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III “In-(unthIIIIIII I.\k.1.I" \ ”I- I NIL-III V)"IIV‘U\\ ‘5 111'" N '1‘ LI 'n \11":‘1'II I1 # I ' I‘: II‘ I1'Iu 714E813 This is to certify that the thesis entitled THE EFFECTS OF MOLD-BOARD PLOWING AND ATRAZINE ON ACARINA AND COLLEMBOLA presented by DAVID MALLOW has been accepted towards fulfillment of the requirements for M. 5. degree in ZOOLOGY Date February 12, 1981 0-7 639 Lass! x.11( Y Michigan State University 4 . l OVERDUE FINES: 25¢ per day per item REIURNING LIBRARY MTERIALS: Place in book return to remove rge from circulation records EFFECTS OF MOLDBOARD PLONING AND ATRAZINE 0N ACARINA AND COLLEMBOLA By David Mallow A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Zoology 1981 ABSTRACT THE EFFECTS OF MOLDBOARD PLOWING AND ATRAZINE 0N ACARINA AND COLLEMBOLA By David Mallow The effects of tillage and Atrazine upon soil Acarina and Collembola populations were evaluated. Soil core samples were taken regularly during one growing season, and the fauna were extracted with Tullgren funnels. Statistical analyses were performed to determine if any quantitative changes occurred in treated plots compared to populations from grassland controls. All treatments yielded similar results: populations of Acarina and Collembola were significantly reduced. Collembolan populations recovered during the growing season, while Acarina did not. ACKNOWLEDGMENTS I would like to thank the members of my guidance committee, Dr. T. w. Porter, Department of ZoologY; Dr. R. J. Snider, Department of Zoology; and Dr. L. S. Robertson, Department of Crops and Soil Science, for their aid in this research project and thesis preparation. I also convey appreciation to Dr. Ivan Mao, Department of Dairy Science, John C. Moore, fellow graduate student, and Doug Ross, Department of Entomology, University of Georgia, for their aid in the statistical analysis of my data. Thanks are extended to Dr. w. C. Melbourn, Ohio State University, Acarology Laboratory, and David Walters, Department of Biological Sciences, California State University, for their help in identifying and confirming certain mites. Thanks again go to John C. Moore for his aid in identifying Collembola. Particular regards go to my wife, Jean, who aided me in the preparation of this thesis, and encouraged me through this work, and to my parents, who stood by me, and supported me through this effort. To Dr. Sigurd 0. Nelson, Department of Zoology, State University of New York at Oswego, I express grateful appreciation for his efforts that provided me with the opportunity which ultimately lead to this thesis. Special thanks go to the University of Georgia for the use of their facilities in the completion of this thesis. ii TABLE OF CONTENTS Page Abstract . ..... . . ................... Cover Acknowledgments . . . .................... . . ii List of Tables ............... . ......... iv List of Figures ........................ v INTRODUCTION . ............ . ............ 1 LITERATURE REVIEW . . ...................... 3 Cultivation Effects ..................... 3 Herb1c1de Effects ...................... 6 METHODS AND MATERIALS ...................... 8 RESULTS . . . ..... . ..................... 20 DISCUSSION AND CONCLUSIONS .................... 44 RECOMMENDATIONS FOR FUTURE STUDY ................. 47 APPENDICES A. Inventory of Arthropods from Grassland Controls ..... 48 B. Inventory of Arthropods from Moldboard Plots . . . . . . . 53 Co Inventory of Arthropods from Moldboard and Cultivated P] Ots O O I O O 00000 O I O O O O 0 O I O O O O O 59 D. Inventory of Arthropods from Moldboard, Cultivated and Atrazine Plots . ...... . . . . . ....... 62 LIST OF REFERENCES ........................ 66 LIST OF TABLES Table Page 1. Percent Extraction Using Tullgren Funnels, Determined by Flotation ........... . . . . . . . . . . l5 2. Soil and Air Temperatures on Sampling Dates ...... 16 iv LIST OF FIGURES Figure Page l. Map of Sample Plots .................. . ll 2. Daily Precipitation . .................. l2 3. Diagram of Core Device . . ............... l3 4. Daily Maximum-Minimum Temperatures ........... 19 5. Mean Population Levels of Collembola . . . ....... 22 6. Mean Population Levels of Brachystomella parvula . . . . 24 7. Mean Population Levels of Tullbergja granulata ..... 27 8. Mean Population Levels of Acarina . . . ......... 29 9. Mean Population Levels of Prostigmata .......... 32 10. Mean Population Levels of Bakerdania sp. ........ 34 ll. Mean Population Levels of Mesostigmata ......... 36 12. Mean Population Levels of Rhodacarellus sp. . . . . . . 38 13. Mean Population Levels of Cryptostigmata ........ 40 14. Mean Population Levels of Opiella sp. . . . . . . . . . 43 INTRODUCTION Studies which involved the role of soil microarthropods in organic decomposition, nutrient cycling, and soil humification, have shed light on their importance in soil ecology (Thompson, 1924; Engelman, l96l; Kevan, l962; Frantz, T962; Nitkamp and Crossley, 1966; Butcher et al., l97l; and Lebrum, T979). Cognizance of this importance has necessitated more precise investigations of current agricultural practices and their effects on soil microarthropod populations. The objectives of this study were to determine the effects of conventional tillage and Atrazine on populations of soil Acarina and Collembola. Quantitative comparisons were made between field populations in grass control plots and treated plots. Summarized below are the treatments evaluated in this study: (1) Conventional tillage, using the moldboard plow, is the principal tillage method in Michigan. This plow breaks soil, turns it over, and completely reorganizes the surface materials. It is usually the deepest and most disruptive tillage practice used in agriculture. When used at medium soil moisture levels, it surpasses all other implements in loosening soil, while incorporating crop residues, manure, fertilizers or lime. Energy requirements are relatively high because of the large volume of soil it moves (Robertson et al., 1977a). 