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LI B R A R Y
Michigan State
University

This is to certify that the

thesis entitled

Spatial Distribution of Aphis omi (Degeer)
and the Predator Aphidoletes Ap i imyza
(Rondani) Relative to Apple Tree Growth

 

presented by

Duane Paul J okinen

has been accepted towards fulfillment
of the requirements for

 

 

M. S . degree in Entomology

 

 

   

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Major pro e40):

Date (0/30 80
I I

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25¢?" rrdwne item

RE:URNING LIBRARY MTERIALS:

Wm“
Chm from circuaonjti ”nah

 

 

 

 

SPATIAL DISTRIBUTION OF APHIS POMI (DEGEER)
AND THE
PREDATOR APHIDOLETES APHIDIMYZA (RONDANI)
RELATIVE TO GROWTH IN THE APPLE TREE
BY

Duane Paul Jokinen-

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Entomology

1980

 

onnytw

ABSTRACT
SPATIAL DISTRIBUTION OF APHIS POMI (DEGEER) AND THE
PREDATOR APHIDOLETES APHIDIMYZA (RONDANI) RELATIVE
TO APPLE TREE GROWTH
BY

Duane Paul Jokinen

Development of an integrated pest management (IPM)
program for the apple aphid, Aphig 29ml requires knowledge
of host plant-pest-natural enemy relationships. In this
study, growth of the apple trees and colonization by A. omi

and the cecidomyid predator Aphidoletes aphidimyza were

 

measured on leaves by population surveys made in 1976 and
1977 in an orchard near Grand Rapids, Micigan. Stratified
samples of these two species from terminals in the tree
canopy (in three divisions: directional, level and inner—
outer) were taken weekly. Host tree growth measurements
including the number of terminal leaf primordia and leaves/
terminal were also taken from each sample quadrant.

Analysis of data, using analysis of variance and linear
regression indicated that g; pgmi responded to actively
growing terminals especially in the inner and upper quadrants
of the apple crown. The distribution and relative abundance
of A. aphidimyza, was closely coincidental with the aphid

population distribution in the apple tree.

Duane Paul Jokinen

Results of these experiments are discussed in relation
to the physiological characteristics of apple crown growth.

The potential role of g, aphidimyza in an IPM program for

 

the control of the apple aphid is discussed as well as

several potential research objectives for further studies.

ACKNOWLEDGEMENTS

I would like to thank Dr. Brian Croft, my Graduate
Adviser for his perserverance and his continual support
on this thesis project. I would also like to extend my
appreciation to the members of my Graduate Committee,
Drs. James E. Bath, Angus Howitt, Peter Murphy, and Mark
Whalon for their help in obtaining this degree.

I wish to acknowledge my peers, whose constant
interaction helped form my perception of the field of
Entomology and in particular Drs. Patrick Logan and Alan
Sawyer, Emmett Lampert, Frank Drummond, and Matt Michels.
This study was supported in part by a grant from the
National Science Foundation and the Environmental
Protection Agency. Financial support was also obtained
through the Department of Entomology at Michigan State
University.

Finally, I would like to acknowledge Drs. Dean Haynes
and Thomas Edens for their contributions. By providing a
source of intellectual challenge in a multitude of areas

they have contributed significantly to my development.

ii

Preface

The appendices included at the end of this thesis are

to facilitate the reconstruction of the data summaries

provided in this thesis as well as document other associated

research efforts initiated.on A. pomi and A. aphidimyza.

 

They were generated from computer files made of the
taken in 1976 and 1977. These files are maintained
magnetic tape at Michigan State University Computer
facility.

Throughout this thesis the term "apple canopy"
to describe the upper foliage area of an individual

However, "canopy" is correctly defined as the upper

data
on

laboratory

is used
tree.

most

‘foliage level of a forest, and in retrospect "crown" would_

be a more appropriate term to designate this area of the

tree.

iii

Table of Contents

 

 

 

 

Page

IntrOductiODOOOOOOOOO ..... O ..... ......OOOOOOOOOOOOO 1
Literature Review........... ................... .... 3

The Biology of Aphis pgmi....................... 3

Biological and Environmental Regulators of

Aphis pomi.................................... 7

The Biology of Aphidoletes aphidimyza........... ll
MethOdSOOOOOOOOOOOO. ..... .....OOOOOOOOOO... ..... 0.. 15

Sampling Procedures ............................. 15

Methods of Analysis.... ....... ...... ...... ...... 19
Results ....... . .................................... 20
Discussion......................................... 46
Appendix A. Weather records for Graham Experiment

Station, Grand Rapids, Michigan. April - August

1976 and April - September 1977....... ....... ... 50
Appendix B; Techniques utilized for establishing

laboratory colonies of Aphidoletes aphidimyza

(Rondani). ..... ..... ..... ....... ........... ..... 57
Appendix C. Frequency class distribution of Aphis

omi and Aphidoletes aphidimyza in 1976 and ‘

1977 ..... ‘ ............... . ....................... 61
Appendix D. Sample site means and variances for

1976 and 1977.......................... ......... 66
Appendix E. Aphid and predator biological

observations............................... ..... 89

iv

Table of Contents (Cont'd)

Page
Appendix F. Sample Size Estimation............... 102
Literature CitationSOOOO......OOOOOOOOOOOOOOOOOOO. 127

Table

2a

List of Tables

Spray schedule for Gibson experiment orchard
block at Graham Experiment Station, Grand
Rapids, Michigan, 1976 and 1977 ....... . .....

Results from the ANOVA for determining
differences in mean aphid densities in three
canopy divisions, l977........... ..... ......

Results from the ANOVA for determining
differences in mean predator densities in
three canopy divisions, 1977...... ..... .....

ANOVA results from testing for mean aphid
differences in the classes of Leaf Primordia
(0-4) and Leaf Classes (1-6), 1977 ..........

Results of modified - LSD tests for sample
dates with significant differences between
aphid densities in leaf primordia classes...

Results of modified — LSD tests for sample
dates with significant differences in aphid
densities in leaf classes.... ...............

Results of curve fitting for leaf primordia
decline and leaf addition: 1976 and 1977....

vi

16

32

41

42

43

44

45

List of Figures

Figure Page
la Leaf primordia mean densities in canopy

levels (1976).......... ..... ......... ..... ... 21
lb Leaf mean densities in canopy level (1976)... 22
lc Aphid mean densities in canopy level (1976).. 24
2a Leaf primordia mean densities in canopy

level (1977) ........ ......................... 25
2b Leaf primordia mean densities in canopy

position (1977).............................. 26

2c Leaf primordia mean densities in canopy

directions (1977)............................ 27
2d Leaf mean densities in canopy levels (1977).. 29
2e Leaf mean densities in canopy positions

(1977)....... ................... . ....... ..... 30
2f Leaf mean densities in canopy direction

(1977)Octcooooovoucovoooohoooooooo. ...... 0.0- 31
3a Aphid mean densities in canopy level (1977).. 34
3b Aphid mean densities in canopy position

(1977) ............... . ...... ................. 35
3c Aphid mean densities in canopy direction..... 36
3b Cecidomyiidae mean densities in canopy level

(1977) ........... ..... ...... . ............ .... 37
3e Cecidomyiidae mean densities in canopy

position (1977) ..................... ......... 38
3f Cecidomyiidae mean densities in canopy

direction......... ........................... 40

’ INTRODUCTION
Integrated pest management (IPM) systems for primary
and secondary pests of apple utilize a variety of control
tactics (Croft 1975, Huffaker and Croft 1976). Currently,
biological control of primary pests is limited (Hagley 1974)
and apple IPM systems use monitoring to predict activity of
these species. For example, adult male codling moth

(Laspereysia pomonella Linn.) activity can be assessed using

 

traps baited with synthetic sex phermones and adult apple

maggot (Rhagoletis ppmonella Walsh) population activity can

 

be assessed by bait and color attractants (Howitt and Conner
1964, Prokopy 1972). Population from these sampling tools
when coupled with computer based phenology models (e.g.
Riedl and Croft 1978, Welch et a1. 1978) provide forecasts
of critical temporal relationships between pest-plant stages
which assist in the implementation of control procedures.
As chemicals are the primary control method for these pests,
improved timing of tactics eliminates unnecessary sprays,
reduces costs and limits the amount of pesticides placed
into the environment.

Secondary pests of apple, primarily the foliar feeding
species such as Aphis pom; Degeer (the apple aphid) Panonychus—

ulmi (koch) (the European red mite), and Typholoyba pomaria

 

McA. (the white apple leafhopper), often reach damaging
population levels in the orchard. These species with

intrinsically high reproductive rates can develop resistance

1

2
rapidly to most conventional pesticides. They are, there-
fore, difficult to control by chemical means alone, and IPM
programs have stressed development of alternative cultural
or biological controls.

Use for control of natural enemies attacking secondary
pests becomes feasible when the pesticides applied for
control of primary (or direct) fruit pests are minimally
disruptive to these beneficial insects. This can be
accomplished by the use of physiologically selective com-
pounds or by minimizing the temporal and/or spatial contact
of pesticides with these natural enemies. Distribution
studies and sampling procedures for secondary pests and
their natural enemies provide population estimates from
which the potential for biological control or the need for
chemical control can be determined. In this study, the
spatial and temporal occurances of A. 29m; and the

cecidomydiid predator Aphidoletes aphidimyza relative to

 

growth features of the apple tree were investigated.

LITERATURE REVIEW

 

The Biology of ApAi§_pgmi

The biological characteristics of A. 29m; have been
reported in several early publications (i.e. Baker and
Turner 1916, Lathrop 1923 and 1928, Matheson 1919, Peterson
1918). The aphid overwinters as a diploid egg (black in
color 0.5mm in length), primarily on shoots or one year old
terminal growth in the apple canopy. Eclosion varies with
latitude from the first week in April in Virginia (Baker and
Turner 1916) to mid or late April in New York (Matheson
1919). Peterson (1916) correlated increased egg to nymph
survival with increased humidity at the time of eclosion.

In their study the average survival through egg hatch was
roughly 25%.

A. REA; has four nymphal instars, the morphology of
each is described in Baker and Turner (1916). The mature
fifth instar female is Wingless, parthenogenic, and vivi-
parous. This stem mother produces nymphs that as adults are
apterous (without wings) or alate (with wings). The ratio
of apterous to alate forms varies throughout the summer; the
environmental and biological factors regulating this process
in aphids has been discussed in Lees (1961) and Johnson
(1960, 1965, 1966). Alate production provides a ready
source of migrant aphids throughout the season within and

between orchards.

4

Lathrop (1923) estimated 5°C. (41°F) to be the develop-
ment threshold for A. pgmi. In 60 experiments during a
single season, he followed the development of aphid nymphs,
24 hours old or less, to maturity and recorded first nymphal
reproduction. From the max-min temperatures reported by
Lathrop (1923), and using the method of Baskerville and Emin
(1969) for calculating cummulative degree days, a mean
generation development time is 230.0 degree days (Jokinen,
unpublished analysis). Adult longevity was estimated to be
19.4 days (Standard Error = 4.6) for aphids in the second to
12th generations (Matheson 1919). The mean daily reproduction
capability is 2.96 nymphs for the same generations (Matheson
1919). A total of 12-17 generations were observed in New
York (Matheson 1919).

In late September haploid (X0) males and diploid females
(XX) are produced in the orchards. Following mating ovi-
position of overwintering eggs occurs about 5 to 7 days
later. Little information has been collected on the longevity
of these adult forms, however, one early study (Baker and
Turner 1916) indicated that it is ca. 25 days. Specht
(personal comm.) noted the survival of both sexes of the
aphid until November in Nova Scotia. Baker and Turner
(1916) report a mean of 4.75 eggs laid per female. Westigard
and Madsen (1964) reported that 90.9% of all eggs observed
_on 40 terminals were laid in the top 20.0 inches of the

terminals.

5

Nutritional and varietal differences of the apple host
in relation to infestations of A. REA; have been described.
Takeda et. a1. (1968) reported on the nutritional quality of
leaves infested by A. 222$! and noted that leaves with high
nitrogen content, low carbohydrate levels, and young leaves
having particular amino acid contents were the most sus-
ceptible to aphid attack. Briggs and Alston (1968) noted
resistance to A. 29m; in the apple cultivars McIntosh,
Baldwin, Northern Spy and Courtland. Wildbolz (1964)
suggested that resistance or susceptibility in apple cultivars
to A. 22m; may be related to the changing practices of
pruning and fertilization which influence growth patterns.

He suggested that temporal differences between aphid egg
eclosion in the spring and varietal variation in the initiation
of tree growth could influence population levels.

Specht (1970, 1972) studied A. 22m; populations on one
year old trees in environmental cabinets. Growth and decline
of aphid numbers correlated well with the growth activity of
the trees. The distribution of aphids on terminal leaves
varied throughout the study which he hypothesized was a
result of the changing physiological condition of the
leaves. Westigard and Madsen (1965) in studying three and
ten year old apple trees in California, noted that the
physiological condition of the tree foliage was a major

factor in influencing aphid densities.

6

Damage to apple from aphid populations includes de-
creased terminal growth, extensive leaf curl, and copious
honeydew accumulation on leaves and fruit (Oatman and
Legner 1961). 'A. pgmgiby intercellular penetration,feeds
directly on the leaf phloem removing plant nutrients. Each
nymphal inStar makes a minimum of five feeding punctures
(Heriot 1944). The collapse of surrounding cells and
subsequent leaf curl reduces photosynthetic capabilities of
the leaf. Quantification for the rate of plant nutrient
removal has not been completed for A. p223, however, a
methodology for estimating such parameters has been developed

for Drepanosiphum platanoides (schr.), the Sycamore aphid

 

(Dixon 1971).

