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This behavior consists of a series of events which are stimulated in a receptive mite by an air speed > 1 mile per hour: (I) initially the receptive mites alter their behavior from a random search movement to a directional movement toward the edge of the arena where (2) they terminate all forward motion, (3) begin to orientate to the air flow and (4) eventually assume an anteriorly raised stance heading downwind of the air flow from which they (5) frequently disperse when the air current is > 6.3 mph. The preovipositing adult females, ovipositing adult females. and to a lesser extent the adult males are the principal life stages that exhibit the behavior and actively disperse. Starvation increases the dispersal behavior of the ovipositing adult females and to a limited extent the adult males and the preovipositing adult females. LABORATORY STUDY OF THE DISPERSAL BEHAVIOR OF AMBLYSEIUS FALLACIS (GARMAN) (ACARINA: PHYTOSEIIDAE) BY Donn Tiffany Johnson A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Entomology 1976 To Paula who made it a reality ii ACKNOWLEDGMENTS I would like to thank everyone in the lab for their encouragement and constructive criticism. I would especially like to thank my advisor. Dr. Brian A. Croft, for his proof- reading and helpful additions that have made this thesis what it is. My greatest appreciation goes to my family, and especially to Paula who all showed great perseverance throughout these last two years. iii TABLE OF CONTENTS Page LIST OF TABLES . . . . . . . . . . . . . . V LIST OF FIGURES. . . . . . . . . . . . . . vi INTRODUCTION. . . . . . . . . . . . . . . 1 METHODS AND MATERIALS. . . . . . . . . . . . “ RESULTS AND DISCUSSION . . . . . . . . . . . 9 SUMMARY AND CONCLUSIONS . . . . . . . . . . . 23 LIST OF REFERENCES . . . . . . . . . . . . 26 APPENDIX . . . . . . . . . . . . . . . . 30 iv Table l. A-Bo Aw. A-7. LIST OF TABLES Wind Tunnel Observations of the Response of Certain Stages of A. fallacis to Air Currents. Pairs of Treatments that are Significantly Different as Determined with Bonferroni Chi Square 2x2 Contingency Tables . . . . . . Amblyseius fallacis Life Stage Characteristics Air Speeds Produced by the Fan Arrangement by Varying Fan Distance from the Stereo- microscope . . . . . . . . . . . . Air Speeds Produced by the Wind Tunnel by Varying Thicknesses of the Cheese Cloth. . . Data of the Observations of the Behavioral Response of the Ovipositing Females to Different Treatment Combinations of Air Speeds (mph), Temperature, and Days of Starvation . . . . . . . . . . . . Data of the Observations of the Behavioral Responses of the Preovipositing Females to Different Treatments . . . . . . . . . Data of the Observations of the Behavioral Responses of the Adult Males to Different Treatments . . . . . . . . . . . . Data of the Observations of the Behavioral Responses of the Immature Stages to Different Treatments . . . . . . . . . 21 30 31 31 32 33 34 35 Figure 1. LIST OF FIGURES The sequence of behavioral events of the response of A. fallacis to air currents; (A) the randOm search movement; (B) dorsal view of mite orientation to the prevailing air flow; (C) lateral view of the stance heading away from the air flow (af); (D) lateral view of the mite immediately before dispersing via the air . . . . Photographs of the stance assumed by A. fallacis ovipositing adult females in response to air currents. . . . . Stance angle of orientation of the ovi- positing adult remale in response to the laminar air current in the wind tunnel . Effects of temperature and mite starvation on the percent response of three life stages of A. fallacis to air currents . vi Page 10 13 1h 19 INTRODUCTION The predaceous mite. Amblyseius fallacis (Garman). a member of the family Phytoseiidae, is found in commercial apple orchards throughout the central and northern United States (Poe et al., 1969; Croft and Brown, 1975) and Canada (Parent. 