IIIIIIUIIIIIIIIIIII I I I n IIIIIII II II I II IIIIII II III II THE-51305303 This is to certify that the dissertation entitled PARAMETERS AFFECTING EMERGENCE AND MANAGEMENT STRATEGIES OF THE APPLE MAGGOT FLY, Rhagoletis pomonella (Walsh) DIPTERA: TEPHRITIDAE presented by Larry Gene Olsen has been accepted towards fulfillment of the requirements for Ph.D. degreeinEntomology C3v¢quwg¥ I\QL3EII\ (I M aon professor Dfie May 21, 1982 MSU is an Affirmatiw Action/Equal Opportunity Institution 0- 12771 MSU LIBRARIES $3.2!!!— RETURNING MATERIALS: PIace in book drop to remove this checkout from your record. FINES wiII be charged if book is returned after the date stamped be10w. PARAMETERS AFFECTING EMERGENCE AND MANAGEMENT STRATEGIES OF THE APPLE MAGGOT FLY, Rhagoletis pomonella (Walsh) DIPTERA: TEPHRITIDAE BY Larry Gene Olsen A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Entomology 1982 ABSTRACT PARAMETERS AFFECTING EMERGENCE AND MANAGEMENT STRATEGIES OF THE APPLE MAGGOT FLY, Rhagoletis pomonella (Walsh). DIPTERA: TEPHRITIDAE BY Larry Gene Olsen Emergence of apple maggot flies, Rhagoletis pomonella, was determined at the Kalamazoo State Hospital (K.S.H.) orchard in 1977-1980 and in the Upjohn orchard in 1977, 1979-80 by four methods. The yellow Zoecon AM trap was determined to be the preferred tool to estimate first emergence. Biotic and abiotic parameters were examined for their influence on timing of first and season long emer- gence. Air temperature degree days was discovered to be the best predictor of first emergence. The biotic parameters of variety of apple reared in, orchard floor culture and location of larval pupation all affect first emergence. The phenological predictive model of emergence developed using the K.S.H. data predicted emergence to within $2.5 days at the K.S.H. orchard and LS days at the Upjohn orchard. Season long emergence expressed as accumulative percent catch can be predicted extremely well by accumulative air or soil temperature degree days or percent soil moisture. Thirty-one commercial orchards were monitored in 1979 and 1980 for the presence of apple maggot flies and percent fruit damage to study parameters associated with different management schemes. The yellow Zoecon AM trap was preferred over the red sticky sphere trap because it caught flies sooner, caught gravid females sooner, and provided more consistent prediction of fruit damage. Yellow Zoecon AM traps should be placed in the perimeter row of the orchard and located near two potential outside sources of apple maggot. Flies were caught in the orchard an average of 8 days after they were caught in abandoned trees outside the orchard but ranged from -17 1x3 35 days indicating flies should be monitored in the orchard and not in abandoned trees. It was shown that fly catch and fruit damage greatly decreases from the perimeter row into the orchard, indica- ting that a feasible alternative management scheme would be to spray perimeter rows when no other pests are present. ACKNOWLEDGEMENTS The author wishes to extend sincere appreciation to Dr. Angus J. Howitt for his support, guidance and patience throughout the course of this study, and for providing the necessary physical and financial resources. Thanks is due also to Dr. Brian A. Croft, Dr. Stuart H. Gage, Dr. Donald J. Ricks, and Dr. Mark E. Whalon for serving on the Guidance Committee and making helpful suggestions. Numerous other people have assisted in this study, without their help this could not have been completed. Dr. Jay P. Brunner helped formulate the objectives and design some of the experiments. Mr. Leonard Larson spent considerable hours assisting in data collection and analysis. Pest Management Field Assistants Phil Schwallier, Scott Ruffe and Pam Cater assisted by monitoring orchards. Many commercial growers let me use a portion of their orchards to carry out studies involving different management strategies. Lastly Mobay Chemical Corporation provided the impetus for the completion and generously gave me time off to finish. Thank you especially to my wife and two sons who have endured my absence with great patience and provided the necessary encouragement and support without which this could not have been completed. iv TABLE OF CONTENTS ABSTRACT 0......O...OOOOOOOOOOOOOIOOOOOOOO TITLE PAGE ..... ........ . .............. . ACKNO‘qLEDGEMENTS OOOOOOOOOOOOOOOOOOOOOO. TABLE OF CONTENTS OOOOOOOOOOOOOOOOOOOOOO LIST OF TABLES .................. . ...... LIST OF FIGURES OOOOOOOOOOOOOOOOOOOOOOOO INTRODUCTION .......................... . LIFE CYCLE OF APPLE MAGGOT ............. PART 1 - PARAMETERS AFFECTING EMERGENCE Emergence Patterns in Michigan ................... First Emergence by Calendar Date .. Parameters Affecting Emergence .... Abiotic Components ........... Air Temperature ......... Soil Temperature ........ Soil Moisture ........ ..... Field Studies ...... Laboratory Studies . Biotic Components ............ Variety of Apple Reared In Two Year Life Cycle ....... Orchard Floor Culture ..... Partial Second Generation Location of Larval Pupation ............ Phenological Predictive Model of Emergence .. First Emergence ........................ Season Long Emergence .................. PART 2 - MANAGEMENT STRATEGIES IN COMMERCIAL ORCHARDS . Determination of Best Method of Monitoring ....... Trap Type Comparison ........................ First Catch ............................ First Gravid Female .................... Most Flies Caught ...................... Last Fly Caught .................;...... Trap Placement .............................. Trap Density ................................ iii iv vii xiv 31 47 50 50 66 78 84 92 95 96 111 118 130 139 149 153 161 167 171 175 173 177 178 180 182 186 Delay in Catch of AM Flies in Commercial Orchards From Outside Sources ........ ..... ........... 190 Identification of Flies on Traps ... ..... ......... 195 Sampling for and Prediction of Fruit Damage ...... 199 Determination of First Oviposition Based on a Biofix of First Catch ... ..... .......... 200 Sample Size to Estimate Fruit Damage ........ 226 Whole Tree Sample ...................... 228 Orchard Sampling ....................... 235 Number of Apples and Trees ........ 236 Edge Effect of Damage ............. 238 Correlation of Trap Catch to Fruit Damage ........ 241 Comparison of Trap Types .................... 244 Comparison to Outside Fly Pressure ......... . 247 Recommendations for Monitoring and Managing AM in Commercial Orchards ..........................249 Timing of First Spray ...... ................. 249 Number of Spray Necessary ................... 251 Last Spray for Apple Maggot Control ......... 252 Perimeter Spraying .................... ...... 253 LITERATURE CITED ...................................... APPENDIX 1: Voucher Specimens ......... ......... .......... 2: Cooperating Commercial Growers............... 3: Degree Day Accumulations at Research Sites... vi LIST OF TABLES PAGE Rating System of Potential Apple Maggot Fly Pressure to Commercial Orchards................... 12 Mean Calendar Dates for Different Propor- tions of Emergence and Trap Catch of the Apple Maggot Fly at Various Locations in Michigan from 1977 to 1980........................ 19 First Emergence or Catch of Apple Maggot Flies in K.S.H. and Upjohn Orchards............... 33 Influence of Years on Different Methods of Measuring First Emergence of Apple Maggot Flies at the K.S.H. and -Upjohn Orchards (Mean iS.E. Date of Emergence and Coefficient of Variation for Combined Data for All Years in Each Orchard) ........ . ....... ............... ...... 37 Influence of Method. of Estimating First Emergence of Apple Maggot Flies at the K . S . H. and Upjohn Orchards Over a Four Year Period (Mean iS.E. Date of Emergence Plus Coeffi- cient of Variation Within the Year)......... ...... 41 Comparison of Yellow Zoecon AM. Traps and Red Sticky Spheres in First Capture of Apple Maggot Flies (Mean :S.E. Date of First Catch) .................... . ....... . ..... . ......... 44 Five Foot Level Air Temperature Degree Day (Base = 48°F; B.E. Method) Accumulations for Different Proportion of Emergence and Trap Catch of Apple Maggot Flies at Various Locations in Michigan From 1977 to 1980........... 55 Influence of Years on Different Methods of Measuring First Emergence of Apple Maggot Flies at the K.S.H. and Upjohn Orchards (Mean iS.E. Air Degree Day Accumulation at Five Foot Level at Base 48°F; B.E. Method Plus Coefficient of Variation Values).................. 58 Influence of Method of Estimating First Emergence of Apple Maggot Flies at che K.S.H. and Upjohn Orchards Over a Four Year Period (Mean iS.E. Air Degree Day Accumulation at Five Foot Level at Base 48°F; B.E. Method Plus Coefficient of Variation Within the Year)0..........OOOOOOOOOOOOOO......OOOIOOOOOOOOOO 61 vii LI... J TABLE PAGE 10. Relationship of Two Inch Degree Days on the South Side of the Tree to Air Degree Days at the K.S.H. and Upjohn Orchards in 1977 - 1980 (Base = 48°F; Method = B.E.)................. 68 11. Influence of Years on Different Methods of Measuring First Emergence of Apple Maggot Flies at the K.S.H. and Upjohn Orchards (Mean iS.E. Soil Degree Day Accumulations at the Two Inch Level on the South Side of the Tree at Base 48°F; B.E. Method)........................ 71 12. Influence: of .Method. of Estimating First Emergence of Apple Maggot Flies at the K.S.H. and Upjohn Orchards Over a Four Year Period (Mean iS.E. Air Degree Day Accumulation at the Two Inch Level on the South Side of the Tree at Base at 48°F; B.E. Method)................. 76 13. Length of Time Necessary to Completely Dry Soil Samples Contained in a 2 Dram Vial in a Drying Oven Held at 30°C. (Weight Loss from Previous Day in Grams at N = North, M = Middle, on S = South Side of the Tree)............ 82 14. Percent Soil Moisture in 1979 at Research Orchards Located in Kalamazoo, Michigan........... 85 15. Percent Soil Moisture in 1980 at Research Orchards Located in Western Michigan.............. 86 16. Season Long Mean Soil Moistures at Various Locations Under Apple Trees in Three Dif- ferent Sites (Oneway ANOVA)....................... 89 17. Comparison of Mean Soil Moisture Per Tree at Different Orchards (Oneway ANOVA)................. 93 18. Coefficient of Determination (R3) of Accumu- lative Mean Percent Soil Moisture with Accumulative Percent Apple Maggot Fly Trap Catch........................................ ..... 93 19. Date and Air Degree Day Accumulation (Base 48°F; B.E. Method) of First Emergence of Apple Maggot Fly Due to Variety 1t Matured in Measured by Seeded Emergence Cages at the K.S.H. Orchard in 1979................................... 101 20. Effect of 'Variety’ on First. Emergence of .Apple Maggot Flies at the K.S.H. Orchard as Deter- mined by Catch. in Cages Placed. Over “Naturally Infested. Ground (Mean. Air Degree Days at Base 48°F; B.E. Method)................................ 101 viii TABLE 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. PAGE Effect of Variety on First Catch of Apple Maggot Fly at the K.S.H. Orchard as Deter- mined by Catch on Yellow Zoecon AM Traps (Mean Air Degree Days at Base 48°F; B.E. Method)........................................... 104 Effect of Variety on First Catch of Apple Maggot Fly at the K.S.H. Orchard as Deter— mined by Catch on Red Spheres (Mean Air Degree Days at Base 48°F; B.E. Method)............ Effect of Variety on First Catch of Apple Maggot Fly at the Fwdium Density Upjohn Orchard in 1979 and 1980 (Mean Air Degree Days at Base 48°F; B.E. Method)........................ Comparison Across Years and Methods of the Influence of Variety on First Emergence or Capture of Apple Maggot Fly at the High Fly Density K.S.H. Orchard............................ Comparison Across Years and Methods of the Influence of Variety on First Catch of Apple Maggot Fly at the Medium Fly Density Upjohn Orchard (Difference from Mean Air Degree Days Base 48°F; B.E. Method....................... ..... Literature References to the Occurrence of a Two-Year Life Cycle of Apple Maggot "Fly with Percent of Carry-Over and Time of Emergence in Relation to First Year Fly Values Re- corded ........................ .................... Effect of Carry-Over Pupae on Timing of Emergence of Apple Maggot Flies at the K.S.H. Orchard in 1980 (Air Degree Days at Base 48°F; B.E. Method)................................ Number and Percent of Apple Maggot Flies Overwintering to the Second Season at the K.S.H. orChard by Variety-..........OOOOOIOOOOOCOO Effect of Orchard Floor Culture on Degree Day Accumulation at the K.S.H. Orchard in 1979 (Degree Days Base 48°F; Ave:age Method)........... Mean Differences in Degree Day Accumulations Between Different Orchard Floor Cultures for Various Time Periods at the K.S.H. Orchard in 1979 (Degree Days Base 48°F; Averaging Method)........................................... ix 105 106 108 110 112 114 116 121 124 TABLE 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. Mean Number of Air Degree Day Units Per Day for Different Time PEriods Throughout the Year at Different Sites in Michigan in 1980 (Five Foot Level Degree Days at Base 48°F; B.E. Method)............................... ....... Mean Number of Grass Litter Zone Degree Day Units Per Day for Different Time Periods Throughout the Year at Different Sites in Michigan in 1980 (Degree Day Base 48°F; B.E. Method) ....... .... .......... . ...... ...... ........ . Occurrence of a Second Generation Apple Maggot Flies from Dutchess Variety in 1979 at the K.S.H. orChardOO......OOOOOOOOOOOOOOOOO0...... Percent Infertile Female Apple Maggot Flies as Determined by Dissections of Trapped Females in 1979.00.00.000000000000 ....... I. ....... Percent Infertile Female Apple Maggot Flies as Determined by Dissections of Trapped Females Caught in 1980.................. .......... First Emergence of Apple Maggot Flies from the South and North Side of the Tree at the K.S.H. Orchard in 1978. ....... . ........ . .......... First Emergence of Apple Maggot Flies from the South and North Side of the Tree at the K.S.H. Orchard in 1979............. ..... .......... Soil Temperatures Measured on the South and North Side of the Tree at the Two Inch Depth Recorded as Accumulated Degree Days Since April 1.... ..... . ............ ..... ................ Differences in Degree Day Accumulations at the Soil Surface and Two Inch Depth at Base 48°F (B.E. Method) on the South Side of the Tree at the Upjohn Orchard............................. Predicted Emergence of Apple Maggot Fly Based on Air Degree Day Accumulations................... Influence of Variety of Apple Reared In on Prediction of First Emergence of Apple Maggot Fly.......OOOOOOOOOOOOOO0......000......00.0.0...O Influence of Orchard Floor Culture on Pre- diction of First Emergence of Apple Maggot FlYOOOOOOOOCOOOOI.0.0.0.0..........OOOOOOOOOOOOOOO X PAGE 127 128 133 134 136 142 144 145 147 151 155 156 TABLE 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. PAGE Influence of location of Pupation on Free diction of First Emergence of Apple Maggot FlYIOOOOOOOO ........ 0...... ..... 0.0.0.0... ........ 157 Predicted First Trap Catch of Apple Maggot Flies with the Yellow Zoecon AM Trap in Abandoned Trees Around Commercial Orchards.. ...... 158 Least Squares Linear Regression of Accumula- tive Percent Trap Catch to Accumulative Air Degree Days at IBase 48°F (8.9°C); B.E. Method... ........ ............ ........ . ........ .... 163 Date of First Catch of Apple Maggot Flies in Commercial Orchards and in Abandoned Trees Around Commercial Orchards on Different Types of Traps ........ . ................... ....... ....... 176 Average Number of Apple Maggot Flies Caught per Trap on Each Trap Type in Commercial Orchards............ ................. . ............ 179 Weekly Average Trap Catch of Apple Maggot Fly Adults on Yellow Zoecon AM Traps in and Around Commercial Orchards ..... .............. ..... 181 Location of Trap Placement in Commercial Orchards in Relation to Different Mean Trap Catch Parameters .................................. 183 Migration of Apple Maggot Flies Into Commer- cial Orchards as Determined by Yellow Zoecon AM Trap Catch (5 Orchard Sample - 4 Reps/Orchard).... 186 Relationship of Trap Density to Catch of Apple Maggot Flies ...................................... 187 List of Tephritidae Flies Caught on Yellow Zoecon AM Traps Placed in Apple Orchards........... ..... . 197 Duration of Pre-Oviposition Period as Deter- mined by the Interval Between a Biofix of First Catch on the Yellow Zoecon AM Trap and the First Stung Apple at the K.S.H. Orchard in 1977........................................... 203 Determination of the Length of the Pre-Ovi- position Period Initiated by a Biofix of Catch on the Yellow Zoecon AM Trap and Terminating with the Finding of Stung Fruit at the K.S.H. Orchard in 1978..................... 205 xi TABLE PAGE 55. Determination of the Length of the Pre-Ovi- position Period in 1979 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the K.S.H. Orchard........................................... 207 56. Determination of the Length of the Pre-Ovi- position Period in 1979 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the Upjohn Orchard......... ...... .................... ........ 208 57. Determination of the Length of the Pre-Ovi- position Period in 1979 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females in Abandoned Trees Around Commercial Orchards ...... ........... ....... 209 58. Determination of the Length of the Pre-Ovi- position Period 1J1 1980 Beginnimg with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the K.S.H. Orchard........................................... 211 59. Determination of the Length of the Pre-Ovi- position Period in 1980 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the Upjohn LJC.1‘.L.I‘o omaa ecu mo Hmuoa Hmzudoucoo xuaameuoz oasu< $I noumx wwm .H ounwam cofimumdmwa uazn< I unmadoao>mn Hm>um4 coaumdnm cofiufim0dw>o +I vowumm cowufim0da>OIoum oocmwumam uaap< 6 PART 1 - PARAMETERS AFFECTING EMERGENCE EMERGENCE PATTERNS IN MICHIGAN Prior to identifying and quantifying variables associ- ated with emergence, emergence patterns have to be estab- lished and data gathered. This should be completed at several locations over several years so that differences that might exist can be shown to be real and consistent. Causes of the variances associated with real differences can then be studied. Literature and Materials and Methods Monitoring Techniques - Throughout this study, four techniques of measuring emergence or flight activity were utilized. The first involved placing an emergence cage over ground that was naturally infested by apple maggot pupae the proceeding fall. One meter square pyramids shaped cages that had a collection devise on the top filled with ethylene glycol were utilized in these studies . This preserved the flies so they could be counted and sexed at each visit. As reported in the literature (Caesar and Ross, 1919; Mundinger, 1930), this method provides a very accurate indication of emergence. However, it does not determine the length of the flight activity period during which female flies can infest the fruit. This may continue forty to sixty days after emergence. 7 In another set of experiments, these same emergence cages were utilized, but they were placed over seeded ground (Herrick, 1912; Allen and Fluke, 1933; Dean, 1942; Lathrop and Burks, 1945; Glass, 1960; Dean and Chapman, 1973). The previous fall, apples infested with apple maggot larvae were collected, and placed on soil where no pupae were previously present. The location was accurately marked, and the cages were placed over this spot the fol- lowing summer. As before, soil was mounded up on the outside of the base of the cage so no flies could escape. This type of emergence provides somewhat unrealistic data because of a high pOpulation of pupae consolidated into a small space, but is useful for answering certain types of questions. Within the past twenty years, visual sticky traps that mimic the foliage have been developed to monitor fly activity in the tree (Still, 1960; Oatman, 1964a; Maxwell, 1968b; Prokopy, 1968a; Moore, 1969; Kring, 1970; Prokopy, 1972a; Buriff, 1973; Trottier, Rivard and Neilsen, 1975; Reissig, 1975; Reissig, 1977). The behavior of the fly that allows this technique to work is that during the pre-oviposition period the flies are searching for potential food sources. Leaves exude substances that the fly feeds on, and supports populations of leafhoppers and aphids that excrete honeydew, another fly food source. Therefore, a surface that resembles a leaf becomes attractive to the fly during this pre-oviposition period when feeding has top priority. Several researchers have tested colors, sizes, 8 shapes, and volatile substances to be added to the trap to determine which combination provides the greatest trap catch (Prokopy, 1968a; Reissig, 1975). Prokopy (1968a) showed the best color was Saturn yellow which reflects a supernormal amount of energy in the 580 mm range. This is the wave length range reflected by green leaves, and is detected by flies as a leaf. A trap size and shape experiment lead to the discovery that the best trap has a rectangle of 8 x 12 cm (Prokopy, 1972a). Odoriferous substances added into the stickum enhance catch, and protein hydrolysate plus ammonium sterrate (Howitt and Connor, 1965) apparently give the best result. These features have been combined into a standard yellow trap that is commercially available as the Yellow Zoecon AM trap (Zoecon, 1980). There are two advantages to using this type of trap. First, many hours of labor involved in seeding cages in the fall, building and repairing cages, setting cages in place in summer, and taking them down in the fall are saved. More importantly, the activity period of the fly in the tree can be more accurately estimated. A fourth monitoring technique utilized was a fruit mimic (Prokopy, 1967b; Prokopy, 1968b; Moore, 1969; Kring, 1970; Prokopy, 1973; Prokopy, 1977). Prokopy (1977) re- ported that an 8 cm red sphere provided a much more accurate indication of fly activity in the tree. His criteria included sooner first catch, more flies, and better predic- tion of the amount of fruit damage. An assumed advantage of these traps is that they trap female flies when they are gravid and ready to oviposite. A catch would indicate that controls should be applied immediately. Research Sites - In 1977, two research orchards were identified for studying the apple maggot. The K.S.H. (K.S.H.) orchard located on the western edge of Kalamazoo, Michigan, was under the supervision of Dr. A. J. Howitt of the Department of Entomology at Michigan State University, and could be used for the duration of this study. The orchard is about 40 years old, and has a mixed planting of approximately 40 acres that includes 15 varieties (Figure 2). It has a minimum level of pesticide applica- tions each year, with usually only a single massive dose of Difolitan(Single Application Technique) applied to control primary apple scab. This maintains foliage for tree growth, but has minimal effect on apple maggot populations. There is a high natural level of apple maggot flies in the orchard, with nearly 100% fruit infestation every year. The second research orchard is located on the Upjohn Chemical Company farm located northeast of Kalamazoo, Michigan. This orchard was used for the duration of the study under a cooperative agreement between Dr. Howitt and the Upjohn Company. The orchard is about 40 years old, has twelve acres, and is mainly composed of McIntosh, Jonathan, and Northern Spy varieties, but also has a row of Snow apples, and two rows of sweet cherries (Figure 3). This site is used periodically for chemical evaluation of single tree replicated designs. Therefore, the resident population of apple maggot flies is under some chemical pressure and is 10 0000000 J 388 DMDSG D JDDDD D J 8668 SSDDDDCCCD C DOC BSDSDDDJ J D DDJ J 8800000 8 SDUTJ JJSTJ J SSSTSSSSSMJDDCCDDD J J J W88 SSDGGMSD CDCCJ S S J 308800 8 GBNDS J 00000 888860 J BBNJ JDDJCDOM J 06680 JGGBGG J JBD J JBNSSMB 83MBJBGGGGSGG J 8 DDJ J J 8663 J J o s :3 s. s JGMB DUTJDUTW J SSSDUTDUT 40x40 Traa Coda m... .. mamamww ...memvww Mum m... .. “mammwm u N Z aflmld ..M ..- ALF--- s.flGSDGTI“ WHSMWGMT. .3 MT.“ J_MmMGJM_ r- l l I l | I l l I I. Kalamazoo Stat. Hoatpltal Appla Orchard Map of the Kalamazoo State Hospital Orchard Showing Study Areas and Variety Composition. Figure 2. 11 Sw Sw W M M J Sn S S S S M M W' M M J J :m: S S S S M M Sw W M M J J Sn S S S M M Sw W M M J J Sn S S S S M M Sw W M J J Sw Sn S S S S M M W’ M M J .1 8w Sn Si 8 S S M M W M M J J Sw Sn S S S S M W M M J J Sw Sn S S S S M Sw W M M J Sw Sn S S S S M M Sw W M M J J Sw Sn S S M 8w W M M J J Sw Sn S S S M W M M J J Sn S S J M Sw W M M J J Sw Sn S S S M M .. w Fan‘s-a‘sn‘s‘rs‘m": Sw w :M M J J 8n 3 s s s M M: SleM M J 13an s s s s M Ml Sw W :M M J 8w Sn 8 S 8 S M MI w :M M J J Sn 3 s s s M M: W IM M J J Sn S S S S M MI w b-r-4.:_-.81_s_2_s..§-r_J Tree Code 40x40 J-Jonathan Sn-Snow M-Mctntoah Sw-Swoot Charry S-Spy W-Woalthy Upjohn Apple Orchard Figure 3. Map of the Upjohn Orchard Showing Study Areas and Variety Composition. 12 not nearly as high as the K.S.H. orchard. However, flies are present and active in the orchard every year, and are present at much higher levels than that experienced by commercial growers. This medium density population might provide data that serves as an important link between high and extremely low population levels of apple maggot. In 1979 and 1980 commercial orchards were also moni- tored. Appendix 2 gives information pertinent about each block studied. Included is the owner's name and location, the varieties and number of traps in the block. Also included is a rating of potential pest pressure to the orchard. After selecting the block to be monitored, a thorough investigation of the surroundings of each orchard was made. Looked for was both the number and distance of abandoned apple trees or other host plants for apple maggot. A rating of pressure to the commercial orchard was made that is found in Table 1. These values for each orchard were determined and are found in the last column of Appendix 2. Flies caught in the commercial orchards were not used in emergence studies, but were used to study factors important to timing for controls. Table 1. Rating System of Potential Apple Maggot Fly Pressure to Commercial Orchards. Rating Criteria 0 No abandoned trees or hawthorns evident 1 One tree in excess of 100 meters 2 One tree adjacent to the orchard 3 Two or more trees in excess of 100 meters 4 Two to ten trees adjacent to the orchard 5 Many host trees adjacent to the orchard 13 Around each orchard abandoned apple trees were located that were assumed to have high populations of apple maggot flies that could disperse into the commercial orchards. Traps were hung in these trees, and flight activity moni- tored. Several factors related to timing for controls were studied, one of which was the normal activity in abandoned tree sites. These trees were monitored at each visit to the orchard in 1979 and 1980. Two additional sites were monitored in 1980 to serve as a validation of predictions based on results obtained from the K...SH and Upjohn studies. Site one had early red variety apples located in the Hofacker yard near the Fruit Ridge Avenue and Four Mile Road intersection northwest of Grand Rapids, Michigan. Site two was a transparent variety tree in the Yabs backyard in DeWitt, Michigan. Site one was an extremely early site due to the sandy soil and mowed yard which allowed for rapid development of the pupae. High populations of flies were present and trapped, so data gathered there should be sound and prove adequate for vali- dation purposes. At site two very few flies were caught, so that data gathered there will not be used for validation purposes. Emergence and Flight Studies - In 1977, 50 yellow Zoecon AM traps were placed in the K.S.H. orchard. Twen- ty-five of these were in early variety trees on the south side of the orchard, and twenty-five were clustered in later maturing varieties on the north side of the orchard. Traps were hung one per tree, and positioned one-third the dis- 14 tance into the tree canopy on the south side of the tree at eye level according to Prokopy (1972a), Reissig (1975) and Neilson et. al. (1976). Apple maggot flies were counted weekly and the traps cleaned off. At two week intervals the old traps were replaced with new ones as per manufacturers recommendations. Monitoring was initiated on June 17 and terminated September 28. Also that year, 15 traps were placed in the Upjohn orchard. Five trees each of McIntosh, Jonathon, and North- ern Spy were monitored. Traps were positioned, checked and replaced as in the K.S.H. orchard. Monitoring was initiated on June 17 and terminated September 28. Emergence cages were placed in the K.S.H. orchard in 1977. Ten cages were placed over natural populations in the early variety section and ten in the late variety section. Five trees were selected in each section that had comparable canopies. Under each tree, one cage was positioned on the south side and one on the north side of the tree. This was done to provide a mean emergence value per tree, per variety, per orchard, and to determine if differences existed in emergence times between the south and north sides of the trees. Cages were monitored weekly, and the flies counted and removed. In 1978, trapping and cage studies were repeated in the K.S.H. orchard. The number and placement of traps and cages was identical to that of the previous year. The only dif- ference in 1978 was that monitoring was initiated on June 23, performed daily for three weeks, twice a week for 15 the next four weeks, and once a week for the last six weeks. Sampling was terminated on September 22. In 1979 the number of sampling locations were greatly expanded. In the K.S.H. orchard, the total number of traps trees was reduced to 15, seven in the early variety section and eight in the late variety section. Each tree was used in a paired test. On one side of the tree was hung the standard yellow Zoecon AM trap. On the opposite side was hung a red sticky sphere. At weekly intervals their po- sitions were reversed. The yellow traps were replaced at two week intervals. Sixteen cages were placed over natural populations of apple maggot flies, eight in the early section and eight in the late section. A cage was placed under the south and north side of each of four trees in each section. Sampling was initiated on June 20 and performed three times a week for four weeks, twice a week for nine weeks, and once a week for five more weeks. It terminated on October 13. » At the Upjohn orchard in 1979, paired tests were also conducted. Three trees of each of three varieties utilized in 1977 were monitored with the yellow trap and red sphere. Traps were checked, reversed, replaced, and taken down the same as in the K.S.H. orchard in 1979. Trees selected were check trees so the effects of experimental insecticides would be reduced on trap catch. Twenty commercial orchards were also monitored in 1979 (Appendix 2). Each orchard had a trap density of 1, 2, 4, or 10 traps per 10 acres. In each trap tree paired compari- l6 sons were made as in the K.S.H. orchard with trap placement, reversal, replacement and removal identical. These orchards were set up on June 19, and monitored three times each week for three weeks, twice a week for the next four weeks, and once a week for the next five weeks. Around each of these orchards were located abandoned trees. Thirteen trees in total were monitored on the same schedule as the adjacent commercial orchard. Each trap tree provided data for paired comparisons, with the methodology identical to previously described. In 1980, monitoring schemes were very similar to 1979. In the K.S.H. orchard, the same trees and methods were used as in 1979 for the trap comparison studies. Cage studies were conducted in 1980, but these were seeded with infested apples in the late summer of 1979 rather than being placed over naturally infested soil. Seven cages in total were seeded on 3-4 day intervals, with the first seeding made on July 16 and the last August 6. They were monitored three times a week for six weeks beginning June 23 and then weekly for eight weeks. In the Upjohn orchard in 1980, the same trapping scheme was utilized as in 1979. Nine trees, three of the main varieties, had a yellow trap and red sphere on them for comparison purposes. Trees selected were check trees in the chemical tests so as to reduce insecticide influences on flight activity and trap catch. Monitoring intervals were the same as those in the K.S.H. orchard in 1980. 17 Different commercial orchards were monitored in 1980 than 1979. Those selected (Appendix 2) were closer to Lansing to reduce travel expenses. Five orchards were monitored three times a week beginning June 23 for nine weeks, then once a week for three weeks. Pest Management Field Assistants in other parts of the state monitored another six commercial orchards on the same schedule. Each orchard had a different density of traps, but each trap tree had a comparative test between the yellow and red sphere traps. In the area around each orchard abandoned trees were located, trapped and monitored identically as the commercial trees. Two new sites were established as validation points in 1980. As previously mentioned, the Yabs site was eliminated because of the limited number of flies caught. At the Hofacker site, one tree of four in the yard was monitored. A yellow trap and red sphere trap were hung on opposite sides of the tree. Positions were reversed weekly, and the . yellow trap replaced every two weeks. Flies were counted and removed starting on June 23 three times each week for six weeks, and then weekly for eight weeks. Results and Discussions Season long emergence and flight patterns of apple maggot fly have been determined for different locations and years in Michigan. Table 2 was prepared to show when on'a calendar basis, different proportions of emergence or trap catch occurred. This compares different years, different 18 locations, and different methods of monitoring. The de- tailed discussion of the average and range associated with each cell in this Table will be delayed until later in this dissertation when parameters associated with these variances are studied. The general purpose here is to show the large variability associated with each event. This is indicated in the last row. The one method of monitoring fly activity that remained the same through all locations and years was trap catch on the yellow Zoecon AM trap. Because of this consistency, season long emergence graphs were prepared (Figures 4-13). These graphs show weekly average trap catch and the accumulative percent emergence. 