Much of the previous literature on economic thresholds
and control of A, pgmi, as for many other orchard pests,
emphasizes preventive or scheduled chemical control methods
(Forsythe and Hall 1973, Glass and Chapman 1955, Pielou and
Williams 1961). Madsen et. a1. (1961) controlled damaging
aphid populations (where aphid honeydew was found on fruit)
when aphid densities greater than 5.0 were found on the
seventh leaf of rapidly growing terminals. This indice was
revised (Madsen et. al. 1975) and chemical control imple-
mented when 50% of sampled leaves were infested; these samples
contained the third, seventh, and fifteenth leaves from ca.

20 rapidly growing terminals.

Biological and Environmental Factors Influencing
'AphiSipomi--

 

Assessment of mortality factors influencing the population
dynamics of aphids i's notably difficult due to their unique
biological features. Specifically, overlapping viviparous
generations, the telescoping of generations within individuals,
varying production of alates, and their short developmental
times limits the ability to separate or quantify the impact
of mortality agents (van Emden 1963). Several studies,
reviewed below, have contributed the identification and
quantification of environmental or biological factors influencing
A. ESTE-

LeRoux (1959) evaluated the effect of frosts, heavy
rainfall, and aestival temperatures greater than 9OOF on
aphid populations in Quebec. He indicated a single heavy
frost in late May killed 96.5% of aphids populations in two
separate orchards. Five rainfalls caused an average of
54.2% mortality. A three day period with maximum temperatures
averaging 90.00F in mid-June reduced populations an average
of 71.5%.

The natural enemies of A. pgmi have been studied in
both Europe and the United States. Bonnemaison (1972)
identified two non-specific fungi attacking A. pgmi_in
France-~Entomophorales aphidus (Hoffman) and A. planchonina

Cornu. .Both required dense host populations and favorable

7

8
environmental conditions to reach epizootic proportions and
exert biological control. The only known specific parasite

of A. pomi is Trioxys angelicae Hal. (Hymenoptera: Braconidae)

 

which is found in Europe and Asia Minor (Evenhuis 1963,
Stary 1966, and Cierniewskii 1973). Parasitism of A. omi
in the U.S. has only been reported from California (Oatman
and Legner 1961, Westigard and Madsen 1965) by an Aphidus
sp. (Hymenoptera; Braconidae) that parasitized less than'l%
of the aphids. Parasitism appears to be of minor significance
in regulating A. pgm£_populations worldwide. 2

Apple aphid predation by species of the families
Coccinellidae (Coleoptera), Chrysopidae (Neuroptera), and
Syrphidae (Diptera) is well documented (Hodek 1966, Niemcyzk
1977, Setti 1973, etc.). Although effective predators,
these natural enemies often are general feeders and require
large numbers of hosts to develop, have few generations per
year, and often experience high mortality from orchard.
pesticides (Croft and Brown 1975). Thus, pesticides usually
limit their effectiveness in controlling aphid populations
in commercial orchards.

Setti (1973) evaluated the effectiveness of syrphid
predation in Italian orchards and noted a complete absence
of these predators where sprays were applied. The effective-
ness of aphid control in non-sprayed orchards varied as the
syrphid species were irregular in their occurance and

abundance. The presence of Chrysopa carnea (Steph)

 

9
(Chrysopidae) in orchards in Poland was related more to the
occurance of spider mite populations than eitherA. pomi

or Dysaphis plantegina (Passerini) (the rosy apple aphid)

 

populations (Niemcyza 1977). Further he concluded that the
limited control of the apple aphids by g. carnea, in com-.:
parison with its effective control of field crop aphids,
may be related to differences in the apple canopy microclimate,
orchard pesticide use, and the high mobility of the adult
predators allowing dispersal to other crops.

Holdsworth (1970) studied the predators of A. pgmi_

in Ohio, citing the species Metasyrphus weidemanni Johnson

 

(Diptera: Syrphidae), Leucopis sp. (Diptera: Chamaemyiidae),

Anatis quindecimpunctata Oliver (Coleoptera: Coccinellidae),

 

Chrysopa sanquinea (L.) and g. nigricornis Burmeister

 

 

(Neuroptera: Chrysopidae) as occasional- predators of A. pom'.

Primary predatores were Orius insidious Say (Hemiptera:

 

Anthocorridae) and Aphidoletes cucumeris (Lintner) (Diptera:

 

Cecidomyiidae). Holdsworth did not evaluate the control
potential for any of these species.

In summary, the literature contains substantial
biological information on A. BEBE! the description of
associated natural enemies, and some information of the
environmental factors regulating aphid populations. Relation—
ships between A. 222i and the apple host have been identified,

however, detailed information on the spatial and temporal

10

distribution of A. pomi and its natural enemies are limited.

The Biology of Aphidoletes aphidimyza

 

Recent taxonomic revisions of the aphidophagous species
in the family Cecidomyiidae (Harris 1966 and 1973, Gange
1973) have reduced species synonymy aiding in the identifi-
cation of their biologies and prey hosts. Specifically,

Aphidoletes aphidimyza (Rondani) is the accepted name for

 

Cecidomya sp., Phaneobremia sp. and A. cucumeris which have

 

 

 

been reported as predators of A. pomi (Bonnemaison 1972,
Evenhuis 1961, and.Holdsworth 1970). The prey range of A.

aphidimyza is exclusively limited to aphids; a list of over

 

60 host species was provided.by Harris (1973).

Utilization of A. aphidimyza for control of glasshouse

 

aphids has been extensively developed in western and eastern
Europe (Markkula 1973, Bondarenko 1975). This use of A.

aphidimyza has required study of its life cycle to facilitate

 

rearing and mass release of pupae and adults. Studies of

aphid-A. aphidimyza interaction have contributed to improving

 

glasshouse aphid control (El-Titi 1973 and 1974, Uygun
1971). The life cycle of this predator in glasshouse crops
has been reviewed by Markkula et. a1. (1976). Adams (1977)
discussed limited aspects of its life cycle in Massachusetts

apple orchards.

 

A. Aphidimyza overwinters as pupae in the upper soil

layer (1.0-2.5cm) beneath the host plant, emerging from May

11

, 12

to June in appleiorchard of Massachusetts (Adams 1977). In
glasshouses, emergent males live approximately 7.0 days and
the females 9.25 days (El-Titi 1974). Following copulation
the adult female lays 50-70 orange eggs (0.3 x 0.09mm)
singly or in clusters from the third to the ninth days. Egg
development under laboratory conditions (21.00C : 10 and
85-95% RH) is about 2.5 days (Uygen 1971). Egg survival
rates have not been determined, however, Uygun (1971) noted
improved egg maturation when females had previously been
provided with aphids and.honeydew. El-Titi (1973) evaluated
ovipositional responses to varying host densities in glass-
houses. He reported that the mean number of eggs laid per

female was proportional to the number of aphids per plant

(Brassica oleracea).

 

There are reported to be three larval instars of A.

aphidimyza which are orange in color and range from 0.4 to

 

3.4 in length (Harris 1973). Development of larvae at 21°C
when supplied with 25.0 hosts per leaf averaged 4.0 days,
survival was estimated to be roughly 75% and the mean number
of aphids killed/larvae during development was 22.5 (Uygun
1971). In feeding the larvae inserts its mouthparts into
interstitial membranes on the aphid's abdomen or legs.

After injecting a paralyzing enzyme (phenoloxidase) from the
salivary glands, the aphid gut contents are liquified and
removed (Mayr 1975). The number of aphids consumed or

killed per predator larvae varies with prey size and density.

13
Increasing the prey size (age) decreases the number of prey

required for development; i.e. 5.4 Myzus persicae adults vs.

 

21.1 numphs (at 21.00C) (Uygun 1971). Several authors (Dunn
1949, George 1957, Nijveldt 1969) have reported that the
number of aphids killed per larvae ranges from 5.0 to 100.0.
Wildbert (1972) has studied larval searching and has

related mortality to prey density. Larval mortality is also
influenced by the ability of the adult females to effectively
find hosts and oviposit eggs close to the colonies (El-Titi
1974). The final larval instar drops to the soil and forms

a small (1.8 x 0.7mm) puparium; emergence of the adults in
the laboratory occurs from 12 to 14 days later (Wildbert
1972). A total generation development time in the laboratory
is 22.5 days at 21°C (Uygun 1971).

Populatiions of A. aphidimyza have been tested for

 

tolerance to insecticides and acaracides (Adams 1977, Markkula
and Tittanen 1976). Adams (1977) reported high egg mortality
from the carbamate carbaryl and the organophosphate (OP)
phosphamidon. Larval mortality was greatest from the OPS,
phosphamidon and demeton-methyl and least from endosulfan.
Differential mortality to the O—P azinphosmethly in population
from an unsprayed orchard (highest) and from a commercial
orchard (lowest) was also detected. Low mortality to six

acaracides for a laboratory colony A. aphidimyza was reported

 

by Markkula (1977).

14

In summary, the biology and life cycle of A. aphidimyza

 

have been studied extensively. .Most experiments were
conducted primarily to facilitate use of the predator for
biological control of glasshouse aphids. Detailed assessments
of orchard populations feeding on A. peg; have not been

made. This information is required to utilize A. aphidimyza

 

more effectively as a biological control agent for aphids on

apple.

 

METHODS

Sampling Procedures

 

Populations of A. pomi and A. aphidimyza were sampled

 

weekly in a single orchard.block during the summers of 1976
and 1977 at the Graham Experiment Station, near Grand Rapids,
Michigan. The block sampled contained 40 Red Delicious
trees. These trees were bound by a 30 tree block of Northern
spys to the north and by a 60 tree block of Rome and Jonathan
trees to the south.

Cultural practices used in the block were similar tox
those applied in commercial orchards. Pruning during the
dormant season included the removal of the previous season's
shoot growth in the inner portion of the tree and limited
pruning of weakened or non-bearing limbs. Spray schedules
and the pesticides applied for control of pests other than
aphids for 1976 and 1977 are given in Table l. The pesticides

applied were selected on the basis of their minimal toxicity

 

to populations of the European red mite predator Ambleysius

fallacis (Garman) (Croft 1976) and Aphidoletes aphidimyza

 

(Adams 1977).

The plant sample unit chosen was an entire terminal
shoot. These were defined as terminals with axilary or
terminal buds formed in the previous season with approxi-

mately 10 leaves or more (Vyvyan and Evans 1932). The base

15

 

Table l.

1976
Date

4-15
4-19
4-30
5-07
5-19
5-27
6-02
6-14
6-29
7-13
7-26
8-10
8-24

1977
Date

 

4-14
4-19

4-25
5-06
5-16

5-25
6-07
6-24
7-05
7—19
8-03
8-17
8-31

16

Spray schedule for Gibson experimental orchard
block at Graham Experiment Station, Grand Rapids,
Michigan 1976 and 1977.

Chemical compounds per loo/gal water

Benlate 2 oz. and 1 gal. Superior oil
Benlate 2 oz. and 1 gal. Superior oil

Cyprex

. Cyprex

Cyprex
Cyprex
Cyprex
Cyprex
Cyprex
Cyprex
Cyprex
Cyprex
Cyprex

Cyprex

% lb.
% lb.
3/8 lb. and 8 lb. Guthion and 2 oz. Plictran
1b.

a

% 1b. and % 1b. Guthion
% 1b. and % 1b. Guthion
% 1b. and 8 1b. Guthion
% 1b. and % lb. Guthion
% lb. and % lb. Guthion
% 1b. and % lb. Guthion
% lb. and % 1b. Guthion

3/8 lb. and 1 gal. Superior oil

Cyprex 1 % lbs. and % lb. Karathane and 1/3 lb.
Systox

Cyprex % lb.

Cyprex 3/8 lb. and 2 pts. flowable Sulfer
Cyprex 3/8 lb. and % b1. Guthion and 1% pts.
Flowable Sulfer

Cyprex 3/8 lb. and % 1b. Guthion

Cyprex 3/8 lb. and g 1b. Guthion

Cyprex 3/8 lb. and % lb. Guthion

Cyprex 3/8 lb. and % lb. Guthion

Cyprex 3/8 lb. and % 1b. Guthion

Cyprex 3/8 lb. and % lb. Guthion

Cyprex 3/8 lb. and % lb. Guthion

Cyprex 3/8 lb. and 8 1b. Guthion

17
of these terminals are demarcated by a whorl of leaves or by
growth rings (Pool 1973). Secondary terminals, from axil
leaf buds of the current season's growth were few in number
and not sampled in this study.

From each terminal the number of leaves, leaf primordia,
aphids, and cecidomyiids were recorded. Leaf primordia were
defined as the number of unopened immature leaves clustered
at the apex of the terminal. Visual counts of primordia
ranged from 0-4, which corresponds to increasing terminal
growth activity. The number of leaves per terminal was
recorded from the top of the shoot to either the first
growth rings or to the whorl of leaves.

During 1976 10 randomly selected trees in the orchard
were chosen on June 3rd and in each a total of 20 terminals
were tagged for repeated observation throughout the season.
In each tree 5 actively growing terminals from 2 directional
axis (N-S or E-W) and in lower and upper tree canopy levels
were sampled per tree. In each axis terminals were selected
from the inner most of the tree canopy to the outer periphery
of the tree. The lower canopy ranged from 4 to 7 feet high
and the upper canopy from 7 to 12 feet high. At each site
on each date the number of leaf primordia, leaves and terminal
length was recorded. Aphid and predator numbers on each
leaf were recorded from the shoot apex to the base. Five of

the 10 trees were sampled on alternate weeks.

 

18

The sampling design for 1977 was expanded and improved
by randomly choosing up to 10 trees in the block on each
sample date. In each tree, counts from 3 terminals located
in the inside, middle and outer region of the canopy, in 4‘
directions and in the lower and upper tree canopy levels
were sampled. As in 1976, the plant variables measured in
1977 included leaf primordia and leaves.

Aphid counts in each subsample included all stages
found on leaf primordia, lower and upper leaf surfaces,
petioles and on stems between leaves. While aphids were
counted cecidomyiid larval locations were noted, and upon
completion of the aphid counts, the entire terminal was
again viewed to count cecidomyiid larvae.