1958; LeRoux, 1961). In these orchards, A. fallacis is an effective predator of plant-feeding mites including the European red mite, Panonychus Elfll (Koch), the apple rust mite. Aculus schlechtendali (Napela). and the two- spotted spider mite. Tetrggychus urticae Koch. Biological control of g. Elfli is possible when there is a prey (P, Agml) to predator (A. fallacis) ratio of ca 211031 (Croft, 1975). Furthermore, it has been shown that A. fallacis is an im- portant predator in commercial orchards due to its additional trait of resistance to several organOphosphorus-related pesticides including parathion, azinphosmethyl (Guthion), phosmet (Swift, 1968; Motoyama et al., 1970; Croft and Brown, 1975) and a carbamate pesticide. carbaryl (SevinfE>(Croft and Meyer, 1973)- Knowledge of the biology of A. fallacis is incomplete regarding the mechanisms responsible for mite dispersal within the apple orchard ecosystem. Adult female A. fallacis principally overwinter in the ground cover beneath the apple 2 tree until May when they begin actively feeding upon ‘1. urticae. Migration of A. fallacis into the tree occurs in mid to late June where they feed on.£..glm; and A. schlechtendaAi. During August and September spider mite densities decrease within the tree, often as a result of the effective biological control by A. fallacis. Subsequently predator populations disperse out of trees and back to the ground cover to feed or locate overwintering sites (Croft and McGroarty, 1973; McGroarty, 1976). The dispersal mechanisms of A. fallacis were deemed an important area for research since it had been shown that there was a direct correlation between the number and timing of predators which migrate into the tree and the degree of successful biological control that was achieved (Croft and McGroarty, 1973; McGroarty. 1976). Furthermore, a dispersal component was an essential element needed to develop a pop- ulation model of the interaction of A, fallacis with plant- feeding mite pests (Croft, 1975b; Croft et a1., 1976). Ultimately, it was anticipated that this knowledge might lead to the discovery of orchard management practices which when implemented would increase the early season effectiveness of A, fallacis as a natural enemy in the tree and decrease the number of acaricide applications which must be applied each season for spider mite control. It has been reported that mites associated with agri- cultural crops disperse by several means. For example. many researchers have observed that P. ulmi disperses by spinning 3 down on a silken thread during wind conditions less than five mph (Newell, 1941; Marle, 1951). Hussey and Parr (1963) reported that migration of T. urticae between host plants occurred by ”roping", a condition where mites congregated in a ball at the apices of the upper leaflets, and descended on silk threads which usually became entwined, forming a rope- like mass. Mites on the rope swung pendulum-like in the air currents and dropped off periodically. Nelson and Jorgensen (1968) observed that the tetranychid brown mite, Bryobia rubrioculus (Scheuten), dispersed within the tree by either walking from leaf to leaf or dropping off the underside of leaves and drifting with air currents. Nault and Styer (1969) indicated that the adult eriophyid wheat curl mite, Aceria tulipae (Keifer), actively initiated its dispersal by migrating to exposed leaf surfaces where they attached their anal sucker and assumed an anteriorly raised position which re- sulted in their dispersal during air currents >‘5 mph. Lastly, Gibson and Painter (1957) reported the phoretic