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When emergence occurs or flies are caught on traps, growers are advised to start pesticide applications in 7-10 days, and continue them at two week intervals until harvest. Because of its importance, studies. were conducted in high and medium density situations to better understand the dynamics associated with first emer- gence. Other studies in low density levels as is exper- ienced in most commercial orchard situations were carried out to determine if those same dynamics are applicable with pesticide pressured populations. A portion of this study contrasts methods of measuring or estimating emergence. Several methodologies have evolved in the past to measure this event. The first method was to place emergence cages over naturally infested ground and monitor emergence. Another was to seed cages in the fall, and emergence monitored the following summer. Later traps were designed to measure populations that are actually present and active in trees. Yellow traps were designed that mimicked foliage and red spheres were used that mimicked fruit. A study of these methods will measure the 32 variability associated with each method, and the result should be to determine which method is best. Best can be defined as that method which catches flies consistently earlier and has the smallest coefficient of variation (C.V.) of first catch, is the most practial to use, and is a better indicator of fly activity in the tree. The C.V. is calcu- lated by dividing the standard deviation by the mean. Materials and Methods Studies were conducted in the K.S.H. and Upjohn or- chards during the years 1977-1980. First emergence was determined or estimated by one or more of four different methods. In all of these cases, the frequency of monitoring is the same as that discussed under Adult Emergence. Results and Discussion Table 3 presents the data on emergence or first catch by each method for all years at the two locations. Included is the range of dates in first catch, and the mean day of first catch for each method. The analysis of the data from which Table 3 was formulated proved to be somewhat untidy. These data were taken for a variety of individual small tests, and when combined a posteriori fit no experimental design. Too 33 TABLE 3. First Emergence or Catch of Apple Maggot Flies in K.S.H. and Upjohn orchards. Mean Date of First Catch Range in Date Year Location Method No of First Catch 1977 K.S.H. Natural Cages 20 07/06 - 07/20 07/11 K.S.H. Yellow Traps 50 06/20 - 07/06 06/21 Upjohn Yellow Traps 15 06/20 - 07/06 06/28 1978 K.S.H. Natural Cages 20 06/27 - 07/11 07/01 K.S.H. Yellow Traps 50 06/27 - 07/12 07/03 1979 K.S.H. Seeded Cages 8 06/17 - 07/04 06/21 K.S.H. Natural Cages 16 06/25 - 07/13 07/01 K.S.H. Yellow Traps 15 06/25 - 07/06 06/29 K.S.H. Red Spheres 15 07/02 - 08/02 07/14 Upjohn Natural Cages 3 07/11 - Never 07/11 Upjohn Yellow Traps 9 07/13 - 08/13 07/24 Upjohn Red Spheres 9 07/16 - 08/13 07/27 1980 K.S.H. Seeded Cages 7 06/28 - 07/02 06/30 K.S.H. Yellow Traps 15 06/28 - 07/09 07/04 K.S.H. Red Spheres 15 06/28 - 07/21 07/10 Upjohn Yellow Traps 9 07/05 - 07/28 07/21 Upjohn Red Spheres 9 07/14 - 08/25 07/25 34 many treatments were missing to have a nested, factorial, or two way analysis of variance design. Therefore, one way analysis of variance tests were performed with each of the methods at the different locations in different years being considered a treatment. The resultant 17 treatment one way ANOVA with unequal sample sizes was appropriate. Three sets of these data were removed from the analy- sis. The 1979 Upjohn natural cage was eliminated because of the extremely small number of flies caught (3), and the few samples units used. The 1977 yellow trap data from the Upjohn orchard was also eliminated. This orchard had been abandoned for several years, and the mean per trap catch for the year was 756. Starting in 1978, insecticide testing was conducted in this orchard, with alternating trees being treated which resulted in a 95% reduction in the population. In 1979 the mean trap catch was 30 and in 1980 it was 34. This greatly differing population within the orchard would cause an extremely large variation in all the data collected. Therefore the 1977 data was not used, and the 1979-1980 data will be used to represent a medium density orchard. Lastly, the 1977 natural cage data from the K.S.H. was eliminated. The rationale for this is that in 1977 eight cages were set on June 29. They were not checked again for one week, and six of them had flies in them. On that day the remaining 12 cages were set, and four of them had flies the following week. All the yellow traps in the orchard had caught flies on them before this date which indicates that 35 the first flies emerged and escaped the cages before most of them were in position. Also, it is highly probable that flies emerged throughout the week, and if recorded on the day of actual emergence, the mean date of emergence would have been earlier. This sampling error was corrected in succeeding years by setting all the emergence cages earlier and by sampling them more often the first two weeks of the season. The one-way ANOVA of the remaining 14 treatments proceeded. as :fiollows (BNPGANOVABAL, 1982). The first attempt was to compare all methods of measuring first catch, across all years and the two research orchards. Extremely high F ratios resulted from the ANOVA and SNK and LSD multiple range tests separated the 16 different treatments into four and six homogeneous groups at the .05 level. However, the test was invalidated because of the extremely heterogeneous variances, which is one of the basic assump- tions for the ANOVA. Thirteen different transformations were made on the data in an effort to meet the assumptions of the ANOVA, none of which were successful. This could be an expected result because of the very different methods of measuring emergence, year-to-year and location differences. The variances should be expected to be very large under these circumstances. The most reasonable approach to reduce this variability would be to analyse like subsets of the data. This was done and the Bartlett's test found the majority of the variances homogeneous. Those that were not were transformed by the 36 square root and/or log transformations to meet the as- sumptions. The SNK and LSD mean separation tests were run at the .05 level. In no cases were the means of the trans- formed data separated any differently than the raw data. The results are found in Table 4. This compares the influence of the year on the different methods of measuring or estimating first emergence at high population levels at the K.S.H. orchard and reduced population levels at the Upjohn orchard. Looking at the data from the K.S.H. orchard, emergence from seeded cages were tested in 1979 and 1980. Emergence was significantly later in 1980 than 1979. Two possible reasons for this could be that 1980 was a cooler year and the influence of variety of apple used in seeding. Both will be examined later. The high C.V. of 53 indicates that this is a highly variable data set, and not one from which predictions should be made. When natural cages were tested, emergence was not significantly different between 1978 and 1979. The influ- ence of temperature is discussed later. Fairly large number of flies were caught (1978 = 459 and 1979 = 791 which helps to stabilize the means. The C.V. of 30 indicates a relatively small variation, so the mean date of July 1 could be used as a good estimator of emergence r-f flies from natural cages in the K.S.H. orchard. 37 TABLE 4. Influence of Years on Different Methods of Mea- suring First Emergence of Apple Maggot Flies at the K.S.H. and Upjohn Orchards (Mean $S.E. Date of Emergence and Coefficient of Variation for Com- bined Data for All Years in Each Orchard). Seeded Natural Yellow Red Year Cages Cages Traps Spheres Kalamazoo State Hospital 1977 - - 6/21 $0.5a - 1978 - 7/01 $1.0a 7/03 $0.7 C - 1979 6/21 $1.9a 7/01 $1.6a 6/29 $1.1 b 7/14 $2.6a 1980 6/30 $0.5b - 7/04 $0.8 c 7/10 $1.7a All 6/25 $1.5 7/01 $0.9 6/28 $0.6 7/12 $1.6 C.V. 53 3O 48 31 Upjohn Orchard 1979 - - 7/24 $4.4d 7/27 $6.8b 1980 - - 7/21 $2.3d 7/25 $4.2b All - - 7/22 $2.4 7/26 $3.7 C.V. - - 27 35 Both Orchards All - - 7/01 $0.8 7/17 $1.9 C.V. - - 62 40 Dates in each column followed by the same letter are not significantly different at the .05 level by the SNK and LSD tests. Yellow traps were tested all four years at the K.S.H. and should provide a realistic appraisal of year to year variation. A large number of traps were set each year (1977 = 50, 1978 = 50, 1979 = 15, and 1980 = 15) and each trap caught a relatively large number of flies. The minimum number of flies caught on any single trap was 1977 = 20, 1978 = 32, 1979 = 137, and 1980 = 142. However, the mean number of flies caught per trap during this period was 1977 = 135, 1978 = 216, 1979 = 330, and 1980 = 248. At first glance this appears to be a competition effect, with the more traps used the fewer flies caught. However, each year 38 there was one trap per tree attempting to catch flies that emerged just under that tree. The difference in the total number of traps is the total number of trees with traps in them. A more plausible explanation for this difference is the varying resource. More or less apples were available for oviposition from one year to another, and that can greatly alter the population size the following year. As is shown the mean date of June 21, 1977 is the earliest year for trap catch, and it was significantly earlier than the other years. The mean emergence date of June 29 in 1979 was the second earliest and was significantly different than the other years. The mean dates of emergence in 1978 of July 3 and 1980 of July 4 were statistically not different, but different from other years. This difference, plus a C.V. of 48 shows that there is a variation component due to year in the first trap catch of apple maggot flies in the K.S.H. orchard. A possible explanation for this of warmer or cooler seasons is discussed later. The red sphere traps were tested in the K.S.H. orchard in 1979 and 1980. Their mean date of first catch were July 14 and July 10, respectively. These dates do not differ significantly. This is different than the other two methods of measuring emergence at the K.S.H. orchard for in both cases 1979 was significantly earlier than 1980. The pre- sumed reason for this must relate to the behavioral response of the flies to the spheres and the number of apples present competing for oviposition and mating sites. The C.V. of 31 is relatively small, indicating that the mean date of 39 July 12 would be a fairly good estimate of first trap catch on red spheres in the K.S.H. orchard. The bottom portion of Table 4 presents mean dates of first emergence or trap catch at the Upjohn orchard which has a much reduced population of apple maggot flies. When the yellow trap was used to estimate emergence, differences between years was not found. The mean date of first trap catch of July 24, 1979 and July 21, 1980, were statistically the same. The mean date for both years of July 22 had a small C.V. of 27 associated with it which indicates a fairly small variance and a good estimator. The red sphere traps had no statistical difference in the mean date of emergence between 1979 and 1980. It is reasoned therefore that when lower populations of apple maggot flies are present, fewer of the extremely early flies are present. When this is the case, the yearly variation is reduced (C.V. = 35), and much more consistent mean date of first trap catch occurs. A comparison between these two orchards is also pre- sented in Table 4. With yellow traps, the mean date of first catch is statistically later in the Upjohn orchard than the K.S.H. orchard. The delay ranges from 26 days in 1979, to 17 days in 1980. The associated overall C.V. is 62 which also shows this very large variation. Reasons for this delay such as variety component and weather will be discussed later. The major reason of population size explains the majority of this difference. In the K.S.H. orchard the mean per trap catch was: 1979 = 330, 1980 = 248, 40 while in the Upjohn orchard the mean per trap catch was 30 in 1979 and 34 in 1980. The greatly reduced population has fewer early individuals, and results in later mean date of first catch. Red sphere traps were also significantly later in the Upjohn orchard than in the K.S.H. orchard. The delay in mean first catch was 13 days in 1979 and 15 days in 1980. The C.V. was 40, a fairly large variation. The possible explanations of weather and variety composition will be discussed later. The population size differences mentioned in the previous paragraph can also explain the majority of the difference. The mean per trap catch in the K.S.H. orchard was 150 in 1979 and 109 in 1980, and in the Upjohn orchard 54 in 1979 and 19 in 1980. This smaller population size results in later first catch. Table 5 was prepared to determine differences between the methods of estimating emergence within each year. The seeded cage method was always the earliest. In 1979 and 1980 at the K.S.H. orchard, the mean dates of June 21 and June 30 were significantly earlier than any of the other methods. This was expected because of the concentration of a large number of flies into a very small area (1 meter square), and every one of the flies being caught (none escaping the cage). Cages placed over naturally infested ground are later in catching emerging flies than are the seeded cages, are the same as yellow traps, and are earlier than the red spheres. In the one year where both cage methods were compared, the mean date of emergence from the natural cage 41 of July 1 was 10 days later than the seeded cage, a signifi- cant delay. The reason for this is the smaller population size sampled, with the mean catch per seeded cage of 125 to that in natural cages of 49. When yellow traps are compared to natural cages, no real differences are evident. In 1978, the mean date of emergence was two days later on the yellow traps, but in 1979 it was two days earlier. Therefore according to this data, these methods are the same. The natural cages are sooner than the red spheres in mean emergence dates. In the R.S.H. orchard in 1979, the dif- ference of 13 days was statistically significant. Yellow Zoecon AM traps are intermediate in measuring the mean emergence dates. They are significantly later than the seeded cages as has been discussed. 'They are no dif- TABLE 5. Influence of Method of Estimating First Emergence of of Apple Maggot Flies at the K.S.H. and Upjohn Orchards Over a Four Year Period (Mean $S.E. Date of Emergence plus Coefficient of Variation within the Year). Method 1977 1978 1979 1980 K.S.H. Seeded Cages - - 6/21 $2.0a 6/30 $0.5a Natural Cages - 7/01 $0.7a 7/01 $1.6b - Yellow Traps 6/21 $0.5a 7/03 $1.0a 6/29 $0.9b 7/04 $0.8b Red Spheres - - 7/14 $2.6c 7/10 $1.7c UPJOHN Yellow Traps - - 7/24 $4.4d 7/21 $2.3d Red Spheres - - 7/27 $6.8d 7/25 $4.2d BOTH ORCHARDS All C.V. 45 23 63 41 Dates in each column followed by the same letter are not sig- nificantly different at the .05 level by the SNK and LSD tests. 42 ferent on the average in measuring first emergence than are the natural cages which is very important in orchard moni- toring. Their responsiveness in comparison to red spheres varies with the population of the flies but is always earlier. In the higher population levels of the K58.H. orchard, they caught flies 15 days earlier in 1979 and 6 days earlier in 1980. These differences were significant. In the medium density Upjohn orchard they caught flies 3 days earlier in 1979 and 4 days earlier in 1980 on the average. However, these differences were not significant. The last row of Table 5 lists the coefficient of variation within each year. This percentage value gives an indication of how variable the data was within all the methods of measuring emergence. Generally a 30 value indicates good biological data. In 1977, the C.V. was 45. This is quite large, especially when only one method was used to determine it. The 23 C.V. in 1978 is very small. This supports the non-significant difference of the two methods used in the K.S.H. that year. The 1979 C.V. of 63 is the largest one in the table. This is to be expected because all four methods are compared for first emergence, and a between orchard component adds variation to this value. Because it is so large, a mean value computed from all the data points would have a large variance and not be useful for predictive purposes. The C.V. in 1980 of 41 is also quite large. ‘Again this should be expected because of the several methods utilized and the between orchard varia- tion. 43 In paired comparisons with traps hung on opposite sides of the same tree, in 38 of 48 trials in both orchards in both years, the yellow trap caught flies first, 8 times they were later, and twice they were the same. Table 6 shows the results from the paired-t test which compared the difference between date of first catch on each trap in each tree (SPSS, 1975). In the Upjohn orchard in 1979, two red spheres never caught flies, so they were given values in this test equal to the last date of first catch on the remaining spheres of August 30. The results in the K.S.H. orchard shows that in 1979 and 1980 the yellow traps caught flies highly significantly earlier than the red spheres. However, in the Upjohn orchard where there is a lower density of flies present, the yellow trap caught flies sooner, but not significantly sooner. When all the trapping locations in the K.S.H. orchard from both years are lumped, the yellow traps caught flies 10 days sooner than the red sphere, a highly signifi- cant difference. This same trend holds in the Upjohn orchard where the yellow traps caught flies 8 days sooner. If all 48 trials are lumped from both orchards and both years, the yellow trap caught flies 9.4 days sooner, a highly significant difference. The fly generally is attracted to yellow traps first as a feeding stimulus. During the next 7-10 days the female is mating and maturing. After mature, she seeks out dark spherical objects on which to-oviposite. Then red sphere traps become attractive as a 44 .mumzmm owm mo c002 0059 0004 Sofiamw wo 0002 ”on 00053 000» ©0Hfi0u 000 :uHB ofiumflu000 ulpmufi0m ooo. o.o 0.0“ 00 0000 0.o0 o 00:0 0. 00000 00000 om. 0.. o.mn om >000 v.00 mm 0000 00 0:000: 000 ooo. o.o0 0.00 00 00:0 0.oH 0 00:0 om .0.0.0 000 00.. 4.. 0..“ mm 0.00 0.00 00 0000 o anoo00 ooo: ooo. 0.0 0.00 o0 >000 0.oH a 00:0 00 .0.0.0 ommfi o.o. 0.00 0.00 o .000 4..“ om 0000 a 000000 mood ooo. m.mfi 0.00 .0 0000 o.on om 0:00 00 .0.0.0 0.00 H®>wd .MMHD mumnnww Umm QMHB 3OHH$W .OZ COHHMUOA Hmmw .cwflm c002 0000 0002 000a 0002 000000 00000 00 0000 .m.00 00020 00000 000002 00000 00 0000000 umuah 0H 0000:00 xxOHum 00m 000 00009 20 coomoN Bodamw mo 000000QEOU .0 mqmme 45 fruit mimic, and catches females when they are gravid and ready to lay eggs. The 9.4 day mean difference between the two traps supports this behavior difference well. Conclusions First emergence of apple maggot flies was measured in two orchards by four different methods over a four year period. Great variability existed between the mean dates of emergence between the two sites. In the K.S.H. orchard a large population of flies existed, and the range of means of first emergence was June 21, 1977 to July 14, 1980, or 23 days. In the Upjohn orchard where a much reduced population was present, the range was from July 21 in 1980 to July 27 in 1979, or 6 days. The small range is because of the limited number of tests conducted in the Upjohn orchard and only over two years rather than four. Between orchard variation indicated that the Upjohn orchard caught flies from 25 to 13 days later in 1979 on the yellow and red traps respectively, and 17 to 15 days later in 1980 than the K.S.H. orchard. These were significant differences, and again shows that the smaller the population, the later the mean date of emergence. The four methods also varied in their estimates of timing of emergence. Seeded cages caught flies 8 to 23 days earlier in 1979 and 4 to 10 days earlier in 1980 than any of the other methods. This was significantly earlier, and if earliest emergence is required, this method should be used. Cages placed over naturally infested ground were inter- 46 mediate in their catch of flies. A lower total population is monitored by this method, which delays mean first emer- gence by not having a higher number of very early indi- viduals present. The yellow Zoecon.104 traps were also intermediate in catching flies. They were no different than the natural cages, but were significantly later than the seeded cages and significantly earlier than the red spheres. If first fly activity in the tree is to be measured, this is the best method to use. The red spheres were significantly later in mean first catch than all the other methods. The 9.4 day mean difference between the yellow trap and red sphere matches very well with the duration of the pre-oviposition reported in the literature. 47 PARAMETERS AFFECTING FIRST EMERGENCE The emergence of apple maggot has long been known to vary considerably and was shown to be true earlier in this dissertation. On a population level, there must be some important underlying basic factors that explain this lengthy emergence period. Figure 14 was prepared to conceptualize the parameters associated with adult emergence. Data on each component can be generated or obtained from the literature. Once quantified, they can be incorporated into a phenological model. Factors associated with emergence have been compartmentalized as either abiotic or biotic. The abiotic factors of air temperature, soil temperature, soil type, soil mositure and rainfall should explain the majority of the variance associated with emergence based on calendar days. The biotic factors of variety reared in, carry over pupae to second year, orchard floor culture, second generation within one year, and location of pupation in the soil, should provide minor refinements to the predictive model. Experiments were initiated to identify and quantify these parameters. Both abiotic and biotic components were investigated. Abiotically, air temperature, soil tempera- ture and soil moisture were the key components investigated Other presumably less important parameters noted were orchard floor culture and soil type. It was assumed the biotic components mentioned in the literature are much 48 more important in the total range in emergence, so they were investigated and their component of the variation of emergence was quantified. The more important parameters measured included: variety the fly was reared in, effect on timing of emergence by pupae that carried over to the second year, orchard floor culture, influence of the proportion of the population that emerged early to complete a second generation in one year, and location of larval pupation in the soil. .0000wu0am 0050002 on 00::uccome w00000Hcoz 0:0 mam uomm0z 0a00< 0nu mo 0oc0w008m maauo0wm< muouo0m .qa 0uswfim 00:0 000000 MNDOHZmUm& DZHNQBHZQE mac 00009 0Haumao> 00009 uw0m 06.50: 00:00 DOHmmm ZOHHHmOmH>OImMm A 49 .om 00:0 H050H> uo0uan _ mafia“: 00u0om _ 00w0u 0w0aaom mxoaum _ 0oa0wu0am 9403mmmm9 Amo 6H nuamn|l— . :ofiu0u0s0u — _ 000000 _ . 0HSDH30 — uooam 0005000 mszmzo0zoo _ wmasm . u0>o 00000 UHEOHm a >00Hu0> — A _0u:u0u00809 ua< _ mozmommzm _ HADQ< _0u:u0n00808 Haem— 0000 H.o0 Q 0u=umuoz HHom MHZMZOQIOU UHBOHQV — Ha0waa0m —_—--—_—-- 5 O ABI OTIC COMPONENTS Air Temperature Earlier in this dissertation, the emergence of apple maggot over several years and in several locations was presented. On a calendar basis, various points along the emergence curve vary by as much as 81 days (Table 2). The abiotic parameter of air temperature converted to physio- logical growth can explain some of this variation. This process involves the calculation of degree days. The prin- ciple involved states that above some threshold temperature, an organism develops physiologically. As the temperature becomes warmer, it grows more rapidly. Therefore, based on some lower developmental threshold, the apple maggot pupae develop and molt into adults. By knowing the temperature at which development starts, measurements of physiological growth can be estimated by accumulating thermal units. Predictions of future events then can be made by recording maximum and minimum temperatures and calculating thermal units each day, and accumulating them through time. The common method of calculating degree days is, daily maximum + daily minimum - base temperature. This 2 method was improved (Baskerville and Emin, 196?) to create a sine wave curve through the maximum and minimum points, and integrating the area under this curve above the threshold. 51 This refinement gives more precise estimates of the daily physiological growth that occurs. The lower developmental threshold (LDT) of the apple maggot has been determined by three separate research groups. Reid and Laing (1976) determined the LDT as 8.7°C (47.7°F). Trottier (1975) said the LDT was 9.0°C (48.2°F). Reissig, Barnard, Weires, Glass and Dean (1979) used his- torical emergence data and air temperatures, and found 6.4°C (43.5°F) gave the best correlation to the first emergence. The differences in these values may be due to genetics of each population studied, errors in calculating thresholds, errors in measuring the temperatures, or the inherent accuracy of each method. The methods reported by Reid and Laing (1976) and Trottier (1975) are the most accurate, and their average value of 8.9°C (48°F) will be used through- out this discussion. At higher temperatures, insects cease to develop. Physiologically they begin to carry out other body functions such as cooling, and do not continue to provide energy for growth. Reid and Laing (1976) have determined this upper developmental threshold (UDT) for apple maggot to be 31°C (88°F) and this will be used in this discussion. Using the LDT and UDT values as limits of physiological growth, one can make estimates of insect growth. This is accomplished by placing thermographs in the field to measure the temperatures in the general habitat where the insect is located. From these readings the degree day totals can be 52 calculated and accumulated to predict key events in the life cycle of the insect. One should note that this technique will give an estimate of a certain event. This estimate may be close and within an acceptible confidence interval, Inn: is seldom accurate. Variables that can influence and alter this prediction are both intrinsic and extrinsic to the organism. Within the population is great genetic variation that allows different individuals 1x) respond 1x) its environment differently. A population has spacial heterogenity, and is found in a variety of microhabitats within the environment. These microhabitats have different microclimates over the long run, and different daily variations. Temperature and humidity may be very different in these microhabitats, and this results in a differential growth rate per day. Lastly, the calibration and accuracy of the thermograph can easily vary just O.5°C which results in estimates that are not accurate. When these variable factors are combined, the resultant prediction based on gross weather records can be significantly different than the real event. Realizing the possible variables associated with the prediction of a biological event based on weather measure- ments is important. However, if they are ignored, predic- tions can still be made within certain realistic bounds that are more accurate than chronological predictions. Materials and Methods Air temperature measured at the five foot level in standard weather shelters is the most convenient and 53 universally accepted. method. of measuring temperature. Predictions based on this method would be most widely applicable becauSe there is a large network of weather 'stations that take this measurement every day. In order to obtain accurate local temperature measurements, three lead Weather Measure recording thermographs were placed in weather shelters in the K.S.H., Upjohn, Hofacker and Yabs orchards each year these orchards were used. The instru- ments were activated April 1, and turned off in September or October. Daily maximum and minimum temperatures were transcribed from the chart and entered into the computer where they were converted to degree days (48°F base) using the B.E. technique. Appendix 3 shows the accumulated degree day totals for each of these locations for the years they were utilized. Results and Discussion Air temperature degree day accumulations for different proportions of apple maggot fly emergence are found in Table 7. This table includes values over a four year period and at four locations which results in a very large range in degree day accumulations at each stage of emergence. The 1979 air dd values for the abandoned trees were taken from the Peach Ridge Agricultural Weather Station which was centralized for the abandoned traps. In 1980 the M.S.U. Hort. Farm Station weather was used as it was a centralized location for the abandoned traps in 1980. Mean values 54 calculated from these various locations would be of limited predictive value, because they represent very different population levels of flies, populations pressured with pesticides, and different methods of estimating emergence. To measure the influence of years on the different methods of measuring first emergence, Table 8 was prepared. The values presented are mean degree day values for first emergence. The data was analyzed by a one way analysis of variance, and transformed by the square root or log trans— formation if needed to meet the assumptions. Student Newman Kuell's mean separation test was then performed at the .05 level. Coefficient of variation values were calculated to determine how variable the data was. The mean date of emergence in seeded cages was signifi- cantly earlier in 1979 than 1980 at the K.S.H. orchard. On a calendar basis this difference was 9 days. The air degree day difference was 181, which corresponds to 8 calendar days at an average accumulation of 25 units per day. Therefore, air degree days was not very useful to explain this varia- tion between the two years. The coefficient of variation however was reduced from 53 to 12 which indicates that the overall variability within the data was greatly reduced, and that the mean value of 943 air dd should be a very reliable predictor of mean fi:st emergence. The mean date of emergence in natural cages was not significantly different in 1978 and 1979 at the K.S.H. orchard. This was also true in Table 4 which showed the 55 mmvm 1000 00aH OHNH Nm0H HmHN Hoam 000 000a HNmH HOHN v0oH mONH v0oH mNHN Hmmm mv0H mmam «HON 000a omHH 0HmH 0NOH mmvm 000m 0mmm Homa I000 000H omHH omma mmva 000a 000 omva mmVH HVNH mm0H I000 000a omHH ommH mmva vaa 000 mmva m0v~ QHNH ~00 omoH ovm 000a coca 000a 0mmH mama NNaH 0ooH ONNH mmma 0NOH 00HH 0m omma IvH0 oaoH 000 Hmmfl mmma mmHH NNO mafia mmva 000a 0H0 omoH mmm omma 00HH 0¢HH momH NOHH vmoa H00 moHH mmma 0mmH mmHH 05 mmmH I000 0HoH 000 Hmma NmNH QHOH NNO QNHH 000 0HOH v00 omoH mam 0omH 00HH 000H m0HH NoHH mmm fi0m H00 mmma 00HH mmaa 50000 00000 N00 0 H HH 0 ma H HH 0 ma 0 m 0H 05 0 ma 0H 0 ma om om 0N ma om 00000 0Hm50m .00 005052 Ham 00000 000000 000500 000 000500 000 000500 000 000500 000 0009 3OHH0» @009 3OHH0» @009 30aa0» @009 30HH00 00000 000000 00000 H000002 00000 H000002 000500 000 000500 000 000500 000 @009 300000 @009 30HH0» 0009 30aa00 00000 a00000z 000B 30aa00 00000 0000002 @009 30HH00 @008 BOHH0> 00:00: 0000 000000 00 00000 .0.0.x 00x00000 000000050 05000: .0.0.0 00500000 000000050 050000 .0.0.0 .0.m.x 050000 .0.m.x 000000050 050000 .mome 000000054 050000 .0.m.M .0:0m0& .0.m.x ..m.m.x 050000 .0.m.x 00000005 .0005 O0 0000 8000 00005002 00 000000004 000000> 00 000H0 000002 0HQQ< 0o 50000 @009 000 000000080 00 0000000000 000000000 000 000fi0 |0H08000< 500500: .0.0 00000 u 00000 000 000000 00000000209 0H4 H0>0A 0000 0>fim o0mH m0ma 00mH 0005 0000 .0 m4m<8 56 00mm I0N00 000m 0000 0000 0000 0000 0mmm 0000 0000 0000 0000 0000 NO0N 0000 0000 0000 0000 0000 0000 000m 000m 0000 00mm 00000 0000 00mm I0mm0 0000 0000 0000 0000 0000 0000 0000 000m 0000 0000 0000 000m 0000 0000 0000 0000 omom 0000 0000 0000 0000 0000 I0o00 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0mmm 000 0 0 00 0 0 00 0 m0 0 m 00 00 0 00 00 0 00 ON om ON 00 om 00000 000E00 00 005502 000 00000 000000 000500 000 000500 000 000500 00m 0009 300000 0000 300000 0009 30000» 0009 300000 00000 000000 00000 0000002 00000 0000002 000500 000 000500 000 000500 00m 0009 300000 0009 30000» 0009 300000 00000 0000002 0008 300000 00000 0000002 0009 30000» 0008 30000» mmmmmm 5.0.000. 0000 000000 00 00000 .=.0.M 00000000 000000050 050000 00000000 000000050 050000 .m.0.x OmomOM 050000 .:.0.0 000000050 050000 .m.0.x 000000050 050000 .m.0.M 0 C 000 mm MDhfi MM 3. m. m. 50 m. 00000000 0000 0000 0000 0000 000» .0 m0m0B 57 flies emergence on July 1 both years. Therefore, air degree days did not improve the estimate of mean date of first emergence from natural cages over that provided by calendar date. However, the coefficient of variation was reduced from 30 to 9, which indicates that the air degree day measurements greatly reduced the variation in the data, and the mean of 1050 should be a much better predictor of emergence than the July 1 calendar day estimate. Emergence as determined by yellow traps provides the best indicator of a possible relationship between the mean air degree day accumulation and mean calendar day of emer- gence because it was used in all locations in all years. Table 4 showed a 13 day range in mean date of first catch in the K.S.H. orchard from 1977 to 1980, and a 3 day range in the Upjohn orchard from 1979 to 1980. The respective air degree day range (Table 8) is 175 in the K.S.H. orchard and 89 in the Upjohn orchard. These values correspond to 7 and 3 calendar days respectively, so the variation is considerably reduced in the K.S.H. orchard, and the same in the Upjohn orchard. The reduced variation within the K.S.H. orchard is also supported by the C.V. values which were reduced from 48 to 9. The mean air degree day accumulation of 1117 associated with a C.V. of 9 should provide a very good estimator of mean first yellow trap catch in the K.S.H. orchard. In the Upjohn orchard there was a three day mean difference in catch by both the calendar and air degree day estimates. However, the estimate based on the air degree 58 .umma mzm man an Hm>ma mo. may no pcmumMMflo >Hucm0HchmHm #0: who Hmuuma mEom map >2 omSOHHOm GESHOU comm :« mammz aH ma .>.o m.mmn N.NHH mama moss cam: guom Hm 5H .>.o m.oefi m.emfl ooVH eemfl cam: cache: 0H m a NH .