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This is to certify that the
thesis entitled
THE EFFECTS OF ATRAZINE

ON NON-TARGET SOIL ARTHROPODS IN NO-TILL
CORN PRODUCTION

presented by

JOHN CHRISTOPHER MOORE

has been accepted towards fulfillment
of the requirements for

_M._S_.__degree in M

 

Major professor

Date November 11, 1981

 

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THE EFFECTS OF ATRAZINE
ON NON-TARGET SOIL ARTHROPODS IN NO-TILL

CORN PRODUCTION

BY

John Christopher Moore

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 ATRAZINE
ON NON-TARGET SOIL ARTHROPODS IN NO-TILL
CORN PRODUCTION
BY
John ChristoPher Moore

The effects of Atrazine on soil microarthropods was
assessed for the first growing season of No-till corn at the
new Crops and Soil Science farm of Michigan State University.

Epigeic Collembola were directly affected by tillage
while edaphic Collembola were not. Atrazine had no negative
effects on Collembola; however two species, Sminthurinus
elegans (Pitch) and Tullbergia granulata (Mills), were found
in greater numbers in Atrazine treated plots. Tillage and
Atrazine decreased Prostigmata and Mesostigmata mite papula-
tions. Astigmatid and Cryptostigmatid mite populations in-
creased as a result of tillage.

Changes in Collembola and Acarina pOpulations occurred
when the habitat changed. It was difficult to determine
whether No—tillage and/or Atrazine directly affected in-
dividuals or if changes in the soil microenvironment and

surface vegetation caused by these treatments affected them.

TO MINDY
+1

ii

ACKNOWLEDGEMENTS

I would like to thank Dr. Ivan Mao of the Department
of Dairy Science and Dr. Robert Boling and his staff for
their assistance with the statistical analysis, and Dr.
Lynn Robertson and Mr. Dallas Hyde of the Departmenn of
Crops and Soil Science for planning and managing the farm
operations. Special thanks is given to my major Professor,
Dr. Richard Snider, for his patience and guidance through-

out this study.

iii

TABLE OF CONTENTS

ABSTRACT . . . . . . . . . . . .
ACKNOWLEDGEMENTS . . . . . . . .
LIST OF TABLES . . . . . . . . .
LIST OF FIGURES . . . . . . . .

INTRODUCTION AND LITERATURE REVIEW . . ...

LABORATORY STUDIES . . . .

FIELD STUDIES-HERBICIDE EFFECTS . . .

FIELD STUDIES-TILLAGE EFFECTS . . . .

MATERIALS AND METHODS . . . . .
RESULTS AND DISCUSSION . . . . .
EXTRACTION EFFICIENCY . . .
GENERAL . . . . . . . . . .
PLOT BIAS . . . . . . . . .
COLLEMBOLA . . . . . . . .
ACARINA . . . . . . . . . .
AGE STRUCTURE . . . . . . .
CONCLUSION . . . . . . . . . . .
RECOMMENDATIONS . . . . . . . .
APPENDICES

13
13
15
16
16
29
41
89
50

A. CHECKLIST OF ARTHROPODS COLLECTED FROM GRASS
CONT ROL PLOTS O O O O O O O O O O O O O C .

B. CHECKLIST OF ARTHROPODS COLLECTED FROM NO-TILL
CONTROL PLOTS O O C O O O O O O O O O O . .

iv

APPENDICES (CONTINUED)

C. CHECKLIST OF ARTHROPODS COLLECTED FROM NO-TILL'
ATRAZINE PLOTS O O O O O O O O O O O O 0 . 6 3

D. MEANS AND 95% CONFIDENCE INTERVALS FOR.ARTHRO’
PODS 2193532391. 0 ‘.' o o o o o o o o o o o o O o 6 8

LITBMTURE RETED O O O O O O O 0 O O O O O O O O O O 8 3

TABLE

1.

LIST OF TABLES

Catalog of Field Preparations and Data
Collection . . . . . . . . . .

O O O O O O O O O O 7
Air and Soil Temperatures on Sampling Dates . . . 8

Percent Extraction Using Tullgren Funnels,
Determined by Floatation . . . . . . . . . . . . . 14

vi

LIST OF FIGURES
FIGURE Page
1. Daily Maximum and Minimum Temperatures . . . . . . 10
2. Daily Precipitation . . . . . . . . . . . . . . . ll
3. Mean Population Levels of Smdnthuridae . . . . . . 18
4. Mean Population Levels of Sminthurinus elegans . . 21
5. Mean Population Levels of Collembola . . . . . . . 23
6. Mean Population Levels of BrachyStomella parvula . 26
7A. Mean Population Levels of Isotoma notabilis . . . 28
B. Mean Population Levels of Lepidocyrtus pallidus . 28
C. Mean Population Levels of Tullbergia granulata . . 28
8. Mean Population Levels of Prostigmata . . . . . . 31
9. Mean Population Levels of Pyemotidae . . . . . . . 33
10. Mean Population Levels of Cryptostigmata . . . . . 35
11. Mean Pepulation Levels of Opiidae . . . . . . . . 37
12. Mean Population Levels of Mesostigmata . . . . . . 39
13. Mean Population Levels of Rhodacaridae . . . . . . 43
14. Mean Population Levels of Astigmata . . . . . . . 45

15A. Mean Population Levels of Juvenile Brachystomella
Rama 0 O O O O O O O O O O O O O O O O O O O O 47

B. Percentage of Juveniles in Brachystomella 2am mla
Population . . . . . . . . . . . . . . . . . . . 47

vii

INTRODUCTION AND LITERATURE REVIEW

Agronomists recognize the need to minimize tillage in
order to reduce runoff and erosion. Many forms of "conser-
vation tillage" have been developed and are being practiced.
The method employed depends on the crop and soil (Robertson
et a1, 1976; Vitosh & Warncke, 1976; Cook & Robertson, 1979).
A form of conservation tillage used for row crop production
on well-drained soil is "No-till” (Nelson et a1, 1976). In
this practice, a narrow slit is made in the soil with a
fluted coulter. Seeds are dropped into the slit and covered.
Pesticides and fertilizer may be added during planting. No
further cultivation is needed. When compared to conventional
tillage, No-till offers labor and energy savings, since fewer
trips to the field are required (Phillips & Young, 1973; Foth,
1978). Because the soil is minimally disturbed and plant
residues are allowed to accumulate, there is a decrease in
erosion and runoff and an increase in moisture retention
(Nelson et al, 1976; Robertson et a1, 1976).

To ensure these benefits however, one must rely complete-
1y on herbicides to control weeds (Chase & Meggitt, 1976).
Many herbicides presist in soil, bound to organic matter and
Clay, long after their application (Talbert & Fletchall,
1963; Bucholtz, 1965; Upchurch, 1966; Williams, 1970; Skipper
5 \holk, 1977). Laboratory and field investigations indicate

that their annual use may adversely affect arthrOpod

1

2
populations responsible for soil formation and nutrient re-
cycling (Edwards, 1970; Galston, 1979). The purpose of this
study was to evaluate the effects of the herbicide Atrazine‘
(2-chloro-4-(ethylamino)-6-(isopropylamino)-s-triazine) on
soil arthropods in No-till corn production.
Laboratory Studies

Several laboratory studies have shown that herbicides in-
duce mutations in non-target plants and animals (Fahmy &
Fahmy, 1954, 1955; Herkowitz, 1956; Wuu & Grant, 1966; Liang
& Liang, 1972; Plewa 8 Gentile, 1976; Gentile et a1, 1977;
Murnik & Nash, 1977). Few investigations on direct effects
of herbicides on soil arthropods have been reported. However,
some of those recently published used Collembola as a test
animal (Eijsackers, 1975; Dwidjasatmoko, 1978)

Eijsackers (1975) observed the reactions of Onychiurus
guadriocellatus (Gisin) to varied concentrations of 2,4,S-T.
Increased activity, leg tremors, and paralysis were observed
at doses of 1.25, 2.50, and 5.00 cc/MZ. Mortality was
related to the concentration of the 2,4,5-T; the higher the
dose, the greater the mortality. Collembola presented with
a choice between treated and untreated food demonstrated a
preference for the latter.

Atrazine and Paraguat, herbicides commonly used in corn
production, may affect the reproductive system of Collembola
(Dwidjasatmoko, 1978). Folsomia candida (Willem) and
Igllbergia granulata (Mills) showed delayed instar durations
and reduced fecundity when fed 600 ppm, 1000 ppm, and 5000

Hum When reared on soils treated with 5000 ppm Atrazine,

3
Collembola showed increased mortality levels.
Field Studies-Herbicide Effects

Since herbicides were introduced to control weeds, many
field investigations on possible non-target effects have been
undertaken. Studies in grasslands and cereal crops have
shown MCPA does not effect the soil fauna (Rapoport &
Cangioli, 1963; Davis, 1965). The same conclusions were
drawn from investigations of 2,4-D and DNOC treated grassland
(Johnson et a1, 1955; Rapoport & Cangioli, 1963; Fox, 1964).
A later study by Edwards (1970) found lower Collembola popu-
lation levels in DNOC treated plots. These populations quick-
ly recovered after levels of the compound in the soil had
decreased.