dispersal of (A. tulipae by aphids. Female adults of A. fallacis were first observed on lids of a mite-rearing unit where they assumed an anteriorly raised stance when an air current was present in the laboratory. It was hypothesized that this response was an integral part of the active dispersal of A. fallacis into and out of the apple tree. The objective of this study was to describe this behavior in detail and to determine the biotic and/Or envir~ onmental factors which influenced its display. METHODS AND MATERIALS Throughout the study, colonies of A. fallacis were reared in units exposed to a constant temperature of 25.50: 1°C. Initially mites were reared in a plastic refrigerator box mea- suring 30x27x10cm where they fed upoan. urticae which were supplied by daily introductions of bean leaves infested with prey. Later, a more convenient rearing unit was used which was a modification of that described by McMurtry and Scriven (1964). This unit consisted of a 20x20x5cm stainless steel pan, a polyurethane foam square which fit inside the pan and a waterproof construction paper arena bordered with strips of Cellucottodg>on top of the foam. Prey mites. T. urticae. were reared on and brushed from bean plants as described by Scriven and McMurtry (1971). Various levels Of starvation of A. fallacis were main- tained by the following procedure: First, mites in the de- sired stage were identified (see Appendix Table A-1) and then transferred via a four-haired brush to a starvation unit and maintained without food for the desired time. The starvation *Cellucotto ------ Kimberly-Clark Corp., distributed by Bauer and Black, a Division of Kendall Co., Chicago. 5 unit, which initially was maintained at 25.50: 1°C.. con- sisted of a bean leaf bordered with xylene-thinned Stickem Speciafgz Units containing ovipositing females were scanned daily for eggs which were subsequently removed. The response of A. fallacis to air currents was initially studied with a 7.50m fan simulating an air current, and a stereomicroscope to observe the response of the mites on a test arena. The test arena consisted of a microscOpe slide with a piece of construction paper (lx3cm) bordered with xylene-thinned Stickem SpeciafE)taped to its surface. Female adult mites from various levels of starvation were placed on the test arena and exposed to various air speeds by al- tering the fan distance relative to the test arena (see Appendix Table A-Z). Observations and photographic records were made of all responses of the mites to the air currents and certain ones were subsequently used in wind tunnel studies as the criteria to measure the effects of starvation, temperature and air speed. A subsequent study was conducted within a wind tunnel comprised of a 1.5 meter galvanized air duct, 18cm in dia- meter. Air entered the duct and became laminar (uni-direc- tional) as a result of a diffusor which consisted of 10cm drinking straws positioned longitudinally in the tunnel. The response of mites was observed through a stereomicroscope positioned over the central acetate window of the tunnel. ”Stickem SpeciaIE) ------ Mapco Products by Michel and Pelton Co. Manufacturing Chemists, Emeryville, California. 6 The observation chamber contained a cover slip arena hori- zontally affixed to a 9cm glass rod upon which the mites were placed and observed under various air speeds. An 18cm fan, at the opposite end from the diffusor, drew air through the tunnel past the mites on the arena at air speeds which were varied by a calibrated rheostat or with various thick- nesses of cheese cloth which covered the diffusor (see Appendix Table A-3). The minimum air speed that stimulated the response of adult females of A. fallacis to air currents was determined with the aid of both the fan and