2 (II) This treatment combined conventional tillage (moldboard plow), with cultivation methods. These methods were used after seeds were planted, or young plants had emerged. The results of cultivation include: break up of crusts, weed control, increased water infiltration, erosion control and facilitation of harvests. With crops of high economic value, cultivation remains a significant input in crop production costs (Robertson et al., l977b). These treated plots were sampled only the last three sampling dates. The decision to take samples from these plots was made after the first five sample dates. (III) This treatment combined conventional tillage (moldboard plow), cultivation and applications of Atrazine (AAtrexe). The general use of Atrazine is to control broad leaf and grassy weeds in corn, sorghum, sugarcane, and turf grass sod. It is absorbed through the roots and foliage, translocated acropetally in the xylem, and accumulates in the apical meristems, and leaves of plants. It acts as a photosynthetic inhibitor, affecting the photochemical activity of the chloroplasts (Hill reaction) (Audus, 1964). The term "cultivation" is used vaguely in the literature. Hence, "cultivation" as used in this thesis is defined as any mechanical manipulation of soil for agricultural purposes. This study was part of a larger effort investigating tillage and herbicide effects on soil fauna at Michigan State University. LITERATURE REVIEW Cultivation Effects The effects of cultivation on soil fauna are more easily recog- nized by comparing grassland and arable land (Tischler, 1955). Arable land is poorer in soil fauna than uncultivated land in numbers of individuals and species (Buckle, 1921; Kevan, 1962; Burnett, 1968; Edwards and Lofty, 1969; and Edwards and Thompson, 1973). However, certain soil animals, such as Symphyla, Protura, Diplopoda, Rhodacaridae (Mesostigmata), sminthurid Collembola, and prostigmatid mites are more abundant in agricultural soils than elsewhere (Edwards and Lofty, 1969). An important reason for this difference is that grassland supports vegetation during a period when arable land is fallow. The presence of vegetation ensures that a food source is always available for a large majority of soil species (Buckle, 1921). Vertical distribution of soil animals in grassland and arable land also differs. In grassland, most animals are within the upper 5.08 cm of the surface, while in arable soils, animals are more evenly distributed through the upper 15 cm (Edwards and Lofty, 1969). Tischler (1955), Sheals (1956), and Edwards and Lofty (1969, 1975) reported reduction of soil animal populations immediately following cultivation. Some taxa are affected more than others. Edwards and Lofty (1969) report Prostigmata, Mesostigmata and Symphyla are least affected, while Cryptostigmata, hemiedaphic Collembola and Pauropoda are reduced after plowing. Burnett (1968) claims that cultivation increased Collembola and mite populations. Aleinikova and Utrobina (1975) report that the method of cultivation has no appreciable effect on the total numbers of group diversity, but considerably effects the ratio of individual groups. Cultivation effects have both advantages and disadvantages. The disadvantages are destruction of soil structure by deep plowing and extensive soil movement. An overall dilution of soil nutrients may also occur when deeper, less nutrient-rich soil is brought to the surface, mixing with, and diluting the nutrient-rich upper layers (Burnett, 1968). Water retention and heat conducting capacity also are affected (Tischler, 1955). The moldboard plow also acts as a forceful disturbance on soil structure, particularly if it is too wet during plowing, thus causing compaction. Another disadvantage of conventional tillage is that when soil is overturned, it becomes susceptible to drying, and vulnerable to wind and water erosion. Also soil organisms brought to the surface become exposed to harsh climatic conditions. In addition, repeated cultivation removes organic matter from the surface and decreases fauna populations by mechanical damage (Edwards and Lofty, 1969). Conventional tillage incorporates and distributes organic matter, making it more readily available to organisms feeding below the soil surface. It also opens up soil, improving aeration and drainage, and increases the volume of a "favorable zone" for soil animals (Burnett, 1968; Edwards and Lofty, 1969). Tillage also reduces crop residues, weeds, and root masses, which become subject to more rapid 5 decomposition (Tischler, 1955). According to Tischler (1955) and Kevan (1962), the moldboard plow is less injurious to soil animals than rotary or surface scraping cultivators. An important indirect effect of cultivation is the change it causes in plant cover. Altering the vegetative cover can greatly change the structure of soil animal populations within one horizon and soil type (Block, 1966; Edwards and Thompson, 1973). Strickland (1947) showed that considerable differences exist between qualitative populations on two plots of similar soil type, largely due to different plant associations on the plots. Changing the vegetative cover will change the type of litter produced, which causes a corresponding change in soil organic matter. This in turn effects the faunal composition. Changes in fauna may affect the quality of humus produced, thus effecting soil fertility (Kevan, 1962; Edwards and Thompson, 1973). Edwards and Lofty (1969) reported that after six months, numbers of soil animals in treated and untreated plots differed only slightly, indicating cultivation effects may not persist into a second growing season. Sheals (1956) showed that reseeding cultivated land resulted in rapid decolonization by Collembola, but populations of Cryptostigmata and Mesostigmata remained low over a period of 17 months. Cultivated land left fallow may result in persistent reduced numbers of soil animals (Sheals, 1957; Edwards and Thompson, 1973). Effects of cultivations on soil animals may persist up to three years (Butcher et al., 1971). Microarthropods with short life cycles become fewer when soil is first cultivated, but they usually recover within a single growing season. By contrast, arthropods that live longer than a year may be affected by cultivation longer than one growing season (Edwards and Lofty, 