>.o H.mmn m.mH H.mHH N.mmfi mama wads omos m.mvm cam: .m.m.x H.mm“ H.mmfi mmmmfi emfiefl ommfi nachos m.HmHH m.maw «mama comma mums :soflmo e.mVH n.5HH o.ew manna onmmHH ammoa cams .m.m.x o.mow e.qHH o.omn «.mMH «mama Mama memos mmmm memfi .m.m.m v.esw m.HNH ammofi mmoofl mead .m.m.x v.fififi omoHH nemfi .m.m.x mmhmflmm mmmth. mmmmv mmmmU Ham? COflfiMOOA Umm 3OHHmM. HGHDHMZ Ummumwmwm ..mmsam> :Oflumwum> wo ucofloflwmooo moam .oonumz .m.m .momv mmmm um Hm>mq uoom m>Hm um coflumasadood xmo mmummo uwfl .m.m H :mmzv .moumnouo csohmb paw .m.m.x on» no mmflam uommmz mamm< mo mocmmuoEm umuflm mcflusmmmz mo moonumz ucoummmeo co whom» mo mocmsHmcH .m mgm.o . Ada spam v.o~HH m.HmHH ummms ummmfi mmumnmm 66m snows: H.mmfl m.maH canes comma manna zoaams anoflm: n.mefi m.~ofi neemfi comma mmumnmm com .m.m.x n.5HH e.¢HH e.vHH e.HHH mNmHH name names mmoflfi mane 3oaam» .m.m.x o.omw m.HNfl semen «woes memo Hmnsnmz .m.m.x o.eH v.mMH mmmofi ammm memo omommm .m.m.x ommfi mean when neafi venom: :oflumooq .ummu xzm may sh Hm>ma mo. mnu um acmnowwflo maucmofiwflcmflm poo mum umuuma mEmm msu wn om3oHH0m CESHOO comm ca mammz asuaH .m mamas 62 method than calendar days because of the total reduced variation. In 1979, the same trends Were evident with estimates made by calendar days and air degree day accumulations. The seeded cage mean emergence was 8 days earlier than the YSllOW' trapl in the K.S.H. orchard. The degree day difference was 142, which corresponds to 6 calendar days. The yellow trap mean first catch was two days earlier than the natural cages (Table 4). The air degree day difference of 43 whiCh corresponds to 2 days was the same (Table 9). The red sphere was the last tool to catch apple maggot flies in K.S.H. orchard in 1979. The mean date was 13 days later than the natural cage on a calendar basis, and 259 degree days later by air degree days which corresponds to 11 days. Therefore, air temperature degree day values reduced the variability again. Zhi the Upjohn orchard, the red trap caught flies 3 days later than the yellow trap, but this was not a significant difference. The corresponding degree day difference was 62 or 3 calendar days, and was not significantly different. When all the data for 1979 was taken as a whole, the C.V. was 63 for the calendar date data. This is a very large variance, and no estimates based on the overall data should be made. If first emergence is based on air degree days, the C.V. is reduced to 23. Therefore, air degree days greatly reduces the variation within the 1979 data set. 'm '.~ v. ' 63 The 1980 year provided the same trends as the other years. The seeded cage had the first mean emergence in the K.S.H. orchard, followed by the yellow trap four days later which was a significant difference. On a degree day basis, this difference was 99, which corresponds to four days, but it was not significantly different by the SNK test. The red sphere mean catch was 6 days after the yellow traps, and 145 degree days later. This corresponds to six calendar days, and was significantly different for both the calendar days and degree days, so the degree day estimate was not any better than the calendar day difference. In the Upjohn orchard, the red sphere mean date of first catch was four days after the yellow trap. The air degree day difference was 104, which is five calendar days. Neither method was significantly different from each other, therefore the degree day estimate was not any better than the calendar day estimate. However, by examining the C.V., the calendar day value is 41 and the air degree day value is only 18. Even though degree days did not alter the significant differences any, they greatly reduced the variability of the data. Conclusions The analysis of the air temperatures converted to air degree days showed that, in general, air degree days was a better estimator and had less variability in the data than existed between calendar day estimates. With the seeded cages, air degree days removed 1 calendar day of the 9 day 64 variation present between the two years. When natural emergence cages were used, the air degree days difference was the same as the calendar day difference, but they did reduce the variation in this method of measuring first emergence. Yellow traps were used to estimate first emer- gence for all years in both orchards. Air degree days did reduce the variance greatly within the orchard and between orchards. In the K.S.H. orchard, the 13 day variation is reduced to 7 days and in the Upjohn orchard the 26 day variation in calendar days was reduced to 5 days. The 17 day average difference between the orchards was reduced to 10 days. Also, the C.V. showed that the data was much less variable. The red sphere mean first trap catch of 4 days was reduced to 1 day in the K.S.H. orchard, but increased from 2 to 5 days when air degree day values were converted to their corresponding calendar day equivalents. Air degree days also reduced by 9 and 5 days the difference between orchards. Therefore, degree days greatly reduced the variability within the data and provides a better base for predictive purposes. When different methods of estimating emergence were compared within each year, degree day accumulations did reduce the variation present in calendar day differences. In 1977, air degree day values reduced the variation in values associated with each method. In 1978, the two calendar day difference was not improved by air degree accumulations, but did reduce the variability of the data. 65 In 1979, the range in values was reduced by two days when air degree day values were used. The 8 day difference between seeded cages and yellow traps was reduced to 6 days, and the 13 day difference between yellow traps and red spheres was reduced to 11 days. The total variability in the 1979 data was also reduced by using degree days. In 1980, the air degree day accumulations did not alter the significant differences, but greatly reduced the variation in estimates over those based on calendar days. 66 SOIL TEMPERATURE As was shown in the last section, the mean day of first emergence based on air degree days was quite variable even though it improved the estimate of emergence over calendar day estimates. One modification that might reduce this variability further and increase the precision of estimating first emergence would be to measure soil temperatures. This approach is reasonable because the pupae are in the soil and develop at a rate relative to the ambient temperature around them. Maximum and minimum temperatures converted to degree days by the B.E. method would provide a method to measure the amount of physiological development that occurs, and might provide an estimate of emergence with less varia- bility. Literature Researchers have used soil parameters to make predic- tions on events of life stages for other insects, but this has not been extensively done for apple mattot. Maxwell and Parsons (1969) said that "the times of first and mean peak emergence of apple maggot may be closely determined for any one site by summing soil temperatures at pupae depth follow- ing a preliminary season's record of soil temperature summations and emergence data. Dean and Chapman (1973) noted that first emergence was very variable and difficult to predict using air temperatures. They felt these predic- tions could be improved if soil factors were studied and correlated to emergence, because that is where the apple maggot is developing. This has been done with Rhagoletis cerasi (Leski, 1963) with great accuracy. 67 Materials and Methods Weather measure three-lead recording thermographs were placed in the K.S.H. and Upjohn orchard from 1977 to 1980. One of the leads was placed in the soil at a depth of two inches on the south side of the tree under its canopy. Maximum and minimum temperatures were transcribed from the charts, and entered into the computer where degree day values were calculated at base 48°F (8.9°C) by the B.E. technique. These values were accumulated throughout the season (Appendix 3). The sampling schemes for monitoring first emergence of apple maggot fly were the same as described earlier. The degree day accumulations on the day of first emergence was recorded for each method at each site. This data was analyzed by the one-way analysis of variance to test for mean differences between methods or years. If the variances were not homogeneous by the Bartletts test, data was transformed by the square root or log transformation and rechecked for homogenity. If significant F statistics were present, Student Newman Kuell's mean separation test was performed. Results and Discussion Ex.mination of the soil degree day data reveals some trends, and also some exceptions to them. Table 10 was prepared to illustrate these points. It presents for each month the accumulation of the soil degree days and the 68 on new oopmHsEDoom mo we op HeOm umsu unmoumm u w H e.mmv ommm Hme eeem wee move emq meme Haw sauna» m.mm omq 40H One mm eHm om Ham mm “whamummm m.mm moo mm eem med Nmm mm mmv mm umsmsm N.om «mm me mmm om «we em mew me ease o.me mam we awe mm eve me oem we maze o.mo mmfi me Hwfi mo emm No mam an em: m.mH OH NH mm em m an mm «H Henna amazomo zmoea: m.mHm qmmm «om. omom Ham meme mom comm mme manmme m.emH mme mMH mam mmfi eee mMH «Hm mHH “wasmummm m.moH How moH ewe eHH ome 0H2 owe Nod umsmsa ~.qm mme em ome em mew OOH .moe em ease o.me wee mm wee mm HHV we can ee mean m.me mam mm 4mm mm owe om eon me em: m.oe He mm moH we am me me we Heum< on sea mo 68 w 68 m we w 66 He sumo: w com: omen memfi mean eeme Omdmomo QBHmmOm WBHucmoemacmam uoc mam amuuma mEmm mcp wc .ummu xzm may en am>ma mo. mmzoaH0m CEsHoo comm me mammz em mm .>.o «.mma m.maa mVOa Nee cam: aaom mm ma .>.o a.moa m.mea maaa emoa cam: accede mm ea ma oa .>.o m.ava N.oaa a.oea v.eaa NaOa mew aee m.mmo cam: .m.m.e m.mma o.mva smeaa oOaOa omma ceoema ~.amaa e.mma nemaa mmmoa mema caoea: o.ova a.eaa e.ma anew umee memo omma .m.m.e «.mma a.maa e.mea e.mma namaa cove ammm meae mema .m.m.e v.Maa m.aaa naoe moem mema .:.m.x N.ma comm eema .m.m.x mwhwfimw WNMHB mwmaMU mwmmo H00? EOHHmOOQ Umm 3OHHTN Hmufiflmz flameuom ..mocumz .m.m «momvummmm um mmae mcu mo moam cusom mcu co am>mq cocH 039 mcu um macaumH585oo¢ wmo mmammo aaom .m.ma :mmz. moamcouo :coflda mam .m.m.x mcu um mmflam uommmz mamd< mo mocmmumam umuam mcausmmmz wo mmocumz ucmammwaa :o mummy Mo mocmzamcH .aa mamde 72 Also, the C.V. is reduced from 12 to 10 indicating that soil degree days reduced the variability of the data as was expected. Examining the data from the natural cage mean emergence reveals some problems. The data in Table 11 says that 1978 was a significantly earlier year for emergence than was 1979. On a calendar basis, the mean emergence was on the same day in 1978 and 1979. Using air degree days as the estimator, the difference between the two years was 25 units which corresponds to one day, which provides a good estimator. However, the soil degree day difference of 215 units which corresponds to 9 calendar days. Also, the C.V. was increased from 9 to 19 units using air and soil degree days. This further supports the belief that one of these years, most likely 1979, was very different. The accumulations for 1979 would have to be reduced by 200 to provide a better estimator of emergence based on soil degree days. “ A discussion of the yellow trap data reveals some highly variable results also. In the K.S.H. orchard, the soil degree day mean first catch is more variable than that based on air degree days as based on the C.V. value that was increased from 9 to 17. On a calendar basis, there is a 13 day range in dates of mean catch. Air degree day accumulations reduced this to just 7 days. The soil degree day difference of 266 units corresponds to 11 days. If the K.S.H. 1979 accumulation was reduced by 200 as suggested, then the range would be reduced to 7 days and be just as 73 good an estimator as air degree days, but no better. In the Upjohn orchard the estimate based (n1 soil degree days increased the variability in the estimates slightly. The calendar day differences were 3 days. ‘This was the same using air degree days. The soil degree day range was 55 units which corresponds to 3 calendar days. Using this data, all three methods provide the same relative range in mean first catch. If the C.V. is examined, it is 27 for the calendar days, reduced to 17 using air degree days, and increased slightly to 19 using soil degree days. Based on this the soil degree day estimate is not better than that based on air degree days. The between orchard comparison reveals that soil degree days increase the variability within the data, but reduces the average daily difference. On a calendar basis, the dif- ference in mean date of emergence was 25 days in 1979 and 17 days in 1980. Using the same years, the air degree day difference converted to days reduced this to just 14 and 11 days. The soil degree day difference was 219 and 245 units or 9 and 10 days respectively. This reduced the difference even further. However, when the C.V. is examined, air degree days reduced it from 62 to 13, and the soil degree days increased it to 23. Therefore, the air degree day values should be used as predictors because of the less inherent variability of the data. Trap catch on red spheres was quite consistent within orchards between 1979 and 1980. In the K.S.H. orchard, the 74 calendar day difference of mean catch was 4 days. Air degree days reduced this to just 1 day difference. Neither of these values were statistically different. However, because of the greater accumulation of soil degree days in 1979, that year was significantly later than 1980 in mean emergence when using soil degree days as estimators. The 238 units corresponds to 10 calendar days, a much larger value than is present when using calendar day or a degree day estimator. The C.V. of 40 with calendar days is reduced to 19 using air degree days, but is increased to 24 using soil degree days. With the warmer 1979 season data used, the soil degree day estimate becomes a poorer estimator than air degree days for predicting mean trap catch on the red sphere in the K.S.H. orchard. In the Upjohn orchard, the soil degree day estimate of mean first catch on the red sphere is a better estimator than air degree day values. The calendar day difference of mean dates of first catch varied 2 days between 1979 and 1980. Air degree day differences of 131 increased this to 5 days. The soil degree day difference was 30 units, or slightly more than one day which indicates it is an estimator. The associated C.V. are 35, 21 and 25. Even though that associated with the soil degree days is slightly larger than that of air degree days, it is good enough to serve as a good predictor. The grand mean value of 1115 soil degree days provides an accurate estimate of mean catch of apple maggot flies on red spheres in the Upjohn orchard. 75 The between orchard variation is improved by using soil degree days. The calendar day difference between the orchards was 13 days in 1979 and 15 days in 1980, with K.S.H. always being earlier. The air degree day differences converted to days was 4 and 10. The soil degree day dif- ferences between orchards was 1 in 1979 and 209 which corresponds to 1 enui 9 days. This appears to reduce the variability between orchards. However, within the data sets ,the C.V. is 40 for calendar days, 19 for air degree days, and 24 for soil degree days. Therefore, even though the soil degree day converted to calendar day differences were the smallest, the air degree day data was less variable and their means should be used as predictors. Table 12 was prepared to determine whether soil degree days could reduce the total variability of the data within a calendar year. Presented are mean iS.E. values for mean emergence or trap catch. The C.V. was calculated to deter- mine the amount of variability in the data. In 1977, the C.V. was 10. This was much smaller than 45 provided by calendar days, but an increase over that provided by air degree days of 7. In 1978, the same trend existed with the C.V. reduced from 23 to 9 from calendar days to air degree days, and then increased to 13 by soil degree days. In 1979, the same trend was present with the C.V. going from 63 to 23 to 24. Lastly, in 1980 the trend remained the same with the values going from 41 to 18 to 23. Therefore, in no case did the soil degree days reduce the variability of the data less than was provided by air degree days. 76 .ummu mZm may en am>ma mo. mcu um ucmammwao waucmoamacmam uo: mam amuuma mEmm mcu >c omonHOM GEDHoo comm ca mammz mm em Ma ca .>.o u aa< em.~ma Neaa ow.emaa mmaa mamaam 66m em.maa caoa oe.mma mooa mane zoaam» oo.oea mam 6a.mma aMaa mameam omm no.eaa mme na.maa mam ma.maa aoe mm.ma omm mane zoaam» ne.mma mam ma.maa oem ammo amasumz me.ma ewe me.mma Nae memo ememmm omaa meaa meaa eema eonumz ..monumz .m.m emomvummmm um mmaa mcu mo mcam cusom mcu co am>mq coca 038 mcu um macaumasesoom hmo mmamma aaom .m.ma cmmzv UOaHmm amm» asom m am>o mmamcouo sconmo mcm .m.m.x mcu um mmaam pommmz madam mo mocmmameu umaam mcaumaaumm mo mocumz mo mocmsamcH euom coflumooq .NH mqmma mo. UN00. pmao. ono. onH. Qmm5v. cmomm. mwom. mvmm. cmmz 00.0 00.0 00.0 No.0 vm.0 m0.0 00.0 mH.H m 00.0 H0.0 No.0 5N.0 0m.0 5m.0 50.0 mm.0 E amcommom H0.0 00.0 0m.0 mm.0 H5.0 H5.0 00.H m0.H Z 00.0 00.0 No.0 NN.0 m0.0 H5.0 m0.H m0.H m mamaw cw wmo m50a>mam anm mmoq ucmamz. Emua m m :a Umcamucou mmHmEmm Haom who wamumamsou ou mammmmomz mafia mo cumcmq H0.0 No.0 m0.0 m~.0 50.0 05.0 MH.H mm.0 Z ccoflma 00.0 No.0 m0.0 mm.0 00.0 05.0 00.H 00.H Z ..mmaB mcu mo moan cuSOmum HO .umme mmcmm mamaaasz m .cmocno 00.0 00.0 00.0 00.0 NN.0 vm.0 M5.0 Hw.0 M .m. 00.0 00.0 00.0 00.0 0m.0 mm.0 M5.0 00.0 m. m. 00.0 00.0 00.0 00.0 ma.0 0m.0 m5.0 N0.0 .m .mHO0HEuE .cuuocuc um .ooom um mama cm>o mcaeaa 6 ca ama> mcp um ucmamwmao maucmoamacmam no: mam amuuma mEmm mcu zc cm3oHHOm mammz CU - :1 e4mrnma mo. 00 m0.5H 00 m0.0H 00 mH.0H .>.U Gmmz meomwom mcu um unmammman maucmoawacmam 0m m0.5m 0m 0m m0.00 0a am m0.00 mm .>.U :mmZ .>.U ccommb . 000a m0.00 m0.00 m0.00 cmmz m.m.x .umme mmcmm mamwuaaz m.:moc50 uo: mam amuuma mEmm mcu xc 0m3oHHow mammz 0N ma.0m 00 m0.00 50 mH.HN 00 m0.00 00 m0.00 50 m0.00 .>.O cmmz dew cmmz ceoeas .m.m.x 050a .A4>oz< wmzmcov mmuam ucmumwwao mchB :a mmmae mammfl amvca mCOaumoOA m30aum> um mmasumaoz H000 :mmz macq commmm eusom maceaz nuaoz .0H mqmflfi 90 Because the variances were homogeneous in the last tests all the data from each orchard were lumped to test for orchard differences. Table 17 shows that in 1979 there were no differences between the orchards. However, in 1980 the Hofacker site was significantly dryer than the other sites. This was expected as the trees were on top of a sandy knoll and the grass was mowed under them, both conditions favoring faster drying after a rain storm. The other two orchards have similar soil types and orchard floor conditions, which account for their similar soil moisture contents. The purpose of this experiment was to determine if soil moisture has an effect on seasonal emergence. An examina- tion of Figure 15 indicates a relationship probably exists. Therefore, accumulative soil moisture values were correlated with accumulative percent emergence (Table 18). When the correlation was made with seeded cage catch, an R2 value of .70 was found in 1979 and .71 in 1980 in the K.S.H. orchard. This indicates a good relationship for predictive purposes, and would be very good for most biological situations. However, it is not as high as found with other monitoring tools used. A correlation performed with the natural cage catch in the K.S.H. orchard in 1979 resulted in an R2 value of .55 which indicates a fairly good relationship. This correlation was also performed for the yellow traps. In the K.S.H. orchard, the 1979 value of .97 and the 1980 value of .96 indicates an extremely high relationship. Soil moisture 91 (ea...) Mag 18:] gonzo dBJl afieJeAv .mema ca camaoao amuammom mumum oonEmamM mcu ca mmmue 20 coomoN soaamw co mmaam uommmz mamm< 00 coumo mmaa adama m0mum>¢ 0cm manumaoz aaom ucmoamm mo GOmHummEoo .00 madman 5:03.. 22. a mm a. a on on c. a 0 Au N o o e .. . Toe m 0 . 9 Nu a. .0 .... . an 0.. mW 01. mn . . m. 5.. . ca 10. a m. . 9 a; . \1 OF L. LT O? H o o ( FF .r 0 NF .r 1.60 92 in this orchard can predict accumulative trap catch almost exactly. Trap catch on red spheres can be predicted just as well by soil moisture in the K.S.H. orchard as indicated by the very high R2 values of .96 in 1979 and .96 in 1980. These same trends were present in the Upjohn orchard where equally high R2 values were found. The yellow trap values of :79 511 1979 and .90 ix: 1980 were very high. However, in each case they were smaller than those found in the K.S.H. orchard. This difference can be explained by the significantly lower population level in the Upjohn orchard which creates a less smooth emergence curve. The R2 values of .84 and .89 for the red spheres were extremely high, but also less than in the K.S.H. orchard. At the Hofacker site, a very high correlation existed between accumulative soil moisture and accumulation percent trap catch. The yellow trap value was .80, and the red sphere value was .78. These R2 values are probably less than in the other orchards because of the very fast drying and significantly dryer soils, and the early emergence of the flies. Laboratory Studies - The results from this experiment were all negative when the original objective of the rela— tionships of first emergence was desired. After 76 days, zero flies had emerged. This equals 2220 degree days at 93 TABLE 17. Comparison of Mean Soil Moisture Per Tree at Different Orchards (Oneway ANOVA). 1979 1282 K.S.H. 28.2a 32.5a Upjohn 24.0a 29.9a Hofacker - 17.8b Means followed by the same letter are not significantly Dif- ferent at the .05 level by the Duncan's Multiple Range Test. 2 TABLE 18. Coefficient of Determination (R ) of Accumulative Mean Percent Soil Moisture with Accumulative Per- cent Apple Maggot Fly Emergence or Trap Catch. 1979 1980 K.S.H. Upjohn K.S.H. Upjohn Hofacker Seeded Cage .70 .71 Natural Cage .55 Yellow Trap .97 .79 .96 .90 .80 Red Sphere .96 .84 .96 .89 .78 base 48°F, and they should have started emerging at 900 degree days. However, some very important information was learned. In the soil, even though only 10% by weight, water must be in a free state readily available for plant and animal processes. As was shown in the previous section, flies still emerged at these low values. In the laboratory, the relative humidity in the air, even though 30 percent, was still too low for insect utilization. The result is that this experiment agreed with data presented by Neilson (1964) when he said that no emergence occurs below 40% relative humidity in the laboratory, whereas in the field 94 seldom is this high level reached during the normal emer- gence time. This also agrees with Dean and Chapman (1973) who say the translation of soil moisture to relative humidi- ty is difficult, as under normal orchard conditions the equivalent of 20-40 percent R.H. rarely occurs, while that is the normal range of soil moisture. Conclusions There were no significant differences found in the percent soil moisture at three locations under an apple tree canopy. The north side of the tree was the most consistent, so if just one sample were to be taken, it should be taken there. The south side was much more variable due to faster rain penetration allowing for faster wetting, and greater sun exposure allowing for faster drying. There were differences across sites however due to soil types and cover crops. The effect of soil moisture on first emergence could not be determined, but it was analyzed for season long effects. Correlations found very high relationships between accumulative percent soil moisture and accumulative percent trap catch on yellow traps and red spheres. Laboratory studies confirmed that adequate soil moisture is necessary during pupation to rear adults. 95 B IOT IC COMPONENTS Several biotic factors in the orchard are involved in the emergence of apple maggot flies from the soil. Those factors presented in Figure 14 will be examined for their influence on first emergence. The variety influence was studied at the K.S.H. orchard for four years, and the Upjohn orchard for two years. This was done with cages and traps. An estimate of the size of the population that carrys over to the second year was made at the K.S.H. orchard, and how that influenced first emergence. The orchard floor culture was noted as to its influence on emergence. If an orchard has early varieties present, then a small proportion of the pOpulation can complete its development in them and emerge as a partial second generation.‘ This was studied to help explain the long duration of emergence by using cages seeded at varying times. Lastly, the location of larval pupation was examined for its influence on emergence. This included location under the tree and depth in the soil. 96 VARIETY OF APPLE REARED IN The variety of apple the apple maggot was reared in may have some role in determining the time of emergence the” following year. Many authors have reported (n1 variety preferences for oviposition, but only a few on the effect on time of emergence. Those that have came up with varying conclusions. O'Kane (1914) said "various other factors, such as the kind of soil, the location of the pupae with reference to shade and the local conditions of moisture, probably have greater influence on timing of emergence than the variety the larval was reared in." Caesar and Ross (1919) seeded cages with infested summer and fall apples, and had the flies emerge one day sooner in the fall apples. They said “so far as summer and fall varieties go there is no difference between dates of emergence." Porter (1928) said "the emergence data indicate the time of maturity of the fruit in which the maggots develop has a definite influence on the time when the flies emerge in the following season. In all cases emergence of flies which developed as larvae in summer fruit rise at an earlier date than do those for the emergence from material which developed in fall or winter fruit." Phipps and Dirks (1933b) reported that flies from Red Astracan and other early fruits began to emerge about a week sooner than those from McIntosh and all later sorts. Chapman and Hammer (1934) said "flies developing from maggots which came from early-maturing varieties tend to emerge on an earlier schedule than flies originating in 97 later sorts," but they presented no data to back this statement. Garman (1934) said "maggots breeding in early varieties may produce flies that emerge earlier in the year," and he used a figure to illustrate this point. Dirks (1935) said "soil temperatures appear to have influenced time of fly emergence to a greater extent than the time of maturity of the fruit in which the larvae developed." He continued "no consistent differences in time of fly emergence has been apparent during the last three years between apple varieties, whether the larvae developed from summer, fall, or winter fruits." Hall (1937) said "the variety of apple in which the larvae feed affects the time of their maturity - those in the early ripening apples maturing first. Only early maturing larvae transform to adults in the current season." Glass (1960) reported emergence from seedings made with Yellow Transparent apples started and reached the peak and 50 percent points 10 to 16 days ahead of those from other varieties such as Wealthy and McIntosh. Glass (1961) said "individuals front early maturing varieties such as Yellow Transparent tend to emerge in early July whereas those from midseason varieties emerge a little later and those from winter varieties such as Baldwin emerge heavily toward the latter part of the month." Oatman (1964b) said "early or late maturing varieties had considerable effect on emergence. Adults tended to emerge earlier, reach peak emergence sooner, and have a shorter emergence period where the larvae developed in early 98 varieties such as Yellow Transparent than did those which develOped in later maturing varieties." Dean and Chapman (1973) performed an analysis of variance on 54 pairs of varieties, and found "there is no strong evidence from this that earliest emergence is associated with an early maturing host variety, although it happened occasionally, and it must be concluded that, in general, the time of maturity of the host variety exerts only a slight effect on the time of fly emergence, but what influence there is usually associates earlier maturity with earlier emergence." Lastly, Reid and Laing (1976) said that the effect of variety reared in had no significant effect on the time of adult emergence. Materials and Methods The first method to determine if there was a difference in first emergence date due to the variety the apple maggot fly was reared in was to use seeded cages. In the K.S.H. orchard in the fall of 1978, eight 1 meter square emergence cages were seeded with a bushel of infested apples, with each cage receiving a different variety. The varieties are shown in Table 19, and they were selected because of their varying harvest dates after fall bloom. Cages were moni- tored three times each week beginning June 14, 1979 to determine first and season long emergence patterns. The second method of measuring differences in emergence due to variety was to use naturally infested emergence cages. During the summers of 1978 and 1979, meter square 99 emergence cages were placed over naturally infested ground under several varieties of trees in the K.S.H. orchard. In 1978 the sampling initiated June 23 and was performed daily for three weeks, and then cut back to twice weekly for the remainder of the season. In 1979, sampling was initiated on June 18 and performed three times each week. Each variety was replicated at least twice, and some four times to pro- vide mean values. The third method for measuring variety difference in first emergence at the K.S.H. orchard was to monitor fly flight with yellow Zoecon AM traps. This should provide the most reliable indicator of actual differences because the traps were used each year, and a greater number of replicates were made with each variety. In 1977, 10 different varieties of apples had traps hung in them. A total of 50 trees were sampled weekly beginning June 20. In 1978, the same trees were monitored as in 1977. Sampling initiated on June 23, and continued daily for three weeks. The sampling plan was altered in 1979 by reducing the total number of trees checked to just 15. Eight different varieties were tested. Sampling was initiated on June 18 and repeated three times each week. These same trees were sampled in 1980, with monitoring beginning June 23 and repeated three times each week. The last technique used to determine variety dif— ferences in apple maggot fly first emergence was to monitor flight with red sphere traps. These traps were placed in 100 the K.S.H. orchard in 1979 and 1980 on the opposite sides of the same trees that had yellow traps. They were monitored the same way as was described for the yellow traps. Variety influence of first catch of apple maggot flies was also monitored in the medium density Upjohn orchard. Yellow traps and red spheres were placed on the opposite side of the trees monitored. In both years, three trees each of three varieties were sampled. In 1979, the sampling was initiated on June 20 and was repeated three times each week. In 1980, sampling started July 2 and was repeated three times each week. Results and Discussion Table 19 shows the range in date of first emergence of apple maggot flies due to the variety they were reared in as determined by seeded cage studies. There are obvious differences in these dates and ‘the air degree day accumulations, however, this was non-replicated data, so no statistics can be performed. In general, the later maturing the apple, the later was the first emergence of apple maggot adults. Table 20 shows the range in first emergence due to variety as determined by natural cages. Because air degree days provided smaller C.V. values than the same data reported as Julian date or soil degree days, they were used 101 Table 19. Date and Air Degree Day Accumulation (Base 48°F; B. E. Method) of First Emergence of Apple Maggot Fly Due to Variety It Matured In Measured by Seeded Emergence Cages at the K.S.H. Orchard in ‘ 1979. Days to Harvest1 Season First Emergence Variety From Full Bloom of Apple Date Air dd Transparent 70-75 Summer June 20 836 Dutchess 90-95 Summer June 17 774 Wealthy 120-125 Early Fall June 17 774 McIntosh 125-130 Early Fall June 19 814 R. I. Greening 135-145 Fall July 4 1,074 Jonathan 140-145 Fall June 19 814 Red Delicious 140-150 Fall June 22 892 Northern Spy 145-155 Late Fall June 19 814 Mean June 21 849 1 Smock and Neubert, 1950. Table 4, pp 14-15. Table 20 Effect of Variety on First Emergence of Apple Maggot Flies at the K.S.H. Orchard as Determined by Catch in Cages Placed Over Naturally Infested Ground (Mean Air Degree Days at Base 48°F; B.E. Method) 1978 1979 Both Years Variety No. Mean No. Mean No. Mean Wealthy 2 943 a 2 943 a Transparent 2 994 a 2 943 a 4 969 a Dutchess 4 1046 a 2 964 a 6 1019 a Northern Spy 4 1007 a 2 1149 a 6 1078 a Greenings 4 1094 a 2 1047 a 6 1082 a McIntosh 2 1080 a 2 1085 a 4 1082 a Jonathan 2 1117 a 2 1117 a Red Delicious _4 1120 a _2 1074 a '_6 1128 a Total 20 1061 16 1040 36 1052 Means followed by the same letter are not significantly different at the .05 level by the Duncan's Multiple Range Test. in this and further analysis. Analysis of variance and Duncan's Multiple Range tests were run on each years' data, and the lumped data for all years. In each run the F value 102 was not significant, so mean separation tests should show no differences. This was the case, as at the .05 level there was no significant differences between the mean date of first emergence for any of the varieties. Even though there are no significant differences, there is a range in the values with the flies emerging sooner in the earlier varieties and later in the late varieties. Data on the influence of first catch due to variety as determined by catch on yellow traps are more variable (Table 21). In 1977 and 1978 at the K.S.H. orchard, there were no significant differences between the air degree day accumulations at first catch. During 1979, the data pro- vided three different subgroups of varieties that were not statistically different within the subgroups. These groups fit the maturity intervals fairly well, with the exception of Northern Spy catching flies earlier than was expected. Data from 1980 showed four subgroups of varieties that were statistically different. This data fits very nicely the maturity dates of varieties, and was what was expected from these