Several factors influencing cecidomyiid larval population
sampling should be noted. Because of first instar larval
size and their tendencies to be hidden beneath larger aphids,
primarily second and third instar larvae comprised the
counts. Uygun (1971) has shown limited movement of larvae
(from eclosion to third instar) from the site of oviposition,
and as such the counts and distribution studies made in this
study were considered to be representative of all instars
even though counts of first instars were limited. Excessive
apple leaf curl can also bias counting methods, however, in
this study aphid densities were not exceedingly high and

only few apple leaves were curled in 1976 or 1977.

 

 

19

Methods of Analysis

 

For data analysis the influence of tree growth on the
pattern of insect infestation was compared in 1976 and 1977.
Mean values per terminal for leaf primordia and leaves
relative to canopy direction, canopy position (inside to
outer periphery) and canopy level were compared on a degree
day (dd) scale (Baskerville and Emin 1969, 5°C base threshold).
Corresponding aphid and cecidomyiid densities per terminal
in each of the quadrats were tested for significant differences
on each sample date by analysis of variance (ANOVA). Prior
to ANOVA, insect counts were transformed by log (X+l) counts.
This is an adequate transformation for populations which
follow a negative binomial distribution (Taylor 1970,
Appendix F).

To determine the type of terminal growth prefered by
aphids, 2 growth features were eValuated and compared. On
each sample date aphid counts in classes of terminals
separated first by leaf primordia (0-4) and then by 6_leaf
classes (10 leaves/class) were tested by ANOVA. On dates
where significant differences were detected, a modified
least significant difference test was used to rank means.

To compare growth characteristics between 1976 and 1977 leaf
primordia and leaves per terminal for both years were
plotted against dd and fit to log and power curves by least

squares analysis.

RESULTS

In 1977 as estimated 279 i 80.3 (i i_SE) terminals and
379 : 141.6 terminals per tree were present in the lower and
upper apple canopy quadrats in the study orchard respectively.
From gross observations there appeared to be no difference
between 1976 and 1977 in terminal growth densities in the
orchard. The preliminary data taken in 1976 is described
first to show the general features of tree growth and aphid
population activity. Figures la-c summarize the 1976 data,
showing means per terminal for leaf primordia, leaves and
aphids in the various quadrats of the apple canopy.

Leaf primordia per terminal declined reapily in early
season in the upper and lower quadrants. However, the mean
for the upper quadrat was initially greater than the lower
quadrant by ca. 1.0 leaf primordia/terminal and remained
greater through the season (Figure la). The appearance of
new leaf primordia ceased at approximately 2400 dd in the
lower quadrat, while in the upper quadrat at ca. 2800
dd. Leaf numbers were roughly equal in both canopies on the
first sample date, thereafter leaves were added more rapidly
in the upper.quadrat (indicated by the slope in Figure 1b).
The rate of leaf addition slowed at 1900 dd in the lower
canopy and at 2550 dd in the upper canopy. Roughly 7.0 and
18.0 leaves were added per terminal in the lower and upper
quadrats over the entire season.

20

 

menu LEHF PRIHORDIH/TERNINRL

21

 

 

O
‘2-
0)
. CRNUPY HEIGHT
O
9- tsunamuwtera
N
q ”PER canon (*1
d
D
9-
d 1
4 b
a ‘ ‘5' ‘
o _ .
O. T I I fi— I T I r T T I l——T"—
900 . 1400 1900 2400

Figure 1a.

DEGREE oars > 41 (F°J 1976

LEAF PRIMORDIA MEANS IN CANOPY LEVELS

(1976).

HERN LERVES/TERI‘I I NHL

22

35.00

I

25.00 30.00

1

20.00

CRNOPY LEVEL

16.00
L

. LONE! b9?!

.. UPPER (*1

 

 

10.00

1 T 1 U i

900 p ' 1150 2000 ZSSO
‘ DEGREE oars > 41 (F°J 1976

Figure 1b. LEAF MEANS IN CANOPY LEVEL (1976).

3100

 
  
  

 

23

Aphid densities in 1976, were greater in the upper
quadrat of the tree at all sample dates (Figure 1c). A
single peak of the population was observed at 1400 dd. At
peak density the mean number of aphids/terminal in the lower
quadrat was 186.54 and in the upper quadrat 433.02. (Figure
1c) . '

Figures 2a-f illustrate the 1977 mean densities per
terminal for leaf primordia and leaves from 3 stratified
sample types. Means per terminal in the lower and upper
canopy levels, in the inner, middle, and outer canopy positions,
and in the four directional axis_are depicted. These three
stratifications are also used in the presentation of the

spatial distributions of both A. pomi and A. gphidimyza

 

(Figures 3a-f).

The mean number of leaf primordia per terminal greatest
in the upper quadrat of the tree until 22 dd; thereafter,
the upper and lower levels were about equal in numbers
(Figure 2a). The decline of upper canopy leaf primordia
began 200 dd later than in the lower canopy. Decreasing
numbers of leaf primordia were recorded from the inner to
outer periphery of the apple tree (Figure 2b) at all sample
dates in 1977. Little difference was noted in the number of
leaves/terminal for each directional component (Figure 2c).

Leaf addition to terminals was most rapid during early
season in all canopy division (Figure 2d—f). Lower canopy

growth, as observed through increase in mean leaf numbers,

HERN HPH I DS/TERH I NHL

24

 

 

 

O
9
D
5—
ID
canon LEVEL
m {-8-}
urm {-‘4
O
D. C'__ .——-——-— ,A
t r I r U I —
°soo . 1450 2000 2550 3100

DEGREE oars > 41 (F°) 1976

Figure 10. APHID MEANS IN CANOPY LEVEL (1976)

MERN LERF PR I NORD I H/TERMINHL

3.00

2-00

 

0.00

.25

CRNOPY LEVEL
MFG-)1

UPPER Ii—J

L

 

1000

l
1500

j

I
2000

V I

I
2600

DEGREE oars > 41 (F°J 1977

Figure 2a. “LEAF PRIMORDIA MEANS‘IN CANOPY LEVELS

HEHN LEHF PRIHORDIH/TERNINRL

26

 

 

 

O
‘1’-
0
i a -' . cnuorv POSITION
. rm («e-1
O
9- amour (+1
N
. aura (+3
4
D
9..
fl
4
. "\
D /_:
O I i —
o. 1 fir I 1 1 j fl 0 l U I —'
1000 - 1500 2000 2500 3000

DEGREE cars > 41 tF°J 1977

Figure 2b. LEAF“PRIMORDIA MEANS IN CANOPY POSITION

 

MERN LERF PRINORDIR/TERHINRL

27

CHNOPY DIRECTION

    
  
  

umuurgsa
our oa-a
omnut~sa

”EST (*1

.00

 

1 U

 

1600 2000 S— 2600 3000
DEGREE DRYs > 41 IF°J 1977

°1DDD

Figure 2C. LEAF PRIMORDIA MEANS IN CANOPY DIRECTIONS

28
terminated at roughly 1600 dd. .Upper canopy growth continued
until roughly 2500 dd (Figure 2d). The relative numbers of
leaves added during the season in these two regions were
6.28 and 10.03 per terminal. The inner, middle, and outer
canopy positions showed cessation of growth at 2700, 1900,
and 1700 dd (Figure 2e),respective1y. The numbers of
leaves added in these positions over the growing season were
12.20, 7.40, and 3.76. As with leaf primordia, directional
differences in leaf addition were not observed in 1977
(Figure 2f).

Figures 3a-f summarize the data on aphid and cecidomyiid
population densities in all canopy divisions on each sample
data. ANOVA procedures were used to test significant
differences in populations within each division. Where
differences ( w.01) existed the largest mean is starred
(note: there were no significant differences in canopy
directions) (see Table 2). Two population peaks were

observed in 1977 for A. pomi and A. aphidimyza. In relation

 

to the previously described growth patterns of the tree,
there was a highly correlated aphid response. Four of the
ten sample dates showed a significant difference in aphid
means for lower and upper canopy divisions. These four
dates also correspond with peaks of growth of the apple host
in these levels and the magnitude of the peaks reflect the

differences in growth activity. The first peak of aphid

 

NERN LEHVES/TERH INHL

20.00 26.00 80.00

15.00

29

 

 

 

1
- CHNOPY LEVEL
. Ixmmtq£1
. m c-a-x

I U ‘ ' U ' U f l
1000 1000 2000 2500 3000

Figure 2d.

DEGREE DRYs > 41 (F°) 1977

LEAF MEANS IN CANOPY LEVELS

 

HERN LEHVES/TERHINRL

30 >

 

 

o
9
o-
G)
J J
o
9
w- ‘
N
o
9
Our
N
s l CRNOPY POSITION
3‘ . mullfikl
-‘ RIDDLE (+1
J :wnm 9+4
a
9
o . j r j
g l a l ' I . ' IR
1000 - 1800 2000 2600 3000

DEGREE DHYS > 41 (Fol 1977

Figure 2e. LEAF MEANS IN CANOPY POSITION

 

 

HERN LERVES/TERHINHL

31

    

 

 

O

‘2

3%

O

‘9

‘0‘

N
J

O

‘3

o—A

N
‘ cauorv DIRECTION
‘ NORTH {-6-}

c, .

‘3

ID— 8881’ (+1

OI!
. 8001’" (*fi-J
.1 "£87 (*3

O

c.’

S T r 7 V T T j —‘r V V I
1000 1500 2000 2600 3000

Figure 2f.

DEGREE oaYs > 41 (F°) 1977

LEAN MEANS IN CANOPY DIRECTIONS

 

32

 

mmHQEwm HHm How HmcHEump umm HEOQ .MH

 

 

ch. NMN.m N cHh. cmv. m Ncc. HHm.m H mm.>H chN nnlmcnc
cvc. ch.m N mcc. cmv. m mvh. ccH. H cn.cH mNcN whthnh
Hcc. mvm.cH N cch. mhv. m ccc. vNH.h H cc.H> cmvN hnlchn
Hcc. th.HH N cwc. NBN. m Ncc. va.> H cc.mm vcHN nhlmHnb
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cNH. qu.N N ch. mvc.N m mcm. ccc. H mm.cN cmmH wthNac
ccv. cmc. N cam. ch. m Ncc. bbv.m H vc.c cme whlmch
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m .cmHm m mm m .38 m mm m :58 m mo 582 no mama
CONuHmom mmocmu coHuomHHo hmocmu Hm>mH mmocmo Ucmuw

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anmfl :mwz CH wmocmnmmmHo mchHEHmumo How 4>Oz< msu scum muHSmmm .N mHnt

 

 

33
numbers showed means of 45.20 and 46.98 for both lower and
upper canopies. The second peak showed the greatest number
of aphids in the upper quadrat, 91.12 versus 42.55 aphids
per terminal in the lower quadrat. This coincided with the
same period of rapid growth in the upper tree canopy (Figure
3a and Figure 2a).

Mean aphid densities in the three canopy positions were
not significantly different on the first four sample dates.
On the remaining dates (Table 2), significant differences
between canopy positions ocurred, with the highest aphid
densities found in the inner canopy position followed by
progressively fewer aphids in the middle and outer periphery
of the apple canopy (Figure 3b). Directional differences
shown in Figure 3c were not significant.

5. aphidimyza populations showed two peaks which were

 

closely synchronized with the temporal, spatial, and numerical
ldistribution of its prey. Predator populations for the

lower and upper canopies peaked at ca. 1900 dd and at 2400

dd (Figure 3d) respectively. The means for the first peak
were roughly equal at 0.33 and 0.28 cecidomyiid larvae per
terminal. The second peak showed greater numbers in the

upper canopy than the lower at 1.35 and 0.85 larvae per
terminal respectively. Of the three canopy positions the
inner canopy contained the largest number of predators for

six of the ten sample dates, similar to the aphid densities

described (Figures 3b and 3e). Directional differences in

 

   

NEHN RPHIDS/TERHINRL

 

 

34

 

 

D
9
O
O—
"
‘ 4
O ‘
Q
:2— CRNUPY LEVEL
. LOWER ('8')
. 00m (*3
D a:
9
D—
ID
O C
9
m-
N
. ¥
¥ '.
O
O. t r 1 T U I r r I
1000 1800 2000 2500

Figure 3a.

DEGREE DHYS > 41 (F0) 1977

APHID MEANS IN CANOPY LEVELS

1
3000

 

. v 1 , ,H,‘

MERN HPH I DS/TERH I NHL

50.00 75-00 100.00 125.00 150.00
1 l l l l l l l l l L l l

l

l

25.00

00

 

unmnrxqra

GNU! h+-)

35

CRNOPY POSIIIBN
aunt‘s?)

 

'4
°1000

Figure 3b.

I I j I

1000 2000
DEGREE oaYs > 41 (F°1 1977

APHID MEANS IN CANOPY POSITION

I
2500

1
8000

HEHN RPH 103/ TERM [NHL

36

  
 
  

 

 

O
9
D
a-
n
a
J
. CflNOPY DIRECTION
O J mm (‘6') . ~
°.
F
a 000m “*3
J an: (*1
a
O
9
Dc-
‘0
J
J
o 1
‘3
m.-
N
J
O
0
5‘1 ' I . T I ‘ ' I I ' I
1000 , 1500 2000 2500 3000

DEGREE DaYs > 41 (F°J 1977

Figure 3C. APHID MEANS IN CANOPY DIRECTION

 

MERN CECIDOHYI IDHE/TERHINHL

37

2.00

CRNOPY LEVEL
um I-e-I

ME! 1*)

1.00

 

 

 

0.00

V j *1 j

I
2500 3000

2300
DEGREE URYS > 41 (F°) 1977

000 1500

Figure 3d. CECIDOMYIIDAE MEANS IN CANOPY LEVELS

 

 

HEHN CEO I DONY I IDHE/TERM I NHL

38

2.60

2.00
l I l I

l

CHNOPY POSITION

o d
“3
..." INNER I-s-I
d
. RIDDLE («A»)
o .. 00m (+1
O
0-4
CI.
d
" 4
.. ,. 4
D " u
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:3 4¥
« 4
. ¥~ ; “

 

 

T

1 "—'——'l
2500 3000

T I l' I

0.00
L

1000 - 11500 ‘2000
» DEGREE DHYS > 41 (F°) 1977

Figure 3e. CECIDOMYIIDAE MEANS IN CANOPY POSITION

 

39
the mean counts of cecidomiids were not observed (Figure
3f). As in the aphid figures, where significant ( «.01)
differences in any canopy quadrat occurred, the dates are
starred (refer also to Table 2a).