the wind tunnel. Initially. adult females were exposed to an air speed of 4.7 mph for a one-minute interval and then the air speed was incrementally reduced from 4.7 mph to h.O mph to 2.9 mph to 1.3 mph and to 1.1 mph at one-minute intervals until the mites resumed a random search pattern. All life stages of A. fallacis were exposed to the con— ditions that initiated the response of adult females to air currents in order to determine the degree that each stage was similarly affected. The life stages tested were immatures (larvae, protonymphs, deutonymphs), adult males, preoviposi- ting adult females and ovipositing adult females. The effects of starvation and temperature on the response of each life stage to air currents were also tested. Ini- tially, mites were starved on a unit for a known time interval at 25.50 t 1°C., and subsequently placed in one of the three following temperature chambers for a four-hour period for 7 acclimation: 180 t 1°C. at 58%R.H.. 24° t 1°C. at #O%R.H.. and 280 t 1°C. at 70%R.H. After acclimation, mites were transferred to the wind tunnel and were exposed to a con- tinuous series of air speeds. Initially, the air speed was 2.3 mph; after a two-minute observation period the air speed was increased slowly for a 30 second interval to 4.7 mph, and so on to 6.3 mph, 10.8 mph. and 13.7 mph. The rheostat could not be regulated to produce specific air speeds such as 2, u, 6, 8, or 10 mph. The response of mites was determined and it was noted if the mites dispersed actively (as they exhibited the response to air currents) or if they were passively blown off without exhibiting the response. Main effects of the factors of starvation and temperature and their interaction were statistically tested for using a chi square method (Blum and Rosenblatt, 1972) and the specific effects of paired treatments were evaluated with a Bonferroni chi square method (Kramer, 1972). Although these methods have less power for detecting main effects compared to an analysis of variance, they were more effective in detecting which treatments specifically affected the behavior. The cumulative number of mites of each life stage of A. fallacis that responded to the air currents and the total tested at each treatment were derived from the data recorded in Appendices A-h. - A-7. This data was used in constructing a 3x5 contingency table where the data was pooled across the rows and columns of temperature (3 levels) and starvation (5 levels) and analyzed by a chi square (df=8) to detect 8 significant effects of the factor interaction in the response of each life stage of A. fallacis. In addition, the overall significance of main effects was tested by a chi square applied to the same 3x5 contingency table. but to each individual treatment cell (df=lh) for each life stage. To evaluate if significant differences existed between 60 paired com- parisons of the treatment combinations as they affected the mites response (R+ or R-), 2x2 contingency tables were con- structed and the Bonferroni chi square was used (Kramer. 1972). RESULTS AND DISCUSSION The following is a description of the response of a receptive adult female A. fallacis to air currents. The characteristic behavioral patterns when exposed to windless conditions were random searching movements, prey attack and consumption, mating, oviposition, and other behavioral patterns which readily were distinguishable from the response to air currents. The random search movements consisted of the mites using the posterior three pairs of legs for loco- motion whereas the forelegs were waved vertically at a level below the horizontal plane of the body and frequently touched the substrate (Fig. l.A). According to Baker and Wharton (1952), the forelegs of mesostigmatid mites functioned as