tests in all years.. When all years were combined, the DMR test showed no differences between any of the varieties in first catch. Red sphere traps were also used to determine dif- ferences in timing of catch due to variety of apple. Table 22 shows that for both 1979 and 1980 there were differences in timing in the K.S.H. orchard. When these two years were combined, differences *were still. present. The three 103 subgroups of homogeneous varieties again. matched the maturity dates quite closely with the earlier varieties catching flies sooner than the later varieties. To test whether variety has an influence in first catch of apple maggot flies in an orchard with some sprays applied, tests were set up in the Upjohn orchard. Table 23 presents the data from this medium fly density orchard. When the yellow trap was tested, no statistical variety difference was discovered in either year. Large ranges in mean degree day accumulations were evident between the varieties, and it was assumed that larger sample sizes would have shown statistical differences. When red sphere traps were tested some differences were found. In 1979, flies were caught on Northern Spy significantly later than the other varieties. In 1980, there were no differences. When both years data were combined the variances were still homogeneous, and the F ratio was significant. The variety influence was present, but not exactly as was expected. McIntosh should have been first, followed by Jonathan and Northern Spy if maturity dates were the predominating factor. However, greatly reduced populations and the pesticide applications greatly influence trap catch, and these factors probably overshadow any possible variety effect. Four different techniques of measuring variety influ- ence on first emergence or catch over a four year time frame were just presented. In order to compare these techniques 104 , mca 0c Hm>ma 00. mcy um ucmamwmam waucmoamacmam 0.00aa 00a 5.a0aa 0a m 0.00aa 00 0 0.0ama m m 0.00aa 0 0 0.50aa m m 0.000a 0 m 0.000a Na co 0.00aa m m 5.00aa aa mo 0.00aa m m 0.50aa 0 m 0.000a 0 non 0.00aa a m 0.00oa 0 ocm 0.000a N m 0.00oa 0a cm 0.000a m m 0.000a 0a m.0.000a N cmmz .oz cmmz .oz mammw HH< 000a um mama mmammn Had :mmz. c.0ee ma he o.mme M: o o.eeea N am m.mee N on e.em0a N e e.eNe a he m.eee N he e.mee N am m.eee N cmmz . OZ eeea .ummu mmcmm mamauasz m.cmocso uoc mam ampuma mEmm mcu 0c nmonHOM mammz 550a 0.050H 00 0.00HH m 0.000H 0H m 0.000H m 0.000H N m 0.500H m 0.000H N m 0.000H m 0.000H 0 m 5.00HH m 0.000H 0 m 0.00HH m 0.N5HH 0 m 5.00HH m 0.H00 m m 0.00HH m 0.000H N m 0.00HH m 5.0HHH 0 m 0.00HH m 0.500H 0 m 0.00HH cmmz .Oz cmmz 050a Acocamz .m.m O 0 Hmuoa I 00NN0000N5| H 0 Z N.0000 mmmm .mmmae Em coomoN BoHamw co coumO 0c GmCaEumqu mm camcono .m.m.x mcu um ham u000m2 mammd mo coumo umuam so mumeHm> mo uommmm ham cumcuaoz cmcumcon mmmmcaz cmoucHoz maceoeama 0mm cwzoamm sesame: ucmummmcmae maecmmao mmmcouso Numaum> .HN magma 105 Table 22. Effect of Variety on First Catch of Apple Maggot Fly at the K.S.H. Orchard as Determined by Catch on Red Spheres. (Mean Air Degree Days at Base 48°F; B.E. Method) 1979 1980 Both Years Varietv No. Mean No. Mean No. Mean Wealthy 1 1047.0 a 1 1140.0 a 2 1093.5 Dutchess 2 1074.0 a 2 1131.5 a 4 1102.7 Transparent 2 1135.5 a 2 1128.5 a 4 1132.0 McIntosh 2 1223.5 a 2 1290.0 abc 4 1256.7 Greening 2 1277.0 ab 2 1187.0 ab 4 1232.0 Northern Spy 2 1338.5 ab 2 1513.0 c 4 1425.7 Jonathan 2 1460.5 ab 2 1513.0 c 4 1486.7 Red Delicious _2 1688.0 b _2 1440.0 bc _4_ 1564.0 Total 15 1296.1 15 1303.1 30 1299.5 Means followed by the same letter are not significantly dif- ferent at the .05 level by the Duncan's Multiple Range test. ab ab bc 106 on» en am>ma me. we» e.Neea ea e.eema no.moea w nem.eeea e em.Neea em.NeNa e :mmz .02 2mm: um ea mN.Ne0a m e eN.eeNa e .02 mamcmm 0mm mmue Bodamw mummy cuom .umma mmcmm mamwuanz m.cmo:so 000a .m.m «mom0 mmmm um mxmo mmamma uefi :mmz. 00amcmo Ezecmz mcu um 0am uommmz mammd 00 coumo amnem so >umaum> mo uommmm .00 magma ucmamwwam xaucmoamacmem no: mam amuuma mEmm mcu 0c 0m3OHHOM mammz 0.N00a 0 H.0a0a 0 5.am0a 0 H.000a 0 Hmuoe IIIIIII I IIIIIII I IIIIIII l .lIIIIII l 000 m0.000a 0 m5.000a 0 £0.000a 0 m5.000a 0 camcuuoz m0.0m5a 0 m0.000a 0 m0.00aa 0 m5.000a 0 cmoucHoz m0.0a0a 0 m0.0a0a 0 m0.aama 0 m0.00aa 0 cmcumcon cmmz .02 2mm: .02 cmmz .oz cmmz .oz Numwnm> mumcmm 0mm WWHB BOHHm» mumcmm 0mm mmae 30HHm> 050a ..eonumz .000a cam 050a ca Unmcoao ccOnQD 107 and years, and provide some data for a phenology model, Table 24 was prepared. For each year and method, the grand mean of first emergence was calculated. From this grand mean, the mean per variety was subtracted. This deviation provides a uniform method of comparing that removes the method and yearly difference in air degree day accumulation at first emergence. The average difference from mean provided data for unbalanced analysis of variance and mean seperation tests. The data was transformed by adding a constant 250 to remove all negative values. The analysis was performed, the variances were found to be homogeneous and the F ratio was significant. From this table, if an orchard with a high density of apple maggot flies was monitored, no matter what method was employed, flies should be caught first from the Wealthy variety. Sixteen degree days later, they should be caught on the Dutchess variety. This trend continues for all 10 varieties listed, with Red Delicious being the last variety to catch flies. Also, if a mean first emergence value per orchard was determined, 96 degree days could be subtracted from that value if Wealthy was present in that orchard. This new accumulation could then serve as a predictor for first emergence in the orchard on the other end of the scale, if first catch on the Jona- than variety was to be predicted, 68 degree days should be added to the mean value of the orchard. These differences between the varieties were significant, and followed very closely the maturity dates of the individual varieties. 108 .Umucmmmaa mammE amsuom c003 mamxamcm H00 000+x 0c 0m8u00mcmau muma .ummu mmcmm maaauaaz mcmocsa mcp 0c am>ma 0o. mcp um ucmamwmac maucmoawacmam po: mam amuuma mEmm mcu ac UmBOHHOM mammz n e0 eeN Ne Ne e- ma- em sea em- mm: 0mm enmnnuoz o em eea Nam Nm me ea em- em em m0 macaoaamn ewe on we eNa eea mm me 0- we ee . mm: cennmaoe one 0 I u n . u eNu 0m . n u mmmmnas one a we- ea- ee- am- 00 mN- e mm mNN meacmmno .a.m nm Na] me- me- Nm ma- Na- NN| me _ ea mm- nmonnaoz one mm s I n . aoa 0N- I I I gazeaen e we- mean eeN- a me- me: em- ee- . me- snnammz 8 ea. eea: NNN- em- ma- m- em- we- ea- me: mmmnonsn 8 ae: eoNu aeau 00: man 0N: ea: em: 00: ma: unmammmnene cmmz mamcam mumcmw amae amae @009 amae mmmo mmmo mmmu Numaam> 80am 0mm 0mm zoaamw Boaamw 30Ham» BoHamw amasumz amasumz vmommm moamnmaean oeea eeea omea eeea meea eeea eeea mema eema .acocumz .m.m «0°00 mmma mma mmamma Had cmmz Eouw mocmamumaa. camcoao .m.m.x xuamcma 0am c000 mcu um 0H0 uo00mz maaafl mo musuamu Ho moammumfim umaaa co >umaam> mo mocmsamca mcu 00 mmocumz 0cm mummw mmoao< QOmwamano .00 macme 109 This same technique of determining grand mean per orchard by method and year, and then calculating deviations from that mean was also performed for the medium fly density Upjohn orchard. Table 25 found significant differences between varieties. Jonathan variety caught flies first, and for predictive purposes, caught them 187 degree days before the orchard mean. McIntosh variety was second, and only 22 degree days after the orchard mean value. Northern Spy variety was last to catch flies, and on the average was 165 degree days later than the orchard mean. Conclusions The variety that an apple maggot fly is reared in has an effect on time of adult emergence. In earlier maturing varieties, the adults emerge earlier the following year. This was varified by using seeded emergence cages, natural emergence cages, yellow traps, and red spheres in the high fly density K.S.H. orchard. These differences were signifi- cant in several trials. Ranges in air degree days from the grand mean were calculated by variety for predictive pur- poses. When tested in medium fly density orchards, the variety still had an effect on first emergence. Both yellow traps and red spheres caught flies first_on the Jonathan variety, followed by McIntosh, followed by Northern Spy. The dif- ference between mean first catch per variety were signifi- cant, and provided degree day values for predictive pur- poses. 110 Table 25. Comparison Across Years and Methods of the Influ- ence of Variety on First Catch of Apple Maggot Fly at the Medium Fly Density Upjohn Orchard (Difference from Mean Air Degree Day Base 48°F; B.E. Method). Differ- 1979 1980 1979 1980 ence Yellow Yellow Red Red From Trap Trap Sphere Sphere Mean Jonathan -228 -103 -211 -208 -187.5a McIntoch 65 71 -253 -207 22.4 ab Northern Spy 164 32 464 2 165.4 b Means followed by the same letter are not significantly dif- ferent at the .05 level by the Duncan's Multiple Range Test. Data transformed by x+260 for analysis with actual means represented. This variety influence on first emergence has an adap- tive advantage to the species. When a variety matures early, flies that have been reared in that variety are mature and ready to oviposite in that variety. Flies that emerge later are then mature and ready to oviposite in later maturing varieties. This ensures species survival by having mature flies present over a long time frame to coincide with fruit susceptibility. 111 TWO YEAR LIFE CYCLE A certain small proportion of the apple maggot popula- tion will not emerge as normal, but will overwinter the second winter and emerge as flies the second summer. For predictive reasons, one needs to know whether these flies emerge earlier than normal first year flies. This was measured at the high fly density K.S.H. orchard by placing emergence cages over the identical spot for two consecutive summers . Literature Many authors have reported that a small proportion of the apple maggot population will overwinter the second year, (Table 26). Generally, it was found that these flies emerge during the same period as first year flies.‘ By going farther north in its range, the fly has a tendency to over- winter for more years. Neilson (1976) in Nova Scotia found .14 to 3.67 percent carried over to the third year, and 0 to .11 percent carried over to the fourth year in seven dif- ferent tests. This may help to insure species survival when suitable oviposition sites are not available every year. A few authors hypothesized on causes for these carry- over individuals. Oatman (1964 b) noted "a 15 percent carry-over population in 1962 and a 5 percent carry-over in 1963. The difference was probably due to below-average temperatures and rainfall during the 1961 emergence period creating a higher carry-over than that caused by above average conditions during 1962." Also, Trottier (1979) 112 Table 26 Literature References to the Occurrence of a Two Year Life Cycle of Apple Maggot Fly with Percent of Carry-Over and Time of Emergence in Relation to First Year Flies Recorded. Percent of Time Source Carry-Over of Emergence O'Kane (1914) Some Ordinary time Caesar & Ross (1919) 6.6 to 18 - Porter (1928) 0.1 to 37.5 Usual emergence Period Phipps & Dirks (1933) 5.2 to 8.3 Height of regular Flies Allen & Fluke (1933) 37 8 days later Garman (1934) relatively small - Dirks (1935) .91 to 2.21 Same period Hall (1937) 1.2 Same Period Glass (1960) few individuals Normal time Oatman (1964,b) 5 to 15 - Dean & Chapman (1973) .97 - Cameron & Morrison Up to 16 Same Period (1974) Neilson (1976) 11.11 to 58.11 - found that a greater percentage of pupae carry-over to the second season.:hi dryer soils. Probably Porter's (1928) explanation is the best. "The two year cycle operates de- finitely txa the advantage (n5 the species, insuring the survival of at least a few individuals over seasons of complete crop failure." Material and Methods All the work related to second year emergence was performed in the K.S.H. orchard. The eight cages seeded with infested apples on uninfested ground in the fall of 113 1978 were monitored in 1979 to determine first and seasonal emergence patterns. In 1980, cages were placed in the identical spot, and monitored again. The dates and air degree day accumulations at first emergence were recorded to determine timing in relation to first year flies. Also the total number of flies were recorded to determine the per- centage involved in carrying over to the second season. These emergence dates were compared with first year emer- gence from seven cages seeded with apples in the fall of 1979 on other patches of uninfested ground adjacent to the second year cages. Monitoring initiated June 23 and was performed three times a week until after all cages had caught the first fly. Results and Discussion The data from this test is presented in Table 27. The average first emergence from first year pupae was June 30 or 1036 air degree days. The second year flies first emerged on average July 3 or 1101 air degree days. Analysis of variance found these two treatments not statistically dif- ferent, which was confirmed by the Duncan's Multiple Range Test. On the average, the delay was three days. This may be an actual difference, and if so, then one that was un- expected. One would theorize that flies passing through two seaSOns would emerge sooner because they have had twice as 114 Table 27. Effect of Carry-Over Pupae on Timing of Emergence of Apple Maggot Flies at the K.S.H. Orchard in 1980 (Air Degree Days at Base 48°F; B.E. Method) First Year Second Year Emergence Emergence Cage Date Air dd Date Air dd 1 6-30 1036 7-5 1140 2 6-28 1016 7-5 1140 3 6-28 1016 7-2 1076 4 6-30 1036 7-7 1187 5 6-30 1036 6-28 1016 6 6-30 1036 7-2 1076 7 7-2 1076 6-30 1036 8 - - 7-5 1140 Mean $S.E. 6-30:.5 1036i7.1 7-3i1.1 1101i21.0 115 much time to develop physiologically. It is possible that this is the case, and the explanation why the delay was present was that a much smaller population was monitored (Table 28). This shows a range of 2.9 to 11.3 percent of the flies carrying over to the second year, and an average of 6.8 percent. With fewer total flies present, it is reasonable to assume that fewer earlier individuals in the population are present, and therefore first catch would be delayed. Conclusions It was found that flies overwintering to the second year emerge an average of three days later than flies emerging the first year. This difference may be real, or may be due to the greatly reduced total population that is sampled the second year. This reduced population sampled may overweigh any real difference that might be present and would be expected. These findings concur with those pre- sented in the literature. It is important to note that in every case a certain proportion of time flies did overwinter the second year. This phenomenon has adaptive advantages to the species. This allows for individuals to be present the second season and continue the species if all the oviposition sites were absent one year. This is a possibility as: (1) spring frosts tend to remove many apples and could destroy an entire crop; (2) diseases such as apple scab could be severe 116 TABLE 28. Number and Percent of Apple Maggot Flies Over- Wintering to the Second Season at the K.S.H. Orchard by Variety. First Year Second Year Percent Cage Emergence Emergence of Total Transparent 34 2 5.6 Greening 66 2 2.9 Wealthy 87 4 4.4 Dutchess 85 3 3.4 McIntosh 55 7 11.3 Jonathan 113 4 3.4 Red Delicious 324 34 9.5 Northern Spy ‘131 10 6.8 ON ON 0\ C CD Total Caught 901 117 and destroy the fruit or make it unsuitable for oviposition; (3) other insect pests such as plum curculio and codling moth could be extremely successful and cause all the apples to abort before apple maggot emerges; or (4) the natural bienniel bearing of trees could result in no fruit alter- nating years. 118 ORCHARD FLOOR CULTURE The orchard floor culture may effect emergence. With deep grass underneath the trees, the sunlight cannot pene- trate to warm the soil as quickly, which can delay emer- gence. If the grass is mowed or a herbicide is applied, then the soil surrounding the pupae warms faster, and earlier emergence should occur. With a clean cultivated orchard, emergence times may or may not be speeded up. The soil warms faster which would result in earlier emergence, but the pupae that survive discing and desication are deeper in the soil and are not exposed as rapidly to the warmth. Also, on cultivated soil fewer flies are obtained than on uncultivated soil which tends to delay emergence (Mundinger, 1930). In this study, temperatures were measured under dif- ferent natural conditions. The environment was not modified by disturbing the soil nor placing artificial objects under the trees. Degree day accumulations were made and compared under these different conditions. It is assumed that degree day accumulations is a good method of integrating all the variables such as depth of grass, amount of shading, and orchard floor culture that effect emergence. Literature Observations relating to orchard floor cultures effect on timing of emergence are scarce in the literature. Garman (1934) said ” research workers have indicated that maggot 119 flies will emerge earlier from sandy soils than from heavy loams." This could be due to the sandy soils warming up faster to cause earlier development, or the soils being looser so the flies can crawl up and out easier. Glass (1960) states that "variations from seeding to seeding due to such factors as shade density of the tree appear to have had very little influence on the time of maggot emergence." His data shows that "the location of the seeding within the test orchard was not a significant factor in time of emer- gence." Materials and Methods Three lead Weather Measure thermographs were placed in the K.S.H. orchard the second week of July in 1979. One probe was in the standard U.S. Weather Service 5-foot shelter, the second was on the soil surface in tall grass directly exposed to the sun, and the third was on the soil surface in tall grass but shaded by thé tree canOpy. These three locations were monitored for the remainder of the season to determine if there were differences in the amount of heat penetrating to the soil surface. In 1980, these same temperature instruments were placed at different sites around the state. Site one was the Upjohn orchard where S-foot air and shaded soil surface temperatures were recorded. Here there was plenty of canopy to shade the probe, and it was placed in thick tall grass for additional protection. Site two was the Hofacker yard 120 north west of Grand Rapids. Five-foot level air temperatures and shaded grass litter zone temperatures were recorded. This grass litter zone probe had indirect sun exposure caused by the shading, but the grass was mowed frequently which would simulate a well mowed or herbicided commercial orchard. This should provide a medium heating and emergence. Site three was the Yabs yard. The S-foot air temperature probe was identically placed in the weather shelter as in the other locations. The grass litter zone prob was in mowed grass, but exposed to the direct sunlight. This should provide the quickest heating and earliest emergence. These locations were monitored from April 1 to the end of July to determine differences in these three types CHE orchard floor cultures. No thermometers were placed in disced soil, as the majority of the apple orchards in Michigan are under one of the above types of sod culture. Results and Discussion There are differences in the accumulation of degree days at the three different sites monitored in the K.S.H. orchard in 1979 (Table 29). Because of the extremely high temperatures which could not be entered into the computer program to calculate degree days by the Baskerville-Emin method, the averaging method was employed to calculate degree days. An analysis of variance was performed on the data broken down into half month intervals (Table 30). The 121 000 0.000 0.000 aoflumaaeaood 00 00 00 00 00 00a 00 H0 H0 H0 hash m.eN Ne mm me me me m.eN ae em on ease m.mm Ne me we we NNa eN 0e we eN ease eN . me me m.mm ee eea m.eN em me eN ease em ee we we ee eaa em 0e ea eN ease m.am ee Ne m.me Ne aNa m.mN ae Ne eN ease mN ee ee eN we we aN me me mN ease eN me am 0.0m ae em eN oe om 0N ease em me ee em Ne eNa me me me mN ease we we 0e m.em em eaa m.eN em om NN ease mm Ne eea me we eNa 0N em me aN ease 0N em ee m.em em maa m.mN em 0e eN ease m.eN em 0e m.ae mm eNa eN we 0e ea ease m.eN em ae m.em we aNa m.eN me me ea ease m.mN em em e.am Nm eOa am ee we ea ease me me we 0.00 Ne aNa m.eN so as ea ease owe ee ea: xez .ee we mmm mmm .ee ee 000 xez enen mmmao flame mmmao Hams “04 uoom m>Hm mmHB 0c cmnmcm cam uomaaa .AnocumE m0mam>¢ “0°00 mmma wma mmumma. 050H :0 mamcouo .:.0.z mcu um mcoHumHSEDood wma mmumma co manuaao HOOHa vamcouo mo uommwm .00 chme 122 005 a50 0.500 cOaamHsESooa 50 00 a0 00 50 00 00 00 00 a0 00 00 00 00 00 00 0.00 00 50 00 00 00 05 00 00 a0 0.00 00 00 00 a0 00 05 0.00 00 00 00 00 05 00 00 00 05 00 00 55 00 00 55 50 0.00 50 05 0.00 00 a0 00 05 05 00 a0 00 05 0.50 a0 00 . 00 00 55 00 0.00 00 05 00 00 00 . 0a 50 a5 00 0.50 05 a0 00 00 55 0.00 00 00 00 00 00 05 0.50 a0 00 00 00 05 00 0.00 00 05 00 00 05 00 00 00 a0 0a 00 00 0.0a 00 a5 0a 00 00 00 5a 00 00 0.0a a0 00 0a. 00 00 0a 0.0a 00 a0 0.aa 00 a0 0 00 00 0a 0a 00 00 aa 50 a0 0.0a 00 00 5a a0 a0 55 0.00 50 00a 5a 00 05 00 0.0a 00 55 0.00 50 00 0.0 a0 00 0a 0a 00 05 0.0a 00 00 0.5 00 50 0a 0.00 00 05 00 00 55 0a 00 05 0a 0.50 00 00 00 a0 00a 0a 50 05 0a 0.00 00 05 0.00 00 00 0a 00 05 Ha 0.00 00 a0 0.00 00 00 0.a0 a0 05 00 0.00 05 00 0.00 05 0aa 00 05 00 0 0.00 05 00 0.00 a5 00 0.5a a0 00 0 0.00 05 00 0.00 05 0aa 00 05 am 5 0.00 a5 00 00 05 0aa 50 00 00 0 50 00 a0 00 50 00 0.00 00 m5 0 a0 a5 50 00 00 00a 0.00 00 50 0 0.00 00 00 0.50 00 00a 00 00 00 0 50 50 00 50 00 00a 00 00 . 05 0 m.aN e0 ae oN me ae m.ea me. me a nesmse 000 00 ea: xmz 000 00 Ca: xmz 000 00 :02 xmz mama mmmau Hams mmmao Hams Rea aoom m>Hm mmae 0c Umomcm cam aomuaa emanannoo neN mamas 123 050 000 0am COHamH:Esoo< 0a 00 00 0.0a 00 05 0.0a a0 05 00 0.0a 00 05 0.00 50 00 00 00 00 00 00 00 05 0.00 50 00 0.00 00 00 00 0.0 00 05 0.5a 50 00 ma 00 00 50 0.0a 00 05 00 50 00 0.0a 00 00 00 0.aa 00 05 0a 00 00 0.00 00 05 00 0a 00 00 0a 00 00 0.0a 00 05 00 0.5 50 00 0a a0 a0 0.0 50 00 00 0 00 00 0a 00 00 0 00 _ 00 00 0.5 00 00 0.0a 00 05 aa 00 00 a0 0.0a 50 00 00 00 00 0.0a 00 05 00 5 50 00 0a 00 a0 0.5 00 00 0a 0.0a 00 05 0.00 50 00 0.5a 50 05 0a 0.0a 00 05 0.00 50 00 00 00 00 5a 0.0a 00 05 0.00 00 50 0.0a 50 05 0a 0 a0 00 0a 50 00 0 50 00 0a 0 a0 00 0.0a 00 05 0a 00 00 0a 0.0a 00 05 0.00 00 05 0.0a 00 05 0a 00 00 05 00 00 00 0.00 00 00 0a 00 a0 05 00 00 00 00 00 00 aa 00 00 05 00 00 00 00 00 00 0a 0.5a a0 05 00 00 00 0a 00 05 0 aa 00 00 a0 50 a0 0 00 00 0 0.0a 00 00 0a 50 00 0 a0 50 5 0.00 a0 05 0.00 50 00 0a 00 00 0 00 00 05 0.0a 00 05 00 00 00 0 0.00 00 a5 0.0a a0 05 0a 00 05 0 00 00 05 00 00 05 00 00 00 0 00 05 05 00 50 00 00 00 00 0 m.mN ee ee m.eN 00 em eN 00 me a .aamm 000 CC CHE xmz o00 00 :02 me 000 00 CHE xmz mama mmmaw aame mmmao dame aaé aoom m>aa mmae 0c cmomcm :50 aomaaa GmDCaacou .00 maame 124 .amma momma maaaaasz m.cmoc5a mca 0c Hm>ma 00. mca am acmamaaaw waacmoaaacmam aoc mam amaama mEmm mca >2 Um3oaaoa mammz c 0.0a m 5.00 m 0.00 c a.00 c 0.00 mmmao Hamelcsm aomaaa m 5.0a m 5.0a m 0.a0 m 0.00 m 0.00 mmmao aamalcmnmcm ne e.ma e 0.0a e N.ON e 0.0N e m.eN mama nae nooe e>ae 00I0a 0ala a0|0a mana a0|0a nmaoaacoz maaaaso aamm aamm amamsd amsmsa >050 HooHa mamcoao .acocamz mcammam>< «mom0 mmma mama mmamma. 050a ca mamcoao .m.m.x mca am mwoaama mEaB msoHam> mom mmasaaao a00aa camcoao acmamwaaa cmmBama chaaeasasoo< 0ma mmaoma ca mmocmamamaa :mm: .00 macme 125 variances were homogeneous for each set of data. The F test was highly significant for three of the five time periods and the Duncan's Multiple Range test separated the means into two signficantly different subsets. During the last half of July and the first half of August, the soil that has no shade is significantly warmer than the shaded soil and that is the 5-foot weather shelter. This fact would in- dicate that pupae in this type of environment would warm up faster, and allow the adults to emerge sooner. In the shaded areas, the soil is cooler, and flies can emerge over a longer time period. Later in the season, from mid-August to mid-September, there are no differences in the tempera- tures in these three sites. It is during this time period that the lavae are leaving the apples to pupate. If the extremely warm surface temperatures were present then as in late July (120+°F, Table 29), it is reasonable to believe that there would be a great mortality of the larvae before they could burrow down into the cooling soil to pupate. During the last of September, the unshaded soil surface was significantly warmer than the shaded soil, but not different than the five foot weather shelter. In all time periods, the mean temperature was warmer on the unshaded soil than in the other two sites. This seems reasonable, and could explain why flies emerge earlier in some locations than others, and why the long emergence period occurs in all sites. 126 Differences were also present in temperatures between the different types of orchard floor culture in 1980. Before analysis of the data on grass litter zone tempera- tures were made, air temperature degree day analysis were performed. Table 31 shows that all three sites did not have statistically different air degree day accumulations for the same calendar day time periods. Therefore, with same air degree day accumulations, any differences that might be present in grass litter zone temperatures must be due to the different type of orchard floor culture where the tempera- ture probe was located. Table 32 shows that differences were present for every time period in the grass litter zone temperatures. The tall grass shaded by the tree at the Upjohn orchard was the coolest site from April 1 to May 15. From then until the end of July, it was no different than the shaded grass shaded at the Hofacker site. The mowed grass exposed to the sun at the Yabs site was always sig- nificantly warmer than the shaded tall grass culture. From the first of May on, the mowed grass exposed to the sun was warmer than the shaded mowed grass. This seems reasonable as the sun should warm up the soil faster when it directly hits the prob, rather than when it is screened out and only indirectly warms the soil surface. 127 s o 00.0 n A00 0000 a amoaaaau .amma mmcmm maaaaaaz m.cmo::a mca 0c Hm>ma 00. mca am acmamawam waacmoaaacmam aoc mam amaama mEmm mca 0a 0m300aom mammz H5.H m 0.00 m 0.00 m 0.00 ~0I0H >090 ea.a me.a ee.e ee.a em.a eN.o ee.aN esae> a e m.mN e m.aN e 0.0a e N.ea e e.e e 0.0 a enea e a.MN e a.ea e a.Na e a.ea e m.e e 0.0 n e.ea aexoeaom e m.mN e a.eN e e.ma e N.ea e m.m e m.m e e.a enoema maua mmuea maua am-ea maua omnea maua mamsem wash mcaw mean 0m: 0m: aaaafi aaaaa 00 :oHamooa .acocamz .a.a “0°00 mmma am mxma mmamma Hm>ma aooa m>aac 0000 :0 cmmacoaz :0 mmaam acmamaaaa am Hmmw mca aaocmsoaca macaamm mEaB acmamamaa new >ma ama mafia: >ma mmamma Ha< mo amcEsz :mmz .a0 magma 128 ~ 0 00.0 n A00 0.00 a amoaaaau .amma mmcmm maaaaasz m.amoaaa mca 0c am>ma 0o. mca am acmamaaam >aacmoaaacmam aoc mam amaama mEmm mca 0a 0m300H00 mammz 00.00 00.00 0.00 00.50 a0.00 00.0a 00.5 0a.00 msam> a 0mm0axanmmmaw c 0.00 c 0.00 c a.00 c 0.0a c a.a0 o 0.0a c 0.5 : 0m3ozumcmw 0mmmcmlmmmao m 0.00 m 0.a0 m 0.0a m 0.0a m 0.0 c 0.0 c 0.0 c 0.0 vmzozuumcomaom cmomcmnmmmao e N.NN e m.aN e m.ea e N.aa e e.e e e.m e 0.a e o.o aaee n gnomes a0|0a 0a|a 00|0a 0a|a a0|0a 0a|a 0010a 0a|a mamamm mash wash mash mesh >mz 0m: HHHQ< aaaaa mo codamooa mwma mmammav 000a :0 cmmacoaz mafia acmamaaaa MOM wma ama .Avocamz .m.m «0600 mmmm am Ca mmaam acmamaaaa am ammw mca aaocmaoaca mUOaama maeca 0ma mmamma mcoN Hmaaaa mmmao mo amafisz can: .00 magma 129 Conclusions The data presented indicates that apple maggot pupae located in short grass or on herbicided soil that are directly exposed to the sun should emerge first as they are warmed sooner. Those pupae that are located in mowed grass but shaded from direct sunlight should emerge significantly later. The last flies to emerge should be in orchard floor cultures of tall grass and shaded from the sun by the tree canopy. 130 PARTIAL SECOND GENERATION The flight pattern of adult apple maggot flies lasts about three months (Figures 4-13). One source of the late season individuals is from.21 partial second generation. Larvae that developed in early maturing varieties develop soon enough to emerge the same year and infest later de- veloping varieties (Illingworth, 1912; Herrick, 1912; Caesar and Ross, 1919; Porter, 1928; Phipps and Dirks, 1933; Hall, 1937; Neilson, 1962; and Prokopy, 1968c; Boulanger, Stanton and Padula, 1969). Seldom do many of these larvae complete development before fall (Chapman and Hess, 1941) and are therefore suicidal. In the more northern regions of its range, no second generation adults appear (O'Kane, 1914 in New Hampshire; Mundinger, 1930 in Hudson Valley; Neilson, 1976 in Nova Scotia). Dean and Chapman (1973) said "bivoltinism undoubtedly occurs in the Hudson Valley, but it is of no consequence so far as the species control in commercial orchards is concerned." Studies were conducted at the K.S.H. and Upjohn orchards to determine if there is a partial second generation in Michigan, and how large a portion it is of the total population. Materials and Methods Emergence cages seeded with infested apples in the summer of 1979 were monitored that fall and the next summer to determine emergence patterns. Seven cages were seeded with the Dutchess variety. Seedings were made on July 16, 131 July 19, July 23, July 30, August 2 and August 6. This sequence of seeding was made to determine how late in the season apples can fall to the ground and still have suffi- cient time to have a partial second generation, and to see if there was an effect on time of emergence the following year due to the time of pupation the preceeding summer. Hall (1937) said that all second generation adults were derived from larvae which reached maturity before August 20, although all larvae that matured by that date did not transform to adults in that same year. Glass (1960) said that, "In 1953 collections from the same group of wealthy trees were made on August 18, August 29, and September 9. Emergence data shows ‘that there 'were no appreciable differences in emergence between cages. This probably means that larval emergence from the fruit was essentially the same regardless of when the fruit was collected. It seems likely the time of collecting infested fruit from any one locality and variety does not influence appreciably the time flies will emerge the following season." Reid and Laing (1976) said that the date the puparia was formed was not significant to the time of emergence of adults the following year. The second method to determine if a second generation of flies was present was to determine the gravidity cf the females. By dissecting females and looking for the presence of eggs in the oviducts, one can distinguish whether the female is mature or not. If no eggs are present, then the 132 female is considered immature and has just emerged. In nature, females very seldom oviposite their full compliment of eggs, so females without eggs are immature and not older individuals with spent ovaries. Also, older individuals appear very ragged and beat up and can readily be distin- guished from newly emerged ones. All flies caught in the K.S.H. and Upjohn orchards in 1979 and 1980 on yellow Zoecon AM and red sphere traps were sexed. The females were then dissected to determine if they were gravid. It was assumed there would be a period of no immature flies caught during late summer, followed by a low percent of immature females that are the second generation flies. As the sex ratio of the flies is very close to 50:50, the results from the dissections of the females should be indicative of the total population. Results and Discussion Emergence cages seeded with infested apples on uninfested ground showed that there is a partial second generation of apple maggot flies in Michigan (Table 33). Between September 22 and September 29 the first of these flies emerged. On October 13 these cages were removed, as it was assumed no more flies would emerge that year. This second generation ranged from 0 1x3 2.17 percent of the emerged flies in each cage, but averaged only 0.25 percent of the total flies that emerged from progeny of the 1979 adults. This is similar to Hall's (1937) data that had 5.4% 133 second generation from Dutchess apples in Ontario. This would have very little effect on a growers control program, and has very little effect on late season emergence. From this table it is evident that larvae that enter the soil as late as July 23 can pupae and complete development yet that fall. Table 33. Occurrence of Second Generation Apple Maggot Flies from Dutchess Variety in 1979 at the K.S.H. Orchard. Percent Seeded Sept. Sept. Sept. Oct. Total Total Second Date 13 22 29 13 in 1979 in 1980 Generation July 16 o o 1 o 1 45 2.17 July 19 0 0 O 0 0 105 0.0 July 23 O O 1 0 1 242 0.41 July 26 0 0 0 O 0 4 0.0 July 30 0 0 O 0 0 247 0.0 Aug 2 O 0 0 O 0 140 0.0 Aug 6 0 Q Q Q Q 18 0.0 TOTAL 0 O 0 0 2 811 0.25 By dissecting the females to determine if they are gravid, one can show that there are late season individuals that are immature. Table 34 indicates that from Septem- ber 13 to the end of the 1979 season at the K.S.H. orchard, about 50 percent of the individuals are newly emerged. These represent the partial second generation. To note, at this high density orchard there is a large preference for the yellow traps over the red sphere by the immature fe— males. At the medium fly density Upjohn orchard, too few flies were caught to make any specific conclusions, but it appears doubtful there are any second generation flies in their orchard. TABLE Date July July July July July July July July July July July Aug. Aug. Aug. Aug. Aug. Aug. Aug. Aug. Aug. Sept. Sept. Sept. Sept. Sept. Sept. Oct. Total NC 134 34. Percent Infertile Female Apple Maggot Flies as Determined by Dissections of Trapped Females in 1979. K.S.H. (n=15) Upjohn (n=9) Yellow Trap Red Sphere Yellow Trap Red Sphere 2. 2121.031. 10 .1139. 2 84.4 37 100 1 NC 0 NC 0 4 86.1 31 NC 0 NC 0 NC 0 6 81.8 45 66.7 2 NC 0 NC 0 9 69.1 67 21.4 3 NC 0 NC 0 11 68.0 66 10.0 1 NC 0 NC 0 13 53.4 126 18.2 6 33.3 1 NC 0 16 26.0 38 16.2 6 50.0 2 100 2 19 21.1 15 5.0 2 NC 0 0.0 0 23 29.1 51 26.5 9 0.0 0 NC 0 26 22.4 22 4.1 2 NC 0 NC 0 30 14.4 39 12.5 10 0.0 0 NC 0 2 17.2 55 82. 