85 2 2 1985 Totals 230 138 10 21 399 06/09/86 1 1 06/24/86 3 2 5 07/07/86 5 3 2 3 13 07/21/86 2 2 09/16/86 2 1 3 10/01/86 10 9 1 2 22 10/17/86 10 9 3 3 25 10/31/86 13 5 18 1985 Totals 45 28 7 8 89 Menistsngiis 06/26/86 1 1 07/09/86 1 1 07/22/86 2 1 3 08/05/85 1 1 09/04/85 1 1 09/17/86 1 -. . 1 10/03/85 12 4 2 18 10/18/85 4 3 3 2 12 11/01/86 1 1 1986 Totals 21 8 7 3 39 118 Table 7. (cont’d.). Page 4 of 4 Site Yellow Board Sweep Net Total Date Male Female Male Female Northport Site 10/03/86 1 1 11/01/86 1 l 1986 Totals 2 2 Walker“ 119 S 11.9. 07/09/86 1 07/22/86 1 08/05/86 1 1 08/21/86 1 09/04/86 1 1 09/17/86 4 l 1 10/03/86 11 7 2 6 2 1 1 MN NHomNi—‘NHH 10/18/86 11/01/86 1986 Totals 25 12 5 4 46