sensors. In comparison, the behavioral response of A. fallacis to air currents usually began as a deviation from the random search movements to a more directional movement toward the edge of the test arena where the subsequent series of be- havioral events occurred. Within a ten minute exposure to air currents the directional movements ceased in receptive mites and they often groomed their legs either by rubbing their walking legs together or drawing their forelegs through their pedipalps. The next event was orientation to the direction of the air flow, which usually occurred in 10 to 9 lO afzg Figure l: af:-_; The sequence of behavioral events of the response of A. fallacis to air currents: (A) the random search movement: (B) dorsal view of mite orien- tation to the prevailing air flow; (C) lateral view of the stance heading away from the air flow (af); (D) lateral view of mite immediately before dispersing via the air. ll 20 seconds, where they turned about at one point (Fig. 1.8) and ultimately assumed an anteriorly raised stance as shown in Figure l.C. Concurrent with the directional orientation, the mites stopped waving their forelegs. As they assumed the stance they gathered in their forelegs and gnathal appen- dages and held their second pair of legs laterad Of the body and off the ground as seen in Figures 1.0 and 2.A-D (arrows). Figure 2.A-D are photographs of various views of the stance, and illustrate the positioning of the appendages as marked by arrows. This stance was maintained until either the air speed was diminished below a minimum threshold or the mite dispersed from the test arena as seen in Figure 1.D which depicts the mite just prior to dispersal. The orientation phase resulted in the mites positioning themselves in a stance with their heads downwind relative to the prevailing direction of the air flow. This is depicted in Figure 3 where adult females of A. fallacis were exposed to a laminar (unidirectional) air current at air speeds from 1.0 to 13.7 mph inside a wind tunnel. The mites maintained a stance nearly parallel (0°) to, but heading away from the air flow (Fig. 3). Once the behavior was exhibited, it was relatively fixed. Directional changes of the air flow, even to the extent of 180°, rarely resulted in mites reorienting, but frequently resulted in their dispersal. A hypothesis suggested by these studies is that as mites respond to the air currents, shifts in the air current direction at a microhabitat level may increase the probability of the mites Ill"? ll‘ I'll-“I'll l l l " '- II I'll .l"ll|.lll|llll|| II: 12 Figure 2: Photographs of the stance assumed by A. fallacis ovipositing adult females in response to air currents 13 14 amass» as“: on» Ca vcom930 has hmsflsmH map op Omsommmu aw MOHMSOH paupm wcfivfimoafl>o on» we nowmeCOfluo mo mamas ooEMPm «m wasmwm Emmy—Gun z: zo_._.6.3 mph. The turbulent air in nature consists of varying air speeds and directions which probably increase the probability of A. fallacis dispersal as it responds to the air currents. The second phase of the study was concerned with the life stages that exhibited the response to air currents and the factors which affected the response. It was shown that the ovipositing and preovipositing adult females exhibited the response equally, the adult males to a lesser extent, deuto- nymphs rarely, and the protonymph and larval stages were never observed to exhibit it. It was shown that there existed some significant effects of the interaction of starvation and temperature on the response of A. fallacis to air currents. In addition, the effects of the factors 24 alone were significant principally for the ovipositing females which were more sensitive to limited food supplies than the adult males or preovipositing females. There were no