1969; Edwards and Thompson, 1973). Herbicide Effects Increased use of herbicides to control weeds in agricultural systems has necessitated investigations of their side effects on soil animal populations. Their use can exert considerable effects by diminishing or increasing numbers of both beneficial and harmful soil animals (Fox, 1964). Rapoport and Canglioli (1963), Edwards and Arnold (1964), and Davis (1965) found MCPA (4-chloro-2-methylphenoxy- acetic acid) had no effects on numbers of mites in treated and untreated plots. Edwards (1965) found 2,4-D (2,4-dichlorophenoxacetic acid) reduced populations of soil animals one-half to one third compared to untreated soil, predatory mites and isotomid Collembola were most effected. Rapoport and Canglioli (1963) found no differences in mite populations in 2,4-D treated plots. Prasse (1975) reported populations of mites and Collembola increased following 2,4-D application. These discrepancies in the results of 2,4-D effects may be explained by several factors. For instance, effects of herbicides on mites and Collembola are proportional to the concentration used (Popovici et al., 1977). Also many studies done on pesticide effects on microarthropods are done in the summer when their populations are low, minimizing effects (Edwards and Thompson, 1973). Experiments with Atrazine show pronounced sensitivity of the soil fauna to the action of this herbicide (Popovici et al., 1977). Subagja and Snider (1981, in press) found that Brewers yeast treated with Atrazine, and fed to FolsOmia candida and Tullbergia granulata caused significant mortalities at 5000 ppm. It also caused longer instar duration and reduced egg production. Fox (1964) and Edwards and Thompson (l973)showed.Atrazine decreased numbers of Collembola and mites. They also reported Collembola were less susceptible to herbicides than mites. Changes in vegetative cover resulting from herbicide application causes indirect effects on the structure of soil animal populations (Fox, 1964). Unlike the blanket destruction that cultivation has on vegetative cover, herbicide effects are often more selective of the plant species it eradicates. This selectivity may diminish the numbers of species of soil animals, but not necessarily the total number of animals (Edwards and Thompson, 1973). Herbicides that effect saprophagus soil animals, which are essential in the breakdown of litter into its organic and inorganic constituents, may ultimately alter soil fertility and structure. The higher susceptibility of predatory mites to herbicides (Edwards, 1965) may positively affect nutrient cycling by freeing saprophagus animals from predation, thus permitting an increase in breakdown of litter and liberation of nutrients (Butcher et al., 1971). An important consideration in herbicide usage is its ability to degrade, and thereby its mode of degradation. The fate of herbicides applied to soil is largely governed by four factors: placement (dominant factor); sorption equilibria; inherent phyto-toxicity; and microbiological effects (Upchurch, 1966). Atrazine degradation is dependent on soil type, concentration, and moisture content (Skipper and Volk, 1972). Harris and Warren (1964) reported Atrazine adsorption depends on soil type. They found a direct correlation 8 between adsorption and content of organic matter, clay, and cation exchange capacity. McCormick and Hiltbold (1966) and Roeth, Lavy and Burnside (1969) found that Atrazine degradation increased two-to-three- fold for each 10°C temperature increase (from 150-350C). The exact process of degradation has three major pathways: hydrolysis at the number two carbon (most predominant); N-deakylation of side chains; and ring cleavage (Kaufman and Kearney, 1970). Accumulation and persistence of herbicide residues in the soil may endanger soil animal biotic processes (Popovici et al., 1977). Considerable changes in community composition of microarthropods persisting for more than a year could be attributed to weed control worsening the living conditions of most soil inhabiting species (Prasse, 1975). Persistent herbicides do not always effect soil animals the most. Simazine (2-chloro-4,6,-diethy1amino-s-triazine) may persist from six months to more than a year, but it is only slightly toxic to soil animals. D-D mixture (1,3-dichloropropene and 1,2-dichloropropane) persists only a few weeks, yet it is extremely toxic to all soil animals and may decrease numbers of them for two to three years (Edwards and Thompson, 1973). Burnside et al. (1965) feund that herbicide carry-over is a greater problem in arid regions than humid regions of the U. S. Popovici et a1. (1977) report that reduction of Atrazine effects on soil animals after fbur months is significant for restoring their equilibrium. METHODS AND MATERIALS The study area is located off Jolly and College Roads, on the north end of section "C" of the new Crops and Soils Science farm at Michigan State University. 9 Samples were taken from a series of plots, each 6.1m x 15.2m, on which corn was grown. The plots comprised seven treatments in four replicates. Each replicate had a grass control (Figure l). The soil is classified as Celina loam (soil management group 2.5a) and supported grass for the past eight years. This soil tends to be high in fertility and available water capacity. Runoff is slow to medium, and permeability moderately slow. There are no limitations that seriously affect its use for farming. Treatments were initiated May 1979 (plowed-May 11, planted-May 15). Atrazine (2-chloro-4— ethylamino-6-isopropy1amino-s-triazine) was applied June 12, at a concentration of .47 cc/m2 and cultivation was done June 22. During the investigation, water was supplemented by sprinkler irrigation five times; 12.7 mm-June 26; 25.4mm-June 28, July 10, July 18; and 38.1mm- July 27 (Figure 2). Samples were taken approximately every three weeks from the following plots (Figure 1): Control—Grass I. Plowed II. Cultivated III. Herbicide Samples, in the form of soil cores were collected using a 15cm deep core device of 6cm diameter (Figure 3). Five cores were taken from each treated and control plot in all fbur replicates. Cores were taken in-row between plants, in the central portion of each plot to avoid edge effects. Samples were placed in plastic bags and taken to the laboratory. Microarthropods were extracted by using Tullgren funnels. Figure 1. Map of Sample Plots 10 11 :26: .h gauged 651:: .5702 .o .232... £2.02 .m 0:33: £2.92 .v 2.3:: 60:02:30 6:81 30! .n. 8:2...50633632 .m. 3.001203 .p. 320133.30 ”3:253... a m. c. .u m o R v m. m m m H 88:9: a E 38.3.: End 2 O m m . no a ..ovm~mms$. n. m H 3.0.301 H 0.00.30: ml 100' M- 10 re 20 80 June aoo~ "n E 100- g * 10 15 20 u so a i Auguet 3 E ii 3 G 10 15 20 25 30 October Figure 2. Daily Precipitation. 