The results of ANOVA comparisons between aphid densities
per terminal for leaf primordia classes (0-4) on each sample
date in 1977 are given in Table 3. For seven dates where
significant differences occurred modified LSD tests showed
terminals with the greatest number of leaf primordia to have
the larger aphid populations (Table 4). Results from ANOVA
comparison of aphid densities in six leaf classes of terminals /
showed almost identical results (Table 3). Table 5 presents
the ranking of leaf class means for each sample date with
significant differences. Again, higher aphid densities were
associated with the larger leaf classes, which are indicative
of more vigorously growing terminals.

Table 6, presents the equations derrived to describe
growth activity in canopy level and canopy position. Leaf
primordia decline and leaf addition rates, indicated by
their slopes in 1976 and 1977, show little difference
between years. Comparison of these growth rates although
influenced somewhat by sampling differences, showed 1977
leaf addition in the lower canopy to be roughly twice the

rate of 1976 and nearly one and a half times the rate of the

1976 in the upper quadrat.

HEHN CEO I DOMYI IDHE/TERHINHL

2.00

1.00

 

 

Figure 3f.

40

   
 
 
 
  
  
   

CHNOPY DIRECTION
mm («B-1
m? {-4-}
mm (+1

180‘! 1*)

 

1500 2000 2000
DEGREE DRYS > 41 IF°I 1977

CECIDOMYIIDAE MEANS IN CANOPY DIRECTIONS

41

mmHmem HHm new

HmCHEHmu Com MNMEHCHCQM .<

 

H

 

 

ch. Hmm.H N mmv. Hcc. m mmc. hcm.v H cmH.c mHmN hhlmclm
ch. hMN.v N nmm. mnc. m mmm. Nmm. H mmm.c MNcN NBINNIN
ccc. mc>.mH N mcm. NNN.H m mmm. Hcv.H H NMH.c cmvN thchn
Hcc. bNH.h N NcN. NmN.H m mnc. ch.m H NvN.c vcHN thMHIh
Ncc. mmc.m N mmn. cam. m cmc. mmc. H HNc.c NcmH bblccln
Hcc. mmh. N 5mm. NvN. m ccm. Nvm. H cmc.c HcNH nhlmNIc
vcm. cmm. N HNm. mmh. m cNm. mHv. H mNc.c cmmH thNNIc
mcm. ccc.H N vmm. ccc.H m mHm. ccc.H H eccsc can nuImHIc
m .33 .m mo m .chme a an C :83 m mo :00: on mama
CoHuHmom hmocmo COHuomuHo NQOCMU Hm>mH >m0Cmu HUCMHU
.bhmH .mConH>HQ NQOCmU mmHCB CH mmHuHmCmo
uoumcmum Cmmz CH mmoCmeHMHQ mCHCHEHmqu Mom <>Oz< Eonm muHCmmm .mN OHQMB

42

HMCHEku Hmm HEOQ .m

 

 

 

 

H
Hoo. oaa.wm a Hoo. mum.om m mm.>H mHmN Enumoum
Hoe. HNN.bm m Hoo. mom.mn m mH.mH mmem Hnsnmnn
Hoo. mHH.om m Hoo. mo>.mH N oo.HH mmam unuomnn
Hoo. mmo.mm m Hoo. amm.om m om.mm vaN huannn
Hcc. oHa.om N Hoo. mmw.mH m mm.ma NmmH nnuoonn
Hoo. HmH.aN N Hoo. mom.vm a am.ma HmHH nnnmmum
Hoe. mmn.NH N Hoo. Nmo.HH a mm.om ommH hnummum
an. ccc.H N mcH. Nmm.H v vc.c cNmH nuImHIc
mmh. mcm. N Hcc. Ncm. a mN.m MhNH thcHIc
mob. mom. N Ham. Nos. 4 NN.H mmHH hunmouo
m .CmHm m mo .m .cmHm . m mo saw: no mumo

mmmmeu mmmH mmmmMHU MHUCOEHHC mmmH UCCHU

H

.HHmH .AmIHc mmmmmHo mama cam Avuoc MHouosHum mama mo
mommMHU 059 CH mmoCmumMMHo CHCQC Cmmz Mom mCHumme Eoum muHmem C>Oz< .m wHCmB

 

43

Hmmch #0: Op umHHomnCm wEmm ms» >2 omsoHHow mmez

muCCoo HHH+xV moH Hc cmEHOMmCmup Eonm mum wommMHo 0:» How Umqummum mCmmE

 

HMCHEnwu “mm HEOQ awH

 

 

nmmv.H nvmm.H Qcmm.H MmmH. hm.hH mHmN hblmclm
£55m.H vam.H QhNN.H mmmc. mh.mH mmwm hhlhmlh
ncmm.m thc.m vav. cc.Hh mmvm chlomlw
QHmm.N nwmv.N nmmm.H mHhm. cm.mm vaN thMHIh
QmN0.H ammo.H thh.H mhbc. mm.m¢ NcmH hhlwclh
nmmc.m Qme.H chm.H Qva.H mmvm. wm.mv HwhH chlmmlw
ammmm. Qmmc. ammm. nbvm. vaN. mc.ON cmmH bcINNIm
v m N H c C002 no mama
mommmHu wHUHOSHHm mme UCmuw

mmmmeo MHUHOEHHC mmmH CH mmHuHmCmo pHCmd Cmm3umm mDOCmummmHQ .

HCMOHuHcmHm CHH3 wmuma meEwm Mew mumwa QmH CDHMHUOZ mo mpHCmmm .v oHnme

44

HmHMHp HOC Op HmHuomme 08mm map an pmonHom

N

 

mCmmz .muCCoo HAH+xV mOHV meHOHmCmuu Eoum UmCkuno mum Umqummum owwz
HCCHEHDH Hmm.mmmm .mH

an.N mom.H ma~.o 04H.o moo.o mm.HH mHmm Hhumoum

moo.o oom.~ nma.H mH~.o 00H.o moo.o mH.mH mmmm HHIHNIH
DHN.H nom.o mvH.o woo.o oo.HH mmam HHIONIH

coo.~ nmm.o ROH.o moo.o om.mm amHm HkImHIH

ammm.o nmv.H «Ha.o mm.ma mmmH HHIEOIH

nHm.H EMH.H mam.o am.ma HmHH Hkummsw

nmm.o mam.o .mmH.o mm.o~ ommH Hknmmsm

m m a m m H cam: on mama

 

mamz mwMHU mmmH

N

. mmmWMHU NMOQ CH

mmHuHmCmQ pHCmd C003Hmm
DamonHcmHm nqu mmuma wHasmm Moo mummeHomu amHmHooz mo muHsmmm .m mHnma

45

Table 6. Results of Curve Fitting For Leaf Primordia
Decline and Leaf Additionwl976‘and 19771

' Logarthimic Leaf Primorida Decline (y = a(log dd) + b)

 

. . . Slope Intercept 2

Year DlVlSlon a . b r
1976 Lower -1.98 15.66 0.90
Upper -3.08 24.75 0.92
1977 Lower -2.46 19.38 0.90
Upper -3.09 24.38 0.91
Inner -3.13 24.96 0.89
Middle -3.02 23.70 0.90
Outer -2.14 16.77 0.91

Power Curve Fit of Leaf Addition (1n y = a 1n (dd) + 1n b)

Slope Intercept

 

Year Division a b r2

1976 Lower 0.31 1.75 0.82
Upper 0.38 1.04 0.73

1977 Lower 0.70 0.12 0.91
Upper 0.53 0.40 0.93
Inner 0.67 0.14 0.93
Middle 0.41 0.86 0.82
Outer 0.22 3.51 0.66

 

 

DISCUSSION

Total apple tree growth includes the subcomponents of
leaf and terminal shoot production, woody tissue enlargement
in both the tree canopy and root zone, bud and fruit formation
and carbohydrate storage. In total, these proceSses are
greatly influenced by the past and current cultural and
environmental conditions to which an orchard tree has been
and is exposed.

More specifically growth of terminal leaf primordia and
subsequent new leaves is regulated by the rate of movement
of soluble carbohydrates and amino acids from roots, bark
and leaves to these sites (Zimmerman 1961). As observed in
this study, these features of tree growth showed yearly
variation over time (Table 6), but varied little in distribution
between years at the same time period dd (Figure 1a, lb, 2a,
2b). This indicated that similar management and environmental
conditions were occurring annually and also that a relatively
controlled process of tree growth was being maintained by
the orchard manager from year to year. Probably due to the
particular prunning practices used, the inner and upper
quadrants of the tree consistently experienced the greatest
amount of growth activity (Figures la, lb, 2a, 2b).

Several workers have observed that aphids distribute

themselves in close association with active plant growing

46

 

47
sites (Kennedy et al. 1950, Kennedy 1958, Dixon 1979).
These responses are related primarily to their ability to.
assimilate and utilize soluble carbohydrates and amino acids
present at these locations of the plant host (Way and Cammell
1970). Takeda et al. (1968) and Specht (1970-72) have
clearly shown that mature growth of apple, which includes
primarily insoluble carbohydrates and amino acids and is
made up of leaves with hardened cuticles, does not support
reproduCing populations of g. pgmi.

In this study 5. pgmi showed little directional preference
for terminal colonization, but as with the distribution of
terminal growth and correspondingly nutrients, the most
dense populations were found in the inner and upper level
quadrants of the trees. Similar relationships (host-plant-
aphid) have been reported for the sycamore inhabiting aphid
Drepanosiphum platanoides (Dixon 1971) and the lime tree
species Eucallipterus tiliae (Dixon 1971a). The methods
utilized in Dixon's studies to obtain information of the
aphid-host nutrient and energetic cycles could aid description
of 5. pgmi impact on tree growth.

Effective natural enemies require a temporal or spatial

distribution that is closly associated with their host. In

this study the distribution of g. aphidimyza larvae relative

 

to that of A. pomi reflected such an association (Fig. 3a-
f). Predator larvae were most commonly found on inner and
upper terminal growth of the apple tree and were always

associated with aphid populations. The delayed colonization

48
and numerical response of the cecidomyiid to the increasing
host density in the upper quadrat were indicative of its
relatively immobile larval state (Wildbert 1972) and the
hoSt searching capacity of the female adult (El-Titi 1978).
Thus, dispersion of this cecidomyiid-in response to changing
host distribution is controlled primarily by adult movement.

In summary, there are several factors which indicate

 

that g. aphidimyza may be an effective natural enemy for
control of 5. Egg; which can be incorporated with existing
IPM systems on apple.

1) It is closely attuned to the spatial distribution
of g. pgmi_and seems to occupy all components of
the habitat used by the aphid.

2) It appears that pesticide resistant or tolerant
strains of this natural enemy commonly occur in

orchards (Adams 1977, Warner, unpublished data).

 

3) g. aphidimyza has a variety of alternate hosts
(Harris 1973) which will support predator populations
in the absence of the pest.

4) It has proven to be a very effective predator of
other aphid species occurring in greenhouses
(Markkula 1973).

Based on these data, there is further need to continue

research on this species, to evaluate its potential for

49

controlling apple aphids in Michigan and to implement an
effective management program compatible with existing apple

IPM stratigies.

 

APPENDICIES

 

APPENDIX A
This appendix contains environmental parameters for
Graham Experiment Station for April - August 1976 and
April - September 1977. Provided are the maximum and minimum
temperatures (OF), the percipitation, and the accumulated
degree days (Base 41°F) estimated for‘A,'pgmi development

(Jokinen unpublished analysis).