factor level effects of temperature on any stages of A. fallacis. In addition to air speed, air flow direction, temperature, life stage, and starvation of the mites, substrate vibration was suspected of being an untested factor that may also have an effect on the response of A. fallacis to air currents in nature. This study has application in the development of an in- tegrated pest management approach to injurious spider mite control. First, the biological data obtained in the study can be used as a guide in developing field studies to deter- mine if these results describe the actual field behavior of A. fallacis. Such a field study should involve the deter- mination of the effects of various prey densities on the dispersal rate of A. fallacis in the apple orchard ecosystem. This field study would involve monitoring the prey densities of Ey.ElTio.é- schlechtendali, and T. urticae, in the apple tree. In addition, determine the life stages of A. fallacis in the air during dispersal, and their population density in the tree and in the ground cover. The results of the field study would be compared to the laboratory study pre- sented here, i.e., which are the principal life stages that disperse in the field. Further studies could be conducted concerning which mechanisms of mite dispersal are involved in the early season movement of A. fallacis into the apple 25 trees and their population increase in the apple tree. All these studies may result in the development of cultural practices that would increase the probability of early sea- son biological control of spider mites in the apple tree and decrease the need for preventative acaricide applications. LIST OF REFERENCES Baker, E. W., and G. W. Wharton, 1952. An Introduction to Acarology. Macmillan Co., New York, pp. 25-26 Blum, J. R., and J. I. Rosenblatt. 1972. Probability and Statistics. W. B. Saunders Co., Philadelphia, pp. 426- 431 Croft, B. A. 1975. Integrated Control of Apple Mites. M.S.U. Extension Bull. E-825, Michigan Co-op. Exten. Serv., 12 pp. Croft, B. A. 1975b. Tree Fruit Pest Management. "Intro- duction to Pest Management." Ed. R. L. Metcalf and W. H. Luckmann, Wiley Intersci., New York, 587 pp. Croft, B. A., and A. W. A. Brown. 1975. Responses of Arthropod Natural Enemies to Insecticides. Ann. Rev. Entomol. 20: 285-335 Croft, B. A., and D. L. McGroarty. 1973. A model study of acaricide resistance, spider mite outbreaks, and bio- logical control patterns in Michigan apple orchards. Environ. Entomol. 2(4): 633-38 Croft, B. A., and J. A. McMurtry. 1972. Comparative studies on four strains of Typhlodromus occidentalis Nesbitt (Acarina: Phytoseiidae). IV. Life History Studies. Acarologia 8(3): 460-470 Croft, B. A., and R. H. Meyer. 1973. Carbamate and organo- phosphorus resistance patterns in populations of Amblyseius fallacis. Environ. Entomol. 2(4): 691-695 Croft, B. A., Tummala, R. L., Riedl, H. W., and S. M. Welch. 1976. Modeling and Management of two prototype apple pest subsystems. Proc. 2nd USSR/USA Symp. on Integrated Pest Management. M.S.U., East Lansing, Michigan. Oct. 15-17, 1974. (In press) Gibson, W. W., and R. H. Painter. 1957. Transportation by aphids of the wheat curl mite, Aceria tulipae (K.), a vector of the wheat streak mosaic virus. J. Kans. Entomol. Soc. 30 (4): 147-53 26 27 Hussey, N. W., and W. J. Parr. 1963. Dispersal of the glass- house red spider mite Tetranychus urticae Koch (Acarina: Tetranychidae). Entomol. Exp. and Appl. 6: 207-14 Kramer, C. Y. 1972. A first course in methods of multivariate analysis. Publisher-Kramer, C. Y., Virginia Polytechnic Institute and State University, Blacksburg, Virginia, Bonfegroni chi square, pp. 246-50., Bonferroni table, p. 32 LeRoux, E. J. 1961. Effects of "modified" and "commercial" spray programs on the fauna of apple orchards in Quebec. Ann. Entomol. Soc. Quebec 6: 87-121 Marle, G. 1951. Observations on the dispersal of the fruit tree red mite Metatetranychus ulmi (Koch). East Malling (Kant) Res. Sta. Ann. Rept. 1950: 155-9 McMurtry. J. A., and G. T. Scriven. 