Results of field studies on the effects of Paraguat and
Dalapon are conflicting. When compared with untreated_fie1ds,
Fox (1964) found no significant difference within the total
fauna in fields treated with either herbicide. Closer
analysis indicated Collembola populations had increased.
Similarly, Edwards et a1 (1971) showed an increase in arthro-
pod populations exposed to Paraguat in slit-seeded (No-till)
wheat fields. Groups demonstrating more pronounced increases
were rhodacarid mites, entomobryid and sminthurid Collembola,
Symphyla, Thysanoptera, and Coleoptera. Curry (1970), on the
other hand, found decreased soil arthropod populations in
fields treated with Dalapon and Paraguat. These changes were
attributed to habitat changes cause by the herbicide rather
than toxicity.

Herbicides that have demonstrated adverse effects are

4

Monorun and the s-triazines. Fox (1964) found a signi-
ficant decrease in wireworms, Collembola, and Acarina in
grasslands treated with Monorun. The same study showed Atrad
zine reduced Collembola populations in grass plots. Popovici
et a1 (1977) treated corn fields with Atrazine at 5 kg/ha
and 8 Kg/ha and found a 68% and 75% reduction in Acarina.
Collembola were reduced 91% and 95% for these concentrations.
Four months following application, Collembola were still re-
duced 59% and 80%. Edwards (1970) and Edwards et a1 (1971)
reported similar results for Collembola populations respond-
ing to Simazine, in that 70% of the Collembola were elime
inated. After 5 months , however, very little difference was
observed between population levels in Simazine treated fields
and untreated controls. The same study reported reduced
numbers of mesostigmatid and oribatid mites, and isotomid
Collembola in plots treated with 2 and 4 Kg/ha of ”Bladex."
Field Studies-Tillage Effects

A problem investigators have when studing the influence
of herbicides on the soil fauna of agricultural soils is
separating tillage effects. The extent of tillage effects
depends on tillage type. Edwards and Lofty (1975) compared
the effects of conventional tillage and slit-seeding (No-
till). Significantly fewer arthropods were found in conven-
tionally plowed plots. The more intensively tilled plots
showed greater reductions. Arthropods most effected were
cryptostigmatid mites and surface dwelling and hemi-edaphic
Collembola. The effects of slit-seeding differed between

different artrhopod groups. Most showed no changes; however,

5
predatory mites, euedaphic Collembola, and larval Diptera
populations were significantly reduced. Loring (1979) com-
pared the effects of No-till, moldboard plow, and chisel
plow tillage systems on soil arthropod populations. Unlike
Edwards and Lofty (1975), Loring (1979) found conventional
tillage systems stimulated microarthropod populations.
However, the authors do agree that surface dwelling and hemi-
edaphic arthrOpods decreased. Loring (1979) found No-till

population levels to be the same as those of 'old fields.”

MATERIALS AND METHODS

The study area was located off JOlly and College Roads,
on the north end of section 'C' of the new Crops and Soil
Science farm at Michigan State University. The field was
maintained as grassland for seven years prior to the present
investigation. The soil was classed as a Celina loam (soil
management group 2.5 a, Aquic Hapuldalf, fine, mixed, mesic) .
In May, 1979, twelve 6.1m x 15.2m plots were established.

The plots were divided equally among three treatments. The
treatments were untilled grass without Atrazine (control),
tillage without Atrazine (No-till control), and tillage with
Atrazine (No-till Atrazine).

On May 12, 1979, corn was planted and fertilized with
lSOkg/ha of 6-24-24. May 18 and June 12, 0.47 and 0.94 cc/n2
of Atrazine was broadcasted. Sprinkler irrigation was
installed over the entire study area June 28, with 12.7mm
supplied June 28, 38.1mm on July 10 and 18, and 25.4 on July

27. The irrigation system was removed August 16 and the

6
corn was harvested October 28.

Soil samples were taken from April 17 through November
8, 1979, at two to three week intervals. Prior to planting,
samples were taken to establish base population levels. On
May 15 and 31 three plots per treatment were sampled. On
those dates 10 samples per plot were taken. On June 14, all
treatment replicates were sampled and five samples per plot
were taken. On subsequent sampling dates seven samples per
plot were taken. A catalog of all field preparations and
sampling dates is provided in Table 1.

At each sampling date, soil (15cm depth) and air temper-
ature (1m height) were recorded using a Yellow Springs Tele-
thermometer (YSI 428C) (Table 2). Daily maximum and minimum
temperature and precipitation were obtained from.the Michigan
State weather Service (Horticultural Garden Station)
located three miles from the study site (Figures 1 and 2).

Samples were placed in plastic bags. To avoid overheat-
ing in the field and during transportation, samples were
stored in a styrofoam cooler. Samples were placed in Tullgren
funnels and extracted for a minimum of three days. A 25 watt
light bulb was used as the heat source. Extracted animals
were preserved in 95% ethanol - 1% glycerol.

Arthropods were identified to species (Collembola).
family (Acarina), and order (Coleoptera, Diptera, Puaropoda,
and Symphyla). Collembola were identified according to
Snider (1967) and Christiansen and Bellinger (1981). Acarina
were identified according to Krantz (1970).

The Collembola were divided into two ecological groups

7
Table 1

Catalog of Field Preparations and Data Collections

Imus-WWW"unusnmnngns:

Dates

 

April 18 48 soil samples were taken to survey the fauna.

May 8 48 soil samples were taken to survey the fauna.

11 Soil was tilled, corn was planted, and 150 kg/ha
of 6-24-24 fertilizer was applied.

15 90 samples were taken, 10 from.each plot. Three
replicates were included.

18 .47 cc/mz of Atrazine was applied to No-till
Atrazine plots. '

31 90 samples were taken, 10 from each plot. Three
replicates were included.

 

June 12 .94 cc/m2 of Atrazine was applied to No-till
Atrazine plots.
14 60 samples were taken, 5 from each plot. Four
replicates were included.
28 Sprinkler irrigation was installed. 12.7mm of
water was supplied.

July 8 84 samples were taken, 7 from each plot. Four
replicates were included.
10 38.1mm of water was supplied.
18 38.1mm of water was supplied.
25 84 samples were taken, 7 from.each plot. Four
replicates were included.
28 25.4mm of water was supplied.

August 16 84 samples were taken, 7 from each plot. Four
replicates were included.

Sept. 8 84 samples were taken, 7 from each plot. Four
replicates were included.

28 84 samples were taken, 7 from each plot. Four
replicates were included.

October 18 84 samples were taken, 7 from each plot. Four
replicates were included.
20 The corn was harvested.

Nov. 8 84 samples were taken, 7 from each plot. Four
replicates were included.

 

 

 

 

 

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Figure 1. Daily Maximum and Minimum Temperature

Figure 2. Daily Precipitation

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12
on the basis of that portion of the soil they inhabit.
Collembolan species of the family Sminthuridae living on the
soil surface were grouped as “epigeic” Collembola. Collemr
bolan species living within the soil were grouped as 'edaphic'
Collembola. One epigeic Collembola species, Sminthurinus
elegans (Fitch) was collected in sufficient numbers to analyze
separately. Four edaphic species were numerous enough to
analyze separately. These species were, in order of their
abundance, Brachystomella parvula (Schaffer), Tullbergia
granulata (Mills), Lepidocyrtus pallidus (Reuter), and
Isotoma notabilis (Folsom).

Acarina were separated into suborders: Prostigmata,
Cryptostigmata, Mesostigmata, and Astigmata. Of the Pro-
stigmata, the family Pyemotidea was analyzed separately. The
cryptostigmatid family Opiidae and mesostigmatid family
Rhodacaridae were analyzed separately.

Extraction efficiency of the Tullgren funnels was
estimated as follows. After extraction, 20 samples taken
June 18, 1980 were immersed in a saturated sugar solution.
.Arthropods previously trapped in the soil, floated to the
surface and were counted.

The following two-way cross classification model with
interactions was used to generate f—ratios, estimate mean
.numbers of selected arthropod groups collected at each sample
date, and calculate 95% confidence intervals for each mean:

ingk1=u+Ti+Sj +P +TS

i:k ij + eji:k1

13

where,
u = Mean common to all observations.
Ti = The effect due to the ith treatment, there are
three treatments.
Sj = The effect due to the jth sample date, there
are 10 sample dates.
Pizk = The effect due to the kth treatment plot repli-

cate, there are four plots in each treatment.

TSij = The effect due to the ith treatment on the kth
sample date. There are 30 treatment-sample
date interactions.

eji:k1= The residual random term corresponding to in:kl-

.All possible linear contrasts were performed with the treat-
:ment, treatment plot replicate, and treatment-sample date
interaction classifications. Treatment contrasts were tested
for differences between treatments in the mean number of
selected arthropods collected over the entire growing season.
irreatment plot replicate contrasts Were tested for differences
among replicate plots of a given treatment in the mean number
(of selected arthropods collected over the entire growing
season. Treatment-sample date interaction.contrasts were
'tested for differences between treatments in the mean number

<3f selected arthropods collected at each sample date.