4 0.0 O 0.0 0 6 0.0 0 0.0 O 0.0 0 0.0 0 9 3.7 6 10.8 4 44.4 4 0.0 0 13 10.7 29 10.2 5 20.0 7 0.0 0 16 .7 1 0.0 0 NC 0 0.0 0 20 3.1 4 6.2 2 0.0 0 NC 0 23 0.0 O 3.8 1 0.0 0 0.0 0 27 0.6 1 0.0 0 7.1 1 0.0 0 30 1.6 3 0.0 0 0.0 0 0.0 0 3 5.6 9 20.0 2 0.0 O 0.0 0 6 2.6 1 0.0 O 0.0 0 0.0 0 10 0.0 O 0.0 0 0.0 0 3.3 2 13 40.0 10 50.0 2 11.1 1 0.0 0 22 57.1 4 NC 0 NC 0 NC 0 29 50.0 2 NC 0 NC 0 NC 0 13 NC 0 NC 9 NC 9 NC 0 650 62 15 Indicates no females were caught on the traps. 135 The dissections of the 1980 season show the same early season trends that were present in the 1979 data, but were not continued late enough in the fall to provide data on what portion of the whole population the second generation represents (Table 35). Early in the season at the high density K.S.H. orchard, 100 percent of the females caught are immature. This indicates these traps are effective in capturing early emerging individuals, and can be success- fully used for monitoring emergence in high population situations. The percent immature flies caught on the yellow traps gradually decreases in number and percent until the end of August. At that time all females caught were gravid. The same trend was present on the red sphere trap. The main difference was that the immature females prefer the yellow traps over the red spheres as indicated by the much larger percents in the yellow trap column. At the medium density Upjohn orchard, trap catch was greatly delayed. .Not until late July was the first immature female caught. Once caught, they were present at about the same percentage as in the higher density K.S.H. orchard. After mid-August, no more infertile females were caught in this orchard. It is reasonable to assume that there is no or a very small second generation in this orchard as the first females caught were in mid-July, and this does not leave sufficient time for the larvae to mature, pupate, and emerge prior to winter. The same preference for the yellow trap by immature females was also present in this orchard“ 136 Table 35. Percent Infertile Female Apple Maggot Flies as Determined by Dissections of Trapped Females Caught in 1980. K.S.H. (n=15) Upjohn (n=9) Yellow Trap Red Sphere Yellow Trap Red Sphere Date % No % No % No % No June 28 100.0 1 100.0 1 NC 0 NC 0 June 30 100.0 1 100.0 1 NC 0 NC 0 July 66.7 2 0.0 0 NC 0 NC 0 July 78.0 32 33.3 3 NC 0 NC 0 July 88.7 70 38.6 7 NC 0 NC 0 July 9 82.0 137 15.8 3 NC 0 NC 0 July 11 81.0 111 0.0 0 NC 0 NC 0 July 14 62.5 110 16.2 6 NC 0 0.0 0 July 16 39.7 29 9.4 5 0.0 0 NC 0 July 18 42.1 32 0.0 0 NC 0 NC 0 July 21 40.7 35 5.2 3 NC 0 20.0 0 July 23 25.9 41 7.4 2 7.4 4 0.0 0 July 25 29.8 31 5.9 1 22.6 7 0.0 0 July 28 20.5 26 5.9 3 8.6 3 0.0 0 July 30 14.1 14 4.5 2 11.8 2 0.0 0 Aug 4 13.8 27 1.3 1 26.7 4 0.0 0 Aug 11 .9 10 0.0 0 9.1 6 0.0 0 Aug 18 3.8 16 0.0 0 0.0 0 0.0 0 Aug 25 4.2 15 5.3 l 0.0 0 0.0 0 Aug 29 0.0 0 0.0 0 0.0 0 0.0 0 Sept 5 0.0 0 0.0 0 0.0 0 0.0 0 Sept 12 0.0 0 0.0 0 0.0 0 0.0 0 Sept 19 18.2 2 0.0 9 NC 0 NC 0 Total 742 39 26 1 NC indicates no females were caught on the traps. 137 Conclusions A partial second generation of apple maggot flies is present in the high fly density K.S.H. orchard. These individuals commence emerging about September 25 as deter- mined by cage studies. This date was a little late as determined by dissections of females from a larger population which found immature females present from September 10 to the end of the season. These flies are few in number, and represent only 0.25 percent of the flies that emerged as progeny from the 1979 summer adults. It was found that larvae that entered the soil as late as July 23 could still emerge that fall. This data affirms that of Dean and Chapman (1973) who claim these flies are of no consequence so far as the species control in commercial orchards is concerned. At the medium fly density Upjohn orchard, it is doubt- ful that even a partial second generation exists. No fe- males were caught in that orchard after September 13, so this could not be confirmed. However, the first individuals were caught two to three weeks later than in the K.S.H. orchard, and this delay is great enough to prevent suffi- cient develOpment of larvae and pupae for the second genera- tion to emerge. In both orchards, there was a definite preference of trap types. The infertile females were caught on the yellow traps 13.5:1 over the red spheres. This supports the belief that infertile females are attracted to yellow traps which 138 serve as a possible site for finding food. After maturation, they search out spherical objects, looking for mating and oviposition sites. Data presented showed that very few infertile females compared to all females were caught on the red sphere which also supports this supposition. The occurrence of a second brood is suicidal to the species. When adults emerge in mid-September, it must take 10 days minimum for their ovaries to mature due to the cool weather. By the first of October when they are ready to deposit eggs, the susceptible varieties, and the ones in which they were reared, are all gone. The apples remaining are the less preferred, hard winter varieties. If an egg is deposited in these apples, it is doubtful they would hatch or survive due to the firmness of the flesh. If they were to survive and hatch there is not sufficient time remaining for maturation of the larvae before freezing weather sets in. If the eggs do hatch and the larvae survive, the apple is probably picked and removed from the orchard. This also interrupts the life cycle and prevents pupation in the soil. All these factors are against a successful second brood, and may be the reason why it is so small. This may also explain why the species does not become established further south and is scarce in the southern limits of its range. 139 LOCATION OF LARVAL PUPATION When an infested apple falls to the ground, the apple maggot larva leaves it and burrows into the soil to pupate. The emergence of the fly the following summer can be in- fluenced by where this pupa is located. If it happens to be on the south side of the tree and close to the surface, emergence is expected to be earlier because the pupa was exposed to the optimum warming and development conditions. If it burrows deeper into the soil, or falls on the north side of the tree where it is shaded, emergenece the fol- lowing summer should be delayed. Experiments were conducted to determine if emergence is actually delayed on the north side of the tree, and to quantify the differences in the soil temperature in these different microhabitats so their influence on emergence will be known. Literature The literature is somewhat contradictory as to the effect of sun exposure on the emergence of flies. Glass (1960) reported that "emergence data showed that the pattern for paired cages was nearly identical under variable shading conditions." He concluded that "the location of a seeding within a test orchard was not a significant factor in time of emergence." Dean and Chapman (1973) said there are no- consistent trends to show differences in emergence from expOsed or shaded cages. However, Porter (1928) said "the flies emerged earlier from cages in the sun than they did 140 from similar material in cages in the shade." And Dirks (1935) said "emergence cages maintained in full sunshine have yielded flies ten to thirteen days earlier and the height of emergence has occurred ten to thirteen days earlier than cages maintained in full shade. It is evident that under orchard conditions infested apples dropping on the south side of the tree and exposed to sunshine would produce flies much earlier than drops completely shaded on the north side of the tree." It is also reasonable to assume the depth at which the larvae pupate would have an influence on emergence next year. O'Kane (1914, p82) found pupae among the upper grass roots, but also discovered they are deeper. His data showed the depth distribution of the pupae to be: 1/2" = 5.7%; 1" = 14.3%; 1 1/2" = 28.3%; 2" = 24.8%; 2 1/2 = 9.0%; 3" = 8.2%; 3 1/2" = 4.1%; 4" = 1.6%; 4 1/2" = 3.2%; and 5" = .l%. At these depths the soil is cooler and the pupae should develop slower. Materials and Methods Emergence cages were set over naturally infested ground on the south and north side of the tree in the K.S.H. orchard. In 1978, 10 pairs of cages were set on June 23 and in 1979 8 pairs of cages were set on June 18. They were monitored daily in 1978 and three times a week in 1979 until emergence occurred in each cage. After first emergence, monitoring was reduced to twice weekly. This will provide 141 data to show whether a difference in emergence times occurs due to the location of the pupae under the tree. To determine differences in temperatures where the pupae were located, three lead Weather Measure recording thermographs were used. The influence that depth of pupa- tion might have on emergence was determined by placing one probe of the thermograph on the soil surface in the grass litter zone, and the other one directly below it two inches under the soil. Any differences due to differential shading between north and south sides of the tree were measured by placing temperature probes at the two inch level on both the north and south sides of the tree. Graphs were made of the temperature fluctuations, data was transcribed and entered into the computer where degree days at base 48°F were calcu- lated and accumulated. Appendix 3 contains the daily degree day accumulations calculated. These values should explain some of the difference in emergence between the different microhabitats. Results and Discussion In 1978, apple maggot flies emerged first on the mean date of June 29 from cages placed on the south side of the tree (Table 36). Mean first emergence was July 2 from cages o; the north side of the tree. By averaging the difference 142 Table 36. First Emergence of Apple Maggot Flies From the South and North Side of the Tree at the K.S.H. Orchard in 1978. ' Date of First Emergence Two Inch Soil ddl Variety South North Diff South North Diff Transparent 6-27 6-28 1 662 615 -50 Dutchess 6-27 7-5 8 662 747 85 Greening Never 7-5 - - 747 - Dutchess 6-27 7-1 4 662 676 14 Greening 6-27 7-5 8 662 747 85 Red Delicious Never 6-28 - - 615 - McIntosh 7-4 6-29 -5 802 636 -166 Northern Spy 6-27 6-29 2 662 636 -26 Red Delicious 7-2 7-11 9 764 867 103 Northern Spy Never Never' - - - - Mean 6-29 7-2 3.9 696.6 698.4 -5.7 1 Degree days at base 48°F; B.E. Method 143 between the two cages from the seven pairs of cages where flies emerged, a 3.9 day delay was present. When the soil degree day accumulations on the date of emergence are recorded for the south and north side, the differences disappear. The mean difference between the two locations is only 5.7 degree days, which is much less than one day. Therefore, in this test soil degree days explained 4 calendar day variation in the emergence dates on the opposite side of the tree. In 1979, the difference in emergence from cages placed on the south and north side of the tree was 7 days (Table 37). In every pair of cages that caught flies, the north side was always later, and the range of the delay was 2 to 16 days. When soil degree day values were recorded on the dates of first emergence, much of the variation in calendar day differences was explained. The mean difference in soil degree days between the south and north side of the tree was 72.8 units, which is equivalent to four days. Therefore, the 7 days calendar day difference was reduced to just three days difference when soil degree days were used. As was expected, the two inch soil temperature is cooler on the north side of the tree than the south side on the same calendar date. Table 38 shows the accumulations 144 First Emergence of Apple Maggot Flies from the South and North Side of the Tree at the K.S.H. Table 37. Orchard in 1979. Date of First Emergence Variety South North Diff Transparent 6-25 6-27 2 Dutchess 6-25 6-29 4 Greening Never 7-2 - Wealthy 6-25 6-27 2 Jonathan 6-27 7-13 16 McIntosh 6-29 7-9 10 Red Delicious Never 7-4 - Northern Spy 7-4 7-11 _1 Mean 6-27 7-4 6.8 1 Degree days at base 48°F; B.E. Method Two Inch Soil dd 1 South North Diff 781 764 -17 781 799 18 - 851 - 781 764 -17 810 1059 249 847 970 123 - 884 - 2.3.9 _1011 -8_1_ 821.7 887.7 72.8 145 Table 38. Soil Temperatures Measured on the South and North Side of the Tree at the Two Inch Depth Recorded as Accumulated Degree Days Since April 1. Date South North South North South North South North 6/20 555 485 542 489 697 647 528 482 6/25 633 567 621 557 781 735 613 571 6/30 719 655 725 656 864 818 751 677 7/04 789 726 802 747 930 884 794 755 7/09 909 844 908 831 1014 970 898 865 7/14 1022 957 1003 919 1128 1083 1012 985 7/19 1143 1086 1103 1017 1238 1190 1131 1109 Mean 64.3 69.7 46. 35.3 Diff. Degree Days calculated at base 48°F; B.E. Method 146 during the emergence period of the apple maggot fly. These degree day differences are equivalent to two to three calendar days each of the four years the temperatures were recorded. These results agree with those found in Tables 36 and 37. Soil temperatures were measured on the soil surface in the grass litter zone and two inches directly below that in the soil. The variation in these two habitats can explain some of the variation in the range in dates of first emer— gence. The mean difference ranged from 29 to 94 degree day units (Table 39), which corresponds to two to five calendar days. Therefore larvae that pupated near the soil surface can emerge as adults two to five days earlier than larvae that pupated two inches deeper in the soil. Conclusions Calendar day differences in first emergence of apple maggot flies between the south and north side of the tree was 3.9 days in 1978 and 6.8 days in 1979. Soil degree day differences at these two sites were 5.7 in 1978 and 72.8 in 1979. In 1978, the soil degrees removed all the calendar day variation. In 1979, the soil degree days removed four calendar days of variation. An evaluation of degree days under the tree was made.. Over a four year period, the mean degree day difference between the soil surface on the south and north side of the Table 39. Date 6/20 6/25 6/30 7/04 7/09 7/14 7/19 Mean Diff. 147 Differences in Degree Day Accumulations at the Soil Surface and Two Inch Depth at Base 48°F (B.E. Method) on the South Side of the Tree at the Upjohn Orchard. 1977 19 Two Surface Inch Surface 683 586 545 759 663 621 848 751 719 921 823 790 1038 945 890 1143 1052 979 1268 1179 1079 94.4 28.9 78 1979 19 0 Two Two Two Inch Surface Inch Surface Inch 515 556 501 445 402 588 621 567 727 500 690 692 639 622 596 762 748 718 695 672 864 815 786 803 781 951 909 886 919 887 1051 995 971 1045 979 38.3 34.1 148 tree ranged from 35 to 64 units, or an equivalent of two to three calendar days. The soil surface and two inch soil degree day accumulations varied from 29 to 94 units in the same four year period, which is equivalent to two to five calendar days. These differences supported those above which help to explain the calendar day variation in emer- gence at the different sites. 149 PHENOLOGICAL PREDICTIVE MODEL OF EMERGENCE In the past, calendar days have been used to determine when to initiate sprays for apple maggot control. For example, July 4 has been the recommended date in West Central Michigan. (Klackle, 1980). By observing Table 3, the range of first capture on traps within the same orchard and in the same year may be as great as 31 days. Table 2 shows that between years and sites, this range may be as great as 41 days. Therefore a more reliable method than calendar date is essential to provide economic control of this pest in specific orchards. Within the past few years, phenological models to predict apple maggot fly emergence have been developed. These models are based on measuring weather factors, and then predicting when emergence will occur based on them. Most predictive models use the daily maximum and minimum air temperatures starting on April 1. From these values a certain amount of development is said to occur based on the amount of accumulation of heat over the developmental threshold. This amount is accumulated each day until the critical event to be predicted occurs. Daily accumulations are made by averaging the maximum and minimum temperatures or by integrating the area under tue curve of the temperatures for that day (Baskerville and Emin, 1968). Several researchers have made predictions for certain events in the life cycle of the apple maggot, and those of 150 first emergence are found in Table 40. Trottier (1975) reported that based on degree days alone, first emergence occurred at 425 :30 degree days at base 9°C using four years of trap data. This predicts emergence to within plus or minus two days at the experimental orchard at Vineland Research Station, Ontario. Reid and Laing (1976) reported that flies would emerge at 550 degree days at base 8.7°C. Lastly, Reissig 33 a1 (1979) found trap catch correlated well with 641 degree days at 6.4°C. The generality of all these predictions to local growers is questionable. If emergence occurred early and growers waited to spray, damage would occur. Contrary, if flies were late emerging but sprays were applied early, they would be unnecessary. Reid and Laing (1976) also noted that all the predictions were based on single orchard data. They found that Neilson (1962) emergence data was off 24 days from their prediction leading them to believe that local populations of apple maggot flies respond differently to the environment. Other researchers have questioned the validity of models to predict the emergence of apple maggot. Dean and Chapman (1973) reported that emergence could not be pre- dicted by use of weather and that average calendar date was not accurate based on more than 40 years of cage emergence data from Hudson Valley, New York, and suggested that cages be set out and monitored to determine first emergence. 151 Table 40 Predicted Emergence of Apple Maggot Fly Based on Air Degree Day Accumulations. Developmental First Location Threshold Emergence Variance Source Vineland, Trottier, Ontario 9.0°C 425 B.E.* : 2 days 1975 Guelph, Reid & Ontario 8.7°C 550 - Laing,1976 Geneva, Reissig, et New York 6.4°C 641 i 3% days al, 1979 * Baskerville Emin (1968) method of determining degree day accumulations was used. Glass (1969), based on six years of caged emergence data in west New York, said that average temperatures in May, June and July effect peak emergence, but that first emergence can not be predicted by weather but must be monitored using caged apples. Even Reid and Laing (1976) said "emergence data will be of limited predictive value since there is such wide variation in emergence dates." These predictions might be improved or more widely applicable if other parameters are included in the model besides air temperatures. Soil factors have been incorpo- rated into an emergence model of Rhagoletis cerasi (Leski, 1963) which increased its accuracy greatly. Soil temperature may be the most important soil related factor to utilize to predict emergence, as this could account for the great difference in emergence due to the microhabitat where the pupa is located. Soil moisture could also increase the accuracy. Mundinger (1930) said "it seems reasonable to believe that soil moisture may be a factor of some im- 152 portance in emergence time since the greatly reduced emergence of 1927 and 1929 paralleled correspondingly dry months." My speculation is that environmental factors must effect the physiological development of the insect, and that by utilizing different methods of accumulating degree days and precipitation, a discrete threshold should be reached that would be a reliable estimate of emergence at a par- ticular site. Several environmental and biological factors are known to effect time of first emergence somewhat, and eventhough they are not easily quantified, will be included in a predictive matrix. Season long emergence of apple maggot fly can be correlated to environmental parameters. Lathrop and Dirks (1945) said "the highly significant correlation is interesting, and emphasizes that temperature is a most important factor influencing the seasonal flow of emergenCe of flies." Materials and Methods First Emergence. The majority of the data that provides a basis for this predictive model has already been presented. The approach will be to start at a mean air degree day value, then add or subtract a specific number of units depending on the parameter measured. All the data was generated at the K.S.H. orchard. Therefore, the predictive model will be most representative of that location. At other locations where different conditions such as 153 population size, etc. exists, this prediction may not be accurate. As was previously discussed, catch on the yellow Zoecon AM traps provides a good estimate of fly emergence. Table 5 showed that flies were caught on traps in the same trees under which natural emergence cages were placed on the same day as they emerged in the cages. Traps are more economical and easier to use. Therefore, trap catch will serve as the indicator of fly emergence in this phenological model. Season Long Emergence. This was correlated with air degree day accumulations. Percent accumulative emergence values were transformed by (flue arcsin transformation to straighten out the curve. Least squares liniar regression was then performed, and the corresponding R2 values calculated. Lastly, a multiple regression of air degree days, soil degree days, and percent soil moisture accumulated after first catch was correlated to the dependent variable of accumulated percent trap catch on the yellow Zoecon AM trap and the red sphere trap. Results and Discussion First Emergence - Mean first emergence of apple maggot fly occurs at 1117 air degree days base 48°F (8.9°C) by the B. E. method (Table 8). Using the yellow Zoecon AM trap as the indicator, this estimate was within :58dd or 2.5 days at the K.S.H. orchard over a four year period. 154 Two other abiotic components were investigated as to their influence on first emergence. Eventhough considered a very important parameter, soil temperature did not increase the accuracy of this estimate as shown by a larger C.V. value (Table 11). The mean emergence occurred at 679 dd (base 48°F) :167 or 7 days at the K.S.H. orchard over the same four year period. The last important abiotic parameter measured was soil moisture. This data was not valuable in predicting first emergence, but might be useful in measuring season long emergence. Several biotic factors were also investigated for their influence on first emergence. The first biotic parameter of importance is variety the fly was reared in (Table 21). When four years' data are combined there is no statistical difference between the first emergence times. However, there is a range of values that can be incorporated into a generalized model (Table 41). By examining these values the deviation from the mean for any variety can be determined. The second biotic parameter investigated was carry over pupae. It was presumed that since a pupa has been in the ground for two seasons, it would mature physiologically the first year and emerge sooner the second year. This did not occur (Table 27) as the mean emergence for second year flies was delayed three days from first year flies. There- fore this parameter can be ignored in this predictive model. The orchard floor culture can have a marked influence on time of emergence. When measuring emergence in one 155 Table 41 Influence of Variety of Apple Reared in on Prediction of First Emergence of Apple Maggot Fly Variety Reared In Air Degree Days1 Dutchess -75 Wealthy -61 Transparent -41 Winesap -22 Greening -14 Jonathan - 7 Red Delicious - 1 McIntosh +38 Northern Spy +47 Baldwin +51 1 48°F; B.E. Method orchard, this is not important. However, between orchard differences may be great (Table 32). This is due to the amount of soil shading in the orchard which directly influences the soil temperature degree day accumulations. The varying amount of heat accumulation can greatly alter the time of emergence of the files. For this particular estimation orchard floor cultures will not be included, but for a more generalized model this parameter should be incorporated. Table 42 presents degree day values to subtract from the mean value of emergence based on the orchard floor culture. These values were calculated from differences in the accumulated heat units from April 15 to June 30 found in the different experimental orchards. Another biotic component studied was the effect a partial second generation could have on time of emergence. This has no effect on first emergence so will not be in- 156 Table 42 Influence of Orchard Floor Culture on Prediction of First Emergence of Apple Maggot Fly. Orchard Floor Culture Air Degree Days1 Shaded by tree tall grass 0 Shaded by tree mowed grass -13O ..- Exposed to sun mowed grass -417 1 48°F; B.E. Method cluded in this model. However, it does influence the late season portion of the season long model slightly, and therefore will be used later. The next biotic component investigated the relationship between first emergence and where the pupae were located. If they were on the south side of the tree and in the grass litter zone, then a normal emergence pattern should appear. If no apples fell on the south side of the tree, but all were on the north side, then a 73 dd or 3.9 day (Table 36) delay could be expected in emergence the following year due primarily to the cooler temperatures on the shaded side of the tree and the resultant slower development of the pupa. If all the larvae burrowed down two inches to pupate or the soil was disced or died out to the extent that all pupae in the upper two inches desicated and died, a delay of 49.0 dd or two days could be expected (Table 39). If pupae were only present at the two inch level on the north side of the tree, then another 54dd or 2 days could be expected before first emergence (Table 38). These values are summarized in 157 Table 43. From this, the relationship of where the larvae pupated can influence emergence of the individual, but in a population sense it would have no effect on first emergence in an orchard. Table 43 Influence of Location of Pupation on Prediction of First Emergence of Apple Maggot Fly. Location of Pupation Air Degree Days1 South Side of Tree Surface ---- South Side of Tree Two Inch Depth‘ - 49 North Side of Tree Surface - 73 North Side of Tree Two Inch Depth -103 1 48°F; B.E. Method All of these parameters investigated are incorporated into the 1117 dd value for predicting first trap catch of the apple maggot fly at the K.S.H. orchard. This provided a $2.5 day estimate over a four year period. However, when these parameters are used to predict emergence at other sites, the accuracy is greatly reduced. Table 44 shows the actual verses predicted first catch at several other sites. All of these sites are either abandoned orchards or abandoned or wild trees around commercial orchards. They all had the yellow Zoecon AM trap placed and changed just as in the K.S.H. orchard. The degree day values presented were 158 TABLE 44. Predicted First Trap Catch of Apple Maggot Flies with the Yellow Zoecon AM Trap in Abandoned Trees Around Commercial Orchards. Actual Predicted Errorl Location Date ddi Date QQE Days dd: Upjohn 1977 06/20 1108 06/20 1110 0 -2 Upjohn 1979 07/13 1102 07/13 1110 0 -8 Upjohn 1980 07/05 999 07/05 1110 0 -11 Hofacker 1980 06/21 705 06/14 625 7 80 1 1979 08/07 1777 07/10 1117 28 660 2 1979 07/12 1185 07/09 1095 3 9O 3 1979 08/07 1777 07/10 1117 28 660 4 1979 08/21 2003 07/12 1164 40 839 6 1979 08/07 1777 07/10 1117 28 660 8 1979 07/27 1525 07/12 1164 15 361 9 1979 09/05 2308 07/10 1116 57 1192 10 1979 07/10 1131 07/10 1117 0 14 11 1979 07/31 1623 07/10 1117 21 506 12 1979 07/24 1457 07/10 1117 14 340 15 1979 07/31 1623 07/10 1117 21 506 18 1979 07/31 1623 07/10 1117 21 506 19 1979 08/07 1777 07/10 1117 28 660 21 1980 07/21 1373 07/10 1117 11 256 22 1980 08/11 1837 07/10 1117 32 720 23 1980 08/06 1718 '07/10 1117 27 601 24 1980 07/16 1245 07/08 1056 8 189 25 1980 08/18 1825 07/15 1117 34 708 26 1980 07/10 1006 07/12 1056 -2 -50 27 1980 07/23 1429 07/11 1117 12 312 28 1980 08/18 1974 07/13 1164 36 810 29 1980 08/08 1786 07/11 1117 28 669 30 1980 07/25 1469 07/11 1117 14 352 31 1980 07/23 1429 07/11 1117 12 312 Range 77 1603 31 539 59 1242 Mean 7/27.5 1521 7/8.8 1099 18.7 426 1 before it was predicted to occur. 2 Negativer sign indicates number of dd actual dd = Accumulated Degree days base 48°F, B.E. event occurred Method 159 calculated at base 48°F; B.E. method from the nearest official weather reporting station. As can be seen, the range of actual first catch in these sites was 77 days or 1603 dd. With the predicted catch the range was reduced to 31 days or 539 dd. This leaves an error that ranged from -2 to 57 days and -50 to 1192 dd. This proves how inadequate a generalized model is that attempts to predict emergence at any location, as was speculated by Reid and Laing (1976). There are several reason that might help to explain the large error in this prediction. First there might be some biological differences in the populations between the K.S.H. orchard and the more northern sites as several authors have suggested. The Upjohn prediction was very close for all three years with the predicted and actual being the same date. The Upjohn orchard is within 10 miles of the K.S.H. orchard, and those flies appear to respond to air degree days in the same manner. The rest of the orchards are 30 to 100 miles north and the flies may respond differently to photoperiods, angles of the sun, air degree days, etc. Another reason could be the proximity of the weather stations to the abandoned trees. The one used for developing the model was located in the K.S.H. orchard, while those at the other sites may have been as far as 20 miles away from the test site. A possible major reason for this very poor predicta- bility of first catch is population size from year to year. In the K.S.H. orchard a very stable mature population exists year after year. In these other sites the population 160 fluctuation can be very great as pesticide drift, host freeze outs, and many other factors influence it. This was tested by running a regression on the number of flies caught per trap against the dd error of first emergence. A R2 of 0.15 reveiled a very small relationship between the popula- tion size and error of prediction. Therefore some other factors are much more important. Another factor is the actual population size. With smaller populations, the tails of the population curve are much closer to the mean. This indicates that there are fewer individuals extremely early or extremely late. This fact alone could account for many days delay in first catch. To test this, the K.S.H. orchard data was reworked to determine the mean 1 and 5 percent trap catch air dd accumu- lations. The mean 1% emergence over the 4 years occurred June 19.7 or 1.1 days later than mean first catch. The air dd accumulations were 1117 for first and 1150 for 1 percent. This did not help reduce the error in the prediction of trap catch in the abandoned trees. Likewise the 5 percent mean trap catch occurred July 4.5 or 1256 air dd. Predicting 5 percent emergence at the abandoned sites also had the same error range. Therefore this did not increase the relia- bility of the model. Lastly, experimental error such as thermometer cali- bration could cause a several day difference in the pre- diction date (all thermometers were calibrated to try to reduce this error). All these factors help to explain why this predictive model is not very accurate at different 161 sites. The results is that it should not be used to predict emergence at locations other than K.S.H. or Upjohn, and at those other sites traps should be used to measure first emergence. Season Long Emergence. Accumulative air degree days is a very reliable tool to estimate the cumulative percent trap catch of apple maggot flies at the K.S.H. orchard. Figure 16 shows the typical sigmoid population curve for the accumulated average trap catch on the yellow Zoecon AM trap in 1978. A least square linear regression of this gave an equation which resulted in a coefficient of determination of .9126. This is an exceptionally good fit for any biological data. When this data was transformed by the arcsin trans- formation which tends to straighten out the ends of accumu- lative percent data and a least squares linear regression performed on the resultant data, a R2 value of 0.9643 resulted. This also indicates that accumulative air degree days at base 48°F (8.9°C) by the B.E. method is a very reliable predictor of percent accumulative trap catch. To test whether this was a fortunate circumstance in 1978 or not, the same procedures were completed for trap catch at another site ‘with another trap and. other' years (Table 45). This does show that air degree days alone accounts for about 95% of the variability between the actual and predicted relationship. This was not significantly improved by transforming the data, so the raw data can be used for predictive purposes. Because the data fit so well, all four 162 .mhoa on ouosooo Hoowomom ouoom oouoemaox may no onus z< .coomoN 3oHHm> may :0 mmflam uommmz waded mo condo mane unmoumm m>wumaseaoom .oH musmwm 82:22 .m.m r73. 233 23o 2:25 .2 ooo.... ooou oopw ooo. oo: ooo. com: oo. . com ..o. ..ow w 0 0 ..oo m x m \\. .Lu? MW .o O 9 3.“... :2. m. «minuxhoosoood m :3 1 O U. ....Om ..oo ..oo. 163 mom. OHHOO. mNm. mHmoo. Hem. mHHoo. mwm. memooo. Ohm. hmooo. mmw. whooo. van. bmmooo. mom. oHoooo. vmm. mHHoo. mmm. oomooo. Hob. moHoo. vmm. mHmooo. mmm. «Hmooo. mam. ammooo. Nvm. NOHOO. vmm. monooo. ©h¢. mmmooo. Hem. mmoooo. mmm. NHHOO. vmm. vumooo. mmm. omhooo. mew. Nmmooo. Hhm. mamooo. mom. mawooo. mmm. ommooo. mom. mmoooo. vow. mNHHoo. Mam. wmbooo. Hum. maoooo. mam. wmvooo. «M m oocuoz .m.m “Aoom.mv momv mmmm um mama mmumma MH< m>flumHDEsoo< mhv.HI moo.HI wom.al wmo.al mmN.HI mmo.HI who.at mmah.l mmv.HI mwom.l mov.HI mvo.HI NNH.HI bmmn.r mHN.HI ammh.l mmo.HI mmmo.) vmm.HI mwmm.l vmom.l mom.l Hmo.HI wmvo.l Hho.HI Hmon.l H©H.HI NMN.) mnan.l Nonv.l d camoud 3mm camoud 3mm camoud 3mm camoud 3mm camoud 3mm :flmoud 3mm camoud 3mm Camoufi 3mm choud 30m :Hm0H¢ 3mm camoué 3mm Camoud 3mm camoud 3mm camou< 3mm cflmoug 30m mama Umm Cmm 66% @mm 6mm @wm soaao» soaflo» zonao» 3oHHo» goaao» soaaos soaaow zonao» Umm 6mm Umm Umm fimm Umm soaao» soaao» zonaow 3oHHo» goaaox zonaos gondo» 3oHHo» soaflo» soaao» NMHH. Hafl Add omma ommH mhma mhma HH< Hdfi omma omma mbma mhma bhma hhma HH< HH< ommH omma mhma mhma HH¢ Had omma omma mhma mhma mhma mhma hhmH fihma Hmmfi anoflmb cache: anonmo anonms agenda enema: gnonma anomma anoflmo anoflmo anoflmo c50an csonms C ..C.