2 8 July 3 to 18 a 28 80 Wt Hue-bu D. Irrigation I- Ian-tel 13 F 40.66"! i) 20cm L ~ ' I 22cm fi-5.7c'm:-"l 15cm +5084. cm Figure 3. Diagram of Core Device. 14 The heat source for the funnels were 25-watt bulbs connected to a rheostat. Low settings were used to prevent the soil from drying too rapidly, trapping the arthropods inside the soil. Soil was extracted four or five days, depending on the initial temperature and moisture levels of the soil (soil too moist or cold required lower heat and longer extraction time). Extracted animals were collected and preserved in a solution of 95% ethanol - 1% glycerine. Inherent biases in the extraction method were considered. Satchell and Nelson (1966) found that the efficiency of Tullgren funnel extraction may vary with the physical characteristics of species, independent of sampled soil type. Haarlov (1962) reported that due to poorly developed locomotory structures, small, slow moving species suffer the greatest losses when extracted with Tullgren funnels. Tamura (1976) found the extraction efficiency of Tullgren funnels to be very low compared with hand sorted samples. He found the deficiency greater in smaller species than larger ones. Satchell and Nelson (1962) found that oribatid mites were extracted more efficiently in Tullgren funnels than by flotation. Extraction efficiency of the Tullgren funnels was determined using dried, extracted soil from 10 controls taken June 18, 1980 (Table l). The soil was immersed and remoistened in a saturated sugar solution, causing less dense organic matter to rise to the surface, where it was collected and identified. Taxa not included in Table 1 were collected in numbers too few for valid conclusions. Air and soil temperatures were taken each sampling date using a Yellow Springs Institute Telethermometer (YSI-42$C)(Tab1e 2). Soil temperatures were measured at a depth of 15cm. No soil temperatures 15 Table 1. Percent Extraction Using Tullgren Funnels Determined by Flotation. Animal % Extracted Brachystomella parvulis 50.0 Tullbergia granulata 60.0 Lepidogyrtus pallidus 25.0 Scheloribates sp. 45.0 Opiella sp. 89.9 Bakerdania sp. 27.4 Rhodacarellus sp. 60.0 Total Acarina 64.2 Total Collembola 48.2 16 Table 2. Soil and Air Temperatures from Each Sampling Date. Date (1979) Air Temperature (0C) Soil Temperature (0C) (15.2cm depth) 9 June 19 July 3 August 26 August 23 September 14 October 1 November 9 December 20. 24. 25. 23. 19. 11. 13. -11. 000 CON 22.5 20.0 23.1 17.5 16.5 * 9.0 * *Temperature meter not functioning. 17 were taken on October 15 and December 9 due to temperature meter not functioning. Maximum and minimum daily temperatures and precipitation during the study were obtained from the National Weather Service, South Farm Station (Figures 2, 4). Following extraction, arthropods were identified to species (Collembola), Genus (Acarina), Order (Insecta, Crustacea) and Class (Myriapoda). Collembola were identified according to Snider (1967), and Christiansen and Bellinger (1980). Acarina were identified accord- ing to Krantz (1978), and from keys provided at the Ohio State University summer Acarology program 1980 (unpublished). Statistical analysis of data utilized a two-way, fixed classifica- tion model and normal equations. These methods generated F-ratios and confidence intervals (Gill, 1978). The classification model used was: y=U+Ti+Sj+P(i)k+TSij+e Where: u = Mean common to all elements T = Treatment effects; i = 1,2,3,4 S = Season effects; j = 1,2,3,4,5,6,7,8 P = Plot replicate effects (nested in treatments); k = 1,2,3, ...15,16 TS = Treatment-season interaction e = error All data was processed by computer using the Statistical Analysis System (SAS) program. 18 Figure 4. Daily Maximum-Minimum Temperatures. 19 e800 .000 so: 3 .30 a L. m. w m. “mu m 56.1.... . mus » 15.. - . . p 4., 1.. 9K 2 ._ . :9 (A w 3 fla. ) . . H I f .. :3?er 2 k ,. . 8 0.. . ,_ L /\.. \ as..." a _ . a) .. . / /L. A a. .,._, ,> .. .8 fl. . ‘ _. ;\. .. . __ ‘ a we . ._. . \4 ‘ S. .. r s. K .. .. . x, N I .3 «P3; .._ _ a . \H H 1. < ._ .\ ,, .2 k I e .z . r b b x 2‘ r0 . \. . \o f . y \ , 2) v y \\ / . ._ . vs x : __ S. I ..t..: .. ; M . a K I ( 7,. . _ . / 1 .1 . i . _... u \ _ x \«c < .. x2. I \ A. .3. Z . Z J .3 ON; , . _ _ x 2‘. _, _ . / \1 A. a _ _. _ L. 2.12 (kc/x ...L. a m ,2. 3 T. 7 . u I ‘ _ .2 . , N 00 K J) , ,_ f I k (g '\ . s .7 I ,, I. .. 3 K .\ 4 \ , k _ . t8 . ,_., v .\ ~ f ._ _ r /. . :8 r... SEES... .. 5:532: . e8... 25:: o (:19) ”humane; RESULTS Quantitative Effects Quantitative analysis of this study was based on two major microarthropod orders: Acarina and Collembola. Of 51 genera identi- fied, only two Collembola and one genus in each of the three major soil inhabiting sub-orders of Acarina (Prostigmata, Mesostigmata, and Cryptostigmata) were collected in numbers large enough to make any statistically valid conclusions. Treatment II data was omitted for the fellowing reasons: Only three sample dates were obtained, and this data analysis produced results nearly identical with treatment I. When compared with controls, overall significant decreases (P 5_.O5) occurred from Collembola populations in treated plots. Individual analysis showed these decreases continued through the first fbur sample dates. No significant differences among treatments were observed on any sample date. Population levels of Collembola in control plots increased steadily from June 9, reaching a maximum August 3. This was followed by a steady decrease, reaching a minimum September 23. Treated plots fluctuated little, within a few data points, over the entire growing season, increasing slightly from October to November (Figure 5). Brachystomella parvula, a hemiedaphic species, (Figure 6), showed overall significant decreases in treated plots compared with controls, and exhibited the same general patterns as total Collembola. 