50

 

 

51

mam wH.c mm mm Hmm

 

 

 

 

5mm mm.c mm mm cmm HmN vm cc cmm
hmc hh.c mm mm mNm th Hm mm va
NHc mm mm cNm ch . cm he va
mmm mv cm mNm ch . mN cm va
Nmm mv cm cNm ch Nm.c 5N mm cNm
Ncm Hc.c Nv mc mNm ch Nm.H cm we va
cmm mm mm va ch vH.c mv Hm va
mvm mm mm mNm NmN Nch mm mm qu
mmm mm on NNm cam cc.c mm Nu NNv
HNm Nc.c mm mm HNm mNN mc.c mm mm HNv
Hcm he Nc cNm vHN mm Nc ch
hmv «m cm mHm ch mc.c cm Nc mHv
mum (mm mm mHm th mm mm ch
va mH.c mv Nh SHm mvH mm mm th
va mm.c mm mm ch hHH cm.c mm cm ch
va Hm mm mHm mm cc mm me
HHv mc.c Hm mm va Hc . we Hc vHv
cmm mm Nc mHm me oN he mHv
Hmm .Hm Hm . NHm mu cN Hm NHv
mmm mH.c Nv Hm HHm mm mH.c SN Hm HHv
Ncm mm mm ch Hm Hm vm ch
cwm mm mm com um wN mv mcv
cmm mN cm mcm mm vN mm mcv
mmm mm.c mN He now Hm Nm mm mcv
mmm mv.H mm mm wcm mN Nc.c mm mm ccv
hHm cw cm mcm cN mN mm mcc
«cm Hc.c mN Nv vcm cH Nc.c mm cc com
com mm.c Nm mv mcm m mm mm mcv
Ncm mm Hc Ncm H , «c.c Hm cw ch
qu Hv vc Hcm H mc.c mm me How
m Hv CH ... m,. m . mumo m He , CH. m m. muma
mmmo mmowum Cmm xmm mwoo mmomum CHm xmm

mHmH mm: mHmH HHumm

mumo Hmnumwz CoHumum Emcmuw

 

 

52

 

 

 

SHSN SH.S NS SS HSN
SSSN SS SN SSN
SSSN SN.S SS SN SNN
HSSN SS SS SNN
NSNN SS.S SS SS NNN
NSNN HS NS SNN
SNNN SS SS SNN
SSHN HS.S NS SS SNN
SSHN SS.S HN SS SNN
NNHN HS NS NNN
NSSN NH.S NS SS HNN
SSSN SN SS SNN
SNSN NS SS SHN
SSSH SS NN SHN
SNSH SS SN NHN
SSSH SS SS SHN
SNSH SS NS SHN
SSSH SS NS SHN
NSSH SS NN SHN
NSSH SS SS NHN
NSSH NN SS HHN
NSNH NS.S NS SS SHN
NSNH SS SS SSN
NSNH HS.S SS SS SSN
NNSH SS SS NSN
SSSH SS SS SSN
HHSH SS SS SSN
NSSH NS SN SSN
SSSH HS SN SSN
SSSH SS NN NSN
NHSH SN.S HS HS HSN
MOHv .QH ho. ho OHMQ
m.oo mHomum H: mm:

SNSH >HSS

GUGQ Hmfluwmz COflflmum EMEMHU

 

NSSH NS SN SSS
SNSH SS SS SNS
NSSH NS.S NS SS SNS
SHSH HS SS NNS
SSSH NS SN SNS
NSSH SH.S SS SS SNS
SSSH SS.S SS NS SNS
SSSH SS HS SNS
SNNH HS SN NNS
HSNH SS SN HNS
SNNH SS NS SNS
SHNH NS.S SS SS SHS
NSHH SS SS SHS
SSHH SS SS NHS
SSHH SH.S NS SS SHS
SSHH SN SS SHS
HNSH SS NS SHS
SSSH SS SS SHS
SSSH SS SS NHS
SSS SS SS HHS
SSS SS SS SHS
SSS SS SS SSS
SNS SS SS SSS
SSS NS HS NSS
NNS NS NS SSS
SSN SS SS SSS
SSN HS SN SSS
SSN SS SN SSS
SNN SS SN NSS
NSN NS SN HSS
moHS SH we no mama
Span mHomum cm: xmz
. ‘ SNSH Scan

53

 

 

 

chm mm Hm cmS
Ncm mN mm mNS
cmm hm SN cNS
mSm Sm HS NNS
mmm Nc.c Nm mm SNS
Hmm NS.c Nm mm mNS
cNm mm Sm SNS
mHm SS mm mNS
mcm NN.c Nm HN NNS
mmN Sc.c cc SN HNS
NmN HS.c mm SN cNS
HmN cm Sc mHS
NcN cm cN SHS
cNH Hc.c cm SN NHS
mmH SS cm SHS
NmH mS Sc mHS
mNH mS HN SHS
NcH mm mN mHS
Hm .mS cm NHS
Nm mm mN HHS
Hm NS mm cHS
NN HN mm mcS
NN SH cm mcS
cN cc.c NN Sm NcS
cN mc.c SH Sm ScS
cN mS.c cm NS mcS
SH mm cm ScS
mH SH.c Hm mm mcS
m NS.H mm mm NcS
c mN NS HcS
m HS CH ,m h mumo
mwao CHbmHm CHm xmm
NNmH HHHmd

mama Hmzummz CoHumum EmanU

NSNS SS NN HSS
SHNS SS NS SSS
SSNS SS.S NS SN SNS
HSHS SS NS SNS
SSHS SS.S SS NS NNS
SSHS SS NS SNS
NNSS HS SS SNS
SSSN SS SS SNS
SHSS HS SS SNS
HSSN SS NS NNS
SSSN NS SS HNS
NHSN SS SS SNS
NSSN NS SS SHS
NSSN SS HS SHS
SSSN SS. SN NHS
SHSN SS SN SHS
SSNN NS.S SS NS SHS
SSNN SS.S NS SS SHS
SSNN HS.S SS SS SHS
SHNN SS.S SS NS NHS
SNSN SS SS HHS
SHSN HS HS SSS
SSSN SS SN SSS
SNSN NS SN NSS
NSSN SN.S NS SS SSS
mNmN mH.o ow om mom
SSSN SS SN SSS
SNSN SS SN SSS
SSSN SH.S SS SN NSS
SSSN SS SN HSS
m HS CH C m . Sumo
Swan cHomum ch xmm

 

SNSH DSSSSS

 

54

mcHH Nm SN HmS

 

 

 

 

SSNH SS SN cmS NScH SS SN cmS
HSNH cH.c SS Sm SNS mScH NS Sm SNS
cmNH cS cS SNS SNcH HS Sm SNS
SSSH SS SS NNS mccH . SS SN NNS
SSSH cS SS SNS NNS SS SS SNS
cSSH Nc.c HS SS SNS SSS HS SS SNS
ScSH . mS Nm SNS SHS mS NS SNS
SNSH SS cm mNS cSS HS.c cS Sm mNS
cSSH SS NN NNS SSS SH.H HS cS NNS
cmSH SS HN HNS SHS NS NS HNS
NHSH mc.c SS NN cNS SNN . SS cS cNS
SSSH NS NN SHS. SSN _ cS SS SHS
SSSH Nm.c mS NS SHS mHN NS SS SHS
cmSH cS SS NHS mSS cS SS NHS
mSmH SS HS SHS cSS NS Sm SHS
cNmH HS NN SHS cNS HS NS SHS
NSmH HS SS SHS SSS SS NS SHS
SNmH SS NS mHS SSS NS SN mHS
SHmH SH.c NS SS NHS SSS cS NN NHS
SSNH SH.c .NS SN HHS SNS Nm SS HHS
mNNH Sm NS cHS SHS mm SS cHS
HSNH NH.c mm HS ScS mHS SN mS ScS
NSNH NS SS ScS ScS Hm SS ScS
mmNH cS SS NcS SSS cS SN NcS
NNNH SN.c mS NN ScS SNS SS SN ScS
ScNH Sm.c NS SS ScS NSS cS.c SS mS ScS
mNHH SS NN ScS SmS cS cN ScS
SSHH Sm SS mcS cNS mS NN mcS
mSHH HH.c SS SS NcS ScS cS mN NcS
NmHH SN.c mS SN HcS Smm HS SS HcS
m HS CH .H .H mumo EH HS CH .H ..u. “...me
Swan mmomum cmm xmm Swan mmomum CHm xmm.

NNSH SSSS NNSH Sm:

mumo Hmsummz COHumum Emnmuo

55

 

SNSS HS NN HSS
HSSS SS NN SSS
SNSS SS.H SS SS SNS
SSSS NN SS SNS
SSSS NS NS NNS
HNSS NS SN SNS
SSSS SS HN SNS
HSSS SS.S SS SS SNS
SHSS SS SN SNS
HSNS SS.S NS SS NNS
SNNS SS SN HNS
HSNS SS HN SNS
SSNS NS SS SHS
SNNS SS NN SHS
SSNS NS.S SS SN NHS
SSHS SS SN SHS
SSHS NS SN SHS
HSHS SS.S NS HS SHS
SSHS HS u.SN SHS
HSSS SS.S NS NS NHS
SSSS SS HS HHS
NSSS SS.S SS SN SHS
SSSS HH.S SS SN SSS
SNSN SS.S NS SN SSS
SSSN SS SN NSS
SHSN SH.S SS NS SSS
SSSN SN.S SS SS SSS
SSSN SH.S NS NS SSS
SHSN SS.S SS SN SSS
SSNN NS SN NSS
SSNN HS.S SS SS HSS
S HS SH S S .mumo
Swan SHSSHS :Hm xmm

 

 

NNSH DSSSSS

mumo Hanummz CoHumum EmCmHO

 

NmNN Sc.c HS Sm HmN
ScNN NN.c cS NN cmN
NNSN cH.c NS Hm SNN
NSSN NS SN mNN
mNSN SS SN NNN
ScSN . SS SN SNN
SSSN Sm.c SS SN SNN
NSSN SS NS SNN
mNSN SS HS mNN
NcSN Hm.c SS Nm NNN
SNSN SS cS HNN
SmSN cN HS cNN
SSmN Sc.H NS cm SHN
SSmN SH.c SS mm SHN
SNmN SH.c SS mm NHN
NSNN Sc.c NS NS SHN
NSNN SN NS SHN
NHNN SS mm SHN
SSHN mc.c SS Sm mHN
SSHN NS NS NHN
cNHN cS Hm HHN
cScN NS NS cHN
NScN cS Sm ScN
cch HS Nm ScN
cch Hc.c SS HS NcN
NSSH cN cS ScN
mNSH Hc.c SS SS ScN
SSNH Sc.N NS SN ScN
SSmH SS cc ch
SNSH SS SN NcN
NcmH SS.c NS SN HcN
moHS CH mo mo mama
m.oo mmomnm CH2 xmz

NNSH SHSS

56

 

MS MS SSS

NS SS SNS

NS HS SNS

NS SS NNS

SS SN SNS

SS.S HS SS SNS

SN.S SS SS SNS

SS.S SS SS SNS

No.0 SS NS NNS

SH.S SS SS HNS

HS SS SNS

SS.H NS NN SHS

SN.S SS HN SHS

SS NN NHS

SS.S NS SS SHS

SS SN SHS

NSSS SS.S NS HS SHS

NSSS NH.o NS SS SHS

SHSS SS SN NHS

ooSm SS SS HHS

SSNm SS HN SHS

SSNm SS SN SSS

SSNS SS NN SSS

NHNm NS SN NSS

SSSS SS SS SoS

SSSm No.0 SS SS SSS

SSSS No.0 SS SN SSS

SHSS Sc.c SS NN SSS

SSSm SS.S SS SN NSS

SSSm Sm.c NS SS HSS

m HS CH m m mumo
Swan mmomum cwm xmm

 

NNSH umnswummm

mumo ngnmmz coHumum Emnmno

 

APPENDIX B
Techniques Utilized for Establishing Laboratory
Colonies of Aphidoletes Aphidimyza (Rondani)
This appendice provides a description of the techniques

and equipment used in the establishment of the aphidophagous

cecidomyiid gphidoletes aphidimyza. The ease with which

 

field samples were collected and laboratory colonies established

facilitates use of £3 aphidimyza to develop information on

 

predator-prey interaction.
A review of literature on rearing methods reported for

g. aphidimyza reveals that several studies describing mass

 

rearing and effective release rates of 5. aphidimyza have

 

been conducted (Bandarenko 1975, Markkula 1973, and Ushchekov
1975). These and studies describing the life cycle (e.g.
Uygun 1971) provide a fairly complete outline of its biology

to initiate development of rearing programs.

Methods
Difficulty in the removal of small cecidomyiid larvae
(0.4 - 3.4mm) from field colonies of the apple aphid, éEEiE
pgmi (Degeer) necessitated development of techniques to
obtain sufficient numbers of predators for experimentation
and rearing. This is in part a modification of the techniques
developed by Markkula and Tittanen (1976). It is not intended

to provide a complete rearing methodology but rather methods

57

 
 
 

58

to combine and manipulate unique populations of'§.'aphidimyza.

 

Prior to field collection of the cecidomyiid, rearing
units were prepared. These units were ca. 23cm x 30cm x 10cm
deep plastic containers filled with approximately 7.0cm of
slightly moistened white sand. Placed upright in the sand were
three rows of five 15cm long test tubes. Each tube was two
thirds full with equal parts of water and Hoaglands nutrient
' solution. Perma—seal ®, and celluloid film material, was
used to cover the test tube top. These units were easily
prepared, reusable, and of minimum cost.

Field colonies of 5. pgmi_on apple terminals

were observed for the presence of g. aphidimyza. The upper

 

30.0 - 50.0cm of reapidly growing terminals found with aphid
and predator were removed and placed into plastic bags. These
bags were immediately set into a chilled polyurethane cooler
for transfer to the laboratory. Upon return to the laboratory,
the terminals were sectioned to ca. 15.0 - 18.0cm and lower
leaves without or with few aphids removed.

These sections, each with leaf primordia, immature
leaves, an aphid colony, and predators were inserted singly
into the test tubes. A 6.0cm diameter cage about 20.0cm in
height was placed over each terminal and test tube. The test
tube solution provided sufficient water and nutrients to
maintain leaf turgor and the aphid colony for approximately

four days at 25°C and 60—80% relative humidity. This was

 

59

sufficient time for the development of larval cecidomyiids
and provided adequate host numbers.

At the completion of the larval development, roughly
five days later, the larvae dr0pped to the sand below and
formed the pupariums. Puparium were found at a depth of
ca. 1.0cm below the sand surface and encased in sand, they were
easily recognized by their size (1.0 x 1.5mm). The puparium
were removed with forceps and placed in petri diShes. The
confinement of the predator within the circumfrence of the
cage further aided the recovery of puparium.

Once in the petri dishes the number of puparium were
recorded and the dish placed at either room temperatures

or at 8°C (in a refrigerator). é. aphidimyza in the petri

 

dishes held at room temperature emerged in roughly twelve
days. Survival of the cecidomyiids following cold storage
was noted at periods up to six weeks later, when dishes
placed at room temperature for twelve days were found to
have adult cecidomyiids.
To determine if these rearing methods were adequate,
the emergent adults were released in screened cages containing

aphid colonies (Acrosiphum pisum) On broadbean plants.