RESULTS AND DISCUSSION
Extraction Efficiency
The results of the extraction efficiency analysis are
provided in Table 3. These values are used for comparing
Tullgren funnel efficiency of other investigations. Analysis

showed that for a 72 hour extraction period, 51.8% of the

14

Table 3

Percent Extraction Using Tullgren Funnels
Determined by Floatation

====mm============= ======a=======a==========

 

 

 

Taxa % Extraction
Total Collembola 51.8
Brachystomella parvula 50.0
Tullbergia granulata 40.0
Total Acarina 35.8
Pyemotidae . 72.8
Opiidae 11.1
Rhodacaridae 40.0

 

Collembola were extracted. Of the Collembola found in num-
bers large enough to analyze, 50% of the Brachystomella
pgrvula and 40% of the Tullbergia granulata were extracted.
Acarina efficiency was 35.8%, with 76.6% of the Pyemotidae,
40% of the Rhodacaridae, and 11.1% of the Opiidae extracted.
Discrepancies in extraction efficiency between investi-
igations stem from different extraction periods, efficiency
<ietermination techniques, and soil characteristics. After
a 42 hour extraction period, Tamura (1976) handsorted dried
soil and obtained funnel efficiencies of 16% and 18% for
(Zollembola and Acarina respectively. Those values are sub-

stantially less than the values reported here. Aside from

15

the various efficiency methods used, differences are a result
of unequal extraction times. Loduvic (1962) reported that at
15 C the majority of Collembola were collected after 4-10
hours, while the Acarina took 80-100 hours. Loring (1979)
estimated Tullgren funnel efficiency with the same funnels
and determination procedures used in this study. The chief
difference between the studies was the soil type. Loring
(1979) had a Spinks loamy sand compared to the Celina loam
of the present study. In his study, Loring (1979) collected
only 3% of the Tullbergia granulata and 95% of the Pyemotidae.

Although results are often inconsistent, studies on the
efficiency of Tullgren funnels have yielded two generalities.
Because efficiency is varied between groups, care must be
taken when estimating population sizes for ecological studies
(Sheals, 1957). Smaller species, particularly those with
poorly developed locomotor structures, have lower effici-
encies than larger, more mobile species (Haarlov, 1962;
Tamura, 1976).
General

Neither tillage nor Atrazine had an effect on the number
of arthropod groups in the field. Differences were observed
in the abundance of certain groups. A complete checklist of
all arthropods collected during the study is provided in
Appendices A, B, and C.

Samples taken to establish base population levels were
:not included in the analysis. These samples were taken at
random from the entire study area before plots were estab-

lished. For this reason, these samplesscould not be compared

16

statistically with samples taken from defined plots.
Plot Bias

Linear contrasts were tested to identify plot biases
using data of taxa collected in sufficient numbers to
analyze. Replicate grass control plots did not differ sig-
nificantly (p 0.05). Differences between the number of
arthropods collected from no-till control plots andlgo-till
Atrazine plots were found (p 0.05). Although differences
between replicate plots were found, plot bias was not con-
sidered an important factor for the following reasons: 1)
’Differences between replicate plots were not consistent
for all groups analyzed. No single plot had high of low
population levels for all taxa. 2) Differences were
restricted to No-till control and lee-till Atrazine plots.
{rhis indicated that undisturbed the study area was probably
tiniform. Differences observed were caused by treatment
and not plot condition of location.
Sggllembola

The epigeic Collembola were affected bypro-tillage.
Significantly fewer (p 0.05) Sminthuridae were collected
from tilled plots than grass control plots. The pOpulations
<1ecreased in each treatment during June and then increased
(iuring July (Figure 3). The Sminthuridae population reached
Inaximum levels on July 25 in the tilled plots and August 16
in.grass control plots. Sminthurinus elegans resembled other
Sminthuridae in that significantly fewer (p 0.05) numbers
'were found in tilled plots than grass control plots on May

15 and 31. After mid-June however, S; elegans populations

17

Figure 3. Mean Population Levels of Sminthuridae

“'1

 

Sempie Meen

 

Sminthuridae \

18

 

 

 

1 e
-—--'" Control
------ tie-tilt Control
‘ ‘ -— -- Ito-tilt Atrazine
sin 7% 7:25 elm 91- (as 10:18 “it

 

 

447

Figure 3.

5:! sins 5:31
Sampling Detes

19

in grass control plots did not differ significantly (p 0.05)
from population levels in No-till control plots-(Figure 4).

Atrazine did not adversely affect the Sminthuridae. No
significant differences (p 0.05) were observed between the
-population levels in no-till control and No-till Atrazine
;plots. Unlike other Sminthuridae, S; elegans populations
.increased only intto-till Atrazine plots during July. On
July 25 significantly more (p 0.05) S_. elegans were col-
lected from plots treated with Atrazine than those that were
ruot. The July 25 increase in the S; elegans population
cuzcurred when much of the original ground cover had been re-
duced to multch in the uo-till Atrazine plots. Increased
available organic matter on the soil surface may have had a
zxositive affect on fungal and nematode populations §;_elegans
feed upon (Snider, R. J., personal communication).

The edaphic Collembola were affected by tillage. Ini-
1:ially, few Collembola were collected from any of the plots.
POpulations increased rapidly in all treatments and were most
numerous by the end of July. In general, populations in each
treatment differed in numbers, but not trend (Figure 5).
Tfirroughout the study more Collembola were found in grass con-
trol plots than tilled plots. Tillage was not found to be
8ignificant (p 0.05) until June 14 (Figure‘S). This indi-
Gated that microenvironmental changes initiated by tillage
1Tather than actual tillage processes contributed to the
declined. The occurance of Brachystomella parvula was very-
Sindlar to that of the edaphic Collembola as a group. ‘BL

Earvula was more numerous in grass control plots than either

20

Figure 4. Mean P0pulation Levels of Sminthurinus elegans

 

 

 

21

30-1.

 

22

Figure 5. Mean Population Levels of Collembola

 

23

 

 

”J _ cm... Collembola
.- ----- Mo-ttll Control
.—~-— tto-tiit Atrezine
«-
.ol
as< l
H
1. \.
sea I , i \
s I o’ '\
e I" |
a 1" ‘t \
. . II t‘
a 2’ ‘ x ‘t \
s ' “ ‘-
0
204 I
. ./'
151 f _/
I" '/
' f
I ./
104 . ..... ,5 fl
./
./
s- ’1
H7 3:. 5:” 5:” .j‘ 7:. 7:” '1‘. oz. 9:2. 10:). "I.

Figure 5.

Semoitne Dot“

 

24
No-till control or No-till Atrazine plots. Tillage was not
found to be significant (p 0.05) until June 14 (Figure 6).
Lepidocyrtus pallidus and Isotoma notabilis were also
affected by microenvironmental changes caused by tillage. L;
pallidus was more numerous in grass control plots than tilled
plots in June, late July, and early fall (Figure 73). I;

notabilis was more numerous in grass control plots than tille

 

ed plots in mid-summer and early fall (Figure 7A).

Edaphic Collembola appeared not to be adversely affected
by Atrazine. The L; pallidus and IL_notabi1is populations
in No-till control and No-till Atrazine plots did not differ

(Figure 7A and 7B) . The _B_._ parvula populations in uo-till
control and No-till Atrazine plots significantly differed
only on July 25 and August 16 (Figure 6). Tullbergia
graulata was not affected by tillage throughout the study
(Figure 7C). Significantly more (p 0.05) T; granulata
were found in No-till Atrazine plots than No-till control
or grass control plots on September 28, October 18, and
November 8 (Figure 7C).

These results reflect differences in niches Collembolan
sPecies occupy and the subsequent changes these habitats
l~11'1derwent after each treatment. The epigeic Collembola were
ill direct contact with farm equipment traffic while edaphic
COllembola were sheilded from much of this activity. This
rue)? explain why fewer epigeic Collembola were found in tilled
Plths than grass control plots immediately after tillage,
While edaphic Collembola were not significantly affected.

Iwaiter differences in epigeic population levels that arose

25

Figure 6. Mean Population Levels of Brachystomella parvula

 

26

45. ._ Control A Brecnyetorneiie pervuil
100-0... mam Controi
=—.— mam Atrezine

 

401 I
.I‘. \
30‘ /\ ‘4

   

 

 

B
3 25* ,. \
I I \ l
5 2o- / ,
a \ 1 I],
\ / I
, pd)- , .’.'
.‘ ‘~ " -.4 I
w .
.' / -
/
iO-l . ‘
5‘ '-
O
E s3e 5:15 5:3: 034 73 7125 site Ce 9:“ :J-ie nit

Semoiing Dates

Figure 6 .

27

Figure 7A. Mean Population Levels of Isotoma notabilis

 

Figure 7B. Mean Population Levels of Lepidocyrtus pallidus

 

Figure 7C. Mean Population Levels of Tullbergia granulata

 

28

5. isotoma notabilis -—— Control
0- ----- tte-tm Control
A --~— tie-tilt Atrazine

 

 

 

F igure 7A.

4 « Lepidocyrtus pallidus

  

31

24

\ -
f...— \/ \‘\_.__.—._..-—.._._.-—--

 

Semole Mean

 

 

 

 

Figure 7B.
sI, Tullbergia granulata
‘l
3.
2.
..
4:" sze 5:5 531 A01“ 75 7:2: alto are ozze Iolie n e

Sampling Dates

Figure 7C.