20 21 Figure 5. Mean population levels of Collembola. 22 mouoo oiEom o'ola'nnr a I. I. I. .AIIIITAII ’5', l I 'III lll“‘|l| ‘- I I... e‘ \e| o .I‘ a‘ OENFZ< 50:03:3065.009.031.111.I 2.2.22. ...... .9. :30 den .32 ..oo .83 63¢ 63¢ :2. OF — vw MN ON a @— 0cm... - . h h p - IN (aw/epueenou; u!) ewmguv to eieqwnu 23 Figure 6. Mean Population Levels of Brachystomella parvula. 24 $30 oEEmw .25 .52 ..oo .38 .92 m3 :2. 0:2. m p VP MN wN m m— m p h p b b h h p 10w 1 ......... 1.11. 1.1.19.1 .1. .. . .91 1 - 11?. 11 11111.11u 1 111 1.1111 111. / \ ./.(.x. / 1mm tom— o:_~u.:<.co:u>:_:o.u.uon2oi1 ........ 9.30030: 1 1 1 1 - .ozcoo (aw/spelpunq ug) Slewguv Io Siaqwn 25 Individual sample date significance (P 5_.05) ended after three sample dates. Maximum and minimum levels were reached on the same sample dates as total Collembola° Population levels of Tullbergia granulata, an euedaphic species (Figure 7), did not exhibit overall significant differences between treatments and control. Analysis of the first two sample dates exhibited no significant differences among treatments, or between treatments and control. On the remaining six dates, significant differences (P §_.05) existed among treatments, with treatment III populations higher than treatment 1. Significant differences (P 5_.05) between treatment III and control appeared on sample dates September 23, October 14, and November 1, with treatment III popula- tions higher than control. A significant difference (P 5_.05) was displayed on November 1 between treatment I and control, control populations lower. Population levels of I. granulata in control plots fluctuated little throughout the eight sample dates, with no definite maximum or minimum. Both treated plots had a maximum on November 1, and exhibited no apparent minimum. Overall, Acarina populations decreased significantly in treated plots (P 5_.05). Individual analysis (Figure 8) showed significant differences (P §_.05) between treatment I and controls, control -populations higher, on each sample date except August 26. Decreases in treatment III populations were significant (P §_.05) on all sample dates except August 26 and December 9. Acarina populations in control plots increased steadily from June 9, and reached a maximum August 3. This was followed by a steady 26 Figure 7. Mean Population Levels of Tullbergia granulata. 27 3qu 0.95m oo .32 ..o ..nom 63¢ .21 :2. 25.. m. . ...o a.“ ...N m e a - 1111:11111111} I x 11 \ 1 11 1 \ \ Tn . \ ./, TWP .\ / .1 .11 11 1 1| .0 ’- / c. \l‘. / x < rmN .\Q\ o .\ 1mm .x. xi. rm? 2:223 .co_.o>_::0.u.ooau.o!.I.1.1.1. 2.2.20: 1 11111 33:00 (,m/epospunu uq) 8|ewguv go eioqwnu 28 Figure 8. Mean Population Levels of Acarina. 29 moaoo macaw 052:;6022230630920! 1111111 .1 632.222 1 1 1 1 35:00 doc .52 .30 ..new .9. .m: > a o .p 3 nm o.~< m< .9“. cam. .. . 10. 10m 100.. (,w/epuesnoqi u” swunguv go SieqwnN 30 decrease, reaching a minimum August 26. Treatment levels fluctuated little over the entire growing season. The sub-order Prostigmata and Bakerdania sp. (Figures 9, 10) exhibited significant decreases (P 5_.05) in treated plots compared with controls. Individual significant differences, maximum and minimum of controls, and treatment population levels, displayed the same pattern as total Acarina, with the exception of Bakerdania sp. on August 3. No significant differences between the control and treat- ments occurred on this date. Mesostigmatid mites (Figure 11) decreased significantly in treated plots (P §_.05), however, individual significant differences occurred only on July 19 and August 3. Reaching a maximum July 19, control populations decreased steadily through the remainder of the growing season, while treatment levels fluctuated little. Individual significant differences (P 5 .05) occurred in popula- tions of Rhodacarellus sp. between treatment I and controls on October 14, and between treatment 111 and controls July 19 and August 3, control levels higher for both. Rhodacarellus sp. did not display overall differences between treatments and controls. Treatment I populations were significantly higher than treatment III on July 19, and exhibited population peaks July 19 and September 23. Treatment 111 reached a maximum October 14. Compared with controls, overall decreases of Cryptostigmata populations in treated plots were significant (P 5_.05). Individual analysis of samples (Figure 13) showed decreases in treatment I which occurred on July 19, September 23, and October 14. Decreases in populations from treatment III were significant (P 5_.05) on July 19 31 Figure 9. Mean Population Levels of Prostigmata. 32 330 038.6 .oeo so: duo ..oom .o: .92 a 2. 23... a . a a.“ mm ..H w. a 12"”...113' \lloriolllllllI-Ib f r 1 .V...‘ I 1.11111”.1\11\11 1 1. 1.61 1 1 OI.11.|/.1L!11./11./111.|11111111119|1\ 110...] T1 111111111 [1.69 On On .oo 6m ec.~2.<.co:e>.::u632.20! ......... 3.2.32... 11111 .choo (.w/epoapunu u.) slewguv io eioqwnu 33 Figure 10. Mean Population Levels of Bakerdania sp. moaoo 295$ 34 doc .32 a _. #1:! .1.1 '1 '1 1 I I '1.l.‘o 1.01 in e1.|l.|Il-| O'Jle'n u".l.u”uo1ellu . 63¢ 63¢ 22. 12. w my 1 .1 ” I11 I” U 01!- |||||| OO ....... 1 I! 11.41””. 1 1. i 1 oE~oz¢.co:o>:_:o.u.oonu.o! 111111111 Boon-:0! 11111 .2200 um" (.ul/epuoonom m) slewguv 10 eioq 35 Figure 11. Mean Population Levels of Mesostigmata. 36 .ooa Soz J00 mouoo 0.955 .36 .21 on oE~o2¢ .co:e>_::o.u.ooau_oz 111111111 2.2.22. ..... 33:00 25.. >01 10.. (,m/opuoenou; In) smuuuv io eioqwnu 37 Figure 12. Mean Population Levels of Rhodacarellus sp° 38 30 3200 oEEom ...om 03¢ nu cm 2.. .5. 2.332 .eo..o>...3o.o.oonu.o! 111111111 2.2.3.3. 11111 .3230 0:3.- uw-Pume a!) tummy so uoqumu 39 Figure 13. Mean population levels of Cryptostigmata. 40 no.2. oEEom con 3 . ...z ....o .3... a... .m. s... a... I b I [IT a 1N n / / / / ‘- / . .1 1 .I 1'! \1 I 1 \I It '- \\.I “““ \ \ \I \ \ I \ \ I I \ *1 \I 11 /I \\ I] \\\\\.\.Il|lllll111|l‘ ........ o a / \ \ \ \ ‘ \l\ I / x . ./ \\ \ 1 / .\ .I. ./ / .\ ./u \ \ /1\.\ to /. 1‘.\ /. 1.1 / / \ \.\ ( .\. O\ \ I l“ ./ \ l .401. 1: 1Q~ 352::60233230630320! 111111111 3.2.3.2. 11111 33.30 (,Iu/epuoenou; ug) elemguv go eioqwnu 41 and October 14. Significant differences among treatments occurred on September 23, with treatment III populations higher. Population levels of controls peaked twice over the growing season, with maxima occurring July 19 and October 15, and a minimum September 26. Treatment levels fluctuated over several data points, but never enough to display a distinct maximum or minimum. Opiella sp. (Figure 14) displayed no significant differences between treatment I and controls on September 23, or between treatment III and controls on October 14. Population peaks in control plots were similar to total Cryptostigmata. Maxima of treatment populations occurred August 3 and November 1, with a minimum September 26. Except for the above two dates, Opiella sp. displayed the same overall, and individual significant differences as total Cryptostigmata. 42 Figure 14. Mean Population Levels of Opiella sp. 43 3.30 3.93.6 3 Eon 33 3 8 9% 33 > 3 25.. n¢ r... / \\ . . . .1..1|.1I.|..I..1l. .|.,“ .1. .\ 1° 03.3.:.co:->.:3o.o.oono_o! 111111111 2.2.3.2.. 11111 3.230 (.w/epuoonouiug) BIBWIUV ‘0 BJquflN DISCUSSION AND CONCLUSIONS In this study, population levels of Acarina and Collembola from grassland controls increased in June, with maxima occurring in July for Acarina and in August for Collembola. However, trends in natural population levels of these soil animals show a great deal of variation with respect to seasonal fluctuations. Thompson (1924), Ford (1937, 1938), and Dowdy (1965) found Acarina and Collembola populations peaked during winter months, November through February. Dhillon (1962) reported peaks in June, with minimum levels in December. Edwards and Thompson (1975) found population maxima during Spring and Fall. Shaddy and Butcher (1977) reported no distinct population peaks. Tullbergia granulata also exhibited variations with respect to its seasonal fluctuations. In this study, I. granulata populations fluctuated little, and did not exhibit any distinct maximum or minimum levels. Usher (1970) found I} granulata peaked September through October, while Shaddy and Butcher (1977) reported peaks in July. Numerous ecological factors, such as variations in soil type, climatic conditions, moisture regimes, and the frequency and quantity of samples taken, may be involved in these discrepancies. Conventional tillage, utilizing the moldboard plow, significantly reduced populations of Acarina and Collembola immediately fbllowing its use. These results are in agreement with studies by Tischler (1955), Sheals (1956) and Edwards and Lofty (1969, 1975). However, Edwards 44 45 and Lofty (1969) reported Prostigmata were least affected by tillage and Cryptostigmata were affected most. In this study, contrary data showed an opposite effect. For example, Prostigmata were significantly reduced over the entire growing season following tillage. Initially Cryptostigmata showed no response to tillage, and exhibited signifi- cant reductions on only three of eight sample dates. After seven months, no recovery to levels comparable to controls were observed for total Acarina or Prostigmata. Cryptostigmata began to recover after five months, while Mesostigmata recovered after just three months. Levels of Rhodacarellus sp. (Mesostigmata) increased initially following tillage, and displayed the same pattern as control populations over the first five sample dates. This increase in Rhodacarellus sp., and quick recovery of Mesostigmata, in general may be attributed to the predatory behavior of Mesostigmata. Being active predators, they range freely through soil. Frequently they are not associated with the soil to the degree of other euedaphic mites (Usher, 1971)o Populations of Collembola, with the exception of Tullbergia granulata, began to recover after four months. I. granulata populations were not effected detrimentally by tillage. Their levels remained nearly equal to control levels, and even increased to levels signifi- cantly higher than controls after five months. The effects of Atrazine on populations of Acarina and Collembola in this study are difficult to assess. The majority of taxa analyzed showed no differences between cultivated plots with and without Atrazine. Tillage, being so great a destructive force on soil animals, has masked any apparent herbicidal effects. Between 46 cultivated plots with and without Atrazine, a few taxa exhibited significant differences. Cryptostigmata, Opiella sp. and Rhodacarellus sp. all had one sample date where these differences occurred. However, differences were probably the result of high natural variation that occurs in soil animal populations owing to their non- random, aggregated distribution (Hughes, 1962; Nef, 1962; and Berthet and Gerard, 1965)° Only Tullbergia granulata displayed consistently distinct differences between cultivated plots with and without Atrazine, thus exposing possible herbicide effects. After the first two dates sampled, populations of I. granulata increased significantly higher than control or treatment I levels. This is indicative of possible stimulation of I. granulata populations caused by Atrazine usage, and is in direct contrast to studies by Subaja and Snider (1981, in press). They reported longer instar duration and decreased egg production for Atrazine fed I. granulata. These results also contradict studies by Fox (1964), Edwards and Thompson (1973). They found Atrazine decreased Collembola populations. Reasons for these discrepancies are not readily apparent, but demonstrate the need for further investigations into possible herbicidal side-effects on soil fauna. Before concluding this study, an important consideration should be made. Although some taxa exhibited levels equivalent to controls in December, the final date sampled, the assumption should not be made that total recovery of Acarina and Collembola populations has occurred. In subsequent months, control populations may increase steadily, while treated plots continue to remain at low levels. Only