1964. The biology of the predaceous mite T hlodromus rickeri. Ann. Entomol. Soc. Amer.. 57: 362- McGroarty, D. L. 1976. Population Studies of Amblyseius fallacis in Michigan apple orchard groundcover. Ph.D. Dissertation, Michigan State University, Dept. of Entomol., 58 pp. (Unpublished) Motoyama, N., Rock, G. C., and W. C. Dauterman. 1970. Or- ganophosphorus resistance in an apple orchard population of Typhlodromus (A.) fallacis. J. Econ. Entomol. 63(5): 1439-42 Nault, L. R., and W. E. Styer. 1969. The dispersal of Aceria tulipae and three other grass-infesting eriophyid mites in Ohio. Ann. of Entomol. Soc. of Amer. 62 (6): 1446-55 Nelson, E. E., and C. D. Jorgensen. 1968. Dispersal of Bryobia spp. and Typhlodromus mcgregori Chant within apple trees in Central Utah. Utah Acad. Proc. 45(1): 168- 81 Newell, I. M. 1941. Bionomical studies of the pacific mite, Tetran chus pacificus McGregor (Acarina: Tetranychidae). In Central Washington. Unpublished Thesis for degree of Masters of Science in Entomol., on file at Washington State University, Pullman, Washington Parent, B. 1958. Efficacite des predateurs sur la tetranyque rouge de pommier, Panon chus ulmi (Koch) dans les vergers du Quebec. Ann. Entomol. Soc. Quebec 4: 62-9 Poe, S. L., and W. R. Enns. 1969. Predaceous mites (Acarina: Phytoseiidae) associated with Missouri orchards. Trans. Missouri Acad. Sci. 3: 69-82 28 Scriven, G. T., and J. A. McMurtry. 1971. Quantitative production and processing of Tetranychid mites for large-scale testing or predator production. J. Econ. Entomol. 64(5): 1255-57 Smith, J. C. 1965. A lab and greenhouse evaluation of T. fallacis (Garman) as a predator of Tetranychus spp. Ph.D. thesis, Louisiana State University Swift, R. C. 1968. Population densities of the European red mite and the predaceous mite Typhlodromus (A.) fallacis on apple foliage following treatment with various insecticides. J. Econ. Entomol. 61: 1481-91 APPENDIX 30 Table A-1: Amblyseius fallacis Life Stage Characteristics Stage Length Width Number Abdominal of legs Red or White (M) (H) Dot Egg 186 138 O --- Larvae 204 133 6 --- Protonymph 207 130 8 --- Deutonymph 270 156 8 --- Adult Male 281 166 8 --- Preovipositing Female 306 179 8 Yes Ovipositing but Egg Free Female 306 179 8 -_- Egg-Ladened Female 383 245 8 --- 31 Table A-2: Air Speeds Produced by the Fan Arrangement by Varying Fan Distance from the Stereomicroscope Fan Distance Air Speed in Inches (MPH) 3 6.2 6 5.7 9 3.9 12 2.6 15 2.0 18 1.6 21 1.2 24 0.8 Table A-3: Air Speeds Produced by the Wind Tunnel by Varying Thicknesses of Cheese Cloth Wind Tunnel Air Speed (MPH) Rheostat Number 4 layers 2 layers No layers 1 0.6 1.8 1.8 2 --- --- 2-7 2 1.; 2.7 3.3 2 1:5 3.; 12°: 7 p... 13 9 5.2 8 ° 5.5 9 2.1 4.5 6.1 10 2.3 5.2 6.6 11 2.9 6.8 8.9 12 4.5 8.8 11.1 13 4.7 10.8 13.7 20% Power Increase in Pheostat 14 6.2 13.6 17.6 A:mv uncommon on» NCHpHanxo p\3 HmuuonnHv o>Humam I: n . A+mv uncommon on» wchHpana oHan Hanuoanv o>Hvo< n: N unsouuso uHa o» oncoamou one wcHzonu .oz :: H ‘32 a o H a a H HH 3H 6H H : NH HN o 6 HH HN o o N HN mN a o n oH 6H o m “H “H o N NH NH 0 m mH oN o o o oN :N a o H 6 NH o N N 6H o o o «H H o a oN H o H HN NH n o a m a o N oH HH 0 H 6 NH o n NH nH o o H mH NN m o N nH oH H m «H. NN o o NH NN n 0 3H nN o o N mN 3N n o H HH HH o N nH nH o N “H nH N H oH NH o o 3 NH NH N o n m o o a nH nH. o H SH SH o N mH 6H 0 o m 6H mN N o A SH SH 6 H «H mH o o o nH o o N mH o o o mH SN N o N HH NN o m nH NH 6 0 NH NN o H o NN o o n mN mH H o o N SH H H m mH o o w mH o o N mH o