29
between treatments were a result of changes that occurred
on the soil surface. Field observations made during the
study, showed surface vegetation in No-till Atrazine plots
had been erradicated and corn emerged by mid-June. The
decaying vegetation and newly emerged corn created a habitat
markedly different from the original grassland. The sharp
increase in S; elegans population in No-till Atrazine plots
during July may have been a response to habitat changes
incurred by Atrazine (Figure 4) .

Edaphic Collembola were affected by changes that occur-
red within the soil. Tillage alters the soil structure
resulting in changes in soil microenvironment (Aleinikova
& Utrobina, 1975) . Collembola living in the soil were
affected by microenvironmental changes rather than actual
tillage processes. The nature of these changes, and means of
their action on edaphic Collembola pOpulations cannot be
determined from these data. Because No-till control and No-
till Atrazine were found not to differ significantly for the
majority of edaphic species analyzed, Atrazine was not an
important factor.

Acarina

The Prostigmata were affected by tillage and Atrazine.
Significantly fewer numbers were collected from tilled plots
til'lan grass control plots. They were least abundant in No-

42111 Atrazine .plots. Populations in No-till control plots
and grass control plots exhibited similar trends. In these
treatments the Prostigmata were more numerous at the end of

the growing season. The early growing season was marked

30

Figure 8. Mean Population Levels of Prostigmata

31

701 Prostigmata

60‘

50+

404

Sample Mean

aoI

204
./'

\.\.\/.

 

‘0‘ ._—.—— Control
- ----- Ito-till Control
—‘— No-till Atrazine

 

 

441 5.4545 son 444 7:0 7525 e314 9:4 9:" IO-ll n-a

Sampling Dates

Figure 8 .

32

Figure 9. Mean Population Levels of Pyemotidae

33

 

 

 

‘0 4 Cum, Pyemotidae
0- ----- NO-NII COM?“
—-— lto-till Atrazine
4
3° 4
c
e
e
3
o
a 20
E
e
to
.l
to 4

447 s-‘eSIls 5:3! 4:44 73a 73: e114 91a 912a l0le ":4

Sampling Datea

Figure 9 .

34

Figure 10. Mean Population Levels of Cryptostigmata

15‘

Sample Mean

Cryptostigmata

,4

 

4:;

Figure 10.

35

o—-—— Control
.. ----- tie-till Control

—— - Ito-tilt Atrazine

5:4 sits 5:31

a." 74 7:25 0:16 9:. '1”

Sampling Dates

V
10°"

Y
"'3

36

Figure 11. Mean Population Levels of Opiidae

37

 

isd °-—- Control 09"le
------ lee-till Cont?!"
~—- Io-tilt Atrazine
ith
B
I
O
3
.2
i
in
51

 

 

4.77 5:65:15 sin 444 71a 7125 tile 9-4 9-2e more "-4

Sampling Dates

Figure 11 .

38

Figure 12. Mean Population Levels of Mesostigmata

 

39

 

iS < lleeoatigmata
R
’ \
’ \
’ \
II \\
I
I \
I \
I \
\
IO; / \
A x .
/ \ I \A
I _/\
C / ' / \ \
I r . \
O \p’ // \x \
’ / 2v" ~. \
. . ,I ‘\ I,
“ / / \~;‘ ‘. ’I’
E / «~32
I \ . .
a) \. ./ ,’ \.
,X. X \
I ‘\ l’ .
II \\ ,’
54 [I ‘g’
d
-——— Control
~----- lie-till Control
—'— tie-till Atrazine
4-17 s-es-ls .5131 6-i4 7-4 7-25 e-le 9-e o-ae lo-ie n-e

I"'I'Lgure 12 .

Sampling Dates

40

with fluctuations (Figure 8). From the end of July through
October significantly fewer Prostigmata (p 0.05) were found
in no-till Atrazine plots than grass control plots. The
Pyemotidae showed an overall trend similar to the Prostig-
mata (Figure 9). This was expected as they were the domi-
nant mite family.

The Cryptostigmata were affected by tillage and Atra-
zine. Populations in each treatment did not differ until
the third sampling date. The Cryptostigmata were most numer-
ous in.No-till control plots (Figure 10). Similar results
showing an increase of Oribatid mites after cultivation were
reported by Shaddy and Butcher (1977). They attributed the
increase to microenvironmental changes incurred by tillage.
An increase after tillage was not found in Noetill Atrazine
plots or grass control plots. Peak abundance in grass
control plots was attained in October. The Opiidae showed
an overall trend similar to the Cryptostigmata as a group
(Figure 11). They were most numerous in No-till control
plots, but, did not sustain high numbers as long as the
Cryptostigmata. The Opiidae population in grass control
plots increased sharply in October, while in No-till Atra-
zine plots they did not appear to recover from tillage.

The Mesostigmata were affected by tillage and Atrazine.
Significantly fewer (p 0.05) Mesostigmata were found in
tilled plots than grass control-plots on May 15 and 31
(Figure 12). In grass control plots three population peaks
‘were observed. These occurred in late May, mid-August, and

Inid-October. Inito-till control plots a single peak in

41
abundance occurred during mid-August. The population fluc-

tuated in No-till Atrazine plots, but no peaks are evident.
The Rhodacaridae were also affected by tillage and Atrazine
(Figure 13). Unlike the Mesostigmata, the Rhodacaridae
recovered in No-till control plots and then followed a trend
similar to those in grass control plots. The Rhodacaridae
responded favorably to Atrazine. Although fluctuations are
seen in No-till Atrazine plots, they are not as intense as
those in grass control and No-till control plots (Figure 13).

The Astigmata were affected by tillage. Significantly
more (p 0.05) Astigmata were found in uo-till control plots
than No-till Atrazine and grass control plots (Figure 14).
The latter two treatments differed significantly only on
September 8.
Age78tructure

An analysis of the effect of Atrazine and.No-ti11 farm-
ing on the population age structure of Brachystomella parvula
was performed. Because no measure of when B; parvula reaches
maturity was available in the literature and attempts to rear
them in the laboratory to obtain this information failed,
it was assumed that members of the smallest size class devoid
of pigment were juveniles. Qualitatively, juvenile 8:. ~
parvula population levels in each treatment were negligable
until July 25 (Figure 15A). On this date an increased number
of juveniles were collected from grass control plots, follow-
ed by No-till control and No-till Atrazine plots. After
July 25 the number of juveniles found in each treatment re-

turned to the previously low levels. Statistically, No-till

42

Figure 13. Mean Population Levels of Rhodacaridae

43

s ‘ Rhodacaridae 3

 

 

 

c
e ' '
i .f, /\” 1 \ //
. v -/ V ' /' '\
g / ; \/
a \ -/ Ill ;\ I 4
\ ,J/ ,' . ,
I, t m [I
I ‘ ‘ I
II ‘ «
2 < . - ‘ ’
or ’ ‘ ,
‘ ’ , .4
\v ’ I ’
I 1
—-—- Control
a- ----- tto- till Control
--—-- lio— tiil Atrazine
4-17 s-e sols 5-31 4-14 74 7:25 the 9:4 elze lolla Illa

Sampling Dates

Figure 13.

44

Figure 14. Mean Population Levels of Astigmata

45

 

 

 

..... 4
n'r ‘
I i
\
3. Astigmata I l
, l
24
C
C
e
a
2
a
E
C
m
‘ a
—— Control
.. ..... uo-ml Control
-—-— Mo-tiit Atrazine
4-17 s-asiis 5:31 41:4 753 7125 Me Hi 9-2a 10-18 11-a

Sampling Dates

Figure 14.

46

Figure 15A. Mean Population Levels of Juvenile Brachystomella
parvula

Figure 15B. Percentage of Juveniles in Brachystomella parvula
population

47

‘4 — Control
0- ----- lto till Control
-—-— Io till Atrazine

‘1

Sample Mean

 

 

 

 

Figure 15A.

 

 

 

1.4
.
3
'5 a.
’
o
a
a
c
e
3 4‘
e
o.
447 5:05;” 5131 4'44 714 7:25 034 934 912: loin "Te

Sampling Datea

Figure 15B.

48
control and No-till Atrazine populations differed (p 0.05)
only on July 8 and 25. Significantly more (p 0.05) juven-
iles were found in grass control plots than no-till control
and No-till Atrazine plots on July 25.

Juveniles were expressed as a percentage of the total
population (Figure 15B). No-till control and No-till Atra-
zine populations had similar trends. Each had peaks occur-
ring in mid-June and late July. During mid-June and late
July juveniles constituted a larger portion of the population
in No-till Atrazine plots. After late July the porportion
of the juveniles accounted for less than 1% of the §;_parvula
population in No-till control and No-till Atrazine plots.
The percentage of juveniles in grass control plots rose and
declined three times at eight week intervals beginning in
late May. The influx of juveniles in grass control plots
occurred two weeks earlier than in tilled plots. The second
influx in grass control plots coincided with that of the '
tilled plots. The third influx occurred in late September.