continuous 47 sampling through subsequent months can reveal long-term effects of these treatments. RECOMMENDATIONS FOR FUTURE STUDY Based on the results of this study, the following recommendations are proposed fbr the future: 1) 2) 3) 4) 5) Evaluate effects of herbicide applications on grass plots without tillage. Examine changes in vertical distribution of soil fauna following tillage. Environmental parameters such as soil moisture, organic matter content and pH should be measured. Continue to monitor soil fauna throughout the entire year. Monitor soil fauna in treated plots in successive year. APPENDIX 48 N o . o — o o o o .am m:.moz mmu..moz m o o o . . . o o mzummo mmpwpwamge< e o o o o o N o N mummmze mwesgucwsmoemuzmo mm o o w o o. oN N. N mwpasaa avowemmcam Nu . N w. o mN N. m m mcmmm.m m.e:;pcwsm mmcwgzgucwsm o. N P o o m P o o mwuwew> esopom. so. N. oN oN N. @— Nm w. m mapoamwo: meopom. mmupsopom. mo. m. N. 3 m. Nm .. N. m magmmemocm mmeswguzco mm o. o. NN m P v e. N mum.=:mem mwmcmnppzh one .2: .2995 nm N o m e m m. e m m..mmgnzcme unsymmmoax: com. mm mm. mON mN mm. mom meN on mF=>Lma m..mEopm>mumcm mmvwgzpmmmonzr momN so. mmN m.m no. mmm mwm «mm mo. mponsm..ou gmuco Page» a . c. mN oN m m. m .uma .>oz .auo .uamm .m:< .m:< 3.33 mesa .mquA .oepcou sage mnoaogzue< yo .gopcm>c. u< xHozm¢m< 49 emsummou .OO mmuonsmooou new .OO Omuomzme NN O O O .. O ON O. O .OOOOOONN. .OO OOOONOOOOOON OO N O N . O O. ON N .OOO..OOOOOO. .OO.ONOOOmgm NOO OO OO NO OO O. NN OO. OO OOOOOOOOOO OOO OO NON NO ON N. OO OO. ON .OOOOLOOOOOOO. .OO OOOOOOOOOO .OO NO ON OO. .O. OO ON ON O .OOOOEOOOOOOO. .OO OOEOOOOOOO OOO. NON OOO .NN OON OO O.. OOO NN. .OOOOLOOOOOOOOO .OO OOOOOOOOOO .85. e .6 229:". 323 3.8:. OOOLOu< gouge O O O O . O O O O OOO...> O._OOOOOLO N . O O O O O O O OOO.OOOO.O.OE ONOOOeOOOO N c o O N m — o o Oaxoumgmn maunxuoanm. .ON ON NO ON N. .O ON N O OOO...OO OOOONOOOOOOO O O . O O O N . O OOOOOOOXOO OOOOOOOOOOOOO N.. O. O O N. NO ON O. N OOOOOOO> O..OO.OOOOOOO mmuwxua03oucm .OOON O O O. ON ON O O. O .umo .>oz .puo .uOmm .m:< .m=< Opzw mean ..OOOO. "O x.OZOOO< 50 O O O O O O O O O .OOOOOOOOOOOOOOLO. .Om szcousuxzomem NN O NO O O O N N O .OOOOOOOOEOOOOOO. .OO OOOOOEOOOOOO OON OO OO OO OO NO OO O O .OOOOOOOOOONO. .OO OOOOOOOOOOOOO OO O O OO O O O O O .OOOOOOOO. .OO.OOONN OOO OO OOO ONN OO NO OOO NOO OO .OOOOOOOO. .OO.mmflmOmmm .OOENOOOOOOUOOm u. OOOEOOOOOOOOOU OOO O OO OO ON O O. N O .OOOOOOOOO .OO OOOOOOOONO ON N OO O O O O N O .OOOOOOOOO .OO OOOOOOOO OOOOOOOOO 1. OOOEOOOOO O O O O O O O O O .OOOOOOOOOOOOO. .OO.OHmONum N O O O O O O O O OOOOOOOOOOO O O O O O O N O O OOOOOOOLOOOOO. .OO OOOOOOOOOO N O O O N N O O N OOOOOOOOOO. .OO OOOOOOOOOOO O O O O O O O O O . = O .OO OOOOOOOOOOOOOO N O O o O o O m o .mOquOmsuOocmzv .OO Ompmmcueocmz OO O O ON O O O O O .OOOOOOEOOOO. .OO OOOOOOOOOOO OOOOO O O OO ON ON O OO O .umo .>oz .poo .pamm .m:< .m:< OOOO 3:33 O.OOOO. "O xOOZOOOO 51 OO O O O O OO OO O O OOOOOOEEO O O O O O O O O O OOOOOOOOOOOO .O0.00000um O O O O O O O O O OOOOOOOOOO>OV .OO.OOOOOOOO O o O O o o o o o OOOOOOOOOOOOV .am OOOOEOmng OO O O O O O N NN O OOOOOOOOOLOOO .OO OOOOOOOOO ON O O N O O O O O OOOOOOOOOOOO .OO OOOOOOOOLOOO OO O N N O O N N O OOOOOOOOOOOO .OO.mOmmOmmxm ON N NO O O O N O N OOOOOOOOOOOOOO .OO OOOOOOOOOO OO O O OO O OO N O O OOOOOOOOOOOOV .OO OOOOOOOOEO O O O O O O N O O OOOOOOOOO .OO OOOOOOOOLO O O O N O O N O O OOOOOOOOO .OO OOOOOOOOOOEOO N O O O O O N O O OOOOOOOOO .OO OOOOOOOOOOO OOO NO OO OO OO OO OO OO N OOOOOOOOO .OO OOOO ONN N NN OO OO OO NO OO O OOOOOLOOOOOOOV .OO OOOOOOOOOOOOO OqumOpmoOmz ONO OO OO OO OOO OO NO OO O OOLOOOEEO O o o o O o o o o OmOuONOumwOOOOV .am OOLOuOmOOOO OO O O O O O O O O OOOOOOOOOOOOOOOOOO OOO OOLOOOOnggaam ON N O m O O O O O OOOOOwsamuopumOv .OO Ozmcawuouqu OOOOO O O OO ON ON O . OO O .owo .>Oz ouuo ouawm om3< .m=< OOOO mesa O.ucoov N< xOozmaa< 52 OO O O o O N O O o OOOOOO< O O O N O N O O O OOOOOOO O O O O O O O O O OOOOOOOOO O O O O O O O O O OOOOOOO OO O N O O O O O O OOOOOOOOO OO O O O O O O O O OO>LOO OOOOOOO OO O O O O O O N O OOOOOOO OO O OO O N OO OO O O OO>OOO OOOOOOOOOO ONO O OO OO O NN OO OO O OOOOOEOO O O O O O O O O O OOOOOOOOOO ONO O OO O O ON NO O O OLOOOOO OOO OO NO OO ON OO ON OO OO OOOOOOOOOOOO OO O O O O N O O N OOOOOOOOEN: OO N O O O O O O O OOOOOOeOI OO O O N O OO OO O O OOOO OO N O O O OO OO OO N OOOOOOOOOO ON N O O O N O O N OOOOOOOOO maomcm _. Omomwz OOOOO O O OO ON ON O OO O .umo o>oz .Ooo .pamm oms< .O=< OOOO mesa O.OOOOO "O XOOZOOOO 53 O O O O O O O O O .OO OOOOOz OOOOmez o o o o o o o o o Ozummu OOOOOOOOOL< OO o o o o OO O o O OOOOOOL OOLOOOOOEOOOOOOOO O o o o o O o m N OOOOEOO OOOOOmmnmm NN o o O o O O N mO OcmmmOw OONOOOOOEO mOOOLOOOOOEO O o o o o o O o o OOOOOO> OEOOOOO om O O O O O OO O m OOOOOOuo: OeouoOO OOOOEOOOOO OOO O ON O ON ON OO O O OOOOOOOOOO OOOOOOOOOO OON OO Om Om ON O N Om OO OOOOchgm OOmngOOOO mOOOOOOguaco o o o o o o o o o OOOOOOOOOOE.OLOOOOmoaO: ONO NO ON O N ON NN OO OO OOO>gumNOOOOEounxzuOLO OOOOOOOOOOOOO: NOO NO OOO ON NN om ON NO OO <40mzm44ou mmoao OOOOO O O OO ON ON O OO O .umo .>oz .uuo .uamm .m=< .m:< OOzc mesa .OOOOO OOOOOOOOz EOOO OOOOOLOOOO OO OOOOOO>OO "O xOOZOOOO LOOOOOOO .OO monomamuuoo OOO .OO OOOONOOO 54 OO O N OO O O O O O OOOOOOOOOV .OO OOOOOOOOOOOO ON O O O O O O O O OOOOOOOOOOOOO .OO OOOOOOOO ONO NO OO OO OO OO O ON O OOOOOOOOOO OO O OO OO O OO O O O OOOOOOOOOOOOOV .OO OOOOOOOOOO ONN OO OO NN OO O N O O OOOOOEOOOOOOOO .OO OOEOOOOLOO OOO OO OOO OO OO O OO ON OOO OOOOOOOOOOEONOO .OO OOOOOOOOOO OpmsmOpOOOO OZOOOOO OOOOO O O O O O O O O O OOOOOO> OOOOOOOOOO O O O O O O O O O OOOOOOOOOOOOE OOOOOOOOOO o o o o o o o o o Ozxovmgma OOOONUOOOOOO OO O N O O O O O O OOOOOOOO OOOONOOOOOOO O O O O O O O O O OOOOsuoxOO OOOOOOOOOOOOO OO O N OO ON NO O O O OOOOOOO> OOOOOOOOOOOOO OOOOOOOOEOOOO OOOOO O O OO ON ON O OO O own .