‘ O 'h 0.. D :1: mmmmmmmmmm 213333323213 mmmmmmmmmm (DUJUJUJUJCD KMKMMKMMMX MXMXKM cofiumooq ou soumu none #cmoumm m>HumH5Esoofi mo cofimmmummm Hmmcaq mmumsqm ummmq .mv mamas 164 years data at the K.S.H. orchard were lumped to generate a generalized predictive equation for that orchard. The resultant equation (4) gave an R? of 0.849 which is an excellent correlation over a four year period. Eq. 4. y = .0005524 x - 0.505 Multiple regression was performed on the 1979 and 1980 K.S.H. orchard data. The dependent ( ) variable was the accumulative percent catch on the yellow Zoecon AM trap or the red sphere trap. The independent variables started with the first catch and were accumulative air degree days (X1), accumulative soil degree days (X2), and accumulative soil moisture (X3). As previously shown, each of the variables can predict accumulative percent trap catch with a great deal of accuracy. This was confirmed in this analysis. When using the yellow trap as Y, x alone gave an 1 R2 = 0.986, X2 alone gave a R2 = 0.989, and X3 alone gave a R2 = 0.984. The multiple regression gave a R2 = 0.982 which is lower than any of the variables alone. In 1980 with the yellow trap as the Y, Xl alone gave a R2 = 0.983, X gave a 2 R3 = 0.985, and X3 gave a R2 = 0.980. The .multiple regression gave a R2 = 0.975 which is again lower than any one of the variables alone. When combining both years data together and Y being accumulative percent catch on the yellow trap, the x1 gave a R2 = 0.949, X2 gave a R3 = 0.961, and X3 gave an R2 = 0.911. The multiple regression shown in Equation 5 gave a R2 = 0.975. Eq 5 y = - 8.91 - 0.06X + 0.12x + 0.67X 1 2 3 When using the red sphere trap as the dependent variable Y, 165 the same analysis as above gave in 1979; X1 - R2 = 0.969, X2 - R2 = 0.976 and X3 - R2 = 0.978. This multiple regression gave a R2 = 0.968 which is again lower than any one of the independent variables alone. In 1980 the same analysis gave: X1 - R2 = 0.983, X2 - R2 = 0.984, and X3 - R3 = 0.978. The multiple regression gave a R2 = 0.975 which is again lower than any variable alone. The regression with both years data lumped together gave the following results: Xl - R2 = 0.939, X2 - R2 = 0.952, and X3 - R2 = 0.903. This multiple regression found in Equation 6 gave a R2 = 0.966. Eq 6 = -14.35 -0.08X + 0.13X + 0.08X 1' 2 3 These equations, or the single linear regression equations, could be used to very accurately predict the accumulative percent trap catch at the K.S.H. orchard. Conclusions When attempting to predict first emergence of the apple maggot fly at the K.S.H. orchard, trap catch on the yellow Zoecon AM trap occurred at 1117 air degree days base 48°F 12.5 days over a four year period. This prediction even held at the Upjohn orchard where trap catch occurred on the predicted day all three years. The incorporation of soil temperature and soil moisture did not improve the predictability. The influence of variety and orchard floor culture were incorporated into the model and it was used to predict first trap catch at 25 different sites. This did not give good results as the actual event ranged from -2 to 57 days from the predicted event. The calendar day range 166 was 77 days. The land 5 percent trap catch prediction also had this same large error. Therefore in other sites traps should be used to determine flight activity and when to initiate controls rather than relying on this predictive matrix. Accumulative air degree days provides an exceptionally good means of predicting accumulative percent trap catch on the yellow Zoecon AM trap and red sphere trap at the K.S.H. and Upjohn orchards. A least squares linear regression on the four years data gave an R2 of .85 for the resultant equations. Multiple regressions performed discovered that it did not improve the predictability by incorporating soil degree days and soil moisture over that achieved by air degree days alone. 167 PART 2 MANAGEMENT STRATEGIES IN COMMERCIAL ORCHARDS Under the current management scheme for apple maggot, commercial growers are advised when to time the first ap- plication by experience, professional advisors, or the COOperative Extension Service (CES). The method employed by the CES staff is to place traps in abandoned trees where known infestations exist, and monitor for emergence. Once it has occurred, they alert growers that in 7-10 days the first application should be made. From then on applications should be made at two week intervals. This regional method is suitable for the CES because it's role is to advise all growers when first flies have emerged. For the great majority of the growers, this strategy would require more sprays than are necessary. Most growers have the pest under control, have limited outside reservoirs of flies, and have a greatly reduced population to manage. A better strategy for them would be to place traps in abandoned trees on their own farm, and monitor emergence there. Based on this approach, control timing would coincide with insect pressure in their own orchard. The best approach however, is to monitor apple maggot flies within the orchard. With little risk on the growers part, this approach should provide them with the most efficient and reliable method of preventing fruit damage. This approach is being inplemented by most IPM programs and private consultants. 168 Figure 17 illustrates these approaches in 29 different test orchards. Line I\ represents the regional approach where after first catch in an area, biweekly sprays are recommended. Using this approach five insecticide applica- tions are needed for apple maggot control. On Lines B and C, the dots indicate first capture of apple maggot flies on the yellow Zoecon AM trap. By trapping in abandoned trees around commercial orchards (Line B), most growers can delay their first maggot spray. The average date of first catch was August 4, indicating that for the average grower two sprays were applied for apple maggot flies that were not necessary. Line C is the preferred method. Growers or consultants can trap flies in grower orchards and wait until flies are caught in the orchard before they spray, assuming other pests are under control. Each dot represents the first catch on a trap optimally placed in the perimeter row of the orchard. If sprays were applied based on this strategy several applications would not be necessary. To note, flies were never caught in seven of the 29 orchards monitored, and in those no applications would have to be made this season for apple maggot. To most efficiently use the in-orchard monitoring scheme to control the population that could potentially infest their orchards, the grower needs to know and understand several factors involved with monitoring the apple maggot. These include the best method of monitoring such as what trap type should be used, where should it be .Apm>o>nsm monocouo may momma Ed coowoN 3oHme do muasp< mo coumu done no Ummmm mmflam pommmz madam mcflaaouucoo MOM mmflmmumuum .ha madman mmmfim Pawm ...maosd >4: ... wZD... mm o. o _ mu m. : c mm a S k on 3 P — _ — _ _ _ _ _ _ _ _ P _ w flD-bOaOOOp—SS3N... O u C 00:. u C”. O.” C 1 . . 845% Ed . ...:....-..-:...n.........u..................-uufluin..uu C O” n O O n u owzoozw>m3m momdzomo mm 92¢... Bog-Am» zo mun—:04 “.0 10.58 a4": 20 owmdm 5.00952 w4¢a< 02......omhzoo mo“. >om._.mH mo. mag um ucmumoman saamoflumaumum no: mm. mm. mo.mH cm.Hm nm.o nfi.o mm. mm. um.mH om.mH nm.o om.o Hm. vm. wo.n cm.o om.H ov.o Hm. mm. MH.n cm.m hm.m mm.m hm. no. an.va m>.> Ho.m Ho.m pom zoaamw pom 30Ham» pom 3OHH0> Anmv am Op pmcoccmnm Eoum m®u9\noumo noumu acne Nmaoo mum Hmpuma wEmm Hm\mo om\mo mH\wo NH\mo mH\mo mo\mo ma\mo HH\mo om\wo mH\mo Umm 3Oaamw noumu umuflm .ummu MEG may >3 pm3oHH0m mommz commam v 30m nausea commam m 30m chase Ummmwm v 30m pcoomm pmmMflm 0H 30m umumefiumm EHOMHca m 30m Hmumeumm .oz coaumooq .mumumfimumm noumo moms com: uamanMflo ou coflumamm CH mpuocouo HMHOHmEEOU cw ucmfimomam moms mo newpmooq .mv mamma 184 spheres. Therefore, this placement scheme is not recom- mended. Equidistant. perimeter trap ‘placement. was greatly improved by placing traps in perimeter trees, but biasing those trap trees selected so they are near potential infes- tation sources. This was tested in 16 orchards, and the overall results are very good. The mean date of first catch was August 11 using the yellow trap and August 19 using the red sphere, the second earliest of the methods tested. The mean catch per trap was the highest of all the trapping schemes tested using the red sphere trap and second highest using the yellow traps. Therefore biasing the traps toward infestation sources can be recommended to determine when controls should be initiated for apple maggot. The last approach is to place the traps at different distances into the orchard. This might be desirable to determine how far in the orchards the flies travel. 'It has been suggested that a grower could spray just the perimeter rows of his orchard late in the season when apple maggot is the only pest present. Table 47 indicates that second row trapping is the best scheme. It caught flies first and had the shortest delay from abandoned trees. However, the sample size is too small to make valid assumptions and recommendations. This practice by all growers will not be recommended . The relationship of trap catch and percent fruit damage is a very important parameter to use when determining which trap type and placement is best. This will be more fully 185 discussed later. To note here is that the scheme that catches the least flies provides the best coefficient of determination. If these small sample sizes actually express the real situation, then one would want to place traps on the fourth row into the orchard and biased toward the infestation sources. However, this probably would not be suitable because that placement caught the first flies last, caught the fewest flies, and had the longest delay from the abandoned trees. This placement should not be considered because control measures were based on trap catch on the perimeter trees in these orchards, and if they were delayed for catches on the fourth row traps, a different set of damage values would have been used to calculate the R3 values. Table 50 indicates that perimeter spraying is feasible if no resident flies are present. The sampling scheme of biasing traps toward the four most likely infestation sources and placing traps in each of the four perimeter rows at each site was replicated in five orchards. A two-way analysis of variance showed no statistical difference between the trap catch from row to row. However, a steady decline in catch occurs, and by the fourth row in the orchard only .05 flies per trap were caught in these five orchards. This indicates that a grower could, with minimal risk to his crop, spray just the perimeter four rows of his orchard for apple maggot when no resident flies are present. This is therefore a feasible alternative to reduce costs for apple maggot control. 186 Table 50. Migration of Apple Maggot Flies into Commercial Orchards as Determined by Yellow Zoecon AM Trap Catch (5 orchard sample - 4 reps/orchard). Row into Orchard Mean Accumulated Catch 1 .45a 2 .30a 3 .25a 4 .05a Means followed by the same letter are not statistically dif- ferent at the .05 level by the DMR Test. Trap Density. Traps were placed in four hectare blocks in an attempt to determine the density or how many trapping locations are adequate. Fewer traps are desired to reduce material and labor costs, but sufficient traps are needed to reduce the risk of missing flies and occurring fruit damage. Table 51 shows that the mean first catch date is sooner on the yellow traps than the red spheres for each density. Also the trend is present that if first fly catch is most important, 16 traps should be placed in the orchard. However, whether by random chance or experimental error, two trap locations gave the very earliest trap catch. There- fore, by proper placing traps near outside infestation sources, two trap locations should be sufficient. The fluctuation in the mean catch per trap is not dependent upon density because these traps are generally located at least 100 yards apart, and their effective range~ is probably limited to a few feet. What this does indicate however, is the relative population of apple maggot in each 187 orchard. As can be seen, these orchards do vary in the average number of flies per trap. In most cases, the red sphere caught more flies than the yellow trap, but this is due to them catching more males. Two traps per 4 hectares appears to be the property density for catching the most flies per orchard. The 10 trap orchards can be ignored because all of them were discovered to have resident populations of flies. This does show however that the trap catches are indicative of the natural population of flies in these orchards. The two trap per block density is also preferred when minimizing the delay in trap catch from abandoned trees. For each trap type, the time difference in catch between abandoned trees adjacent to the orchard and in traps on the perimeter row nearest the source was the shortest. Table 51. Relationship of Trap Density to Catch of Apple Maggot Flies. Den- No. Mean Mean Mean Delay sity Orchards First Catch Catch/Trap from Abandoned Yellow Red Yellow Red Yellow 33g 1 6 8-12 8-27 2.33 6.17 10.7a 22.0b 2 7 7-30 8-14 5.09 6.92 4.8a - 3.0a 4 6 8-15 8-22 0.75 1.83 11.0a 24.5b 10 3 8-7 8-15 12.87 4.63 12.5a 5.0a 16 5 8-6 8-9 0.26 0.32 8.4a - .25a Means followed by the same letter are not statistically dif- ferent at the .05 level by the DMR Test. 188 Conclusions Trap Type Comparison. Based on the data gathered from the 31 commercial orchards, the yellow Zoecon AM trap is the preferred trap to use for monitoring apple maggot flies in commercial orchards. They catch flies sooner, catch gravid female flies sooner, and catch females later in the season than the red sphere trap. Trap Placement. Yellow Zoecon AM traps should be placed in the perimeter row of the orchard, and biased toward the outside orchard sources of infestation. With this placement, flies will be caught sooner than most other placement schemes, more flies will be caught, and the delay in catch from abandoned trees will be reduced over other placement schemes. By placing traps in consecutive rows into the orchard, it was found that flies do not disperse widely throughout the orchard, but generally stay in the outside rows. This can greatly aid management and reduce costs by spraying just ”the perimeter rows late in the season. Trap Density. Two yellow traps are sufficient to monitor a 4 hectare block if the traps are located near outside orchard infestation sites. This density provided the earliest mean date of first catch, the largest mean catch per trap, and the shortest delay in catch from abandoned trees. To reduce the risk associated with missing earlier flies, more traps could be set, but this risk 189 avoidance is accompanied with larger material and labor COStS. 190 DELAY IN CATCH OF AM FLIES IN COMMERCIAL ORCHARDS FROM OUTSIDE SOURCES As has been discussed previously in this dissertation, the arrival of apple maggot flies in the commercial orchard :us a very important parameter to measure for efficient control. Few orchards have resident flies, and the flies that invade the orchard must come from outside sources. These sources are generally abandoned and neglected trees in backyards, fencerows, or orchards. Most often there is sufficient food sources and ovipositional sites in these abandoned sites for fly develOpment. However, as over crowding and competition for available resources increases, the flies expend energy, disperse and risk the chance of finding other unexploited resources. Once they reach the commercial orchards, then some control measures need to be taken by the grower to prevent crop loss. Literature The dispersal and ovipositional drives in apple maggot can be very strong. ProkOpy (1978) found that flies in Dorr County, Wisconsin, left a fruitless orchard, found a green tomato he had hung in a birch tree one-half mile away and oviposited in it. This fruitless condition or lack of oviposition sites is one of ‘zhe factors involved with dispersion. It should be noted that flies reared on wild and abandoned apple as well as several species of hawthorn 191 can infest apples in commercial orchards (Reissig and Smith, 1978). Therefore all these sources should be trapped. Disperson is an evolutionarily stable strategy achieved by the apple maggot. Frequently apple and hawthorn flowers or young fruits are destroyed by early summer frosts. Also, many varieties of apple are naturally biennial and diseases can be severe and leave no fruit for oviposition when the females emerge. These factors tremendously reduce the potential oviposition sites, therefore, making it beneficial for a portion of the population to expend energy and disperse 1x) find untapped ovipositional sites. This is apparently what happens when flies enter the commercial orchard. Once there, they can oviposite and maintain the gene pool. An important criteria growers should know is the lag time between first emergence in abandoned trees and first arrival in commercial orchards. Reissig and Tette (1979) noted first trap catch in mid-July in abandoned trees and in early August in commercial orchards. Prokopy (1979) found in 6 orchards that this delay was 3-6 weeks. Practically this meant that growers who were advised to spray 7-10 days after first trap catch in abandoned trees and then every 14 days thereafter would have applied 1 or 2 sprays for a pest that was not even in the orchard. This delay in arrival in commercial orchards was estimated by trapping flies in outside sources near the commercial orchard and in the perimeter row of the orchard nearest to that outside source. 192 Material and Methods In each orchard monitored, both the yellow Zoecon AM and red sticky sphere traps were placed in abanonded trees and on the perimeter row adjacent to the abandoned trees in the commerical block. These were monitored and cleaned off, and the flies counted and saved for sexing. The females were dissected to see if they were gravid on their arrival 1J1 the orchard. Thirty-one commercial. orchards ‘with potential or known apple maggot damage were monitored. Data from three of these orchards was discarded after it was discovered there was a resident population of flies in them, hence the delay in movement into the orchard could not be measured in them. Results Table 46 shows that the mean date of first fly catch in abandoned trees around commerical orchards was August 1 on the yellow traps. When red sphere traps were used, this was delayed by six days to August 7. In the commercial orchard, the mean date of first catch on the yellow trap was August 10 and on the red sphere was August 15. The mean delay from outside sources to the orchard was 8.2 for the yellow trap and 7.9 for the red sphere. This was not significantly different. This indicates that on average, a grower or consultant could place a trap in an abandoned tree, and expect flies to be in his orchard 8 days later. However, the range in delay of catch on the yellow trap was from 17 days in the orchard before catch on the abandoned 193 tree to 35 days later on the traps in the orchard than in the abandoned trees, and with red sphere the respective range was from 14 days before to 29 days after. These extremely large ranges of about 7 weeks presents too large of a risk, and indicates the grower should monitor for apple maggot in his orchard, and not assume a mean of 8 days delay in being trapped in the orchard. Different trap placements were evaluated for their effect on the delay between fly catch in abandoned trees and commercial orchards. Table 49 clearly shows that traps should be placed either in the first or second row of the orchard, even though the differences in mean delay are not significant. When using either trap type, the second row is preferred by this data, but the small sample size precludes its general recommendation. The first row with traps uniformly distributed around the orchard was the best method when using yellow traps, and the first row with traps biased toward the outside sources was best when using the red Spheres. When all the criteria are considered for best placement, the first row biased scheme is preferred. Traps were also placed in orchards at different densi- ties to determine density effect on delay of catch. Ta- ble 51 showed some unexpected results. One assumes with greater densities, earlier catch would occur, and with smaller densities later catch would occur. This was the general trend, with the 2 trap density being the only outlayer. With both trap types, the 2 trap density provided 194 the smallest delay in catch from that in the abandoned trees. This may be due to randomness, but because of the other criteria used and the favorable results with this density, it is recommended for apple maggot monitoring in commerical orchards. Conclusions On the average, adult apple maggot flies were caught 8 days later in commerical orchards than in abandoned trees outside the orchard. However, the range in dates was -17 to 35 days using the yellow trap, and -14 to 29 days using the red sphere. Therefore, trapping in abandoned trees is not recommended for timing sprays in commerical orchards. Instead, traps should be placed in the perimeter row of the orchard biased toward the outside infestation sites. Two traps per 10 acres when biased toward the two likeliest sources provide the greatest chance of trapping flies as soon as they arrive in the orchard. 195 IDENTIFICATION OF FLIES ON TRAPS An important criteria to any control program is to be sure of the identification of the insect under inves- tigation. Using a visual trap with very low specificity like the yellow Zoecon AM trap or the red sticky sphere, very many different species of insects can be attracted to them or accidentally caught. An attempt was made in this study to collect representative specimens that might be confused with apple maggot. These specimens were then ' identifed and photographed. The final goal is to have these published in an Extension Bulletin so fieldmen and growers will have a identification tool at their disposal to cor- rectly distinguish the apple maggot from other picture winged flies that are caught in apple orchards. Literature Several researchers have attempted to identify insects caught on bait traps placed in orchards. Howitt and Connor (1965) studied different baits attractive to the AMP. The baits were used in conjunction with the yellow panel. They broke down the trap catch to orders, and discovered that the most abundant insects trapped by all baits were Diptera. However, seven other orders of insects were caught. Moore (1969) placed sticky-coated baited and unbaited red wooden spheres and yellow panels in apple trees to determine their effectiveness in catching AMP and beneficial flies. He identified the Diptera to family and discovered the great majority of the flies caught on all trap types were 196 Table 52. List of Picture-Winged Flies Caught on Yellow Zoecon AM Traps Placed in Apple Orchards. Genera Species Common Name ANISOPODIDAE Sylvicola alternata (Say) BOMBYLIIDAE Ogcodocera leucoprocta (Wiedeman) CLUSIIDAE Clusia czernyi (Johnson) OTITIDAE Delphinia pigta (Fabricius) Pseudotephritis vau (Say) Pseudotephritina cribellum (Loew) Seioptera vibrans (Linnaeus) PLATYSTOMATIDAE Rivellia viridulans (Desvoidy) TEPHRITIDAE Euaresta bella (Loew) Euleia fratria (Loew) Eutreta s arsa (Wiedeman) Icterica seriata (Loew) Paroxyna albiceps (Loew) Rhagoletis basiola (Osten Sacken) Rhagoletis c1ngulata (Loew) Eastern Cherry , Fruit Fly Rhagoletis fausta (Osten Sacken) Black Cherry Fruit Fly Rhagoletis pomonella (Walsh) Apple Maggot Rhagoletis suavis (Loew) . Rhagoletis tabellaria (Fitch) Dogwood Maggot ' TETANOCERIDAE Euthycera arcuata (Loew) Tetanocera valida (Loew). Host Wild Cherry, Pin Cherry' Wild Cherry, Pin Cherry Apple. Hawthorne Dogwood 197 Tachinidae. Leeper (1978) went further and included photographs of six different Rhagolitis flies that can be caught in orchards that could possibly be confused with the apple maggot. Methods Specimens were removed from traps throughout the four year study and mounted or preserved in alcohol. At the conclusion of the field study, these flies were mounted, labeled, identified, and left in the Michigan State University Entomology collection as voucher specimens. Results Table 52 presents a list of species of picture winged flies that were caught on the traps placed in commercial orchards. To the novice, many of these could be confused with the apple maggot. Three of the Rhagolitis should be mentioned, because they emerge before apple maggot, are very similar in appearance, and can be found in the commercial apple orchards. The eastern cherry fruit fly, Rhagolitis cingulata, is a principle direct pest of tart cherries. It also has been recorded from wild black cherry and pin cherry, both of which are common species. Quite frequently they will appear _on yellow AM traps in early July when their adult population is peaking. In some orchards, due to the presence of a large number of wild cherries, they will be much more 198 numerous than apple maggot flies. Generally they do not feed on apple, so no controls need to be initiated for them. The black cherry fruit fly, Rhagolitis fausta, is also a direct pest of commercial cherries, but of less importance than the eastern cherry fruit flies. Its hosts are the wild black cherry and the pin cherry. It generally appears on the yellow traps 10 days before the eastern cherry fruit fly, and 3 1x) 6 weeks before apple maggot. One should recognize this species and be sure not to initiate sprays for it because it has not been reported to feed in apples. A third Rhagolitis that appears on yellow traps in apple orchards and is very similar in appearance to the apple maggot, it is the dogwood maggot or R tubellaria. As is indicated by its common name, its hosts are the dogwoods. It is not as common as the other Rhagolitis, but can easily be confused with apple maggot. It does not infest apples, so no sprays should be initiated when it is found on the traps. Conclusions There are many picture winged flies that are trapped in commerical apples. One needs to be careful when identifying flies on the traps, as many are very similar in appearance to the apple maggot, but do not infest apples. 199 SAMPLING FOR AND PREDICTION OF FRUIT DAMAGE The goal for any monitoring system should be to more efficiently and effectively detect the pest being monitored. With apple maggot, two trap types were evaluated for thier use in detecting the pest. Once detected, then control programs were initiated. The detection in this case provid- ed the biofix for determining when oviposition occurs,and when controls should be initiated. One method of evaluating the success of this detection is to determine the amount of damaged fruit in the orchard. Experiments were carried out to determine how large of a sample size was required and where to sample to efficiently estimate the percent fruit damage. This involved both within and between tree measurements. Once sampling procedures were known, then estimates of fruit damage were made. Lastly, correlations of the trap catch to fruit damage were made. With these parameter measured, one could determine the usefulness of the trapping scheme to measure fruit damage and in managing apple maggot. 200 DETERMINATION OF FIRST OVIPOSITION BASED ON A BIOFIX OF FIRST CATCH There is a need to know when first oviposition will occur after a certain biofix such as first trap catch, so that growers can efficiently control the AMP. The timing of this event was studied, and a pmediction was determined based on air temperature degree day accumulation. Factors that influence this such as fruit availability and fruit susceptibility will be ignored, as once the threshold is reached, the female will likely find a susceptible site and initiate oviposition. With this predictor available, pest managers can then recommend controls based on trap catch within an orchard. This firm level monitoring should reduce the risk to each grower from apple maggot damage. Literature Oviposition is the key event in the life cycle of apple maggot that growers attempt to pmevent. Once eggs are deposited in the fruit, damage has occurred and the apple is unmarketable. Neilson (1978) reported that the first oviposition occurred 21 days after first catch. This time frame is much longer than is the general understanding. Hall (1937) reports that after emergence, the adults pass through a pre-oviposition period of 4-14 days during which time they feed, mature, and mate. At the end of this time period they are capable of laying eggs. The critical event to measure in the past was first catch, and then sprays were advised in 7-10 days. If the 21 day period is correct, then 201 further delays in spraying could occur and the pest could be more economically managed. Methods Experiments were initiated in 1977 to obtain an esti— mate of the length of the pre-oviposition period. Flies were captured on yellow Zoecon AM traps on 10 trees at weekly intervals at the K.S.H. orchard. This catch served as a biofix for initiating the pre-oviposition period. These same trees had 25 apples tagged each. The apples were examined weekly for the presence of oviposition punctures. Once found, they signaled the end of the pme-oviposition period. From the results obtained in 1977, it was realized that the weekly monitoring was too long of an interval, and the experiment was modified in 1978. Assistance was available to help monitor, so checks were made twice weekly. Again 10 trees had yellow Zoecon AM traps to serve as a biofix, and 25 apples were tagged and checked for oviposition stings on each of those trees. During 1978, females were observed probing apples with their ovipositor. These exact spots were marked, and the apples picked and examined under magnification. The new stings were not easily distinguishable. This indicated that most of the stings counted previously were several days old. This time period allowed for apple growth and a slight depression to be formed which was more evident. Therefore, the experiment was modified in 1979 to determine a better 202 estimate of the time interval between first catch and oviposition. The method chosen was to examine ovarian develOpment. Fifteen trees in the K.S.H. orchard, nine in the Upjohn orchard, and twelve abandoned trees around commercial orchards were used as experimental units. Biofix of adult activity was determined by placing yellow Zoecon AM and sticky red sphere traps in each tree. The traps were monitored three times each week. They were cleaned at each visit, and their positions reversed every two weeks when the yellow traps were replaced. All flies were saved, sexed, and the females were dissected to examine for ovarian development (Neilson et. al., 1976). If eggs were found in the oviduct, then they were considered mature and capable of oviposition. In 1980 these same two trap types were used to serve as a biofix for determining first flight activity. Fifteen trap trees were set in the K.S.H. orchard, 9 in the Upjohn orchard, and 15 in abandoned t'rees around commercial orchards. Traps were checked daily for the first two weeks of flight. Female maturity was checked by dissecting all the females caught on all the traps until eggs were evident in the oviducts. During each year, data was separated by the variety of apple in which the traps were located. This enabled analy- sis of any significant differences in the duration of this interval due to variety. 203 Results Weekly sampling in 1977 proved to be too long of an interval to determine the length of time between first catch and first oviposition. Table 53 shows the mean delay was 15.6 days or 388.8 degree days at base 48°F. This value is much larger than what is normally considered the true length. The weekly fruit sampling by Neilson (1978) prob- ably explains why his value was 21 days. However, valuable information was gained on the number of new stings per apple (Figure 18) per week. A nice curve was present that matched quite closely the mean weekly trap catch, which indicates that when more flies are present more oviposition will occur. Table 53. Duration of Pre-Oviposition Period as Determined by the Interval Between a Biofix of First Catch on the Yellow Zoecon AM Trap and the First Stung Apple at the KSH Orchard in 1977. Biofix First Length of Period Tree Variety Date Sting Days DD 48° F 1 Transparent 6-22 6-29 7 170 2 Red Delicious 6-20 7- 6 16 395 3 Dutchess 6-20 7- 6 16 395 4 Dutchess 6-20 7- 6 16 395 5 Greening 6-20 7-13 23 581 6 Jonathan 6-29 7-13 14 380 7 Red Delicious 6-20 7- 6 16 395 8 Red Delicious 6-20 7- 6 16 395 9 Northern Spy 6-20 7-13 23 581 10 McIntosh 6-20 6-29 9 221 Mean 15.6 388.8 204 .Ammmne oH now owns mom mmamm< mmv oumnouo Hmuwmmom mumum ooNMEmamx on» no uommmz magma on one xmmz Hod mamm< Mom mmCHum coflufimoma>o 3oz mo Honezz com: .mH musmflm EB ta «a; 33 3:. 