These data suggest that juvenile B; parvula were
affected by tillage but not Atrazine. Significantly fewer
(p 0.05) juveniles were found in tilled plots than grass
control plots. Although No-till control and No-till Atra-
zine populations differed significantly on selected dates
overall the populations did not differ significantly
(p 0.05). Furthermore, when the number of juveniles was
expressed as a percentage of the total population, No-till
control andtlo-till Atrazine populations followed similar

trends while grass control populations followed a different

49

one .

CONCLUSION

No-tillage significantly reduced Collembola, Mesostig-
matid, and Prostigmatid mite pOpulations. Only epigeic
Collembola were directly affected by No-tillage. Alterations
in Edaphic Collembola, Prostigmata, and Mesostigmata poPue
lations were the result of changes in the soil microenviron-
ment. Astigmata and Cryptostigmata papulations increased
after tillage. These result are in agreement with Edwards
and Lofty (1975).

Atrazine directly affected only the Prostigmata. Fewer
Prostigmata were collected from Atrazine treatediplots:after
the herbicide was applied and for the remainder of the study.
Other changes in arthropod populations occurred after surface
vegetation had died and corn had emerged. This suggested
that habitat alterations caused by Atrazine and not direct
toxicity were involved. Similar conclusions were drawn by
Curry (1970).

The population age structure of §L_parvula was altered
by No-tillage. Fewer juvenile E; parvula were collected from
up-till control and uo-till Atrazine plots than grass control
plots. It was difficult to accurately assess the impact
of the alteration since conclusions'would have been based
on data collected from one growing season. However, if this
trend were to continue, fewer E; parvula would reach maturity

to reproduce in No-till plots.

50
RECOMMENDATIONS

This study and others performed at Michigan State Uni-
versity have determined that NOFtillage and herbicides
affect arthropod populations to varying degrees. Further-
more, with the implimentation of more SOphisticated statis-~
tical analyses we are now able to determine when during the
growing season arthropods are affected by different treat-
ments. If the following recommendations are executed, a
better understanding of how herbicides affect these animals
and the consequences of the effects could be obtained.

1) Before the soil is tilled or treated with herbicides the
sampling area should be defined. This would enable research-
ers to determine immediate treatment effects. As it was, in
this study, no definitive conclusions could be drawn on the
initial disturbances.

2) More environmental parameters should be measured. If
done frequently, a regression could be developed to determine
the relative importance of each parameter.

3) The soil should be analyzed periodically for herbicide
concentration. A relationship between arthropod population
levels and herbicide concentration could be established.

4) Continue this project for many growing seasons and fole
1ow community succession. Huhta (1979) has provided a sum-
mary of the effectiveness of a few indiCes of succession.

By following community succession, changes in species com-

postion could be monitored.

APPENDICES

Appendix A. Checklist of Arthropods Collected From No-till
Control Plots

51

 

 

 

 

mom mH me Hm mm em mm SH
«H H m N o m H H
o o o o o o o 0
NH H H m o m H o
How on mm mm hm mm om mm
H o H o o o o o
OHom mm hmH mHH mm eom NHNH mum
«um mm mm mm mm mm mm mm
m mH mm m pH mm m
Houoe >02 uoo ummm ummm 05¢ >Hoh sth o

 

 

 

mH

mvm

«H
can

hm

How

 

HH oucoHoH> oHHochoooomm

 

H ouoHoooxom mHHochoomSmm

 

o moooonHoH> mouH>00UHmoH

 

o moxooouom mouu»00pHmoH

 

m mooHHHmm mouHNooonoH

 

o ouoHOmMMHuHoE chunosoucm

mmowmuoosoucm

 

mm oHobncm oHHmEoumwnooum
oopHHHoEoumazomum

dqomZMHHOU

 

 

 

om mmqmzdm ho mmmSDz

3
as: means oquezam

 

 

 

mEOHm HomBZOU mmflmw 20mm QMBUMHHOU mDOmOmmem< m0 BmHHMUmmU

4 NHDZWQQfi

52

OHN
hm
mom

Hm

mmm

50H

mm

mm

omm

NvH

NH

mm

Hv

me

vH

me

0N

mm

mH

vH

Hm

mH

HH

mH

Hm

mm

mm

vm

mm

mm

mm

mm

mm

av

mH

mH

mm

mm

om

NH

hm

VH

hm

HHH

oH

ov

HH

Hm

 

mm nHHHsse naeennmamn

 

5H .mm mocHuoBuCHEm

 

mOH mammoHo mscHHorucHEm

 

m snowman mononucHEmouousmo

 

o .mm mouHHomoruud

mneenseeeesm

 

Hm ouoHocon onHmnHHoB

 

m mouomuooco mouoHsoado

omofluoflnomco

 

e monocHE msHooz

mooHHooz

 

o mHoHHH> oEou0mH

 

vH mHHHnmuoc oEouomH

wooHeouOmH

 

H mHHoHHnscoE ousuumomomww

mooHHouumnmomkm

 

 

AQOSCHUCOOV 4 xHszmm<

53

mvm

NmN
mm
Hmm
mmN
mumN

mmN
NHm

mH

NHH

NH

0H
NH

moH

me
o

hm

50H

NNH
Nm
mhh

«NH
HvH
o

mm

on
om

Hom

v0

mm

HH

«H

NH

mm

 

ON

mN
He
no
vv
HNN

mm

Nm mH m v

H N o H

o o o 0
NH N 5 0H
m H H H
Hm N v NNH

mm vN h h

NmN mu va mom
m o o o
N o H N
HH H N mN

HOH NM mv No

m mnonoxe

o mneHHeHQEone

o mooHno>couuoB

mH wooHEocomuoB
e mneemnsmeem
mNH mooHuoomusom
mm mneeeeeenem
HMN omoHuoEmNm
H mneHmHnaunme
m mneHaoHHsreomz
NN mooHumonouocoz
mm mneHeoesm
N mneHHHmem

aaazoeemome

€2HM<U¢

 

AemaeHucoov a xHozmmma

54

MNv
mwm
NHH
mmH
Hem

mH

mm

mON

Hmm

Hv

Nv

mH
mH

NN

mh

we
HOH

mN
Nv

omN

mH
vN
HH
5N
mm

we

HNH

HH

me
mm
NH
vN
Nm

HH
MH

N¢H

mm
mH
NN
Nm
Hh

mH

mm

 

hm

hH

hv

0H

mN

mH

«H

mmH
ow
NH

OH

OH

mN

mN

HomscHueoo. < xHozmaa<

mN ounEmHumomoz ousuoEEH

an mnoHuooooorm
HN mooHHmmou>sm
«H moonoHooH
MH mooHom<

dadonBmOmmz

H camEmHumouQ>HU muoumEEH
m mmoHonmooouoma
m mcoHHoumoHHo
mOH mooHHmo
H mooHuoooHnnumom
H mooHccmaroHHmm ousuoEEH
OH mnoHccoErOHHmu

NadszBmOBmme

H ouoEmHumoum onouoEEH

 

55

hMH
Nm
mOH
hm
oH

HmN

Nmm
Hv

«H

vH
HVH
mm

NH

HH

mm

mm

 

VH

OH

NH

om

mm

wH

MH

HH

mm

em

Nm

0H

MH

mm

mm

mm

 

AemscHucoov a xHozmmma

MH

NN

Hv

me

mH

mN

pH

5N

mN

oH

5N

Hv

flmmamozmzwm
flmmBmozom
m<>m<q <mMBmHQ
md>m¢H dmmEmOMHOU
dmmamomqoo
dmbfiomm
4mDHmHQ
daummZH
dqwmmzwm
<00m0m94m
NOOHQHQ
doomOHHmU
<m¢z<m<

¢B¢20H8m4

nunsmHunonmz emHuHocmeHca

56

Nm

bHv

mwN

HH

NN

Hv

om

0H

0H

HN

oN

vm

Nv

em mm mv

o o o
o o o
m o o

HN VH HN

mm

mm

mm

dnomOmH
NMUNBMDmU
dmmemOZNmMmB
fimmBmOUmm
dmmBmomBmo

m<>m¢q <mMBmOQOmmH

onoHoHEuom

 

 

 

AemseHucoov « xHozmmma

Appendix B. Checklist of Arthropods Collected From No-till
Control Plots

57

 

mH

 

 

 

 

 

 

 

 

 

 

 

mNH m NH mH mN «H cH N « m mucoHoH> oHHochoooomm

« o o o c N H o H o o oumHsooxmm oHHmchooosmm

H o o o o o H o o o o nsomonHoH> msumnooeHmmH

m H H o o o N H o o o moxooouom moumnooonoq

omH e H Hm e eH oe mH eH m e nseHHHem neusnooeHmmH

m o o o o o m o o o o nunHonanuHsa nannosoucm

mooHNHQOEoucm

«mHH NH OH «H m« mm mm« mHN th on mm oHs>unm oHHoEoummnooum

mooHHHoeoumwsocum

<Hom2mHHOU

th mN mN mN NN NN mN mN ON on mN mmHmz<m mo mmmzaz
m mH mN m mH mN m «H Hm mH

Houoe >02 poo umom umom ms< NHsn NHDh mosh an: an: mmefia oqumz<m

 

 

 