 

Following three days of development and oviposition the
confined adults were removed with an aspirator. In the next
four days aphid colonies were observed for egg and larval
develOpment. The experiment was concluded when second and

third instar larvae were observed.

The methods presented here allowed the establishment of

laboratory colonies of Aphidoletes aphidimyza from field

 

colonies of the apple aphid Aphis pomi on apple terminals.

 

The relative ease with which predators were manipulated

using these methods could aid in mass rearing programs,
determining pesticide tolerances of field populations by
limited sprays to individual terminals, and in manipulating
colonies in the laboratory to describe the predator-prey
interaction. Finally, use of these methods to obtain accurate

density and survival information of the predator is recommended.

60

 

APPENDIX C 7 . S
Frequency Class Distribution of Aphis pomi and
Aphidoletes Aphidimyza for 1976 and 1977

 

 

The following provides the frequency distribution of

aphid and predator counts for 1976 and 1977 on each sample

date. The sample dates are coded as:
A 6-10-76 A 6-03—77
B 6-18-76 B 6-10-77
C 6-25-76 C 6-15-77
D 7—02-76 D 6-22-77
B 7-09-76 B 6-29-77
F 7-16-76 F 7-06-77
G 7-23-76 G 7-13-77
H 7-30-76 H 7-20-77
I 8-06—76 I 7-27-77
J 8-13-76 J 8-03-77
K 8-20-76 K 8-10-77
L 8-27-76 L 8—l7-77

61

62

OH

 

 

H S
S
N N
S

S H S HS SNS u HSS
H H SS SSS u HSS
S . H N SN SSS . HSS
S H S SN SSS u HSS
S H N H NN SSS n HNS
N H H SN SNS u HSS
N H SN SSS u HSS
S_ S N SN SSS u HSS
S H N SN SSS u HSS
S S H NN SSS u HNS
N H H HN SNS a HSS
S H S SN SSS u HSS
N H S H SH SSS : HSS
S H S N SH SSS s HSS
S H H N H NH SNS u HNS
S N S SH SNS : HSS
S H N H N SH SSS u HSN
S H S S H SH. SSN u HSN
HH S S S SH SSN . HSN
S N N S H NH SSN : HNN
SH . N N S S HH SNN : HSN
SH S H S N SH SSN r HSH
S H N N S S SSH n HSH
NH S S S S H S SSH . HSH
SH H S S S S N N SSH s HNH
NN H S S H N S SNH n HSH
SN . H S S S S N S SSH u HS
SS H N N N S N S S SS a HS
SN H H S S N S S S SS s HS
HN N N H H NH S S SN SN N SS a HN
SSH H H S S SH SH NH SH HH S. HN HS H SN u H
LEN PS E SN Sm NN SS N NH S SN S S
Houoa a x S H = o S S S o S < «SSHS SSSHo
.Soum SHSSS

Sumo oHdEuS scam you m:0HuanuuSHo SSSHU Socwsvoum SH£Q< SNSH

 

 

63

1976 Predator Frequency Class Distribution

 

 

 

Predator Frequency ‘ 'Sample‘Date
Count‘ Class ' D E F G Total

0 0 96 39 90 94 320
l 1 3 l4 2 20
2 2 1 ll 1 3
3 3 6 3 9
4 4 6 3 l 9
5 5 3 l 3
6 6 3 4
7 7 5 6;
8 8 4 4
9 9 1 l 3
10 10 2 2
11 ll 3 3
12 12 1 l
13 13 2 2
14 14
15 15
16 16
17 17
18 18
19 19
20 20
21 21
22 22
23 23

N
:5:
N
J)

 

 

64

S
H H
H
H
H H
H H
H H N
H H
H H
N N N
H N H H
S N N H
H
H H H H S
H N H N H
H H H S H
H H H H S H H
H S H S H N H
N H S H
H N N S S
H S S S S S S N
H H H S H N S OH S
H S N N S N HH H S H N
N N N H N S S S S H N
H H S S H S S HH SH HH S S
H NH S N S N SH SN SN S S H
S S 0N SH SH SH SS NN SN HS SN SS HH

SSN SSN SSN NSH NSH SSN SNH SSH OS NS SoH SON SSH SNN

Z 2 H x S H m U k m G U m <
mmumo mHmEmm

HS HSS
SS SSS
SN SSS
SN SSS
NN SSS
SN SNS
SN SSS
SN SSS
SN SSS
NN ISS
HN SNS
0N SSS
SH SSS
SH SSS
NH SSS
SH SNS
SH SSS
SH SSN
SH SSN
NH oSN
HH SNN
SH SSN
S SSH
S SSH
N oSH
S SNH
S SSH
S SS
S SS
N SS
H . H
o
mmmHU
.vmum

mmuma mHmEmS HHS mom mcoHuanHumHo Scamsvmum mucsou UHnmd NNSH

I HSS
I HSS
I HSS
I HNS
I Hom
I HSS
I HSS
I HSS
I HNS
I HoS
I HSS
I HSS
I HSS
I HNS
I HOS
I HSN
I HSN
I HSN
I HNN
I HON
I HSH
I HSH
I HSH
I HNH
I HSH
I HS
I HS
I HS
I HN
I H

o

mucsoo
SHsm<

 

 

65

 

H
H
S
H
H
H
H
S H
S
H
S N
H N N N H
S S S H H H
N S N S H S .
S S S N N S S H
S N S SNN NHH NNN SSN SSN
b H m 0 h m D U

mmumo mHmEmw

coHuanHumHo Nocmsvmum Houmwwum NNSH

chanLnuiNcnc\Serq
HIHr4r4Hr4r4~thvoaN

O
QHNMQ‘LOUDI‘mar—I

 

wmeU
Nocmsvmum

Or-‘IN
NNN

Or-levlnkcbmm
OHNMVmWPmer-{HHHHHHHH

unsou

uoumwmnm

 

   

APPENDIX D
Sample Site Means and Variances for
1976 and 1977
Appendix D contains the means and standard deviatidns
for the variables observed for 1976 and 1977. The

variables in 1976 were length of each shoot, number of

leaf promordia, leaves, and g, pomi. counts, an in 1977

 

leaf primordia, leaves, a, pomi and g, aphidimyza counts.

 

The results are sorted by sample site in a tree, thus,
for each date 10 site means are shown for 1976 and 24

in 1977 (refer to the methods section of this thesis).

66

 

 

7
6

 

 

o chcmmm¢

SS.SS 0S.ON SS.N oo.SH NS.S SS.N OS.N SH.SH ON OH
SS.NH SS.S 0S.N SS.SH SN.S OS.N SS.N SS.S ON S
SN.SN SS.SH NS.N SN.NH SS.S HH.S SSo.S SN.SH SN S
No.0S SS.NN NS.SH SS.SH SS.S SH.S NS.S SS.SH ON N
SH.SN SN.HH SSWN SO.SH HS.S SS.S HS.S SS.SH SN S
SN.NS OH.MN SS.H SN.NH SS.S So.N NS.H SH.S SN S
NS.NH OS.OH SN.S OS.NH SS.S SS.N SS.S SS.S SN S
SS.OH SH.N SS.W SN.SH SN.O OS.N SN.S SS.HH ON S
SN.HH SS.S SS.H SS.HH HS.S SS.N SS.N SS.S ON N
SHH.SH SH.S HH.N SS.NH SN.S ON.N NS.N SH.SH SN H

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68

 

 

a xHScomm<

SH.NN SS.SS SS.N SH.NH NS.H SS.N SS.N SS.SH SH SH
SH.SS SS.HS SS.N SS.SH SS.S SN.N HH.S SS.SH SH S
NS.SSS SH.SSS SS.N SN.NH SH.H SS.N SN.S SS.SH SH S
SH.SS SS.NSH SN.S SS.SH NS.S SN.S SS.S SN.SH SH N
SS.NS SS.SHH SH.S SN.SH SS.S SS.S NS.S SS.SH SH S
HS.SH SS.N SS.N SN.SH SS.S SS.S SS.N SS.NH SH S
HS.SH SH.SN SS.S SS.SH SS.H SS.H SS.S SS.SH SH S
HS.SN SS.NS SS.N SN.NH SN.S SN.N SS.N SS.NH SH S
SS.NSH SS.NS NN.N SS.SH SS.H SS.N NS.S SS.NH SH N
SN.SS SS.Hm NH.S SN.SH SN.S SN.N SS.S SS.SH SH H
.>GQ .fium Emmi .Gum cam: .>wD .Quw :mm: .>@Q .Cum cum: 2 wUHm
HEom mHzm< mm>mmq MHSHOEHHS mama AZHS. zumcoq SN\SH\S
wumo

 

69

 

 

NS.SNS SS.SSS NS.S SS.SH NS.H SS.H NS.S SS.NH SH SH
SH.NSN SH.HNS SS.N SH.SN NS.S SS.N SH.S SH.SN SH S
SH.NSH SS.NSS NS.H SS.SN SN.S SS.S NS.N SH.SH SH S
.HS.SSS SS.SHS NS.N SS.SN SS.S SS.S HNwS SS.NN SH N
NS.SSN SN.NSS SN.N SN.SN SN.S SH.S SS.S SS.NN SH S
SS.NS SS.NS” NH.S SS.SH SS.S SN.S SS.S SS.HH SH S
HSNSS SNWSS. NS.N SS.SH NS.S SH.S .SN.S SS.S SH S
SS.SSS SS.SNN NS.S SN.SH NS.H SS.H SS.S SS.SH SH S
SN.NSN SS.NSS SN.S SS.SH SH.SH SH.N SS.S SS.SH SH N
SS.SSSH SS.SSS SH.S SS.NH SH.H SH.N SS.S SN.SH SH H
.>mO .fium cam: .>mo .Gum cmwz .>mO .mum :mwz. .>mo .Sum :mm: 2 mun
HEom mHnm< wo>wmq menoeHum Smog AZHS zumcmq SNNSN\S
came

a xHScmeQ<

 

70

 

 

Q wacwmm<

NS.SSH SS.SHN SS.S SN.HN SN.H SS.H SS.S SN.SH SH SH
SS.SSN SH.HSS SS.N SS.NN SS.H SS.N SH.S SS.HN SH S
SS.SSN SH.HSS SS.N SS.NN SS.H SS.N SH.S SS.HN SH S
NS.SSN SN.SNS SS.S SS.SN SN.S SS.N NS.S SS.SN SH N
HS.SSN SH.SSS SH.S SH.SN. SS.S SS.N .SS.S SS.SN SH S
SS.SS SS.NH_ SS.N SN.SH SS.S SS.S SS.S SS.NH SH S
HS.SS SS.SS NS.S SS.NH SS.S SS.S SN.S SS.SH SH S
NS.SSH SS.SSH NS.S SS.SH NS.S SS.S SS.S SS.SH SH S
SN.SNNH SS.NSS SSSN SS.SN SS.S HH.H SS.S SH.SH SH N
SS.HNN SS.SSN SH.S SS.SH SS.H SS.H SN.S SS.NN SH H
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HEOQ mHzm< mm>woa MHSHOEHHS mama AZHS. :pmcoq SN\NS\N
mama

71

 

 

o XHccmmm<

mo.Nm mm.mm Nm.m mN.NN NN.H NN.H No.N NN.ON NH OH
ov.HOH Nm.NOH NH.S NH.mN NN.H NS.H mm.N mc.mN NH N
mN.NHH No.OHH NN.N Nm.mN NS.H mN.H mm.s om.vN NH m
Hm.mNH NH.mmH om.m om.SN SS.H oo.N mm.m mm.NN NH N
Nm.mm Nm.HHH HH.N mN.NN NH.H mm.N NS.S om.mN NH 0
HN.NS Nm.mH HN.N NH.SH oo.o oo.o om.m mo.HH NH m
mo.v4 mN.H mo.m om.SH oo.o oo.o sv.m oo.HH NH S
wH.mm Nm.mm HS.S NS.oN NN.H Sm.c NS.OH SH.ON NH N
NN.wOH Nm.mN_ mo.m NS.HN NN.H mN.H NS.N NN.HN NH N
SN.SN om.om NN.ON NN.SN NN.H .mm.H SN.N NS.HN NH H

.>ma .cum cam: .>mo .ouw :moz .>mo .oum cam: .>mn .cum saw: 2 mHHm

nHeom mHgm< mm>mmH «chomwum SmmH HzHc. zumcmH SN-NOIN

.. mama

 

72

 

 

O chcmmma

mm.H OO.O OO.O OO.ON NN.H ON.H OO.N OO.HN OH OH
OO.O OO.N ON.O OO.ON mm.H OO.H NO.O Om.HN OH O
OO.OOO OH.HOH OO.O ON.ON mm.H Om.H OO.O OO.NN OH O
HO.OO OO.ON Nm.N OO.NN ON.H OH.N -HONOH OH.ON OH N
OO.ON ON.NH Om.N ON.Hm OO.O ON.N Nm.N ON.NN OH O
OO.O OO.O HO.N OO.OH OO.O OO.O mH.O OH.NH OH O
O0.0. OO.O. ON.N ON.OH OO.O OO.O ON.O NN.OH OH O
OO.O ON.O ON.O ON.ON OO.O ON.O mm.N OO.ON OH O
OO.O OO.H. OO.O OH.ON OH.H OS.O OO.O OO.ON OH N
NH.NN OH,OH OO.O OO.HN OO.O ON.O Oq.HH OH.ON OH H
.>mO .Oum cam: .>mo .oum cam: .Owa .num cam: .>ma .cum saw: 2 mHHw
«Eon mHzm< mm>mmg MHSHOEHHS mqu. AZHV zumcoq SNISHIN
.. mama

 

73

 