9: am; I; 2.”; _ :3 _ a? _ 9...: _ o: _ «a; .v F. . m .... N B U i 0. W" m a. 3 v. m o 1.. no I, m. S. w. 9 CW. 01 A : 5. ~58. em. 205 In 1978 the sampling interval was shortened to twice each week. This shortened the mean length of the pre-oviposition period to 10.9 days or 217.2 air degree days at base 48°F at the 5 foot level (Table 54). This value is much closer to that reported in the literature. Table 54. Determination of the Length of the Pre-Oviposition Period Initiated by a Biofix of Catch on the Yellow Zoecon AM Trap and Terminating with the Finding of Stung Fruit at the KSH Orchard in 1978. Biofix First Length of Period Tree Variety Date Sting Days DD 48°F 1 Dutchess 7- 5 7-11 6 113 2 Transparent 6-30 7-11 11 204 3 Greening 7- 5 7-11 6 113 4 Dutchess 7- 6 7-11 5 88 5 Northern Spy 7- 4 7-11 7 136 6 Red Delicious 6-27 7-18 21 421 7 McIntosh 6-27 7-18 21 421 8 Northern Spy 7-14 7-18 4 85 9 Red Delicious 7- 8 7-21 13 273 10 Northern Spy 7- 6 7-21 15 318 Mean 10.9 217.2 The number of new stings per fruit per week is found in Figure 18. The mean number of stings per fruit was 1.47 in 1977 and 3.76 in 1978. Corresponding to this there were 223 flies caught per trap in 1977 and 235 per trap in 1978 in the same trees where the fruit damage ratings were made. One possible explanation for there being twice as many stings per fruit in 1978 than 1977 would be fruit availabil- ity. If more apples were present on the trees in 1977, then more oviposition sites would be available which would reduce the number per fruit. However, no data was taken on the number of fruit per tree, so this is only speculation. Both 206 these values are quite low when compared to the 20 punctures found on many apples and 46 found on one apple reported by O'Kane (1914). However, his mean stings per apple on the 22 trees rated was 3.41 which corresponds well with my data. In 1979, all flies caught on these traps were sexed (n=6963 in K.S.H. orchard and n=662 in the Upjohn orchard, n=2622 in abandoned trees) and the females were dissected to determine ovarian development. The first gravid fly caught on each trap served as a termination of the pre-oviposition period, as those flies were capable of laying eggs and causing fruit damage. Table 55 shows that at the K.S.H. orchard the average length after first catch on the yellow Zoecon AM trap to mature females was 3.5 days or 64.8 air degree days at base 48°F. On the red sphere trap this interval was 2.5 days or 55.7 air degree days at base 48°F. These were not significantly different. In the Upjohn orchard (Table 56), the mean values were 2.9 days or 61.4 dd base 48°F for the yellow trap and 7.3 days or 154.5 dd base 48°F for the red sphere trap. Again these means were not significantly different even though the interval was longer. The data from the abandoned trees around commercial orchards found in Table 57 is very similar. Using the yellow AM traps as indicators for the initiating of the pre-oviposi- tion period, its length was foun? to be 1.0 days or 22.7 dd base 48°F. The length using the red sphere trap was 4.9 207 Table 55. Determination of the Length of the Pre-Oviposition Period in 1979 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the KSH Orchard. Biofix Gravid Length of Period Method Variety Date Date Days DD 48° Yellow Trap Transparent 1 7-2 7-9 7 123 Red Sphere Transparent 1 7-9 7-9 0 0 Yellow Trap Transparent 2 7-2 7-2 0 0 Red Sphere Transparent 2 7-6 7-9 3 69 Yellow Trap Greening 7-2 7-2 0 0 Red Sphere Greening 7-13 7-13 0 0 Yellow Trap Dutchess 7-2 7-4 2 27 Red Sphere Dutchess 7-6 7-6 0 0 Yellow Trap Greening 7-4 7-6 2 27 Red Sphere Greening 7-13 7-13 0 0 Yellow Trap Wealthy 7-2 7-9 7 123 Red Sphere Wealthy 7-2 7-11 9 178 Yellow Trap Dutchess 7-2 7-2 0 0 Red Sphere Dutchess 7-2 7-9 7 123 Yellow Trap Jonathan 7-6 7-9 3 69 Red Sphere Jonathan 8-2 8-2 0 0 Yellow Trap McIntosh 1 7-2 7-6 4 54 Red Sphere McIntosh 1 7-13 7-16 3 73 Yellow Trap McIntosh 2 7-2 7-4 2 27 Red Sphere McIntosh 2 7-9 7-23 14 337 Yellow Trap Red Delicious 1 7-4 7-4 0 0 Red Sphere Red Delicious 1 7-30 7-30 0 0 Yellow Trap Red Delicious 2 7-2 7-2 0 0 Red Sphere Red Delicious 2 7-30 7-30 0 0 Yellow Trap Northern Spy 1 7-2 7-9 7 123 Red Sphere Northern Spy 1 7-23 7-23 0 0 Yellow Trap Northern Spy 2 7-2 7-9 7 123 Red Sphere Northern Spy 2 7-9 7-11 2 55 Yellow Trap Jonathan 2 7-4 7-16 12 276 Red Sphere Jonathan 2 7-9 7-9 0 0 Yellow Trap Mean 7—3 7-6 3.5a 64.8a Red Sphere Mean 7-14 7-16 2.5a 55.7a Means followed by the same letter are not significantly different at the .05 level by the DMR test. 208 Means followed by different at the Table 56. Determination of the Length of the Pre-Ovi- position Period in 1979 Beginning with a Biofix of Trap Catch and Terminating with a Catch of Gravid Females at the Upjohn Orchard. Biofix Gravid Length of Period Method Variety Date Date Days DD 48° Yellow Trap McIntosh 1 7-30 8-2 3 66 Red Sphere McIntosh 1 7-16 - - - Yellow Trap McIntosh 2 8-6 8-6 0 0 Red Sphere McIntosh 2 7-16 7-19 3 42 Yellow Trap McIntosh 3 8-13 8-13 0 0 Red Sphere McIntosh 3 - - - - Yellow Trap Jonathan 1 7-13 7-13 0 0 Red Sphere Jonathan 1 7-16 7-19 3 42 Yellow Trap Jonathan 2 7-13 8-2 20 421 Red Sphere Jonathan 2 7-23 8-6 14 318 Yellow Trap ’Jonathan 3 7-13 7-13 0 0 Red Sphere Jonathan 3 7-16 8-9 24 525 Yellow Trap Spy 1 8-13 8-13 0 0 Red Sphere Spy 1 8-30 8-30 0 0 Yellow Trap Spy 2 7-13 7—16 3 66 Red Sphere Spy 2 8—13 8-13 0 0 Yellow Trap Spy 3 8-6 8-6 0 0 Red Sphere Spy 3 - - - - Yellow Trap Mean 7-27 7-30 2.9a 61.4a Red Sphere Mean 7-27 8-3 7.3a 154.5a the same letter are not significantly .05 level by the DMR test. 209 TABLE 57. Determination of the Length of the Pre-Oviposition Period in 1979 Beginning With a Biofix of First Trap Catch and Terminating With a Catch of a Gravid Female Orchard 10 ll 12 Mean Mean in Abanonded Trees Around Commercial Orchards. Method Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Biofix Date I00 I mflflflmmm II \INI—IWQQQ Gravid Date N I \IWNI—‘fll—‘Q I—‘uD-N I N |._.| £00m“ I (I) I ooqxloooooooo I I (~me Length of Period Days r-u—a \IO I OQOWOOOOOUJO I O I OQNNOU'IO 1.00a 4.89a DD 48° 0 94 0 248 272 166 0 U1 Ch ' NO I OQOUOOOOONO I OI ....I O 9...: N 22.7a 94.7a Means followed by the same letter are not significantly different at the .05 level by the DMR test. 210 days or 94.7 dd base 48°F. These values were not signifi- cantly different. To summarize this, after first catch on the yellow trap, females have eggs in their oviducts 2.5 days or 49.9 dd base 48°F later. Using the red sphereas the indicator, 4.2 days or 87.1 dd base 48°F after first catch, the females have eggs in their oviducts. This indicates the red sphere would give a slightly longer period to prepare for initiating controls. However, in these three sites the red sphere caught the first fly a mean of 4.7 days later than did the yellow trap. Therefore, flies caught on the yellow traps would have eggs in the oviducts before flies are even caught on the red sphere. This suggests the yellow trap would be a better indicator for establishing a biofix and predicting when oviposition could occur. In 1980, all the flies caught were again sexed and the females dissected to determine ovarian development. The number of flies checked was: n=5386 in the K.S.H. orchard, n=483 in the Upjohn orchard and n=536 in the abandoned trees around commercial orchards. At the KLS.H. orchard, the yellow traps caught flies first as its biofix date was July 4, six days sooner than with the red sphere trap (Table 58). The length of the pre-oviposition period was 5.3 days or 133 dd base 48°F for the yellow trap and 2.1 days or 51 dd base 48°F for the red sphere. These are significantly different values. This shows that this year in this orchard the length of the pre-oviposition period was 211 TABLE 58. Determination of the Length of the Pre-Oviposition Period in 1980 Beginning With a Biofix of Trap Catch and Terminating With a Catch of Gravid Females at the K.S.H. Orchard. Biofix Gravid Length of Period Method Variety Date Date Days DD 48° Yellow Trap Transparent 6-30 7-5 5 104 Red Sphere Transparent 6-28 7-5 7 124 Yellow Trap Transparent 7-5 7-7 2 47 Red Sphere Transparent 7-9 7-9 0 0 Yellow Trap Greening 7-2 7-2 0 0 Red Sphere Greening 7-7 7-9 2 54 Yellow Trap Dutchess 6-28 7-5 7 124 Red Sphere Dutchess 7-2 7-2 0 0 Yellow Trap Greening 6-30 7—5 5 104 Red Sphere Greening 7-7 7-14 7 188 Yellow Trap Wealthy 7—5 7-7 2 47 Red Sphere Wealthy 7-5 7-7 2 47 Yellow Trap Dutchess 7-2 7-5 3 64 Red Sphere Dutchess 7-7 7-7 0 0 Yellow Trap Jonathan 7-7 7-9 2 54 Red Sphere Jonathan 7-16 7—16 0 0 Yellow Trap McIntosh 7-7 7-18 11 304 Red Sphere McIntosh 7-16 7-18 2 51 Yellow Trap McIntosh 7-5 7-18 13 351 Red Sphere McIntosh 7-5 7-11 6 155 Yellow Trap Red Delicious 7-5 7-14 9 235 Red Sphere Red Delicious 7-16 7-16 0 0 Yellow Trap Red Delicious 7-7 7-14 7 188 Red Sphere Red Delicious 7-16 7-21 5 146 Yellow Trap Northern Spy 7—7 7-14 7 188 Red Sphere Northern Spy 7-16 7-16 0 0 Yellow Trap Northern Spy 7-9 7-9 0 0 Red Sphere Northern Spy 7-21 7-21 0 0 Yellow Trap Jonathan 7-7 7-14 7 188 Red Sphere Jonathan 7-5 7-5 0 0 Yellow Trap Mean 7-4 7-10 5.33a 133.2a Red Sphere Mean 7-10 7-12 2.07b 51.0b Means followed by the same letter are not significantly different at the .05 level by the DMR test. 212 determined to be shorter when using the red sphere trap. This has the disadvantage of allowing for a shorter time period to initiate sprays after the first fly is caught. In the Upjohn orchard the biofix date for the yellow trap was July 21, 4 days sooner than that of the red sphere trap (Table 59). Because of this date being later in the season, the females caught on it were already gravid. With the red sphere trap, gravid flies were caught 2.6 days or 56 dd base 48°F later than its biofix. Therefore, the red sphere caught gravid females 7 days later than the yellow trap which was hung in the same tree. This indicates that if one wanted to estimate when damage occurs, the yellow trap was far superior in this orchard. In the 15 abandoned trees around commerical orchards checked in 1980, the yellow trap was better (Table 60). The mean biofix dates were July 24 for the yellow trap and July 30 for the red sphere. The lengths of the pre-oviposition period were 4.2 days for the yellow trap and 4.5 days for the red sphere. These correspond to 113 and 116 degree days base 48°F, respectively. These differences are not significant, but the yellow trap has a seven day lead period over the red sphere traps for catching gravid females. To summarize the 1980 data for these three locations, the yellow trap had a mean biofix date 5.3 or 116 dd base 48°F days sooner than did the red sphere trap. The 213 Means followed by the same letter at the .05 level by the DMR test. TABLE 59. Determination of the Length of the Pre-Oviposition Period in 1980 Beginning With a Biofix of Trap Catch and Terminating With a Catch of Gravid Females at the Upjohn Orchard. Biofix Gravid Length of Period Method Variety Date Date Days DD 48° Yellow Trap McIntosh 1 7-25 7-25 0 0 Red Sphere McIntosh 1 7-23 7-23 0 0 Yellow Trap McIntosh 2 7-23 7-23 0 0 Red Sphere McIntosh 2 8-25 8-25 0 0 Yellow Trap McIntosh 3 7-23 7-23 0 0 Red Sphere McIntosh 3 7-25 8-11 17 384 Yellow Trap Jonathan 1 7-23 7-23 0 0 Red Sphere Jonathan 1 7-21 7-23 2 37 Yellow Trap Jonathan 2 7-23 7-23 0 0 Red Sphere Jonathan 2 7-14 7-14 0 0 Yellow Trap Jonathan 3 7-5 7-5 0 0 Red Sphere Jonathan 3 7-14 7-14 0 0 Yellow Trap Northern Spy 1 7-23 7-23 0 0 Red Sphere Northern Spy 1 7-21 7-25 4 83 Yellow Trap Northern Spy 2 7-28 7-28 0 0 Red Sphere Northern Spy 2 7-30 7-30 0 0 Yellow Trap Northern Spy 3 7-16 7-16 0 0 Red Sphere Northern Spy 3 7-25 7-25 0 0 Yellow Trap Mean 7-21 7-21 0.0a 0.0a Red Sphere Mean 7-25 7-28 2.6a 56.0a are not significantly different 214 TABLE 60. Determination of the Length of the Pre-Oviposition Period in 1980 Beginning With a Biofix of First Trap Catch and Terminating With a Catch of a Gravid Female in Abanonded Trees Around Commercial Orchards. Biofix Gravid Length of Period Orchard Method Date Date Days DD 48° 1 Yellow Trap 7-23 7-25 2 49 Red Sphere 7-28 7-28 0 0 2 Yellow Trap 8-18 9-3 16 435 Red Sphere 8-27 9-3 7 194 3 Yellow Trap 8-8 8-8 0 0 Red Sphere 8-8 - - - 4 Yellow Trap 7-25 7-25 0 0 Red Sphere 8-6 - - - 5 Yellow Trap 7-23 7-23 0 0 Red Sphere 8-6 8-6 0 0 6 Yellow Trap 7-21 7-21 0 0 Red Sphere 7-28 7-30 2 43 7 Yellow Trap 8-11 8-11 0 0 Red Sphere - - — - 8 Yellow Trap 8—6 8-6 0 0 Red Sphere 8-11 8-11 0 0 9 Yellow Trap 7—16 7-16 0 0 Red Sphere 8-6 8-6 0 0 10 Yellow Trap 8-19 8-19 0 0 Red Sphere - - - - 11 Yellow Trap 7-10 7—15 5 140 Red Sphere 7-31 7-31 0 0 12 Yellow Trap 6-21 6-24 3 80 Red Sphere 6-23 7-9 16 397 13 Yellow Trap 7-25 - - r Red Sphere 8-22 8-22 0 0 14 Yellow Trap 7—5 7-18 13 347 Red Sphere 7-5 7-5 0 0 15 Yellow Trap 7-5 7-25 20 531 Red Sphere 7—5 7-30 25 638 Mean Yellow Trap 7-24 7-28 4.21a 113.0a Mean Red Sphere 7-30 8-4 4.54a 115.6a Means followed by the same letter are not significantly different at the .05 level by the DMR test. 215 mean length of the pre-oviposition period was 3.7 days for the yellow trap and 3.0 for the red sphere or 94 and 73 degree days base 48°F, respectively. Therefore, the yellow trap should be used as a nwmitoring tool to determine a biofix for apple maggot flight. Because the 1979 and 1980 tests were performed identically, so both years data were lumped to provide a greater number of replicates for general recommendations (Table 61). At the K.S.H. orchard, a total of 30 replicates showed that the mean first catch or biofix was 8.5 days sooner using the yellow traps. The length of the period lasted 4.4 days for the yellpw trap and 2.3 days for the red sphere trap, or 99 and 53 degree days base 48°F. zu: the Upjohn orchard where there were 18 replicates, the biofix was 2.0 days sooner on the yellow trap. The length of the period was 1.4 days on the yellow trap and 4.5 days on the red sphere, or 31 and 105 degree days base 48°F respectively. In the abandoned trees around commercial orchards there were 27 replicates. The mean biofix date was 4.5 days sooner using the yellow trap. The length of the period was 2.7 days with the yellow trap and 4.7 days with the red sphere, or 71 and 106 degree days base 48°F, respectively. When all these locations are lumped, the yellow trap caught flies 5.0 days sooner than the red sphere trap. The length of the periods were 3.1 days with the yellow trap and 3.5 days with the red sphere, or 73 and 77 degree days base 48°F, respectively. This indicates that 216 TABLE 61. Determination of the Length of the Pre-Oviposition Period Beginning With a Biofix of First Trap Catch and Terminating with a Catch of a Gravid Female. Length in Period Location Year Method Days dd 48°F K.S.H. 1979 Yellow Trap 3.5 65 Red Sphere 2.5 56 1980 Yellow Trap 5.3 133 Red Sphere 2.1 51 Upjohn 1979 Yellow Trap 2.9 61 Red Sphere 7.3 155 1980 Yellow Trap 0 0 Red Sphere 2.6 56 Abandoned 1979 Yellow Trap 1.0 23 Red Sphere 4.9 95 1980 Yellow Trap 4.2 113 Red Sphere 4.5 116 All All Yellow Trap 3.1a 73a Red Sphere 3.5a 77a Means followed by the same letter are not statistically different at the .05 level by the DMR test. 217 the yellow trap should be used to monitor for the biofix, and that controls should be initiated within 3 days after first catch. The data generated was recorded by variety to determine if there were any differences related to variety of apple the traps were hanging in. Data from 1977 and 1978 at the K.S.H. orchard are presented in Table 62. Only the yellow trap was used these two years. In each case the unbalanced design of the analysis of variance and the DMR test were performed. In all cases the variances were homogeneous by the Bartlett's test at the .05 level. On the 19 trees examined 511 detail, there was rub statistical difference due to variety in the biofix or first trap catch date. The same is true for the length of the pre-oviposition period. However, the number of stings per apple does vary. The earlier varieties have less stings due to their maturing sooner and falling off the tree. This indicates that they are suitable for larval develOpment for a much shorter time frame. The later varieties are susceptible and hang on the tree longer thereby being exposed longer which results in their having significantly more stings. The flies caught per tree are statistically different. for' the different. varieties. One jpossible explanation might be that the trees had varying number of apples the preceeding year which would result in there being more or less flies under each tree to be caught on the traps. 218 TABLE 62. Mean Differences in the First Trap Catch Date, Length of Pre-Oviposition Period, Number of Stings Per Apple and Number of Flies/Trap Due to Variety at the K.S.H. Orchard in 1977 and 1978. No. Biofix Length Stings/1 Flies/ Variety Trees Date in Days Apple Trap Dutchess 4 6-28a 10.75a 0.42a 208ab Transparent 2 6-26a 9.00a 1.50ab 293ab McIntosh 2 6-24a 15.00a 1.52ab l61ab Greening 2 6-28a 14.50a 1.66ab 94a Red Delicious 5 6-25a 16.40a 3.11 b 311 b Northern Spy 4 7-3 a 12.25a 6.63 b 186ab 1 Mean of 25 apples/tree Means followed by the same letter are not significantly different at the .05 level by the DMR test. 219 In 1979 and 1980, the two trap types were compared for these same parameters at the K.S.H orchard (Table 63). Using the yellow trap there was no statistical difference in the biofix or first trap catch date due to variety, but there was a range of five days present. Using the red sphere trap, the early varieties Transparent, Dutchess and Greening had flies caught significantly earlier than did the later varieties, Jonathan, McIntosh, Red Delicious and Northern Spy. The length of the pre-oviposition period was essentially the same for both trap types among the vari- eties. The number of flies caught per trap on the yellow trap did not differ statistically, but had a range in the means from 224 to 389. The catch on the red spheres was likewise nonsignificant between the varieties, but the means were lower and ranged from 37 to 253. Comparisons between the four years at the K.S.H orchard were not made. This is due to the different sam- pling intervals with 1977 being weekly, 1978 twice a week, and 1979 and 1980 three times each week through the termina- tion of the pre-oviposition period. The means generated on the biofix date and length of the pre-oviposition would have a considerable amount of sampling variance associated with them. In the Upjohn orchard in 1979 and 1980, traps were placed in three trees of three different varieties each year. They were monitored three times per week to quantify the parameters found in Table 64. When measuring the biofix TABLE 63. Variety Transparent Greening Dutchess Jonathan McIntosh Red Delicious Northern Spy Orchard Method Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Yellow Trap Red Sphere Biofi Date X m n: W U‘ U‘ 0 O I I—‘U’INUII-‘th-‘mbI—‘I-‘NUIN \I I—' IIII w \I\I\I\I\I\I\I\I\I\I\I\J\I\I I WWW W a Q) Q) 22 U‘U‘U‘ O bc 0" 00 U‘ 0 0 O bcd (1.. (D D: (D CL (0 Length in Days 3.50abc 2.50abc 1.75abc 2.25abc 3.00abc 1.75abc 6.00abc 0.00a 7.50 c 6.25 bc 4.00abc 1.25ab 5.25abc 0.50ab Mean Differences in the Biofix Date, Length of the Pre-Oviposition Period, and Number of Flies Per Trap Due to Variety During 1979 and 1980 at the K.S.H. (4 Replicates). Flies/ Trap 257.7abc 69.2ab 235.5abc 102.7ab 329.2 bc 168.7abc 271.0abc 45.2a 313.7 bc 145.2abc 224.0abc 37.2a 389.2 c 253.2abc Me an Mean Yellow Trap Red Sphere 7-12 289 117 Means followed by the same letter are not significantly different at the .05 level by the DMR test. 221 TABLE 64. Variety Effect on the Biofix Date, Length of Pre— Oviposition Period, and Flies Per Trap at the Upjohn Orchard in 1979 and 1980. Biofix Length Flies/ Variety Method Date in Days Trap McIntosh Yellow Trap 7-25ab 7.5a 17.8a Red Sphere 7-27ab 5.0a 11.2a Jonathan Yellow Trap 7-15a 3.3a 58.3 b Red Sphere 7-17a 7.2a 93.3 b Northern Spy Yellow Trap 7-27ab .5a 20.7a Red Sphere 8-5 b .8a 6.5a Mean Yellow Trap' 7-22 3.8 32.3ns Mean Red Sphere 7-27 4.3 37.0ns Means followed by the same letter are not significantly different at the .05 level by the DMR test. 222 or first trap catch with the yellow trap, there were no statistical differences between the varieties even though the range in mean dates was 12 days. With the red sphere traps, flies were caught significantly earlier on JCnathans and significantly later on Northern Spy, with McIntosh intermediate. This indicates that the yellow trap gives a less variable first catch, and in all cases the mean first catch was sooner on the yellow trap, with the overall mean being five days sooner. Statistically there was no differ- ence in the length of the pre-oviposition period between the varieties for either trap type. The grand mean indicated that the difference was only half of a day between the trap types. Therefore, either trap could be used with equal reliability to predict the length of the pre-oviposition period in this orchard. There was a difference in the mean number of flies caught per trap due to variety. For both trap types, the traps placed in the Jonathan trees caught significantly more flies than traps placed in McIntosh or Northern Spy trees. When these data are summarized, al- .though not statistically significant, the Jonathan variety would be the best one in which to place the traps in this orchard. This is because the biofix date was first in this variety and the most flies were caught in it. Also, the I yellow trap would be preferred because its biofix date was five days sooner than the red sphere trap. 223 Conclusions In 1977, weekly monitoring determined the pre-oviposition interval to be 15.6 days or 388.8 air dd base 48°F. This length was much longer than reported in the literature, and the sampling interval had to be shortened. When the interval was shortened to every three days in 1978, the length of the pre-oviposition period was reduced to 10.9 days or 217.2 dd base 48°F. It was discovered in 1978 that the method of determining the end of the period (looking for stings) was not the best as many early stings could be missed. Therefore, these data will not be used as predic- tors. Useful information was generated from these experi- ments however. There were found 1.47 stings per apple in 1977 and 3.76 in 1978. These mean values compare well to that reported in the literature. Also, the curve of new stings per fruit followed very closely the trap catch numbers of adults, which indicates as more adults are present more damage is being done. During 1979, the yellow trap was found to be a better indicator of the biofix than was the red sphere trap by catching flies 4.7 days sooner in three separate tests. Using the yellow trap, the pre-oviposition period was found to last 1.0 days or 23 dd at base 48°F. The red sphere had a length of 4.9 days or 95 dd at base 48°F. In 1980, the yellow trap again caught flies sooner in all three tests, with the mean value being 5.3 days sooner. The mean length of the pre-oviposition was found to be 3.7 days with the 224 yellow trap and 3.0 days with the red sphere trap or 94 and 73 dd base 48°F respectively. When both years data were lumped over all locations, the yellow trap caught flies 5.0 days sooner, and had gravid flies on them two days before the red sphere caught any flies. This indicates it is the preferred monitoring tool, and controls should be initiated with 3 days after first catch. In tests conducted from 1977 to 1980 at the K.S.H orchard, there was no statistical difference due to variety in the biofix date nor length of pre-oviposition period when the yellow trap was the monitoring tool. However, there was a difference in the number of stings per apple, with the earlier maturing varieties having fewer as they fall off the tree sooner and are not exposed as long as the later maturing varieties. There were also some differences in the mean number of flies caught per trap. These differences are probably due to the size of the apple crop the preceeding year which would regulate the population size available for trap catch the following year. When the red sphere trap was used in 1979 and 1980, the early maturing varieties caught flies significantly sooner than did the later maturing varieties. However, there were no differences between varieties in the length of the pre-oviposition period nor the mean number of flies caught per trap. In the Upjohn orchard in 1979 and 1980, there were no differences between varieties in the biofix date nor the length of the 225 pre-oviposition period when using the yellow trap. There were some differences when the red sphere trap was used, but in each case the red sphere trap was inferior due to the later first catches. Even though not statistically signifi- cant, the yellow trap should be placed in the Jonathan variety in this orchard to measure the biofix date. 226 SAME SIZE TO ESTIMATE FRUIT DAMAGE An important consideration in any experiment is to know how large of a sample size is required to measure the variables investigated. The Optimal sample sizes can seldom be taken because of time and cost constraints. However, several different experiments were conducted to determine what sample size should optimally “be taken to obtain a reliable estimate of fruit damage in research and commercial orchards. In one test, every apple was picked off trees that had a moderate infestation and rated for damage. With the absolute mean damage known, probability equations could determine how large of a sample is needed to be within reasonable ranges of the true value. This test also re- vealed the spacial distribution of the damage within the tree, and determined where apples should be picked to obtain more reliable estimates. To determine how many trees per orchard to sample, intensive sampling was conducted to find estimates of percent fruit damage on many trees. This data can be used to determine the fewest number of trees that needed to be sampled in the orchard. Finally, the edge effect was studied to determine if it is a.reelity with apple maggot. If so, then biases in the estimation of damage in the entire block can be controlled by understand- ing where the damage is most likely to occur. 227 Literature In attempting to estimate fruit damage within a tree Cameron and Morrison (1974) examined 20 apples per tree. Neilson (1978) picked 25 apples from trees that had traps in them. He felt this was a suitable sample size to obtain an estimate of the mean damage in the tree. The fruit damage rating sample size for estimating apple maggot damage utilized by the Michigan Apple Pest Management project was to randomly select 20 apples per tree and 15 trees of each major variety in the orchard (Olsen, Unpub- lished). Reissig and Tette (1979) sampled 100 apples from the top and middle of the tree, and all drops from 5 trees per block. To calculate optimum the sample sizes, equations from Southwood (1978) were used. Methods Whole Tree Sampling. In 1979, every apple was examined for damage on four trees. Three of these trees were at the Upjohn orchard, and the fourth was a backyard tree that was routinely sprayed. After recording all other damages, thin slices were removed to look for apple maggot larval tunneling to confirm its presence. The number of apples with damage and the number of stings per apple were recorded. Each apple was given a coordinate to the nearest inch off the ground and north or south and east or west directions from the center of the tree. The sampling was 228 performed in late September and early October to assure no further damage would occur. Orchard Sampling. In five orchards in 1980, fifty trees at random were rated for apple maggot damage. Trees were assigned rows and numbers in that row, and then a random number table was consulted for determining sample trees. This large sample assured all varieties and all areas of the orchard would be checked. Fifty apples were examined per tree. This was performed in mid-September so apples would be present on all varieties, and no new damage would occur. Edge Effect. In five orchards in 1980, a sampling scheme was devised to detect an edge effect. Five trees each in the four perimeter rows were evaluated for apple maggot damage. Again, 50 apples per tree were carefully examined. The fruit damage rating was performed in mid-September for reasons already discussed. Results Whole Tree Sampling. Table 65 shows the data from the whole tree sampling for apple maggot damage. The first three trees are in the Upjohn orchard where every other tree was sprayed. As shown elsewhere, this spraying scheme reduced the apple maggot population by 95% as determined by trap catch. The percent infested level is high, but the mean stings per fruit is quite low. The last tree is in a backyard situation with many sources of apple maggot 229 TABLE 65. Whole Tree Sample for Apple Maggot Damage. No. Apples Percent Std Mean Stings Sample Size Variety Sampled Infested Dev Per Fruit i 10% i 5% McIntosh 348 37.1 .484 .95 46.7 187 McIntosh 290 24.8 .433 .46 37.3 149 Jonathan 335 54.9 .498 1.55 49.5 198 Red Delicious 377 14.8 .356 .15 25.2 101 230 surrounding it. The tree was sprayed weekly to hold the damage down to a satisfactory level. These data show differences between trees in the frequency of damage. In the Upjohn orchard this probably is due to the variety preference as the insect is established in the orchard and equally distributed throughout. By observation, and with this data, Jonathan seems to be preferred over McIntosh for oviposition. The Red Delicious tree is in another locality where the population is very low (1 female/trap for the entire season), resulting in the lower frequency. Also shown in this data is the relationship between frequency of occurrence or percent fruit damage and the mean number of stings per apple. A least squares linear regression analysis found a coefficient of determination (R3) of 0.998. Therefore, using these four trees as the data base and having the percent fruit damage in the range of 15 to 55%, a very reliable estimate of the number of stings/fruit (y) can be found by plugging x (the percent fruit damage) into equation 7: Eq 7 Y = 0.035 x — 0.386 To calculate the optimum sample size from each of these four trees to estimate the percent fruit damage, the 2 equation N = E71239 from Southwood (1978) was used. This assumes a frequency of occurance estimate is to be made. The p is the probability of occurence found in a preliminary 231 survey, here the true value. The q equals l—p, the t is the Student t of standard statistical tables and approximates 2 for samples of greater than 10 at the 5% level, and D is the predetermined half-width of the confidence limits of the mean. To be 95% (alpha = .05) sure that the sample estimate is within confidence limits of 110% of the true frequency, sample sizes of 25 to 50 need to be taken on these trees. If one wanted to be more precise and be within t5% of the true mean, larger sample sizes in the 100 to 200 range would need to be taken for each tree. This data can also give an indication of the spacial distribution of the damage within the tree. A priori it was assumed that at the very low frequency of occurrence in the commercial orchards the damage would be clumped or aggregated. As there are very few flies that successfully enter the orchard and sting fruit, those fruit that were damaged would be clumped. The female would not have much time to search out oviposition sites and lay eggs before she would be killed by the insecticides routinely applied. This distribution would likely approximate the negative binomial where the variance is greater than the mean. As will be shown, this distribution fit very closely the negative binomial distribution. In these medium density orchards the flies have very limited insecticide pressurn to kill them and they have much more time to search out new unexploited oviposition sites. This would suggest a random or uniform spacial distribution. There are two aspects of their 232 behavior would encourage a uniform distribution. After laying an egg in an apple, the female deposits a deterrent pheromone to prevent other eggs from being laid in this same fruit (Prokopy, 1972b; Prokopy' et. .al., 1976). This behavior tends to even out spacially the damage. Also, females show aggressive behavior toward each other in a territorial defense. These aspects have been noted and a uniform distribution shown to occur by Levoux and Mukerji (1963), Cameron and Morrison (1974) and Boller and Prokopy (1976). Reissig and Smith (1978) however found a random spacial distribution of eggs in the tree which was described by the Poisson distribution. They reasoned that under heavy fly pressure the females did not respond adequately to the marking pheromone, and they laid their eggs anywhere they found a host. To test the hypothesis that there is no significant difference in the frequency of damage in any part of the tree at the medium density Uphohn orchard the trees were artifically layered at 1 meter levels and quartered in the north-south and east-west plains. Analysis of variance was performed with each of the thin gradrants of the tree representing a different treatment. If significance was found, the DMR test at P = .05 separated the means. On the two McIntosh trees there was no sta;istical difference between the treatments in the frequency of damaged apples (Table 66). These trees were believed to have a random or TABLE 66. Height 1-2 Meters 2-3 Meters 3-4 Meters 475 Meters P Value Analysis of Spacial Distribution Damaged Apples Within the Tree. Quadrant NE NW SW SE NE NW SW SE NE NW SW SE NE NW SW SE I2 ONONO 90 17 37 10 61 44 wqoo 1 233 Mac 1 IX! .25 .50 .40 .44 .47 .47 .30 .25 .39 .36 .57 .33 .27 NS of the Apple Mac II 8. ‘2': 10 .30 3 .33 10 .60 6 .17 3s .14 29 .10 16 .25 53 .18 49 .27 4 0 13 .54 39 .18 4 .25 1 0 2 1.0 13 .31 1.95 NS Maggot Red Del 11 2E 43 .09b 24 .04b 10 0a 73 .03b 65 .23b 43 .31b 22 .23b 85 .16b 4 .50b 0 .. 2 .50b 1 0a 0 _ 0 _ 0 .. 0 ... 3.66 ** Means followed by the same letter are not statistically different at P=.05 by the DMR test. 234 uniform distribution. and this cannot be refuted because there were no statistical differences in the spacial distribution of their damaged apples, even when the two trees were lumped and analysed as one (F = 1.17 NS: critical .25(11,587) 1'32 and F.5(11,587) = '96)' However, the Red Delicious tree did have significant value F differences. The bottom SW and top SE quadrants had zero damage which was significantly less than the other quadrants in the tree. To determine if the clumping effect is real, the variance (.1268) was divided by the mean (.1485) to give 0.85 which is not indicative of a clumped population. This more closely fits the random distribution approximated by the Poisson H:= sumo .3: 3:02 05 5 uHmoawc smmHo mama manna mca mooEHomom pmumHH w>onm ecu pm>Hmuwm MINmmH .oz pocoso> :59»: A3982 m.u0uwwHumm>cH Azummmmumc NH muwwcm H~¢0Hquea away .D.m.z H H Hz ..00 oonEmme mumsoum unwounuom mmonmUozaamB "mmmemHo .D.m.z N m 1 TmDMG a mcoHumooH Hz mDOHHm> mcmHopHuH> mHHHm>Hm mmoHBASOBmwB mumuQOHmm .D.m.z H ofimH mHon mH Hz ..00 coucHHo om> mHuHuamuuoosmmm .a.m.z H mamH msa mm Hz 00 name HmnHuo mcHuHuadmuooswmm .D.m.z HH NH Woump a mGOHumooH Hz Hmuw>mm muon MHcHzmHmo MdoHBHaoummmamHo .D.m.z H m mmump a mcoHumooH Hz mDOHum> chumuo uHmoHU adoHHmDHo admmBmHo .D.m.z H aan >HohHH .Hz ..00 :mmmHH< muooumooomH mumooooomm mmoHHHMmzom admmBmHo .:.m.z H mamH >Hsn m .Hz ..oo oon86me mumcumuHm mHoOH>Hmw m so moumnouo mHmmm :H unmsmo ouoz mmHHm HHd u b: L.:—~:_—Z III, 1,.I Ia! lelI .l I .mmmuu Z4 :ooo ON onHw> :o mpumcouo onam :H ucmsmo mum: muHHN . HH< APPENDIX 1.1 Voucher Specimen Data Pages 2 2 of Page some bayou :mm/asa? 3 SuHmuo>Hca ououm cmmHonz any :H uHmooop pow mooEHoodm voumHH o>ono onu pm>Houmm m «N am.” . oz Aerosol, NmmH qHN an: 36: Accessc Azummmooo: «H muoonm HmcoHqupm omsv AHNJHHAHHHHJWMHHH. Amvmsmz m.u0umeumo>cH .D.m.z H mmmH mod n .Hz ..00 ucom ANpHunmnv mm mHumHommam .D.m.z H HH mmump a mCOHumooH .Hz mDOHum> uHHMHHonou mHuoHomonm .D.m.z 6H mN mmumo w mQOHumooH .Hz m50Hum> mH>msm mHuoHommmm .D.m.z m Hm mwumo a mcoHumooH .Hz msoHHm> MHHocosom mHuwHommnm .D.m.z . H msmH >Hsh m .Hz ..oo cousHHU mumomm mmuoHommmM .D.m.z H o mouuo a mGOHuuooH Hz msoHuu> mumHsmmHo mHuoHommnm .D.m.z N N mmumo a mGOHumooH Hz mSOHuo> MHOHmmn mHumHommnm .D.m.2 H mhaH 09¢ mN .Hz ..00 usmx mmonnHm mswxoumm .D.m.z H man 09¢ 5 .Hz ..00 MHcoH MDMHHom ooHuouoH .a.m.z m msmH one a .Hz ..oo mzmuuo mmummm mumuusm .D.m.z H man Doom m .Hz ..oo MHcoH mHuumnm onHmm .D.m.z H m moumo a mcoHumooH Hz mDOHum> mHHon mummumnm m¢DHBHmmmm9 u co moumsouo onmm :H unmsoo ouo3 meHw HHd o ouommoum mo mCHumm oH oH mmmue mo .02 .UmuouHcoz mooHoHHmo pom mooHOHHmo cmoHow oHoo seam msoHoano.emm cmzumc0h amouaHoz Naumm msoHoHHmQ pom mam :uosuuoz amoucHoz msoHOHHmQ pom wmm :uohuuoz smoucHoz mDOHOHHmQ pm; wmm sumnuuo: amoucHo: mooHoHHoQ pox ham cumzuuoz emoucHoz moHumHum> CH6: mpumnouo uHonu Ho coHumHuomoa a 6:6 mum3ouo mcHumuonou NH mH om ow ow wmm cowaHuU xuoum copHow snow 5 zoom pcoum HHHm QESQ QEQZ XUOHm manhmmchq muonmmCHmH muohmmcHMH HHS: HHS: HHDE HHS: coHumooq >1umq .mchh >HumH .mmCOh opwHU .mmcon mHocmum .mmu mHocmum .000 em .mzusHm em .mnusam mama mEmz mumczo new» mo umaa .N xaecmdda .. .4” 1:: :thLLLC EHozu no :OHuQHuOmva m Dcm mumzouo mcaumMQQOOU MO umd . .H .N XHDCTQW mooHOHHmo pom >mm suscuuoz cmsumc0h m OH mcoucmm om Essa mHHH>mquoou comm .HmHHHmanom 980m 0 N CMSDMGOb mH wEom CHHxsou ummnom .uwumHmm H v mooHOHHmQ :moHou wH cmoHow sHchoo uuonom .HoumHmm mooHOHHmQ pom smoucHoz smcumsoo N N mooHoHHmo copHoo om Eumm mpHmmm ocmuw ccmHU .mcHmem m v mam auocuuoz mmImH oEom moHQmm ocmuo scmHO .mCHxHom mEom mooHoHHmQ pom v e wmm cumnuuoz NH Hmcuou muummm Hohom .chHx mcmcmm HmucHz mommcHB >Qm :Hocuuoz swechoz m N :mzum20n om mEom muummm moo .chHm ammH ouommmum mmmue mmHumHum> mmm oEmz xoon coHumooq mEmz mumczo How» wo mcHumm mo .02 chz A.ucoov omuouHcoz moumcouo uHmzu mo coHumHHomoo m can mum3ouo mCHumummooo Mo umHH .N xHocmmm< H H musmmmum mmmue mo mCHumm mo .02 mooHOHHmQ.omm wmm cumnuuoz smoucHoz cmnumsoo U:MHHHOU msoHOHHmo pom wam cumnpuoz assumCOH msoHoHHmo 60m amoucHoz CMSDMCOH msoHoHHoQ pom conumc0h smoucHoz casuMCOh mDOHoHHoQ pom CMQDMCOU pom 33H moHumHum> C H flow.