 

mBOHm HomBZOU HHHBIOZ 20mm QMBUWHHOU mQOmommBmd mo BWHHXUmmU

m xHazmmmd

58

ONH
HvH

Nm

N«N

mN

mH

NHH

HNH

mm
mm

m«

mH

m«

 

h«

HH

H«

AUODGHUCOOV m xHQmemd

 

.mm mocHuocucHEm

 

mcommHo mocHuosucHEm

 

ouommou msusrucHEmoumuooa

 

.mm mouHHomonuu<

mneHnsruaHsm

 

ouoHscon nHmumnHHoa

 

mounmwoocm mouoHnowco

mmoHunHromco

 

monocHE monmz

mneHHmmz

 

mHoHHH> o80u0mH

 

mHHHnouoc oEou0mH

mooHEOHOmH

 

mHHMHHnocoE ououummmomhm

wooHuouumomomam

 

59

OON
O«

mn«
mmN

NNNN

HON

Hm

O«

mm

«OH

mh

N«
OH
NNH
«H

HHm

N«
HOH

«N

mm

«N

OMN

HO
mm

OOH

mN

MN

mo

h«

mmH

hm

hm
«H
mm
mm

HH«

HH
Hm

NOH

ON

O«H

mH

OO

MH

mm

HO

MH

m«N

mH

mH

«N

ON

mm

 

 

HOOSCHUCOOV m XHQmem<

oopHHoHoeouB
wooHnoacouuoe
mMOHEocomuoB
mmeHmmsmHum
mooHumoouoom
mnoHHeHmnam
wooHpofimwm
mooHoHnnucom
mneHnoHHandomz
ooOHummronocoz
mooHoomsm
mooHHHoom
<B¢ZOHBmomm

<2HM¢U<

 

mHHHEom nHoHuoommm

60

MON
Om
MHH

OOH

mH
mO
OOH
ONHH

OOH

MN

NHM

OH

OH

ON

ON
NO

ON

ObH

OM

MN
5N

MH
OH
ONH

MN
OH

OH

OH
NN
NHH

OH

«M

ON

OM

OH

NN

OOH

H«

OH

OO

«H

OH
NM

HNH

OH

OH

N«

OH

OH

H«

NH

OON

MH

NH

OO

OH
NO

ON

«H

 

HUODCHUCOOV m XHDmemfl

mopHHoooponm
mooHHmm0u>nm

moonoHooH

oopH0m<

«deoHBmommz
muoEOHumouQNHU manuoEEH
manmrmmoouooe
oooHHouonHuo

omoHHoo

oooHHoooHHnumsm
oopHccnenoHHmm ououoEEH
mooHcccfinoHHmm
daflszBmOBmme
ouoEmHumoum mHouoEEH

meeHmeae

 

61

MMH

hMH
OO
NM.

OO

«NN
ON
om
MN
«M
hMM
««H
OON

OH

MM

HH

ON

OH

OM

HO

HO

OH

 

Hm

NH

ON

ON

m«

HN

HH

NN

«m

Hm
5N

MO

OH
Oh
HH

MN

MH

HN

«H

M«
MH

OM

MH

ON

OM

HH

 

ApmscHueooO m xHozmmm<

NH

OmmBmOZOm

m<>m<H (mmBmHQ

m<>m<H fimmBmOMHOU

dmmBOmOMHOU

dmseomm

deHmHD

dBUMmZH

<memzwm

<Qom0maflm

daomOHmHD

dQOQOHHmU

¢m42¢m<

<B<ZUHBO€

nucEOHumomoz oonHucopHcO

oumEOHumomoz ououoEEH

 

62

OO

m«M

«OH

000

OH NH

 

NN

SN

Om M« NO HO
O O O O
O O O O
O O O O

ON

OH

NDOmOmH

¢m0<9m3m0
<mmem02¢m>mfi
fimMBmOUmm
flmmamomfimo

m<>m4H deBmOQHmmH
ooOHOHEH02

<mm9mozmzwm

 

 

HGODGHUCOOV m XHQZNQQN

Appendix C. Checklist of Arthropods Collected From No-till
Atrazine Plots.

63

MO

O«

HNOH

 

 

 

 

 

 

 

 

 

 

 

NON

Hence

 

m m mH m mm oH OH N m « muemHoH> MHHmaHmoememe

o o o o m o o H H o eumHsooxmn eHHmchoomsma

o o o o e o o o o o nsomonHoHe nsaauooeHmmH

O O O O O H O O O O moxooouom msuwuooonoq

o o H H e on N mH m m nseHHHnm naewnooeHemH

o o o o o o o o e o nunHonanuHss nannosouam

mmonunoeoucm

mH m OH om eeH ems omm omH en es an>enm nHHmeosmmeonum

ooOHHHwaoumhnomum

«HomzmHHoo

mm em em mm em mm ON on on em mmHazam no mmmzaz
m mH mm O OH mm m «H Hm mH

>oz poo ummm seem msa NHsn NHsn mean an: an: mme<o ozHHmzem

 

mBOHm mZHudmefl HHHBIOZ 20mm QmBUmHHOU mQOmommBmd m0 BmHHMUmmU

U XHQmem<

64

Oh
OON
MN

Hm«

NNH

OO

HN

NO

OOH

OO

MH

ON

OH

OO

M«

OH

OH

ON

ON

ON

m« m«

ON O

NM

mMH

ON

mm

 

 

ON

NH

«N

NH

NN

OH
Mm
b

O

MN

 

om .mm nacHnsaueHEm

 

m« mcoOmHo mocHuonucHEm

 

OH ouommou mononucHEmonouooo

 

O . .mm mouHHomonqu

mneHnsneaHsm

 

MN mumHoccuO oHOHooHHoe

 

O monomuooco msustowco

mooHuonoaco

 

m mHHHHHCflE mDHOOZ

omoHHooz

 

O mHOHHH> nEOOOmH

 

MH mHHHnouoc MEOQOmH

onoHfiouomH

 

N mHHoHHnscoE ououuuomommm

mooHHoHumomommm

 

AUODCHUCOOV U xHszmm¢

65

«ON
OON
MOH
OOH

OOOH

O«
«ON

ONH

N«

MN

ON

OO

«N

O«
ON
OM
MN

HOH

OO

ON

OO

HN

OOH:

MM

OH

HM

O«
OM
OH
OM

me

HH

O O O O
O O O O
O N O H
O H O O
M O OH M

OH OH O O

«H OO OO OON

O O O H
O O O O
O « O O
O HH MH HM
O O H O
«O « O OH

OH

OOH

OH

OOH

OH

moOHHOHoEOHB
oooHrowcmuumB
mopHEOGOmHoB
manooEOHum
mopHHooouoom
mneHHeHmnam
wooHuosmOm
mmonHonucmm
mneHnoHHmamomz
mooHumorouocoz
mnoHo0dsm
ooOHHHoom
dadszamomm

dZHm¢U¢

 

mHHHeom oHOHHoonmm

 

 

 

HomseHueooO o xHozmmma

66

OO

O«
OO
ON

«O

OO
OH
NM
ON
OH
«OH

MOH

ON

NH

NH

NH

OO

OH

OH

HH

O « M O M
O O O O O
O « O HN M
H O H O OH
«H N H N O
«H H « «H NN
« N O HH O
M H O O O
M O O « N
N H H O O
H M M H N
OM M HN O O
ON O OH H O

 

HN

fimmemozom
m¢>m€H dmmBmHD
m¢>m<H 4mmBmOMHOU
<mmBmOMHOU
deeomm
GmDHmHD
NBUMmZH
<memzwm
NDOmOmbflm
<DOmOHmHQ
GQOQOHHEU
¢m¢2<m4

<B¢20H8m<

nunEOHumOmmz ououoEEH

 

 

AGODCHUGOOV U xHQmemd

67

«H

OOH

OON

OH

NN

OH

«H

O H O
O ON ON
O O O
O O O
M O O

MH MN ON

NO

ON

OO

OO

NDOmOmH

«WUNBmDmU
<mmam02<m>m9
flmmemoomm
4mmBmOEBmO

m¢>¢¢H (mmBmOQHQMH
mooHoHEuom

dmmfimozmzwm

 

HeoscHucooO o xHozmmm¢

 

 

Appendix D. Means and 95% Confidence Intervals for Arthropods
analysed.