 

o XHScmmm<

NS.H SS.S NH.N SS.SN SS.S SS.S HS.S SS.SN SH SH
NS.H SS.S SN.S SS.SN SS.S SS.S SS.S SN.SN SH S
SS.N SS.H SH.N SS.SN SS.S SS.S NH.S SN.NN SH S
HSSSS SS.SH SN.S SS.HS NS.S SS.S SS.S SS.NS SH N
SS.SS SN.NN SH.S SH.SS SS.H SS.H HS.N SS.SS SH S
SS.S .SS.S SS.S SS.SH SS.S SS.S SS.S SS.HH SH S
SS.S SS.S NN.N SH.SH SS.S SN.S, SS.S SH.S SH S
SS.S SS.S NH.S SH.HN HS.S SH.S SN.HH SN.SH SH S
SS.S SS.S SS.N SH.SS SS.S SS.S SN.HH SS.SN SH N
SS.N SS.N NN.N SS.SN SSS.S SS.S NS.S SN.SN SH H
.>mo .wum cam: .>m0 .cum :mmz .>mo .cum cmwz .>mo .vwm cum: 2 auHm
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74

 

 

Q XchQO<

SN.SH SS.S NS.N SS.SN SH.H SN.S NN.SH SS.SN SH SH
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HS.N SN.H SS.S SS.NS SN.S SS.S SS.S SS.SS SH S
SN.NN SS.HH NS.S SS.SS SS.S SS.H SS.NH SS.NS SH N
NN.H SS.S SS.N SS.NS NS.S SS.S SS.HH SN.SS SH S
SS.S SS.S NS.S SH.SH SS.S SS.S SH.S SN.NH SH S
SS.S SS.S. SH.S SSNSH SS.S SS.S HS.S SH.NH SH S
SS.SS SS.HH HS.S SS.SN SS.S SS.S SS.S SS.HN SH S
SS.S SS.S NS.N SN.HN SS.S SP.S SN.S SS.SH SH N
SS.S SN.S HS.S SS.SN SS.S SN.S SN.SH SN.SN SH H
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; mama

 

 

75

 

 

o xHSCmmm<

SS.S SS.S SS.S SS.SN NS.S SH.S SN.S SS.SN SH SH
SS.S SS.H HS.N SN.SN SS.S SS.S SS.S SN.SS SH S
SS.S SS.S SS.N SH.SN NS.S NN.S NN.S SS.NN SH S
SS.S SS.H SS.S SN.HS SS.S SS.S SS.S SS.NS SH N
SN.S .SS.N SS.N SS.SS MS.H SS.S SN.S SS.NS SH S
SS.S .SS.S SS.S SS.SH \SS.S SS.S SS.S SS.HH SH S
SS.S SS.S NN.N SH.SH SS.S SS.S SS.S SH.S SH S
SS.S SS.S SS.S SN.HN SS.S SS.S SS.NH SS.SH SH S
SS.S SS.S. SH.S SN.SN SS.S Sfl.S SS.NH SN.SN SH N
SS.S SS.S SN.S SS.SN SS.S SS.S SQ.S. SS.SN SH H
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_HEom mHzm< mo>wmq mHSEOEHum Smog. Asz. sumcmq SNISSIS
_. wbwo

 

76

 

 

o prcmmm<

SS.SS SS.NN SN.SH SS.SN SS.S SS.S SS.HH SN.SN SH SH
SS.S SS.S NS.N SS.SN SS.S SS.S HS.N SN.NN SH S
Sm.S SN.H HSNN SS.SS NS.S SH.S NS.N SS.NS SH S
NN.SS SS.NH SS.S SN.SS SN.S SS.S HH.SH r SHWSS SH N
SS.S HH.S SN.S SS.SS SS.S SS.S SHNS SS.SS SH S
SS.S SS.S NS.S SH.SH SS.S SS.S SH.S SN.NH SH S
SS.S SS.S HN.S SN.SH SS.S SS.S HS.S SH.NH SH S
SS.S SS.S SS.N SS.HN SS.S SS.S SS.SH SS.SN SH S
SS.S SS.S NS.N SN.HN SS.S S¢.S SS.S SS.SN SH N
SS.S SS.S SS.S SH.HN SS.S SS.S HSuSH. SH.NN SH H
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_. obmo

 

77

 

 

SS.S SS.S SN.N SH.SN NS.S SH.S SN.S SS.SN SH SH
SS.S SS.S SS.N SS.SN NS.S SN.S SS.S SN.SN SH S
SS.S SS.S SN.N SS.SN NS.S SN.S SS.S SS.SN SH S
SS.S SSrS SS.S SN.HS SS.S SS.S SS.S SS.NS SH N
NS.S SH.S SS.N SS.SS NS.S SN.S SHqSH SN.SS SH S
SS.S -SS.S NS.S SN.NH SS.S SS.S SS.S SS.SH SH S
O0.0 OO.O ONLN OH.OH OO.O O0.0 OO.S SH.N OH O
SS.S SS.S SS.S SN.HN SS.S SS.S SSSNH SS.SH SH S
SS.S SS.S SS.S SS.SN NS.S SH.S SS.SH SH.SN SH N
SS.S SS.S NS.N SS.SN NS.S SH.S SNuSH SH.SN SH H

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. muco

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‘78

 

 

SS.S SS.S SS.SH SS.SN SS.S SS.S SS.HH SN.SN SH SH
SS.S SS.S NS.S SS.SN SS.S SS.S SN.S SS.SN SH S
SS.S SS.S NS.N SS.SS NS.S SH.S SS.S SS.HS SH S
SS.S SS.S SN.SH SN.SS NS.S SN.S HS.SH SS.SS SH N
NS.S SH.S NH.S SS.SS SS.S SS.S SS.S SS.SS SH S
SS.S SS.S NS.S SH.SH SS.S SS.S SH.S SN.NH SH S
SS.S SS.S HNVS SN.SH SS.S SS.S SS.S SH.NH SH S
SS.S SS.S HS.S SS.SN SS.S SN.S SS.S SS.HN SH S
SS.S SS.S NS.N SN.HN SS.S SS;S SS.S SS.SN SH N
SS.S SS.S SSJSH SS.NN SS.S. SS.S SS.SH SS.NN SH H
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79

 

 

o .o SN.SN ON.S SSS.N .ow.SH mo.H ON.N OH SN
6 o SH.N om.m SSS.“ ON.NH Sm.c OS.N 0H SN
0 o . o o SSS.N ON.mH ON.O OS.N OH NN
o O SS.H ON.O SSS.N CH.NH NS.O o.N OH HN
o o o O SSM.N om.mH mm.o om.N OH ON
C. o o o SNN.H OS.SH Nm.c om.N OH SH
0 o o o NSN.H ON.NH mm.o om.H OH SH
0 o O o SmS.N OO.SH Nm.c OS.N OH NH
o o o O NSS.N om.SH SO.H OS.N OH SH
0 o Sm.cm OS.NH NSN.H om.NH Sm.c OO.N OH SH
0 o o o mmm.H oo.mH mo.H oo.N OH SH
0 o o o HHo.N OS.SH ON.O OS.N OH SH
0 o Sm.c ON.O OHN.N ON.mH SH.H cm.H OH NH
o o NN.H OS.O oom.N om.SH SH.H oo.N OH HH
o o o o OMS.H OS.SH NS.O OS.N OH OH
o o o o NSS.H OS.SH Sm.c ON.H OH S
o o. O o SHS.H OS.SH Nm.c Om.N OH S
o O HH.S om.H NSS.H ON.SH Sm.c om.N OH N
o o o o NHm.H OS.SH NS.H om.H OH S
o o .o o mSm.H om.SH Sc.c OS.N OH m
o o o o oom.H om.mH Sm.c om.N OH S
o o SS.N o¢.o SHN.H omth ON.H OH.H OH S
o o o o HHN.m om.MH mfi.o. ON.N OH N
O O .O O HHO.N ON.OH Nm.O OH.O OH H
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S chcwam<

 

 

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O O SS.N OS.O SS.N OS.SH hH.H OS.H OH HN
O O O0.0 O0.0 SN.H ON.SH NS.O OS.N OH ON
O O SQ.H ON.O SS.S OS.SH NN.H O¢.N OH SH
O O HQ.SH OS.H NS.N OV.SH SH.H OH.N OH SH
O O bv.v OS.H SH.N OS.bH SS.O OS.N OH hH
O O O0.0 O0.0 SN.S OS.SH SO.H OS.N OH SH
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O O SS.S OS.O SS.Q ON.hH VS.O O¢.N OH VH
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O O SN.N Ob.H SS.N Oh.SH NS.H ON.N OH OH
O O SmuH OS.O HS.H SS.SH SN.H ON.H OH S
.0 O OH SSaN OO.H SS.N OH.¢H NS.H OS.H OH S
8 O O Oh.m ov.v hH.N OS.mH hm.O OS.N OH 5
O O SN.SH ON.¢ VS.S OS.VH NN.H OS.O OH S
O O HN.HH OS.S HO.N OS.VH NS.H OH.N OH S
O O SS.HH ON.S vH.S OS.SH SS.O OS.N OH v
O O SH.S OO.H SN.N OS.dH SS.H ON.O OH S
O O SN.SH OH.S SS.N OO.¢H VHtH. OS.H OH N
O O HSJSH ON.S SS.S OS.SH ON.O OV.N OH H
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anecwmmm‘qumdwvflnmw HEOQ mwnmfi mm>mmA mficuofiflnm wweq ONIQHIS
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D xficcmmmd

 

 

 

 

Q XHocmmm<

O .O SS.SN ON.S SN.N OS.SH NN.H OO.N OH ON
O .O SS.v VS.N vv.S .Om.hH hS.O OS.N OH SN
O O O0.0 O0.0 SN.N OH.hH S¢.O Oh.N OH NN
O O OSsH OS.O vN.S OS.NH SN.H OS.H OH HN
O O O0.0 O0.0 HN.S OH.bH SS.O OS.N OH ON
O O hH.Oh OS.NN NN.N OS.SH v5.0 OS.N OH SH
O O O0.0 O0.0 mv.N OO.SH Sv.H OS.H OH SH
O O Hh.hH OS.S SN.S ON.SH VH.H OS.N OH NH
O O SquN OS.S NN.S OS.SH hH.H Ov.N OH SH
O O SS.H OS.O vm.N Oh.mH SN.H OS.N OH mH
O O O0.0 OOuO vS.S OS.NH bH.H O¢.N OH «H
O O O0.0 O0.0 OH.S. OS.hH SO.H OSwN OH SH
O O SS.N OS.S ON.S OV.QH SH.H OS.H OH NH
O O QN.HH Oh.m SS.N OH.SH O¢.H ON.N OH HH
O O OS.VH OS.v NN.N Ov.hH hm.O OS.N OH OH
O O HN.h OS.N SS.N OmeH NS.H OS.H OH S
O O Sv.m OH.H NN.N OH.mH NS.H OS.H OH S
O O VSOSN OV.OH SO.S OS.SH NS.O ON.S OH 5
M O O SS.O OH.O Sv.S OO.VH NS.H OS.O OH S
. O O SS.SH OS.H OS.S O¢.mH SN.H OS.H OH m
NS.O OH.O SH.vSH OS.v HS.S OS.SH NS.H ON.N OH v
O O O0.0 O0.0 SS.N OO.mH SS.O OS.O OH S
O O .ON.S ON.v OS.N ON.SH SN.H Oh.H OH N
O O OO.SN OS.SH Hh.N Oh.mH HW.O. OS.N OH H
.>00 .fium cam: .>wO .vuw :mwz .>wD .Cum C662 .>wO .Gum cum: 2 muHm
mnwscwnmm mmwmmvcflnmfl HEom mflnmm mm>wmq .wHUHOEHHm Mdmn hhlmalm
mama

82

 

 

O .O OO.OHH OO.OO ON.O ON.OH O~.H OH.~ OH ON
O O OH.OO Om.HO OO.O OO.OH OO.H OO.H OH OH
O O mOsmm OO.OO OO.O OH.ON OO.H OO.~ OH NH
O O OO.Om OO.Hm OH.O OO.OH ON.H OO.H OH HO
O O OO.OOH OH.OO OO.O OO.ON OO.O OH.~ OH OH
mO.O OO.O OO.HO OO.OH OO.O O~.H~ mm.H Om.m OH OH
O O OO.OH OH.OO OO.H OO.OH OO.H OO.H OH OH
O O ON.OH OO.O OH.O OO.OH Nm.H OO.H OH OH
O O OO.OH ON.OH OO.O OO.OH OO.O OO.~ OH OH
O O ON.OH OO.O OO.O OO.ON m~.H OO.O OH OH
O O OO.O OO.O OO.O OO.OH OH.H ON.O OH OH
OO.O OH.O OO.OH OO.ON ON.O OO.- OO.O OH.O OH OH
O O OO.HH OO.O OO.O ON.OH OO.H OO.O OH NH
OO.O OO.H OO.OH OH.ON OO.O OO.OH OO.H OO.H OH HH
O O ON.OO OO.- OO.O OO.ON OO.H OO.O OH OH
O O OO.OH OO.O OH.O ON.OH OO.H Om.H OH O
O O OO.O OO.O OO.O OH.OH m~.H O~.H OH O
O O OH.ON OO.OH OO.O OO.OH ~O.H O~.~ OH O
O O OO.O OO.H OH.O OH.OH OO.O OO.O OH O
O O OO.ON OO.OH OO.O ON.OH HO. Om.H OH O
O O OO.OO OO.OO OO.O OO.OH OO.O OO.~ OH O
OO.O OH.O mO.O OO.O OH.O ON.OH mO.O OO.O OH O
O O OO.ON OO.OH OO.O OO.OH OH:H Om.H OH H
O O .OO,~O OO.HH HO.O OO.OH O~.H Om.H OH H
.>mO .Oum cum: .>mO .cum cam: .>mO .cum cams .>mo .Oum cum: 2 mHHm
quangmm mmumHocHgmH Haom mHOOH mm>mmH .mHOHoEHum ummH OnummuO
. mama

O chcmmmd

 

r w w

83

 

 