>oz .puo .pamm .m=< .m:< OOOO mczn O.OOOOO HO OOOZOOOO 55 mm OOO mm OOO NO q-MLOOOOO oO OO 0 0000000 N OOOOOOO oO mm OO NoO (\l r—OOOOOO l\ r—I—OOOOO OO OO Om r—OQ’OOOO OOOOOOO ONOOOOO O OOOOOOOOOOOOOOOOOO .OO OOOcopsuxzuOgm OO OOOOOOccOEOOOOOOV .am OOOOOEOOOOOO N OOOOOOOOOOOOOV .OO OOOOOOOOOOOOO O OOOOOOOOOO .OO.OOOOO ON OOOOOOOOOO .Om.mflflmmmm OOOELOOOOOOOLOO uv OOOEOOOOOOOOOO o OmOOONOuoz OO .OOO mm OOOOO OOzw mcan O.OOOOO "O xOOszOO 56 MQ'Q'OmOr—NNQ' (no r—m N OmO ON NLOu—OOOOOOOOO m NNOOOMOOOOr-O Lnd'r—NOOF-Or—OOO OOOOOOOOF-OO 03 m 01 N r—OF-ONMOOOOMI— MONOOOOOOON LO (*3 OOONOOOOOr—r— O l\ v '- OmOOOpOOOLOOV .OO OOOOEOOLOO OOOOOOOOOOOOV .OO OOOOOOOOO OOOOOOOOOO>OV .OO.mOmmONNO OOOOOOOOOOOV .OO OOOOOOOOOOO< OOOOOOOOOOOV .OO.MHmmmmmxm OOOOOOOOOOOOOV .OO.OOOOOOONON OOOOOOOOOOOOOV .OO OOOmwaOe< OmOuOuOoz OO .OOO ON .OOOO ON .m:< m .m:< OO OOOO m mcaw O.OOOOV HO xOOzOOOO 57 l O O O O O O O O O OOOOOOOOO O O O O O O O O O OOBEO OO O O O O O O O N OOOOOOOOO OO O O N O O N O O OO>OOO ONOOOOO O N O O O O N O O OOOOOOO OO N N N O N O O O OO>OOO O.OOOOOOOOO NN O O O O O O N N OOOOOEOO O O O O O O O O O OOOOOOOOOO NO O O O O O O N O OOOOOOO ON O N O O O N O O OOOOOOOOOOOO O O O O N O O O O OLOOOOOOEO: OO O N NO O O O O O OOmuaoeo: OOO O O O O OO O O O OOO< OO N O N O O N OO N OOOOOOOOOO OO O O O O O O O O OOOOOOOOO OOOOzoz .OuO .pamm .OO< .O:< OOOO OOOO O.OOOOV HO OOOZOOOO 58 O O O O O O O O O OOOOOLO OO O O O O N O O O OOOOOOO OOOOO O O OO ON ON .O_ OO O .omO .>oz .puo .OOOO .O:< .O:< OOOO OOOO OOOOOOO nO XOOZOOOO 59 APPENDIX C: Inventory of Arthropods from Moldboard and Cultivated Plots Oct. 14 Nov. 1 Dec. 9 Tota] ORDER COLLEMBOLA Hypogasturidae Brachystomella parvula Hypqggstura manubrialis Onychiuridae Tu11bergia granu1ata Onxphiuris encarpatus Isotomidae Isotoma notab01us Isotoma viridis Sminthuridae Sminthuris elggans §phaeridia pumulis Deuterosminthuris russata ArhopaIites caecus Nee1idae Nee1us sp. En tomobryi dae Pseudocine11a violenta Pseudocine11a sexoculata Lepidocyrtus pallidus Lepidoqxrtus paradoxus Entomobnya multifasciata 0rchece11a vi110sa ORDER ACARINA Prostigmata Bakerdania sp. (Pygmephoridae) Tarsonemus sp. (Tarsonemidae) Scutacarus spa (Scutacaridae) 00000 (A) OONOWW 42 10 0 O 000‘ OO—‘OOW 87 32 2 GOOD OOO-HON 39 38 17 34 78 27 OOO-b O (A) 0000-me 169 80 19 60 APPENDIX C: (Cont.) Oct. Nov. Dec. 14 1 9 Tota] *Eupodidae 22 32 22 169 Rhagidia sp. (Rhagidiidae) 3 5 O 8 Triophtydeus sp. (Tydeidae) 0 7 1 8 Eustigmaeus sp. (Stigmaeidae) 0 3 0 8 Nanorchestes sp. (Nanorchestidae) 11 12 10 33 SpeIeorchestes sp. (Nanorchestidae) 1 2 O 3 SpinibdeIIa sp. (BdeIIidae) O 0 0 0 Ba1austium sp. (Erythraeidae) 0 0 0 O Brxobia sp. (Tetranychidae) 0 1 O 1 Eriophyidae 0 0 O 0 Astigmata (= Acaridei) Schwieba sp. (Acaridae) 0 3 7 10 Tyrophagus sp. (Acaridae) 0 1 O I Cryptostigmata (= Sarcoptiformes) OgieIIa sp. (Oppiidae) 28 42 28 _98 93913 sp. (Oppiidae) o 3 o 3 Sche10ribates sp. (OribatUIidae) O 13 1 I4 EphiIohmannia sp. (Epilohmanniidae) 7 11 5 23 Brachvphtonius sp. (Brachychtoniidae) 0 I 1 2 Tectocepheus sp. (Tectocepheidae) 0 7 3 10 Euphthiracarus sp. (Euphthiracaridae) 1 0 0 1 PaIaeacarus sp. (PaIaeacaridae) 0 O 0 0 Immatures 8 0 2 10 Mesostigmata RhodacareIIus sp. (Rhodacaridae) 11 4 4 19 A§23_sp. (Ascidae) I 0 0 1 *Eugodes sp. and CocceupOdes sp. together APPENDIX C: (Cont.) 61 Oct° 14 Nov° 1 Dec. 9 Total Cheiroseius sp. (Ascidae) Gamaselloides sp. (Ascidae) Arctoseius sp. (Ascidae) Amnyseius sp. (Phytoseiidae) Phxtoseius sp. (Phytoseiidae) ngoasgis sp. (Laelapidae) Androlaelgps sp. (Laelapidae) Allighis sp° (Eviphididae) Parasitus sp. (Parasitidae) Petgamasus sp. Parasitidae) Urogoda sp. (Uropodidae) Immatures MISCELLANEOUS DipIOpoda CoIeoptera Ants Homoptera Hymenoptera Thysanoptera Dip1ura Psocoptera Symphyla Coleoptera Iarvae Diptera Diptera Iarvae Pauropoda Protura Chilopoda Isopoda Araneae OOOOOOO-‘OOOO c: c: -a —a J: _. c: as u: _. -4 N m -> -a -O c: c: OOOOOOU‘IMOOOO OOOOOOOONVO 1 N OOOON OOOO-‘OOOOOOO OOOOOOO—‘OOOOOO-J—‘O OOOO—‘OU‘l-hOOOO Oddd-bflOOSONKO—I' 3 00 -b -O -a c: n: 62 NN Om OO OOO OOO 0000 d” N ON ON 0 FOOD OOON N 0 GOOD 0 N USP-VD O N Q‘l—F-r— F OO NOOO N O NN kOr—r—O O O OOOOOOOOOOOOOOZV .OO Owummgugocmz .3. mmzlwz mOOOOmmz OOumOu OOOOOOOOOO< OOOOOOO OOLOOOOOEOOOOOOOO OOOOEOO OOOOOOOOOO OcOmuOm OOOOOOOOEO OOOOOOOOOOEO OOOOOO> OsouoOO OOOOOOOOO OeouoOO OOOOEOOOOO Ozpmmgmocm OOOOOOOOOO OOOOOOOOO OOOLOOOOOO OOOOLOOOUOOO OOOOOOOOOOE OLOOOOOoqam OOO>OOOOOOOOEOOOOOOOLO OOOOLOOOOOOOO: <4002m44ou «memo OOOOO O .qu O .>oz OO .OOO ON .OOOO ON .OO< O .OO< OO OOOO O OOOO .OOOOO OmpOmLO OOONOOO< OOO .umuO>OuO:u .OLOOOuOoz EOOO muoaogcug< mo Ogoucw>cO “a xOoszO< 63 NOO OO NO O OO N OO O O OOOSOEO O O O O O O O O O OOOOOOOOOOOOOOV .OO OOLOUOOOOOO O O O O O O O O O OOOOOOOOOOOOOOOOOO .OO OOOOOOOOOOOOOO OO N O O O O O O O OOOOOOOOOOOOOOOV .OO OsmsnmoopomO O O O O N O O O O OOOOOOOOOOONOOOOOO .OO OOOOOOOOOOOOOO ON O OO OO O OO O O OO OOOOOOOOOEOOOOOOO .OO OOOOOEOOOOOO OOO OO ON OO OO OO ON N O OOOOOOOOOOOOOO .OO OOOOOOOOOOOOO NN O O O O O O O O OOOOOOOOOV .OO.OOOOO ONO OO OO OOO OO NN OO ON OO OOOOOOOOOO .OO.OOOOOOO OOmELOOOpOouOOO uv OOOEOOOOOOOOOO O O O O O O N O O OOOOOOOOOO .OO OOOOOOOOOO OO N O OO O N OO OO O OOOOOOOOOO .OO OOOOOOOO OOOOOOOU< uv OOOeOOOO< O O O O O O O O O OOOOOOOOOOO O O O O O O O O O OOOOOOOOOOOOOOO .OO.OOOONum O O O O O O O O O OOOOOOOOOOOOOO .OO EOOOOOOOOO O O O O O O O O O OOOOOOOOOOO .OO OOOOOOOOOOO OO O N O O O O O O OOOOOOOOOOOOOOZV .OO OwummsugomOmOO OOOOO O O OO ON ON O OO O .umO .>oz .uuo .OOOO .OO< .OO< OOOO OOOO O.OOOOO HO xOOZOOOO 64 ON OO O \OFOOOMr—NOOOOOO O Mr-OOOFOMOONOOO f—OONOOr—l—‘OOOOO £0 <- N LDNOOOr—l—NOF-I—OOI— O LDNr—OOOr—OOQ'OONO N NNOr-Nt—OLDOMMOOED F F f—OOOC‘r—Of—‘l—OOOI— O N CDQ'OOF-OOr—OF-OOOP OLOOOOOOOO OOOOOOOO OOOOzOO .OO.OOOOOOOO OmOOOOOOmOOV .OO OQOOOOOOLOO< OOOOOOOOOOOV .OO.mmmmmmmxm OOOOOOOOOOOOOV .OO OOOmOouxmm OOOOOOOOOOOOOV .OO OOOOOOOOE< OmOOOuOoz OO .OOO ON .pamm ON .OO< O .OO< OO OOOO O OOOO O.u=ouv no xOoszO< 65 O O O O O O O O O OOOOOOO O O O O O O O O O OOOOOOO OO O O O N N O O O OOOOOOOOO N O O O O O O O N OLOOOOO O O O O O O O O O OOOOOOOOO OO N O O O O O O N OO>OOO OOOOOOO O O O O O O N O O OOOOOOO OO N O O O O N O O OO>OOO OOOOOOOOOO O O O N N O O O O OOOOOEOO N O O O O O O O O OOOOOOOOOO OO O N N O O N O N OOOOOOO OO O O O O O O N O OOOOOOOOONOO N O O O O O O O O OOOOOOOOEO: OO O O O O O N N O 2388: OOO O N O OO OO O O OO OOOO OOOOO O O OO ON ON O OO O .umO °>oz .Ouo .OOOO .OOO .O:< OOOO OOOO O.O=OOO HO OOOZOOOO LIST OF REFERENCES LIST OF REFERENCES A1einikova, M. 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