68

or» up Houucou mmouu or» ECHO

 

 

 

 

«a

 

«a

.nHm>mH NHHHHnnnone Haas mo.o can H.O eH.o

 

 

«m.H m~.H mm.H e~.H e~.H e~.H e~.H
mo.~ om.H we.H mm.H «e.m me.OH oo.m
. «a
«~.H e~.H e~.H «m.H e~.H e~.H «~.H
Ne.~ «~.m ~a.~ oo.m oe.m eo.a ee.m
«~.H «~.H e~.H e~.H m~.H e~.H e~.H
ee.H «e.~ mm.H ne.H mm.e ~e.m mm.~
m mH mm O OH mm m
>oz poo omen seem one HHse NHen
H<>mmeze

D XHQmem<

an

an

 

moocmuomme ucoonHcOHm OHHooHumHuoum ouocoo mmenoumo one

 

 

 

 

 

 

ee.H e~.H m~.H +
am.~ me.« om.e new:
mzHNamea HHHenoz

an «a I

H«.H e~.H a~.H +
OO.« O0.0 O«.m coo:
Homezoo HHHeaoz

ee.H ee.H m~.H H
eo.« No.5 eo.e new:
Homezoo mmamo

«H Hm mH

mean On: an: mmeaa ozHHmzam

MUZEOQOZOU.OOO QZN MHm2€m mmm mfionDmEZHZm ho mmmZDz 24m:

69

.mHm>mH OuHHHnmnoum Harv O0.0 can Hrv OH.O
on» um Honucou mmouo may Eoum mmocmnmmmHo acconHcOHm OHHMOHumHuoum muocoo mmeuoumo one

 

 

 

 

 

 

 

 

 

 

 

«a «a an an I
«m. mm. mm. em. em. we. «a. am. we. em. +
Ho.H mm. mm. mm. mH.H em.m mm.H eH.H ee.~ OH.~ new:

mzHeamea HHHeioz

«a «a l

«m. «m. an. am. we. em. «m. mm. mm. em. +
me. He. we. on. em. mo.~ o«.~ mm.H mm.~ em.H new:
Homezoo HHHeioz

mm. «m. «m. «m. em. «m. «m. am. me. mm. H
em. we. me. me. «a. «e.H aH.H eH.~ me.« mH.e new:
Homezoo mmamo

m «H mm m eH mm. m «H Hm mH

>oz poo seem ummm O54 NHse HHsn page an: an: mmeaa ozHHazam

 

 

H¢>MMBZH mUZMDHmZOU wOO 02¢ MHmzdm mmm mZOQNHm mDZHmDmBZHZm ho mmmZDZ Z<m2

AGODCHHCOOO Q xHozmmm<

70

.Hm>wa muflaflnmnoum A«*V mo.o can A«v OH.O

ms» um Houucou mmmuw ms» Eoum mmocmumuwwo acmOwMficmfim madmowumflumum muocmo mxmwumumm 0:9

 

 

 

 

 

 

 

 

«« «i I

mm.o mm.m um.o em.m em.o hm.e mm.m oH.m ma.m no.5 +
~m.m H5.m mm.m ¢~.n mH.¢H 5H.mm pm.na ~m.~a Hm.m e~.¢ cum:
mzHN¢m9< quauoz

«« «« at l

wm.m em.m «m.o em.o hm.m mm.o «m.m oH.m mm.n oH.~ +
mo.oH mo.HH NH.HH ¢~.ma mm.~a om.om ~H.«H ~m.na mm.¢ mo.m cam:
aomezoo aqHauoz

¢m.o mm.e mm.m mm.o ~H.~ mm.o mm.o oH.m mm.» m~.m H
oo.oH mm.HH ~v.m mm.m ~m.m~ mm.om mm.m mm.a~ hm.oa mn.m cam:
qomazou mmmmw

m ma mm m ma mm m «a an ma

>02 poo ummm ummm msa wash xasn mean an: an: mme<o wqumz<m

 

A<>mmBZH mUZmloZOU wmm 92d mum2¢m mam dqomzmdaou ho mmmZDZ z¢m2

Aomscflucoov o xHozmmm<

71

 

man an Houucou mmmuw may scum

.mHm>mH auflflflnmnoum A««. mo.o can ««v OH.O

mmocmumMMAc unmowmwcmfim waamowumwumum ouocmn mxmwumumm mne

 

««

««

 

««

 

 

 

 

 

mm.¢ ~m.¢ ~m.v mm.v mq.q ~m.q mm.¢ v~.m m¢.q o¢.¢ +
oo.o mm.m -.m m~.m mm.oa oo.mm mo.ma mm.HH sh.m hm.v cum:
mzHNmmea queuoz

« «a «a «a I

mm.v mm.¢ mm.¢ mm.v mq.q mm.q mm.q ma.m mq.v Hm.¢ +
mm.m m~.m ~e.m mm.m ~m.oH m~.m~ an.mH pm.oa mm.m om.m cam:
nomezoo aqunoz

mm.q mm.q mm.¢ mm.v mm.¢ mm.¢ mm.v mH.m oo.m om.¢ H
~o.m m¢.m on.m He.m m>.H~ qu.mv av.oa om.mH mm.oa o~.o cum:
aomazou mm<mw

m «H mm a ma mm m «a an ma

>02 poo mmmm ummm ms¢ wash wash mean >mz an: mmaco wqum2¢m

II
II
II
I
II
II
|
l
I
I

 

A¢>mmBZH NUZWQHhZOU mmm 02¢ Hamidm mmm ¢AD>m4m dAAmSOBmMmodmm ho mmmZDZ z<m2

Aomscflucouv a xHozmmmm

72

 

«4

.mam>ma wuflflflnmnoum A««V mo.o can A«v oa.o
0:» pm Houucoo mmmuo may Scum mmocmumMMfio pancamwcmwm haamoflumfiumum muocwc mxmflumumm was

 

a;

 

 

 

 

 

 

mm. pm. pm. mm. hm. hm. ma. ~H.H hm. ma. +
Ho.H mm. on. mp. v~.H mo.H mm. mm.a mm. o~.H ammz
f . mzHN<m9¢ aqunoz

. a; a; I

mm. mm. mm. mm. hm. mm. ma. ~H.H hm. am. +
an. as. Hm.H -.H on. om. om. am. me. am. cams
aomezou queaoz

hm. m¢. ma. mm. mm. mm. mm. NH.H HH.H «m. H
Hm. mh.a -.H on. Hm.~ o~.~ -.H vm.q mm.a Hm. cam:
nomezou mm¢mu

m ma mm a ma mm m «a an ma

>oz poo ummm ummm ms< wash >Hsn mcsn am: an: mmado wzHamz<m

 

A<>mWBZH MUZWQHEZOU mmm

 

AUmSCHuGOOV Q anzmmm¢

OZfl mam2¢m mam mHAHm¢BOZ <ZOBOmH ho mmmzbz Z¢WZ

73

.mam>ma aufiaflnmnoum A««v mo.o can A«V OH.O
may um Houucou mmmuw on» EOHM mmocmummmww unwowMficmwm maawo«umfiumum muocmc mxmwumumm one

 

 

 

“l""l'|'ll'U "|"|‘

 

 

 

 

 

 

 

«« «a «t 1* «t I

«a. mm. mm. em. mm. mm. em. mm. mm. mm. . +
mm. mm. om. om. mm. om.a no. H~.H Hm. mm. cam:
mszmmem nganoz

«« ** 1* 4* I

«o. «m. co. «0. mm. «m. we. mu. mm. so. +.
mm.a mH.H «m.a ¢~.H hm.a mm.~ mm.H mm.a mH.H mH.H cam:
nomezou AAHeuoz

mo. em. we. we. no. em. em. on. mu. «m. H
ov.H mm.m mm.a hn.m m~.q mm.m m¢.H mm.a am. am. cum:
qomazoo mm<mw

m ‘mH mm a ma mm m «H an ma

>oz poo ummm ummm mam masn wash mcsn an: an: mmaaa oqumzcm

 

M
I
I
I
I

 

A<>mmBZH HUZMOHmZOU wmm 92¢ mamim mmm mDQqudm mDBmMUOQHm—MA .mO mum—:52 Zfimz

Aumacfiuaoov Q xHozmmm4

74

.mHm>mH aufiaflnmnoum A*«V mc.o can AIV OH.O

 

way an Houucoo mmmuu 0:» scum mmozwummmwn unmoflwficmwm adamoHHMflumum muonmv mxmflumumm was

 

*«

*i

*k.

 

 

 

 

 

mo.H mo.H mo.a no.a mo.a mo.a mo.H m~.H ma.H oa.a +
mm.m «m.¢ em.m m~.~ om.~ «m.a m~.H Ho.H om.a om.H cam:
mzHN¢ms< aquuoz

SA SA SA 84 mo; 84 84 mo; 34 :4 H
mo.m o~.m mm.H mv.~ e~.~ m¢.~ mo.a Hm. mm.a wo.a :mmz
aomezoo queuoz

mo.H no.H no.H no.H HH.H no.H no.H mm.H ¢~.H mo.H H
H>.H mm.~ mo.~ hm.a mm.~ om.~ mn.~ hm.H m~.~ mm.~ cam:
aomezou mm¢mw

m ma mm m ma m~ m «a an ma

>oz poo pamm ummm mam masn mash wash an: an: mmeco quqmz¢m

 

A£>mmBZH mozmn—HKZOU wmm 02¢ mqmfim mmm £95520 <memmAADB .mO mmmzbz 24m:

Acmscfiucoov a xHozmmm¢

75

.me>wH hUHHHQMQOHQ Acyv mo.o can A*V OH.O
map um Honucou mmwuw mnu Scum mmocmummmwo unmowmficmwm madmowumwumum muocmc mxmwumumm was

 

 

 

 

 

 

 

 

 

 

«« «a «a «a t I

ma.” nm.~ hm.n «5.5 ca.n no.» ~m.» m~.a mm.» wo.m +
ma.e~ H~.m~ mo.e~ om.vH Hg.mH Ha.ma m~.oH mm.>a ~F.H~ Hm.o~ cum:
mzHN¢m9¢ ngBIQZ