o o mH mN H o H NH NH o N 3H oN H n mH :N o H NH mN H o N 0N SN H o N NH 3N o H oH mN o N oH NN o o oH NN H 0 NH NN NH 0 o H m NN o m NH NN o o o NN o o o NN o a n NN NN o o o a on o H o Hm o o m Hm o o a Hm o o o Hm :N o o H m 0H o N «H 6N o N NH mN o H OH oN o H w on NH swam. Mes. .. Ham .. Has .. Has a EA“ 1 Ea :mma :mHn, :me, :mHn -mHn pawn N.nH m.oH n.c 5.: n.N upooam hH< m p< mcova>hmmno ovscuz N o>Hp=oomcoo .COva>hmvm no ammo cam .musvmpoasos .Anasv momoam LH< mo chvacHnEoo 9:05pmoua unchowhwo o» mOHmsmm mcfivwmonw>o on» no momcoamom HNHOH>Ncom 0:9 ho wcoHpm>hmmno on» yo mama .du< OHnms 33 OH S o H S S o N HH H 0 0H NH n N a NH N H N 0N NN S o H N N N n N NH 6 N m SH 6 H N mH o o S mH SN S o H N . SH 0 m NH 0H 0 o OH oH o H 0 ON 0 o N 0N NH n o H NH nN o N NH NN o 0 NH NN o H NH NN N H HH Nn NN n o H N N o S m HH H N CH SH 0 N 0H NH H N S oH SN m o o N MH 0 N N NH o o N NH O o N NH 0 o N NH NH N o S nH NH 0 M NH NN o N NH SN N n oH 0N S o NH mm NN N o S HH NH o m mH NH o 0 SH NH 0 H NH NH o o N NH SN N o o N NH H N N NH 0 H N NH 0 o N NH o o 0 NH NH H o 0 NH mN H N NH NN o N NH on o o NH on H o S Hm NN H o n N a o N SH NH 0 0 SH NH o H NH NH N o m 0N SN H o N HH HH H N nH SH 0 H 0H NH 0 H a NH 0 o m NH NH 6 o H 0 NH N H N NH 6 o N “H o o H “H o o o “H SN 0 H n SH NN o m 0N NN o o NH NN o H SH NN o o S NN SN 0 o m NH ON 0 S HN SN 0 o HN SN H 0 NH mN N o N NN NH Smfiflm -9...“ 2 .2 Shaw“ 2 .2 Haw.“ 2 .2 He...“ 2 .2 New. 2 .2 new” :mHQ umHQ :mHa ImHo ImHQ vmww N.nH mcomnm un.pr< mCoHpm>ummmmwmpscH£ N O>Hpnmwmcoo n.N .mp:msvmou9 pcmummmHo op moneOm mchHmonH>omHm any we mmmCogmmm Hmu0H>mcmN as» ho chHpm>pmmpo on» No Name .m:< oHnma 34 S o o S N N H N N H N N NH 6 N N SH 6 o H SH NN S o o N N H H N 6H 6 N H NH 6 o N NH 6 o H NH SN S o o N HH 6 o S HH 6 .o N HH 6 o H HH 6 o 6 HH NH N o H N N H o S N H o H N o o o N o o o N NN N H N S N N N N NH 6 o N NH 6 o 6 NH 6 o 6 NH SN N o N N SH H N N .NH 6 N HH SN 6 6 6H SN 6 o . N oN NH N o o N NH H S N NH H N N SN 6 N S NN o o H NN NN N H o N NH H S N NH H o N NH 6 o H NH H o 6 NH SN N o o N N o o S N o o o N o o H N H o H 6H NH H H H N N N o N oH H o H HH H H H NH 6 o 6 NH NN H o H N N N N N NH H o S SH H o N NH 6 o 6 NH SN H o o S NH H H N oN o o S SN 6 o N oN o o o oN NH 6 o o N NH N o N NH N o H NH H o H NH 6 o 0 NH NN o o o N N H N N HH 6 o N HH 6 o N HH 6 o 6 HH SN 6 o N oH NN N N NH HN o 6 HH HN o o N HN o o N HN NH SWNNN. New“ 2 .2 Haw“ 2 .2 NEH“ .2 .2 Nam“ 2 .2 New“ 2 .2. 5a -NHN. :NHN -NHN :NHN -NNN pmmm N.mH mcoonm NHW.MHP< n:0HpN>ummmmNevscHz N N>prmwmcoo n.N .mpcmspmoua unouwmuHa ow mmHNE PHso< N59 no mmmcoamom HNhoH>NnmN 0:9 no mCOva>Homno on» Me Name .N|< oHnNa 35 uncommon NEH) d NNHpHmano NOHNsoH ho NNHNE nnshcovsoo... . S O H ON NH O H tn nH O O ON nH O O UN MH H O O SH NN S O O O NH. N O O SH H O O NH O O O NH O O O NH SN S O O O HH O O O HH O O O HH O O O HH O O O HH NH n O O O SH O O O SH O O O SH H O O NH O O O NH NN n O O O NH O O O NH O O O NH O O O NH O O O NH SN n O O O nH O. O O nH O O O nH O O O MH O O O nH NH N H O O OH N O O NH O O O NH O O O NH O O O NH NN N O O O N O O O N O O O N O O O N N O O 9 SN N O O O HH O O O HH O O O HH O O O HH O O 0 HH NH H N O O OH O O O OH O O O OH H O O HH S O O NH NN H O O O m H O O OH O O O OH O O O OH O O O OH SN H O O O OH O O O OH O O O OH O O O OH O O O OH NH O O O O NH O O O NH O O O NH O O O NH O O O NH NN O O O O NH N O O SH O O O SH H O O NH O O O NH SN O O N tn NH N O tn SH N O O NH O O O NH O O O NH NH 5%”.an New“ 2 .2 Han” 2 .2 Ham... 2 .2 Haw“ 2 .2 Han”. 2 .2. $me :NHO numb, :nwn. Ian. INHni, Wmmm N.nH acooam un.mHv< m:0Hva>ummmmwstcHx N o>prmwmcoo n.N .uvsospwoua acououth on NNNNPN mauvmesH an» no noncoamom HNNOH>NSNN 0:» no msoHvN>Nompo on» me Nada .NI< NHnaa “"IITI'I‘IJL‘HHJLEMIMLiflirflfyifiiiu’y‘wiflfflim“