“ mm ov mm mm mv NH omd HmcomMHQ ommmoq ucoum xomm oEom .0>< seem mEmz xoon monmm pcmuo mpHmmm compo muuomm muummm mHHH>mHmaoou mHHH>mquoou coHumooq >uuoh .mEmuHmm >Huob .mEmuHmm mHHH>uO .Nuumznom mHHH>HO .Nuumzcom omua .umflHamsaom womb .umHHHmszom oEmz mquBO A.ucoov pmuouHcoz moumzouo uHozp mo :oHumHuome m can mumzouo msHumquooo mo ume .N mhmH Hum» prsmmmd AoHVSCUV UQWQJACCE ECLUZUHQ EH32; ~3 :3.H«LHHCI.JZ D 12.3 DHU3C~3 DCfia-wkmuQGCU HQ drum; ..N XH~42315~< H ouommmum wo mcHumm A.u:oov oH 0H wH N mmmue mo .02 UmuouHcoz 30cm msoHoHHmo our wmm sumnuuoz smoucHoz cmnumcoo mooHoHHmo cooHoo Hmcmmz mooHoHHmQ pom wan cumnuuoz cmnumcoo mooHoHHmQ cooHow pcmHauou mooHoHHmQ pom ham cumsuuoz smoucHoz mooHOHHoQ copHoo ucmummmcmua mchmouo .H.x msoHoHHoQ pom xmm cumsuuoz smoucHoz msoHoHHmQ :oUHoo nmuu moHumHum> :Hmz om mm mN (\I H :21 chucsoz oHQm< WWQHB UHO mpooz zuuoz .em cmEoum mEmz xoon UQMHmoum mason .um moron .um mmemco coHDMOOH snow .Nch anon .xomm coo .xowm HHHm .me04 ommH oEmz muoszo How» mpumnouo uHmzu mo coHuQHuomma m pom muo3ouo mcHumuomoou mo umHH .N xHocmmm< N muommmum mo msHumm A.ucoo. 0H 0H mmmuB mo .02 pmuouHcoz Nam cumsuuoz smoucHoz canam:oo msoHoHHmo pom wmm cumnuuoz msoHoHHmQ com >Qm cumzuuoz amoucHoz conumcon mooHoHHmo cooHoo mooHoHHmQ pom Nam :umnuuoz amoucHoz mmHumHum> :Hmz mv ov mH ow wmm eumnouo 6H0 660a mcoonE 466m acoum mEmz xoon muuoHsmao mmewco pcmHomum MHz: coHumooq chumm .mmccme mmuooo .uuonm room .cmEmoH mHocmum .mwo ommH @EMZ MHWCS HMWM mpnmnouo HHmzu mo :oHumHuome m can mumzouo mcHumummooo mo umHH .N XHpcmmm< whommmum mo mcHumm A.ucoov 0H mmmue m0 .02 pmuouHcoz msoHoHHoQ pom ham cuonuuoz smOHGHoz mooHoHHmo pom smoucHoz pom moH msoHoHHmo 60m 60m mHsma smoucHoz cmhumcon Umx MUH mooHOHHoQ cooHoo mmHumHum> chz moumnouo uHonu mo :oHuQHuomoo m can mumzouu mCHumummoou mo umHH mN mH mmm mmHmQ< xoon emxaz HMHccouch mEmz roon UcmHmoum cox UCMHWOH& mmfiflb GWWOHHMSU QHCHQQ .umcuse .cmmuona .mmcch ommH :oHuMOOH oEmz mumc30 use» .N prsme¢ AC Ka. D II) Ll -.4H+—- ‘ob—«Ommqmmhwmp‘ NHHt—‘I—JF—‘o—Jr: OKDODVGUI-IAUJI (wrumm #6me WNNNNM OKDCDNmU'I w H APPENDIX 3-1 Accumulation of Air Degree Days At Base 48°F Kalamazoo State Hospital Orchard in 1977 (B.E. Method). U 33 K NHI—‘I—‘I—‘b—‘Hr—‘l—‘HH Okoooummbwmr—aomooqowawmw NNNNNNNN mummwaI—i uww H00 APR 1 8 ll 12 12 12 13 14 15 28 47 7O 87 100 116 134 159 185 210 230 252 263 268 270 270 273 284 293 300 312 312 MAY 324 333 342 349 370 389 397 402 404 409 417 429 450 470 488 509 .534 560 586 611 636 661 685 711 737 756 775 798 816 835 857 JUN 865 871 880 897 918 925 930 937 945 956 967 976 986 1003 1021 1046 1076 1098 1120 1135 1151 1170 1190 1220 1242 1265 1292 1320 1340 1359 1359 JUL 1377 1394 1417 1454 1492 1533 1563 1592 1621 1651 1674 1702 1729 1756 1792 1823 1855 1887 1927 1965 1990 2012 2038 2063 2080 2096 2117 2143 2164 2192 2216 AUG 2236 2258 2286 2312 2338 2364 2391 2417 2440 2466 2480 2497 2518 2535 2552 2577 2588 2598 2608 2622 2637 2656 2671 2682 2696 2917 2752 2783 2801 2820 2847 SEP 2873 2893 2914 2941 2959 2984 2994 3016 3036 3046 3056 3067 3075 3087 3098 3117 3138 3161 3171 3177 3186 3201 3213 3230 3246 3258 3267 3280 3294 3302 3302 (8.9°C) at the OCT 3314 3318 3320 3325 3331 3337 3361 3361 3368 3372 3376 3376 3376 3378 3382 3383 3383 3386 3386 3388 3391 3399 3404 3404 3412 3412 3412 3412 3412 3412 3412 APPENDIX 3-2 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Kalamazoo State (B.E. Method). Hospital Orchard in 1977. DAY H OkOCDflmU'Io-WUJNH I 12 l3 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 bWNNI—‘H {DI rOU1oLuxJwcnU1w+Hc>o<3c>ooco m an \IONONOO‘ONUIUIQ HKDQQAHCDUJCD \I\I\I UlU'le MAY 77 81 85 87 94 102 109 112 114 117 120 124 133 140 149 157 169 181 193 205 217 229 241 254 267 279 291 303 315 327 339 JUN 349 357 365 373 385 395 403 411 419 426 434 442 451 461 472 486 503 521 538 555 568 582 597 614 633 651 668 687 705 719 719 JUL 734 751 767 789 813 837 862 886 909 932 952 975 999 1022 1045 1072 1096 1122 1148 1177 1206 1231 1255 1273 1295 1318 1338 1358 1379 1399 1422 AUG 1447 1471 1493 1515 1537 1558 1581 1602 1622 1644 1666 1685 1704 1723 1744 1763 1784 1802 1820 1836 1852 1868 1886 1903 1920 1936 1955 1978 2003 2026 2048 as: 2072 2097 2120 2143 2166 2187 2209 2229 2249 2267 2284 2299 2313 2328 2340 2354 2370 2388 2405 2419 2432 2446 2460 2475 2491 2506 2519 2534 2548 2560 2560 OCT 2573 2583 2593 2601 2609 2618 2626 2631 2637 2641 2645 2648 2649 2651 2653 2655 2656 2656 2657 2658 2659 2662 2664 2665 2667 2667 2667 2667 2667 2667 2667 APPENDIX 3-3 Accumulation of Two Inch Soil Degree Days on the North Side of the Tree at Base 48°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1977. (B.E. Method). 281 88.13 LAX >111“. 9.111: as SEPT 992 1 0 34 258 671 1379 1988 2493 2 0 36 266 687 1400 2011 2505 3 0 38 273 704 1423 2033 2516 4 0 40 282 726 1445 2056 2524 5 0 44 294 750 1467 2077 2532 6 0 51 304 771 1488 2098 2540 7 0 59 314 796 1508 2119 2549 8 0 63 324 820 1529 2139 2556 9 0 67 333 844 1549 2159 2563 10 0 69 342 867 1571 2177 2570 11 0 71 352 890 1591 2194 2576 12 0 74 363 913 1610 2209 2581 13 1 78 375 935 1630 2224 2584 14 3 82 389 957 1650 2239 2587 15 5 86 404 983 1668 2252 2590 16 7 91 422 1008 1688 2266 2594 17 9 95 439 1033 1706 2283 2596 18 12 101 456 1059 1722 2302 2597 19 15 108 471 1086 1738 2319 2599 20 18 116 485 1115 1754 2334 2602 21 19 125 499 1141 1771 2348 2605 22 21 135 514 1165 1788 2363 2608 23 22 146 531 1188 1805 2377 2612 24 24 159 549 1210 1821 2393 2615 25 26 171 567 1232 1837 2409 2618 26 27 184 585 1251 1855 2424 2618 27 29 196 604 1271 1877 2438 2618 28 30 209 623 1292 1901 2452 2618 29 32 221 639 1313 1922 2467 2618 30 33 234 655 1335 1943 2480 2618 31 33 246 655 1357 1965 2480 2618 APPENDIX 3-4 Accumulation of Air Degree Days At Base 48°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1978 (B.E. Method). 228 EBAAIMEEEAIEEEQEE 1 0 82 505 1074 1719 2364 2880 2 0 85 520 1088 1736 2385 2886 3 8 90 531 1108 1754 2409 2893 4 11 92 544 1127 1765 2426 2894 5 14 92 558 1150 1778 2448 2898 6 16 96 578 1193 2474 2898 3007 7 20 99 602 1202 1815 2502 2898 8 20 111 607 1220 1838 2535 2898 9 20 114 619 1240 1862 2565 2902 10 29 122 641 1254 1879 2593 2911 ll 29 131 666 1263 1898 2624 2915 12 34 149 688 1275 1921 2637 2922 13 34 160 694 1296 1946 2647 2925 14 34 166 700 1317 1973 2667 2928 15 34 176 717 1342 2001 2683 3928 16 35 187 736 1358 2024 2701 2925 17 37 203 764 1376 2048 2722 2925 18 37 223 788 1402 2074 2741 2928 19 38 243 808 1432 2095 ’ 2769 2929 20 38 248 831 1464 2108 2799 2934 21 39 257 845 1493 2124 2808 2945 22 42 267 859 1525 2145 2814 2959 23 42 286 877 1542 2170 2821 2959 24 48 304 902 1560 2197 2831 2959 25 52 324 924 1583 2219 2837 2960 26 58 349 951 1614 2242 2845 2960 27 64 375 981 1636 2266 2854 2962 28 71 400 1008 1649 2290 2857 2962 29 80 428 1033 1669 2309 2864 2962 30 82 456 1059 1683. 2328 2871 2967 31 82 477 1059 1699 2345 2871 2970 APPENDIX 3-5 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1978 (B.E. Method) m: flmwitaaesgiegz 1 0 15 274 745 1396 2048 2633 2 0 17 289 764 1417 2069 2645 3 O 19 302 783 1436 2091 2657 4 0 20 315 802 1454 2111 2669 5 0 20 328 821 1471 2132 2680 6 0 21 341 842 1490 2154 2689 7 0 23 356 865 1509 2177 2696 8 0 26 370 887 1528 2201 2703 9 0 29 382 908 1550 2225 2710 10 0 33 395 929 1569 2249 2719 ll 0 38 411 947 1589 2273 2729 12 O 45 426 953 1609 2293 2739 13 0 53 439 964 1630 2209 2747 14 0 59 449 1003 1652 2329 2754 15 0 65 462 1023 1675 2348 2759 16 0 72 475 1042 1698 2369 2763 17 O 81 492 1060 1720 2390 2767 18 0 91 509 1081 1744 2411 2772 19 0 103 525 1103 1766 2435 2776 20 0 116 542 1126 1786 2460 2781 21 0 126 557 1151 1806 2482 2787 22 0 134 572 1176 1826 2501 2795 23 0 144 587 1200 1848 2518 2801 24 l 155 604 1222 1874 2535 2804 25 2 167 621 1245 1896 2552 2807 26 3 180 641 1269 1919 2567 2810 27 6 195 662 1292 1941 2582 2813 28 8 209 683 1313 1964 2595 2816 29 11 225 704 1335 1986 2607 2818 30 14 242 725 1356 2008 2619 2820 31 14 258 725 1376 2028 2619 2823 APPENDIX 3-6 Accumulation of Two-Inch Soil Degree Days on the North Side of the Tree at Base 48°F Hospital Orchard in 1978 (B.E. Method) 991 £13,841 1 0 26 2 O 28 3 0 3O 4 0 32 5 0 32 6 0 34 7 0 35 8 0 38 9 0 41 10 0 45 11 0 48 12 0 55 13 0 63 14 0 68 15 0 73 16 1 79 17 1 86 18 1 95 19 1 , 106 20 1 117 21 2 126 22 3 134 23 3 143 24 5 152 25 7 163 26 10 175 27 13 188 28 17 202 29 20 216 30 24 231 31 24 245 JUN 261 274 284 295 306 318 330 341 351 362 376 391 401 409 419 430 445 460 474 489 501 512 525 541 557 575 595 615 636 656 656 JUL 676 693 711 728 747 767 789 810 831 850 867 882 900 919 939 957 975 995 1017 1040 1063 1088 1110 1131 1153 1177 1200 1220 1242 1252 1282 AUG 1304 1324 1344 1363 1380 1399 1418 1439 1462 1481 1501 1521 1543 1566 1590 1615 1639 1663 1687 1709 1732 1755 1781 1808 1830 1853 1876 1900 1925 1947 1972 (8.9°C) at the Kalamazoo State SEP 1996 2023 2051 2077 2104 2133 2162 2194 2223 2252 2282 2300 2315 2334 2355 2378 2400 2420 2445 2472 2498 2519 2538 2557 2577 2594 2160 2624 2636 2649 2649 OCT 2664 2677 2689 2699 2708 2715 2721 2727 2731 2740 2748 2757 2765 2771 2773 2774 2776 2779 2782 2786 2792 2799 2804 2805 2807 2808 2809 2811 2812 2813 2816 APPENDIX 3-7 Accumulation of Air Degree Days at Base 48°F {8.9°C) at the Kalamazoo State Hospital Orchard in 1979 (B.E. Method) 25: 9.115 mwflééfléflfl 1 0 138 446 1034 1730 2250 2700 2 0 148 458 1047 1751 2272 2723 3 0 150 476 1062 1774 2298 2734 4 0 150 497 1074 1798 2322 2744 5 0 152 516 1087 1820 2346 2760 6 0 164 539 1101 1843 2371 2772 7 0 183 560 1122 1872 2395 2780 8 0 209 587 1148 1896 2412 2788 9 0 236 615 1170 1920 2434 2793 10 0 264 636 1198 1936 2455 2794 11 0 286 649 1225 1945 2465 2798 12 12 286 660 1254 1959 2470 2799 13 16 292 674 1277 1971 2478 2799 14 17 298 695 1304 1976 2498 2799 15 17 303 724 1327 1983 2517 2799 16 18 309 751 1350 1995 2537 2799 17 20 317 774 1366 2007 2562 2799 18 24 338 796 1383 2020 2570 2799 19 30 352 814 1402 2041 2576 2799 20 40 362 836 1425 2052 2586 2799 21 44 366 865 1450 2065 2600 2799 22 51 373 892 1479 2085 2617 2799 23 63 381 909 1507 2108 2622 2799 24 73 383 917 1534 2120 2632 2799 25 87 389 929 1558 2134 2644 2799 26 92 394 940 1584 2149 2650 2799 27 97 400 957 1606 2165 2656 2799 28 103 405 980 1632 2181 2668 2799 29 115 417 1000 1658 2199 2679 2799 30 125 423 1020 1688 2215 2690 2799 31 137 435 1020 1711 2232 2690 2799 APPENDIX 3-8 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 84°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1979 (.B.E. Method). U 3’ K1 \OCDNO’NU'IbbJNF-‘l {D "U 55 NNI—‘t—‘H \meomooooooommooooooooooo p.50.» CDNLAJ \oooqqoxm meONm 102 MAY 104 109 115 117 121 129 140 156 174 194 211 219 227 236 245 254 263 276 288 299 308 317 326 333 340 347 354 361 369 377 386 JUN 395 405 417 429 443 456 472 488 508 529 544 558 572 589 609 627 647 664 680 697 714 734 753 768 781 795 810 828 847 864 864 JUL 880 896 912 930 945 961 977 994 1014 1035 1057 1080 1103 1128 1153 1176 1197 1218 1238 1258 1280 1302 1327 1351 1375 1397 1421 1445 1468 1495 1520 AUG 1541 1563 1585 1608 1629 1652 1672 1696 1719 1742 1761 1778 1794 1815 1829 1845 1857 1872 1889 1904 1922 1941 1961 1980 1998 2011 2029 2047 2065 2087 2107 SEP 2125 2144 2167 2190 2213 2236 2256 2277 2299 2317 2332 2351 2369 2389 2408 2428 2444 2464 2481 2499 2518 2533 2550 2567 2479 2592 2606 2621 2636 2656 2656 OCT 2673 2684 2691 2697 2705 2713 2724 2733 2740 2747 2754 2759 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 2764 APPENDIX 3-9 Accumulation of Two Inch Soil Degree Days on the North Side of the Tree at Base 48°F (8.9°F) at the Kalamazoo State Hospital Orchard in 1979 (B.E. Method). 25: flflflflkwéflfl 1 0 76 352 835 1486 2082 2620 2 0 79 361 851 1508 2103 2635 3 0 84 372 866 1530 2125 2652 4 0 87 384 884 1553 2148 2666 5 0 89 397 900 1575 2172 2680 6 0 95 410 916 1598 2196 2696 7 0 104 425 933 1622 2219 2707 8 0 116 443 950 1647 2240 2718 9 0 131 462 970 1670 2262 2728 10 0 146 481 990 1693 2284 2738 11 0 162 496 1011 1715 2304 2744 12 0 172 510 1034 1733 2320 2750 13 0 180 524 1059 1751 2335 2755 14 1 189 540 1083 1766 2352 2755 15 1 198 558 1105 1781 2370 2755 16 1 207 577 1128 1796 2388 2755 17 1 215 597 1150 1810 2408 2755 18 1 226 614 1170 1826 2424 2755 19 2 238 630 1190 1844 2439 2755 20 4 249 647 1210 1861 2454 2755 21 6 259 667 1231 1879 2469 2755 22 9 267 686 1253 1898 2486 2755 23 14 277 704 1276 1918 2499 2755 24 19 285 720 1299 1938 2512 2755 25 26 293 735 1322 1957 2527 2755 26 34 301 749 1345 1976 2539 2755 27 41 308 764 1367 1990 2551 2755 28 49 316 781 1391 2005 2564 2755 29 57 324 799 1414 2020 2578 2755 30 65 332 818 1437 2042 2600 2755 31 75 342 818 1463 2062 2600 2755 APPENDIX 3-10 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1980 (B.E. Method). 0 mcn\umuwncunna I» K: 10 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 APR H OU‘HHi—‘O 18 25 25 25 26 26 26 26 26 26 29 36 47 59 70 92 105 105 105 108 110 111 115 118 118 MAY 124 135 150 165 190 207 212 212 215 225 231 241 254 258 264 272 280 288 303 314 330 352 372 396 416 433 448 468 490 512 527 JUN 543 560 576 586 599 616 630 637 644 649 662 678 694 708 722 739 755 770 781 789 809 837 870 900 933 964 990 1016 1026 1036 1036 JUL 1055 1076 1096 1119 1140 1159 1187 1216 1241 1266 1295 1321 1346 1375 1409 1440 1467 1491 1521 1557 1586 1611 1630 1650 1675 1700 1719 1740 1762 1780 1805 AUG 1835 1858 1881 1908 1936 1961 1989 2021 2052 2078 2101 2121 2138 2163 2183 2197 2213 2233 2255 2283 2314 2336 2351 2369 2394 2419 2449 2479 2510 2538 2565 SEPT 2590 2616 2636 2661 2684 2706 2725 2747 2772 2784 2797 2816 2841 2863 2879 2893 2902 2911 2927 2948 2974 2997 3009 3017 3027 3030 3036 3046 3058 3071 APPENDIX 3-11 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F Hospital Orchard in 1980. DAY \omummbwmr—I APR r—u—w—w-I QQ'WNCDUIUJF-‘OOOOOOOOOOOOOOOOOO NNNNN 0000me MAY 30 32 36 39 42 46 49 53 55 58 61 65 69 74 79 85 92 99 108 119 129 142 154 168 181 194 208 222 237 251 266 JUN 280 293 307 320 334 347 361 374 387 399 411 423 435 448 460 474 487 501 514 528 542 558 575 594 613 634 654 675 695 715 715 JUL 735 755 775 794 815 835 856 877 898 921 943 966 988 1012 1035 1059 1083 1107 1131 1155 1180 1205 1230 1255 1280 1305 1331 1356 1382 1407 1434 (8.9°C) at the Kalamazoo State (B.E. Method). AUG 1461 1486 1511 1537 1563 1589 1615 1641 1667 1692 1717 1742 1766 1789 1812 1835 1858 1881 1904 1929 1953 1977 2001 2026 2051 2077 2102 2129 2155 2182 2208 912.22 2234 2260 2286 2312 2338 2363 2388 2413 2437 2461 2485 2508 2531 2553 2575 2596 2616 2636 2655 2674 2692 2711 2729 2745 2760 2774 2787 2799 2810 2821 APPENDIX 3-12 Accumulation of Two Inch Soil Degree Days on the North Side of the Tree at Base 48°F (8.9°C) at the Kalamazoo State Hospital Orchard in 1980. (B.E. Method). 21:: A35 m 199 9.99 9.09 _SEPT 1 0 45 270 696 1441 2245 2 0 49 282 715 1466 2274 3 0 54 294 734 1491 2303 4 0 60 306 755 1516 2331 5 0 66 318 776 1542 2359 6 0 72 331 794 1568 2386 7 0 77 345 818 1597 2413 8 0 80 357 842 1625 2439 9 0 82 368 865 1652 2466 10 0 84 378 889 1677 2491 11 0 88 387 912 1702 2516 12 0 91 397 836 1725 2540 13 0 95 407 960 1748 2564 14 0 99 418 985 1771 2587 15 0 102 429 1010 1795 2609 16 0 105 439 1036 1818 2631 17 0 110 450 1060 1842 2651 18 1 116 461 1085 1865 2670 19 2 126 471 1109 1889 2689 20 5 136 482 1135 1914 2707 21 9 147 493 1161 1940 2726 22 15 159 509 1187 1966 2744 23 20 169 528 1213 1990 2761 24 25 182 549 1238 2015 2777 25 29 194 571 1264 2041 2791 26 31 206 594 1290 2069 2805 27 33 216 616 1315 2098 2818 28 35 226 638 1339 2127 2830 29 38 237 657 1364 2157 2842 30 41 248 677 1389 2186 2854 31 41 259 677 ‘1415 2216 .lala‘a‘fl‘a‘fi‘33 APPENDIX 3-13 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Upjohn Orchard in 1977 (B.E. Method). 9511 flflflflflflfl 1 1 273 859 1357 2205 2790 3167 2 8 282 865 1376 2224 2812 3179 3 14 291 874 1397 2249 2829 3188 4 15 298 882 1433 2279 2847 3193 5 15 321 893 1468 2307 2861 3197 6 15 343 899 1508 2334 2875 3198 7 17 350 904 1542 2361 2894 3198 8 17 357 911 1572 2385 2915 3203 9 18 360 919 1601 2406 2935 3203 10 29 367 929 1627 2433 2943 3207 11 46 376 940 1652 2445 2955 3207 12 61 389 949 1680 2460 2962 3207 13 77 413 959 1706 2480 2970 3208 14 85 431 976 1733 2496 2979 3211 15 98 450 995 1771 2514 2986 3211 16 112 472 1019 1802 2536 3002 3212 17 131 498 1049 1835 2547 3021 3213 18 152 525 1071 1864 2556 3040 3213 19 172 553 1093 1902 2565 3057 3215 20 192 583 1108 1940 2577 3063 3218 21 211 613 1124 1969 2588 3070 3224 22 219 639 _1143 1993 2604 3083 3224 23 223 665 1165 2018 2615 3093 3224 24 227 693 1195 2042 2625 3108 3232 25 227 722 1216 2058 2638 3122 3232 26 231 741 1241 2072 2655 3131 3232 27 241 763 1270 2090 2686 3137 3232 28_ 250 788 1297 2112 2713 3143 3232 29 254 807 1318 2135 2727 3150 3232 30 261 829 1335 2162 2744 3161 3232 31 261 851 1335 2186 2768 3161 3232 Accumulation of Two Inch of the Tree at Base 48°F (B.E. Method). 1977. DAY H oxoooqmmhwmt—I I 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 3O 31 APR oomari—‘ooooooooooo WNWWWWMWWNNHHH mmmmmmmwwqwmwr—a MAY 39 43 44 46 55 63 68 70 72 76 82 91 101 111 122 136 151 168 186 204 221 238 256 274 290 306 321 337 352 367 381 APPENDIX 3-14 Soil Degree Days on the South Side (8.9°C) at the Upjohn Orchard in JUN 390 398 405 414 423 434 442 450 457 464 470 479 487 498 509 522 538 555 572 586 599 613 628 645 663 680 698 718 735 751 751 JUL 768 784 801 823 848 874 900 922 945 967 986 1009 1030 1052 1079 1103 1128 1152 1179 1207 1232 1252 1272 1292 1312 1330 1347 1365 1384 1404 1426 AUG 1444 1461 1481 1501 1521 1542 1563 1585 1606 1628 1647 1663 1680 1697 1712 1730 1745 1755 1763 1772 1784 1795 1806 1817 1827 1836 1850 1870 1887 1903 1921 9923. 1940 1959 1976 1992 2008 2024 2041 2059 2076 2089 2101 2111 2122 2132 2140 2150 2164 2180 2194 2205 2214 2225 2236 2249 2260 2270 2278 2290 2301 2312 2312 OCT 2321 2327 2331 2335 2340 2341 2341 2344 2344 2345 2345 2345 2345 2345 2345 2345 2345 2345 2345 2345 2345 2345 2345 2346 2346 2346 2346 2346 2346 2346 2346 .1ula‘a‘r‘a‘fiz33 APPENDIX 3-15 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1977. (B.E. Method). 293 9P_R 11.29: m 299 ABE _SEPT 99? 1 0 127 481 865 1555 2123 2560 2 0 131 489 882 1574 2143 2569 3 2 135 496 899 1595 2162 2576 4 3 137 504 921 1617 2181 2583 5 4 147 513 945 1639 2201 2590 6 4 160 525 970 1660 2219 2594 7 5 171 533 994 1682 2238 2596 8 5 179 541 1016 1705 2258 2600 9 6 184 549 1038 1726 2277 2603 10 7 188 556 1059 1749 2293 2605 11 9 193 564 1079 1770 2308 2606 12 11 201 572 1101 1789 2322 2606 13 14 211 581 1122 1808 2334 2606 14 18 222 591 1143 1829 2347 2607 15 23 232 602 1168 1847 2358 2607 16 29 244 616 1193 1866 2370 2607 17 36 257 633 1217 1885 2384 2607 18 44 271 651 1241 1899 2400 2607 19 53 287 668 1268 1911 2416 2607 20 63 303 683 1297 1923 2428 2607 21 74 319 696 1324 1936 2439 2608 22 84 335 710 1347 1949 2452 2608 23 94 351 725 1369 1963 2463 2608 24 104 367 742 1390 1976 2476 2609 25 111 383 759 1413 1989 2489 2609 26 116 399 775 1432 2003 2503 2609 27 120 413 794 1451 2023 2513 2609 28 122 428 813 1470 2045 2525 2609 29 123 443 832 1490 2065 2538 2609 30 125 458 848 1511 2083 2550 2609 31 125 471 848 1535 2102 2550 2609 Accumulation of Air Degree Days at Base APPENDIX 3-16 Upjohn Orchard in 1978 (B.E. Method) DAY \OCDQONU'l-thH APR 2 2 8 11 13 14 19 19 19 28 28 33 33 33 33 34 36 36 37 37 38 41 41 46 51 56 63 70 78 80 80 MAY 81 84 88 90 90 95 98 110 112 119 128 146 157 163 169 179 191 206 225 245 251 260 270 289 309 333 360 386 413 439 459 JUN 485 501 512 524 538 556 579 584 595 613 636 658 664 669 686 705 730 755 775 797 812 825 842 866 888 915 943 968 993 1018 1018 JUL 1036 1051 1071 1090 1112 1135 1161 1180 1200 1214 1225 1239 1260 1280 1303 1319 1337 1361 1391 1422 1451 1482 1499 1519 1542 1572 1597 1610 1631 1645 1661 48°F (8.9°C) at the AUG 1683 1702 1723 1735 1749 1769 1788 1811 1837 1854 1874 1898 1925 1953 1982 2006 2031 2058 2081 2095 2112 2135 2162 2189 2212 2236 2262 2288 2308 2327 2346 SEP 2365 2386 2410 2426 2449 2474 2503 2534 2564 2592 2622 2635 2644 2665 2682 2701 2721 2742 2769 2798 2809 2816 2825 2837 2844 2852 2862 2866 2874 2882 2882 OCT 2892 2901 2910 2912 2915 2916 2916 2916 2920 2931 2936 2944 2948 2948 2948 2948 2949 2952 2954 2961 2972 2985 2985 2986 2988 2988 2991 2991 2993 2999 3003 APPENDIX 3-17 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1978 (B.E. Method). m wwwflwflw 1 0 9 264 709 1333 1963 2474 2 0 10 280 726 1353 1982 2484 3 0 11 293 744 1374 2002 2495 4 O 11 305 762 1391 2019 2503~ 5 0 11 317 781 1406 2038 2510 6 0 12 330 801 1424 2058 2515 7 0 - 12 345 823 1443 2079 2518 8 0 15 356 845 1463 2102 2524 9 0 16 367 864 1484 2126 2526 10 0 20 379 883 1503 2149 2532 11 0 23 393 898 1522 2173 2539 12 0 30 408 914 1543 2193 2547 13 0 39 418 932 1565 2206 2552 14 0 45 426 951 1588 2223 2553 15 0 50 437 971 1612 2240 2553 16 0 56 449 990 1636 2258 2553 17 0 63 465 1008 1658 2276 2553 18 0 74 482 1028 1681 2296 2555 19 0 86 498 1051 1705 2318 2555 20 0 99 515 1074 1723 2342 2558 21 0 107 528 1099 1741 2359 2563 22 0 114 541 1125 1760 2374 2571 23 0 124 555 1147 1781 2387 2573 24 1 134 571 1168 1803 2400 2573 25 1 146 588 1190 1825 2413 2575 26 2 159 607 1214 1847 2424 2576 27 3 175 628 1237 1868 2435 2576 28 5 192 648 1256 1891 2444 2577 29 8 211 669 1276 1910 2452 2577 30 9 229 690 1295 1928 2463 2577 31 9 246 690 1314 1946 2463 2579 APPENDIX 3-18 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1978 (B.E. Method). w flwflflflfllfiéflfl 1 0 33 291 738 1367 2056 2684 2 0 36 306 754 1388 2079 2698 3 0 41 318 772 1409 2104 2711 4 0 42 329 790 1427 2127 2720 5 0 42 342 809 1444 2152 2728 6 0 45 354 828 1463 2177 2734 7 1 46 369 850 1483 2205 2739 8 1 49 381 871 1504 2234 2743 9 1 52 391 890 1527 2262 2748 10 1 58 403 910 1547 2290 2758 11 1 61 418 926 1567 2318 2766 12 1 69 435 942 1589 2337 2775 13 1 78 446 960 1612 2351 2784 14 1 84 455 979 1637 2370 2787 15 1 89 466 999 1663 2390 2790 16 1 95 479 1018 1688 2412 2790 17 2 103 495 1036 1713 2433 2793 18 2 114 513 1057 1738 2453 2798 19 2 126 528 1079 1763 2477 2800 20 2 138 545 1103 1784 2504 2806 21 2 147 561 1127 1805 2523 2815 22 3 155 575 1153 1828 2541 2824 23 3 164 589 1176 1853 2560 2830 24 5 174 605 1197 1878 2579 2833 25 8 184 621 1219 1901 2597 2835 26 11 196 640 1243 1923 2615 2837 27 16 210 660 1266 1946 2631 2839 28 21 224 679 1286 1969 2645 2842 29 25 241 699 1308 1991 2658 2844 30 30 258 719 1327 2012 2671 2846 31 30 274 719 1347 2034 2671 2849 APPENDIX 3-1 9 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Upjohn Orchard in 1979. (B.E. Method). 992 9113 11.4.: 9.1m & .499 SEPT 992 1 0 80 374 880 1512 2070 2478 2 0 88 385 893 1528 2095 2489 3 0 91 403 912 1547 2120 2491 4 O 91 419 923 1568 2141 2498 5 0 92 431 934 1600 2163 2499 6 0 103 454 947 1621 2180 2501 7 0 119 472 963 1643 2201 2506 8 0 143 499 982 1674 2218 2506 9 0 168 526 1004 1698 2229 2508 10 0 193 553 1026 1721 2243 2510 11 O 214 563 1050 1738 2256 2510 12 7 214 573 1076 1748 2274 2510 13 11 219 586 1102 1758 2291 2510 14 13 224 605 1127 1770 2315 15 13 229 633 1147 1775 2327 16 15 236 659 1168 1783 2331 17 18 244 674 1180 1798 2338 18 21 263 684 1194 1809 2349 19 27 277 707 1209 1823 2364 20 33 287 728 1227 1842 2373 21 41 291 750 1245 1853 2379 22 48 298 759 1269 1872 2386 23 56 303 765' 1297 1892 2392 24 66 313 773 1324 1915 2397 25 76 319 784 1343 1925 2404 26 78 324 802 1363 1939 2415 27 79 327 822 1390 1955 2425 28 79 332 845 1412 1970 2435 29 79 343 859 1435 1986 2446 30 79 346 872 1461 2014 2464 31 79 357 872 1489 2046 2464 APPENDIX 3-20 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1979. (B.E. Method). .129: 542 1193. 229 119 25% SEPT .0C_T 1 0 29 220 652 1220 1782 2192 2 0 30 229 687 1240 1802 2210 3 0 30 238 703 1258 1822 2228 4 0 30 248 718 1277 1842 2242 5 0 30 262 729 1297 1862 2252 6 0 31 278 741 1317 1880 2259 7 0 34 298 754 1337 1898 2266 8 0 41 318 769 1359 1916 2270 9 0 53 339 786 1380 1927 2275 10 0 66 351 803 1401 1936 2280 11 0 80 363 822 1421 1947 2283 12 0 89 374 843 1437 1962 2284 13 0 96 388 864 1451 1977 2285 14 0 105 406 886 1465 1994 15 0 113 424 906 1478 2009 16 1 118 442 924 1489 2019 17 2 122 456 941 1501 2029 18 3 133 469 956 1512 2040 19 5 143 484 971 1526 2052 20 8 151 501 987 1542 2062 21 11 156 519 1005 1557 2069 22 15 162 535 1023 1573 2079 23 19 168 546 1044 1589 2088 24 23 171 556 1065 1609 2095 25 27 174 567 1084 1625 2103 26 28 177 579 1102 1641 2112 27 29 182 594 1122 1657 2120 28 29 186 609 1141 1685 2138 29 29 193 626 1159 1710 2155 30 29 201 639 1179 1736 2174 31 29 210 639 1197 1764 2174 APPENDIX 3-21 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1979. (B.E. Method). w 893 MAX 1119 9.99 2% SEPT fl 1 0 43 273 704 1243 1808 2341 2 0 46 282 718 1262 1828 2362 3 0 50 293 733 1281 1849 2385 4 0 50 306 748 1300 1870 2399 5 0 51 320 760 1320 1891 2410 6 0 56 336 772 1339 1911 2417 7 0 66 354 785 1359 1931 2424 8 0 80 373 799 1381 1951 2429 9 0 . 96 394 815 1404 1965 2432 10 0 114 406 832 1424 1987 2436 11 0 129 417 850 1442 2009 2438 12 0 135 428 869 1459 2025 2439 13 0 140 442 888 1474 2042 14 1 146 460 909 1489 2060 15 2 153 478 929 1503 2078 16 3 160 495 948 1514 2092 17 5 167 510 966 1526 2105 18 8 179 524 981 1537 2119 19 11 190 539 995 1551 2134 20 15 200 556 1011 1566 2151 21 19 207 572 1027 1579 2165 22 24 214 588 1044 1596 2178 23 29 222 599 1063 1613 2191 24 34 227 610 1083 1632 2202 25 39 230 621 1104 1652 2215 26 41 235 633 1122 1669 2229 27 42 240 648 1140 1684 2246 28 42 245 663 1160 1711 2263 29 42 250 678 1180 1736 2282 30 42 256 692 1200 1761 2302 31 42 263 692 1220 1789 2321 APPENDIX 3-22 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Upjohn Orchard in 1980 (B.E. Method). 991! flflmflwfla 1 0 84 397 918 1676 2380 2 2 96 415 938 1691 2400 3 2 109 426 959 1711 2427 4 2 126 440 981 1737 2447 5 4 133 461 999 1758 2469 6 10 133 480 1027 1783 2485 7 18 133 485 1060 1813 2506 8 29 133 493 1084 1842 2528 9 29 135 496 1109 1868 2540 10 29 142 504 1137 1887 2552 11 29 145 519 1160 1905 2570 12 29 153 540 1183 1921 2595 13 29 160 563 1214 1939 2615 14 29 166 579 1252 1957 2627 15 29 170 586 1278 1970 2642 16 29 176 595 1299 1983 2649 17 29 180 609 1320 2006 2659 18 31 189 628 1350 2028 2675 19 37 198 639 1386 2056 2698 20 45 210 654 1412 2086 2724 21 54 227 676 1438 2106 2746 22 64 245 701 1455 2123 2754 23 79 263 729 1475 2142 2762 24 80 273 754 1499 2166 2774 25 80 281 783 1521 2191 2780 26 80 295 812 1539 2220 2787 27 80 318 839 1556 2250 2797 28 80 341 860 1578 2281 2812 29 81 356 876 1595 2307 2826 30 81 365 895 1623 2332 2826 31 81 381 895 1650 2356 APPENDIX 3-23 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1980. (B.E. Method). 942 99.1.3 11A: J_UN & .4153. SEPT 1 0 11 209 614 1212 1880 2 0 13 218 630 1231 1903 3 0 18 225 652 1251 1927 4 0 24 226 672 1273 1948 5 0 31 226 691 1294 1969 6 0 33 230 711 1317 1988 7 0 36 237 739 1341 2008 8 0 36 248 760 1366 2027 9 0 36 262 781 1391 2045 10 0 40 275 801 1413 2063 11 0 44 284 819 1434 2080 12 0 48 292 840 1453 2098 13 0 51 300 863 1472 2116 14 0 51 309 887 1492 2131 15 0 53 320 907 1509 2147 16 0 55 336 925 1530 2160 17 0 58 353 940 1548 2176 18 0 63 372 959 1568 2195 19 0 69 389 979 1590 2214 20 1 79 402 998 1612 2233 21 2 89 419 1017 1632 2253 22 5 100 435 1036 1652 2265 23 8 109 457 1052 1673 2279 24 9 116 479 1070 1695 2294 25 10 124 500 1085 1719 2303 26 10 135 521 1099 1745 2312 27 10 149 542 1111 1770 2323 28 10 163 562 1126 1794 2334 29 10 173 580 1146 1817 2349 30 10 187 596 1168 1838 2459 31 10 197 596 1190 1859 APPENDIX 3-24 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Upjohn Orchard in 1980. (B.E. Method). ELY 9P_R 1493. .4911 199. A119 SEPT 1 0 23 222 639 1322 2011 2 0 24 234 658 1341 2035 3 0 29 245 676 1361 2059 4 0 36 257 695 1383 2081 5 0 38 267 714 1404 2104 6 0 42 279 734 1427 2124 7 0 51 292 754 1451 2146 9 0 51 302 778 1475 2166 8 0 53 313 803 1499 2183 10 0 56 322 826 1521 2200 11 0 59 331 849 1542 2216 12 0 63 340 871 1562 2237 13 0 67 351 894 1582 2256 14 0 69 364 919 1603 2272 15 0 72 379 945 1623 2290 16 0 75 392 971 1642 2303 17 0 78 404 996 1661 2321 18 0 84 417 1020 1683 2338 19 1 91 431 1045 1707 2358 20 '3 99 445 1070 1730 2380 21 7 107 458 1094 1751 2401 22 12 116 474 1115 1772 2415 23 19 127 491 1137 1794 2429 24 21 136 509 1158 1817 2446 25 21 144 527 1179 1841 2454 26 21 152 546 1200 1867 2462 27 21 163 565 1221 1894 2474 28 21 176 583 1241 1920 2487 29 21 188 603 1260 1945 2503 30 21 199 622 1279 1967 2503 31 21 211 622 1300 1989 APPENDIX 3-25 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Hofacker Site in 1980 (B.E. Method). 241 A_PB. 19! 29.11 99.1: 999 1 0 234 591 1037 1757 2 2 243 605 1053 1779 3 3 264 616 1075 1801 4 14 282 630 1098 1829 5 28 295 645 1116 1851 6 43 302 659 1135 1877 7 57 302 678 1166 8 77 302 688 1190 9 87 302 691 1210 10 90 311 695 1237 11 94 315 703 1265 12 102 324 716 1289 13 120 326 735 1315 14 141 328 755 1341 15 160 339 759 1363 16 164 342 765 1383 17 165 347 777 1408 18 172 356 794 1435 19 180 366 804 1465 20 191 380 807 1502 21 206 394 822 1522 22 221 410 843 1538 23 222 429 868 1555 24 222 452 895 1579 25 222 471 923 1600 26 223 483 947 1619 27 224 497 965 1640 28 225 518 988 1661 29 226 539 1002 1681 30 228 561 1016 1707 31 228 578 1016 1734 APPENDIX 3-26 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Hofacker Site in 1980 (B.E. Method). PA! A_P_P m 999 £11. .499 1 0 146 360 689 1301 2 2 150 371 704 1321 3 9 158 383 719 1343 4 20 166 395 736 1365 5 32 175 406 754 1386 6 43 181 421 771 1408 7 58 183 431 790 8 68 183 435 811 9 73 183 439 830 10 77 183 439 852 11 78 187 443 874 12 82 192 450 895 13 91 194 460 917 14 103 196 471 942 15 117 200 480 967 16 127 203 487 989 17 127 206 494 1009 18 127 212 505 1027 19 128 218 510 1049 20 131 225 518 1072 21 136 231 528 1093 22 142 238 542 1111 23 143 248 557 1127 24 143 262 575 1144 25 143 275 594 1163 26 143 286 614 1181 27 143 296 631 1200 28 143 308 648 1220 29 , 143 321 662 1239 30 144 335 675 1259 31 144 349 675 1279 APPENDIX 3-27 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Hofacker Site in 1980 (B.E. Method). 999. 91:13 999 999 999 999 1 0 231 516 899 1556 2 0 245 528 915 1576 3 0 262 539 935 1598 4 11 280 551 954 1622 5 23 298 562 973 1644 6 34 311 580 990 1667 7 52 314 591 1013 8 63 314 595 1036 9 67 319 600 1056 10 72 327 607 1080 11 76 334 612 1103 12 ‘ 82 346 622 1126 13 93 349 630 1151 14 108 352 647 1180 15 125 358 654 1206 16 132 360 661 1229 17 132 364 670 1250 18 143 372 682_ 1270 19 154 379 687 1295 20 168 387 698 1322 21 182 393 712 1343 22 198 401 730 1365 23 206 413 748 1382 24 206 427 768 1400 25 211 440 791 1417 26 213 447 812 1435 27 216 454 830 1454 28 217 465 850 1474 29 219 479 867 1492 30 223 492 882 1513 31 223 504 882 1535 APPENDIX 3-28 Accumulation of Air Degree Days at Base 48°F (8.9°C) at the Yabs Site in 1980 (B.E. Method). 999 99E. 999 999 J_UL. 999 1 0 129 681 1346 2272 2 0 148 702 1375 2297 3 0 168 722 1408 2327 4 0 192 741 1432 2361 5 0 215 759 1459 2390 6 0 233 784 1486 2419 7 0 238 804 1517 2451 8 0 243 821 1548 2485 9 0 251 833 1578 2516 10 0 262 845 1601 2543 11 0 279 862 1629 2569 12 0 294 883 1658 2596 13 0 307 907 1692 2621 14 0 317 927 1729 2648 15 0 329 944 1767 2672 16 3 344 964 1801 2698 17 8 355 987 1832 2717 18 14 367 1006 1862 2745 19 23 387 1021 1896 2773 20 37 408 1042 1935 2804 21 54 430 1064 1963 2832 22 74 455 1090 1994 2861 23 89 479 1119 2022 2890 24 89 503 1148 2053 2923 25 97 525 1180 2086 2952 26 99 545 1210 2111 2986 27 103 570 1238 2136 3021 28 107 597 1269 2160 3056 29 114 621 1299 2186 3091 30 119 645 1320 2209 3124 31 119 667 1320 2239 3153 APPENDIX 3-29 Accumulation of Two Inch Soil Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Yabs Site in 1980 (B.E. Method). 999 9P9 LA: at! 999 999 l O 77 487 1043 1831 2 0 90 506 1062 1852 3 0 107 518 1088 1874 4 0 134 530 1113 1904 5 0 152 546 1136 1931 6 0 161 570 1155 1960 7 0 161 584 1189 1991 8 0 161 592 1214 2021 9 O 164 598 1238 2047 10 0 170 601 1261 2070 11 0 179 611 1288 2093 12 0 190 631 1310 2110 13 0 195 656 1332 2133 14 0 198 688 1369 2156 15 O 203 695 1401 2169 16 0 215 704 1431 2189 17 0 226 720 1454 2207 18 0 235 739 1483 2228 19 12 247 753 1516 2256 20 20 260 769 1552 2288 21 35 277 789 1577 2314 22 54 298 814 1600 2334 23 57 324 841 1618 2355 24 57 347 869 1639 2381 25 59 357 900 1667 2407 26 59 366 929 1689 2437 27 61 386 950 1710 2469 28 63 412 979 1731 2501 29 66 441 1003 1751 2535 30 69 466 1018 1774 2562 31 69 479 1018 1803 2588 APPENDIX 3-30 Accumulation of Ground Litter Degree Days on the South Side of the Tree at Base 48°F (8.9°C) at the Yabs Site in 1980 (B.E. Method). 919 913.13 999 1 0 79 2 0 92 3 0 107 4 0 125 5 0 142 6 0 155 7 0 161 8 0 165 9 0 169 10 0 177 11 0 189 12 0 200 13 0 208 14 0 216 15 0 225 16 0 235 17 0 246 18 0 257 19 7 272 20 20 288 21 33 305 22 46 324 2 53 344 24 53 365 25 59 383 26 59 398 27 63 415 28 65 436 29 68 458 30 72 479 31 72 498 JUN 512 531 547 563 579 601 619 633 644 653 665 680 698 718 735 750 766 784 799 815 835 855 877 901 928 954 978 1003 1025 1045 1045 JUL 1067 1087 1111 1135 1160 1181 1207 1232 1257 1282 1310 1334 1361 1392 1422 1451 1477 1504 1534 1566 1594 1621 1645 1670 1697 1721 1746 1771 1795 1818 1847 AUG 1876 1901 1928 1958 1988 2018 2049 2080 2109 2136 2163 2188 2213 2241 2263 2287 2308 2333 2360 2389. 2415 2441 2467 2495 2523 2553 2585 2617 2650 2680 2708 GRN STATE UNIV. LI