«« ¥a . «4 I

no.5 mm.n mm.> em.> om.> «a.» mm.> m~.m ma.~ ma.m +
mm.¢m mo.m¢ mo.om ~m.m~ mm.>~ mm.om ~¢.H~ mm.o~ mm.n~ mn.ma cam:
aomazou quauoz

om.» mm.n ~m.> mm.» va.m Na.“ ~m.~ o~.m oa.m mp.“ H
HH.m~ ma.~m -.~q Hm.ma mm.am HH.vm m~.¢a mm.m~ mm.mm Hm.om cam:
aomezoo mm<mw

a ma mm a ma mm m «a an ma

>oz uoo ummm ummm ms< wash mash mean am: am: mma¢o ozHamzcm

 

 

A¢>mmBZH MUZmQHhZOU wmm 02¢ QO2<m mmm dBfiZUHBmOmm m0 mmmZDZ dez

.omscflucoov a xHozmmmm

76

man an Houucou mmmuw may 502m

.mam>02 20222000000 2«*v mo.o 000 2«V OH.O
mmocmHmMMflc unmoflMMcmflm >HamowumfluMUm muocmc mxmfiumumm was

 

 

 

 

 

#2 «2 «« «0 I

00.0 00.0 00.0 20.0 m¢.q 00.0 ov.¢ H~.m 00.0 mm.v +
ho.m mo.~H «H.~H m>.m ~m.u 00.0 mm.m nv.m >0.ma mm.~H 000:
22HN2292 aquuoz

fit. flan . I

20.0 20.0 «0.0 20.0 «v.0 00.0 20.0 H~.m 00.0 nm.¢ +
mo.qa mm.m~ Hmuoa oo.m~ om.ma mm.m~ ~0.~H H~.HH no.0H pm.m 000:
2029200 aquuoz

me.¢ 00.0 ov.e 00.0 mm.v 00.0 ov.q NH.m NH.m mm.v H
mm.HH ~m.mm mm.m~ >«.m Hm.mH >¢.ma «p.02 q>.ma mn.ma m~.mH 0002
2029200 mmmmo

m ma m~ a ma mN m «H Hm ma

>02 000 000w 000m 00¢ 2200 2200 0:00 >02 20: mmaao wqumzmm

 

A4>KHBZH HUZMOHKZOU wmm 02¢ mam2<m mmm mdnHBOZHMm ho mmmZDZ Zflmz

Anmscflucoov o xHozmmm<

77

.mflm>02 20222000000 2««v mo.o 0:0 2.2 oa.o

way an Honucoo mmmuw ma» 502m mmocmuommwo unmoflmacwwm >HHMOfiumfiumum muocmc mxmwumumm mne

 

.1“

 

+
CMGS

 

 

 

he.m mm.m
¢m.v mv.¢
5v.m h¢.m
mm.m mN.vH
mm.m h¢.m
HN.m Nm.hH
m ma
>02 #00

 

 

22HN<KB< AAHBIOZ

+
c002

 

AOfiBZOU AAHBIOZ

+
:mmz

 

AOfiEZOU mm¢mw

mmfifln 02HAAZ¢M

 

 

A¢>mm92H WUZmQHEZOU wmm D24 mqmzdm 2mm dfidszBmOBmme mo mmmZDz 2¢m2

20000200002 0 xHozmmm<

78

1%

022 um 2020200 00020 0:» 8020

 

.020>02 20222000020 .000 20.0 000 200 02.0

00020202020 acmoww2cm2m 2220022020000 000200 020220000 029

 

 

 

 

 

20.2 20.2 20.2 20.2 20.2 20.2 20.2 20.2 00.2 20.2 +
00.0 22.2 20.2 00.0 22.0 20.0 20.0 20.0 22.2 22.2 0002
02200020 2222-02

«a 00 «0 |

20.2 20.2 20.2 20.2 20.2 20.2 20.2 20.2 20.2 02.2 +
00.0 00.0 02.0 02.0 20.0 00.22 20.2 00.2 02.0 02.2 0002
2002200 2222-02

20.2 20.2 20.2 20.2 02.2 20.2 20.2 20.2 00.2 00.2 H
02.0 02.02 02.2 20.0 02.0 20.0 00.0 20.0 22.0 00.0 0002
2002200 00000

0 02 02 0 02 22 0 02 20 22

>02 000 000m 000m 000 2200 2200 0000 202 202 00200 02220200

 

n

A¢>2292H muzmaHhZOU wmm Q2< Hamidm mmm wdoHHmO 20 mmmZDz 2<m2

20000200000 0 22020000

79

«0

20 2022200 00020 022 8022

00

.020>02 20222000000 2000 20.0 000 200 02.0

 

«0

«0

00020202220 22002222020 2220022022020 020200 020220200 022

 

 

 

 

 

 

20.2 00.2 00.2 00.2 00.2 00.2 20.2 00.0 00.2 00.2 +
20.2 00.2 22.2 02.0 00.0 00.0 00.0 00.0 02.0 00.0 0002
02220020 2222-02

«0 0 . «0 «a I

00.2 00.2 00.2 00.2 00.2 00.2 00.2 00.0 00.2 00.2 +
02.0 00.0 02.2 22.0 02.22 00.0 02.2 20.0 02.0 00.0 0002
2002200 2222-02

00.2 20.2 20.2 20.2 00.2 20.2 20.2 00.0 20.0 00.2 H
00.2 00.02 02.0 20.22 00.22 20.22 00.0 22.2 00.22 00.0 0002
2002200 00000

0 02 02 0 02 22 0 02 20 22

>02 000 0000 0000 000 2200 2200 0000 202 202 00200 02220200

 

 

 

A<>2282H MUZmOHWZOU wmm 02¢ mamz<m mmm dfidszfimOmmz mo mmmSDZ ZQMZ

20000200000 0 22220000

80

.020>02 20222000000 200. 20.0 000 200 02.0

022 20 2022200 00020 022 8022 00020202220 22002222020 2220022022020 020200 020220200 022

 

 

 

 

 

 

 

 

 

 

«0 «a 02 I

00.2 22.2 22.2 00.2 22.2 22.2 00.2 02.2 22.2 22.2 +
20.2 20.0 22.2 00.0 00.2 02.0 00.0 00.2 20.2 00.2 0002
02200220 2222-02

00.2 00.2 00.2 00.2 22.2 00.2 00.2 02.2 22.2 02.2 +
20.2 22.2 02.2 00.2 20.0 20.2 02.2 22.2 00.2 20.2 0002
2022200 2222-02

22.2 00.2 00.2 00.2 02.2 00.2 00.2 02.2 22.2 00.2 2
02.2 00.0 00.2 20.2 22.0 00.2 02.2 00. 22.2 20.0 0002
2022200 00020

0 02 02 0 02 22 0 02 20 22

>02 000 0000 0000 000 2200 2200 0000 202 202 00200 02220200

 

 

 

2¢>KNEZH MUZMDHmZOU wmm 02¢ m2m2¢m mum mdaHmdoénomm m0 mmmZDZ dez

2005:22200v Q xHozmmm<

81

.mam>ma huflHwnmnoum A*«v mo.o can Agv OH.O
man an Houucoo mmmuw ms» Eoum mmocmuwMMflc u:mo«wa:mflm >HHm0flumflumum muocmv mxmwnmumm was

 

 

 

 

an. om. cm. mu. om. am. an. mm. om. om.
mm. om.H om.~ Ho.H wh.a em. mH.H vb. mm. cm.

+
Cm”:

 

MZHN<MB¢ AAHBIOZ

«« «a
mm. mm. mm. ms. cm. an. an. mm. om» Hm.
mH.H am.H mN.N mm.H HH.m mc.m mm.H vb. NH.H ms.

+
CMGZ

 

AOMBZOU AAHBIOZ

cm. as. ms. mp. am. m5. m5. mm. Hm. ms.
nu. mm.H mm.a mN.H oo.~ mm.a os.a NN.H mm. mo.H

+
fimmz

 

AOfiBZOU mmdmw

 

a ma mm a ma mm a VA Hm ma

>oz poo ummm ummm ms< mash >Hsn mean an: >mz mma<a wqumz¢m

 

A¢>mmBZH WUZMQHEZOU wmm 02¢ mam2<m mmm deflszBmd m0 mmszz dez

Acmscfiucoov o xHozmmm<

 

82

.mam>ma apflaflnmnoua A«.V mo.o can A.v oa.o

may um Houucou mmmuw may Eoum mmocmummmww unmowMMcmwm waamoflumflumum muonmo mxmfluwumm mna

 

 

 

 

mm. mm. mm.
mm. mm. mm.

we. mme mm.
vm. em. vm.

mm. hm. mm.
oo.c oo.o vw.

 

 

 

 

 

 

 

 

m ma mm
>02 uoo ummm

«« I

oo. hm. mm. on. an. mm. mm. +
mm. mm. ~n. ma. mo.H mm. mm. :mm:
mzHN¢ma< quauoz

*« «« |

mm. mm. mm. we. an. mm. mm. +
hm. «w. ah.H Ho. mm. mm. we. saw:
qomszou quanoz

om. mw. we. oc.o mp. 55. mm. H
Ha. Ho.H m~.m oo.o om. mg. 5H. cam:
gomazoo mm«mw

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LITERATURE CITED

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