Q Xchmmm<

O .O HS.vS OS.NV SS.S .OS.SH hS.O OV.O OH «N
Nv.O ON.O bH.vv ON.Sv HN.m OS.NH SO.H OS.O OH SN
SS.O ON.O S¢;vh O0.00H hv.v OO.SN SS.H OS.H OH NN

O O SS.SS OO.vN SS.S ON.SH SN.H OS.O OH HN
SS.O ON.O HS.SOH ON.SS NS.v OH.SN SN.H OS.H OH ON

O O S0.0h OS.Sm SS.S OO.hN SS.H OS.H OH SH

O O SS.S ON.OH Nm.v OS.SH SH.H OO.H OH SH
SS.O ON.O OS.Sm ON.SS NH.S OS.ON SO.H OS.O OH NH

O O SN.HS ON.NS SH.v ON.SN NS.H OS.H OH SH
NS.O OH.O OS.NS OS.SH h¢.v ON.HN SO.H OS.O OH SH

O O NS.Sh OH.mS Oh.m OS.SN NS.H OS.H OH VH
SS.O OS.O HS.SS OS.SS SH.S OO.mN hS.H OH.N OH SH

O O SS.S OO.S HS.S OS.SH O0.0 O0.0 OH NH

O O SS.OS OS.Sv NN.¢ OS.ON db.O OS.O OH HH
bh.H OS.O SS.SS OS.SB NS.S OS.NN SS.H OS.H OH OH

O O OS.N Oh.H Nv.N OH.HN ON.O O¢.O OH S

O O. vHsHv O¢.¢N OS.S OS.SH Sv.O OS.O OH S
Hh.O m.O SH.SvH OS.OSH Sh.N O¢.SN Sh.O ON.N OH 5

O O HN.m OS.N Hh.S OS.ON NS.O OH.O OH S

O O SS.SN OS.NS HO.S ON.ON SS.O ON.O OH m
vS. Ov. SS.SS OS.SS SS.S ONJVN SH.H OS.H OH H

O O HN.N OO.H NS.S ON.ON O0.0 O0.0 OH S

O O SS.SS OB.SN SS.S OO.HN SN.H. ON.O OH N
SS.O ON.O S0.00H OS.Sh hv.S ON.HN Hv.H OO.H OH H

.>ma .cum cam: .>mo .cum 2mm: .>mo .cum cam: .>mo .Oum cum: 2 muHm
mumewflamwzmmumwoonmfi _HEom mflsmd wm>mmq .wHOHOEHHm Mqu hulmmlm
oumo

 

84

 

 

a wacmmm¢

o o SS.SH om.NH Sh.o om.mm mv.o ON.O m vm
Hm.o OH.O mo.mm oo.mv H¢.~ om.m~ om.o OH.O m mm
vm.H OS.O mn.m¢H OO.HvH NS.H OS.SN OO.H OO.H m mm

o o mmna ON.O SS.H om.o~ mv.o ON.O m Hm

o o om.mH ov.mH mo.m om.m~ mm.o OH.O m cm

0 o mm.hv o~.m> mm.m om.hm Nm.H ov.H m mH

o o Nm.c OO.H NS.H OH.ON vw.o OS.O m mH

o o mv.SH oo.~H Hm.m OS.NN om.o OH.O m OH
m>.H om.o ~m.om oo.mm om.o om.mm mm.o OS.H m SH

0 o mm.w OO.H NS.H om.- mm.o OH.O m mH

o o mm.H~ O0.0H Sc.c om.m~ mm.o OH.O m «H
mm.m OS.H Sm.o5 oo.mOH mo.e om.- OO.H OO.H m MH

o o Nm.cm oo.mH mm.v ov.mH O0.0 O0.0 m NH

o o mm.m~ ON.OH m~.m OS.SH oo.o oo.o m HH
mv.o ON.O Hm.mm oo.mmH H~.m o¢.m~ OH.H om.H m OH

O o mo.mm oo.vH mm.m ON.ON vm.H OS.O m m

o o Hm.mH oo.mH mm.v OS.SH mm.o ov.o m m
SS.N o~.H ov.mm OO.HHH vh.v oo.m~ Hb.o OO.H m h

o o oo.o oo.o OS.H ow.mH oo.o oo.o m m
mm.m OS.H «m.SOH ov.mm H~.m OS.NN mm.o OH.O m m
mh.H om.o oo.mm oo.mOH Sw.m ON.HN OO.H OO.H m w

o o oo.o O0.0 Nm.H OS.HH oo:o oo.o m m

o o vm.mH ON.O so.m om.mH O0.0 O0.0 m N
mv.o om. Nm.o> ow.mm mH.m ov.v~ Hm.o cS.c m H
.cum cam: .>wo .oum :mm: .>mo .Oum :mmz .>mo .Uum saw: 2 muHm

mumEOHnmm mmumHOUHSQ< HEom mH£m< mw>mmq MHOHOEHHm mama thSOIO
mama

85

 

 

O O O0.0 O0.0 MN.m Om.OH O0.0 O0.0 OH ON
O O OH.mN O0.0 OO.m ON.OH O0.0 O0.0 OH mN
m0.0 ON.O Om.HOH O0.0HH OH.O OO.HN O0.0 Om.O OH NN
O O mm.mm om.NH mm.m om.mH O0.0 O0.0 OH HN
Nm.c OH.O mm.mOH O0.0m OH.H OO.NN O0.0 O0.0 OH ON
O O mN.HHN OO.MMH OH.O OO.mN O0.0 OH.O OH OH
O O NO.HH om.m NN.H Om.OH O0.0 O0.0 OH OH
O O mm.mN om.NH O0.0 Om.NN Nm.O OH.O OH NH
m0.0 ON.O OH.OOH O0.00 ON.O om.vN m0.0 ON.O OH OH
O O HO.mv ON.NN OO.H OH.mN O0.0 O0.0 OH mH
O O Hm.NHH OO.NH Hm.m om.HN Nm.O OH.O OH OH
Nm.c O mHanH ov.hm NO.» .ON.ON SH.H O0.0 OH mH
O :O O0.0 O0.0 OH.m OO.mH O0.0 O0.0 OH NH
Om.H om.O ON.HN O0.0 mH.O ON.ON O0.0 O0.0 OH HH
OO.H O0.0 OO.HO OH.Om mm.v om.NN Nm.O OH.O OH OH
O O O0.0 O0.0 SH.N om.HH O0.0 O0.0 OH O
O O Nm.c OH.O NN.H O0.0H O0.0 O0.0 OH O
Nm.H O0.0 mm.mm O0.00 NH.m ON.HN HbuO om.O OH O
O O Om.vOH OO.mm ON.O ON.OH m0.0 ON.O OH O
O O O0.0 OH.O ON.m OO.mH O0.0 O0.0 OH m
O O OH.ONH ON.mO O0.0 om.NN O0.0 OM.O OH O
O O O0.0 O0.0 mv.N om.mH O0.0 O0.0 OH m
O O OO.H OOuHH mO.m O0.0H O0.0 O0.0 OH N
Om.H om.O Hv.mm O0.0H OO.m om.HN Nm.O OH. OH H
.>mo .wum cam: .>ma .Uum‘ cmwz .>ma .cum 2mm: .>mo .Uum 2mm: 2 muHm
mustcHnmm mmumHowH£m< HEOQ‘OHnmm mm>me mHOHOEHHm mama hptMth
mama

a xHOnmmm<

86

 

 

O0.0 O0.0 O0.0 ON.O. HH.m O0.0N O0.0 O0.0 m «N
ON.N OO.H O0.00 O0.0H O0.0 O0.0N O0.0 O0.0 m mN
ON.O O0.0 OO.Nmm O0.00H OO.H ON.ON O0.0 O0.0 m NN
ON.N OO.H O0.0HH O0.00 O0.0 O0.0N O0.0 O0.0 m HN
OH.O ON.O O0.00H OO.HO NO.m O0.0N O0.0 O0.0 m ON
O O O0.00H O0.00 NO.m O0.0N O0.0 O0.0 O OH
O O O0.0 O0.0 HN.O O0.0H O0.0 O0.0 O OH
OO.H O0.0 NN.OHN O0.00H O0.0 O0.0N O0.0 ON.O O OH
OH.O O0.0 OH.OOH ON.OOO O0.0 ON.Nm O0.0 O0.0 O OH
O0.0 ON.O OH.N OO.H Hm.v O0.0N O0.0 O0.0 O OH
HO.m OO.N ON.Om O0.0N OO.N O0.0N O0.0 O0.0 O OH
O0.0 OO.H NH.OON O0.00H NO.H ON.HO O0.0 O0.0 O OH
O O OH.HOH OO.NO Nm.m O0.0H Nm.O OH.O OH NH
O O O0.00 O0.0H O0.0 O0.0H O0.0 O0.0 OH HH
HH.¢ OO.H O0.0HH OO.HO O0.0 OO.NN O0.0 O0.0 OH OH
O O O0.0 O0.0 O0.0 ON.OH Nm.O OH.O OH O
O0.0 O0.0 OO.NO O0.0H OH.O .Nm.OH Nm.O OH.O OH O
OO.N OO.H OH.OO OO.HO OH.O OO.NN Nm.c OH.O OH O
Nm.O OH.O O0.00H OO.NO O0.0 O0.0H O0.0 O0.0 OH O
O O O0.0 O0.0 O0.0 O0.0H O0.0 O0.0 OH O
OO.N O0.0 HO.HOH ON.Om O0.0 ON.NN NN.O OH.O OH H
O O O0.0 O0.0 O0.0 O0.0H O0.0 O0.0 OH O
O O O0.0 O0.0 O0.0 O0.0H O0.0 O0.0 OH N
ON.O O0.0 O0.00N ON.OOH O0.0 O0.0N Nm.c OH.O OH H
.>ma .Oum 2mm: .>mo .cum cum: .Oum :mmz .>ma .Oum cmmz z oaHm
MOOEHUHQQO mmumHocHamd HEom mHnm< mm>me MHOHOEHHO wmmH OOIONIO

O xHUcmmmd

mama

O0.0

 

 

O O O0.0 O0.0 O0.0 O0.0N O0.0 OH ON
Nm.c OH.O OO.NOH OO.NO O0.0 O0.0N O0.0 ON.O OH ON
OO.H O0.0 OO.HHH OH.OH O0.0 O0.0N O0.0 O0.0 OH NN
O O O0.0 O0.0 O0.0 ON.OH O0.0 O0.0 OH HN
OO.H O0.0 O0.00 O0.0m O0.0 O0.0N O0.0 O0.0 OH ON
OO.H O0.0 O0.0v O0.0N OO.HN O0.0m O0.0 O0.0 OH OH
O O O0.0 OO.H O0.0 O0.0H O0.0 O0.0 OH OH
O O O0.0 ON.O HN.O OO.NN O0.0 O0.0 OH OH
O O Hv.mHH O0.0m ON.O OH.ON Nm.c OH.O OH OH
O ..‘O O0.0 OO.H O0.0 OH.ON O0.0 ON.O OH OH
O0.0 O¢.O H0.0m OO.HN O0.0 ON.ON O0.0 ON.O OH OH
Nm.O OH.O O0.0m O0.0H NN.O O0.0N O0.0 O0.0 OH OH
Nm.c OH.O O0.0 O0.0 O0.0 O0.0H O0.0 O0.0 OH NH
O O O0.00 OO.NN NH.O .O0.0N Nm.O OH.O OH HH
W O0.0 ON.O OH.HOH O0.00 O0.0 OH.ON OO.H O0.0 OH OH
O O O0.0 ON.O O0.0 O0.0H O0.0 ON.O OH O
O O O0.0 O0.0 O0.0 O0.0N O0.0 ON.O OH O
O O O0.0H O0.0H O0.0 O0.0N O0.0 O0.0 OH O
O O O0.0 O0.0 Hv.m O0.0H O0.0 O0.0 OH O
O O O0.0 O0.0 HO.N O0.0H O0.0 O0.0 OH O
N0.0 OH.N NO.mNH OH.OO Nm.O O0.0N O0.0 O0.0 OH H
O O O0.0 O0.0 NO.N O0.0H O0.0 O0.0 OH O
O O O0.0 O0.0 OO.H O0.0H O0.0 O0.0 OH N
OO.H O0.0 HN.OO O0.00 O0.0 O0.0N O0.0 O0.0 OH H
.>wD Auw .. :mmz .>0Q .Oum 2mm: .>ma .Oum :mmz. .>mo_.wum saw: 2 muHm
mumaHOHnmm mmumHOOHsm< HEOQ wH£m< mw>me MHUHOEHHm mmmH OOIONIO

Q xHUcwmmd

u-
.1

J“

 

88

 

 

O0.0 O0.0 ON.OO O0.0N OO.N O0.0H O0.0 O0.0 OH ON
O O O0.0 O0.0 OO.H O0.0N O0.0 O0.0 OH ON
O O O0.0m O0.0H O0.0 O0.0N O0.0 O0.0 OH NN
O O Nm.OH O0.0 O0.0 O0.0N O0.0 ON.O OH HN
O O OO.HO O0.0H ON.O O0.0N O0.0 ON.O OH ON
O0.0 O0.0 O0.0v O0.0m O0.0 O0.00 OO.H O0.0 OH OH
O0.0 ON.O OO.HOH O0.00 O0.0 OO.HN O0.0 O0.0 OH OH
O O OO.vv O0.0H O0.0 O0.0N O0.0 O0.0 OH OH
O O OO.HO OO.HH O0.0 O0.0N O0.0 ON.O OH OH
O O O0.0 O0.0 ON.O O0.0N O0.0 O0.0 OH OH
O0.0 ON.O O0.0HH OO.NO ON.O O0.0N O0.0 O0.0 OH OH
ON.H O0.0 O0.00H OO.NO O0.0 O0.00 OO.H O0.0 OH OH
O O ON.O OO.H O0.0 O0.0N O0.0 O0.0 OH NH
O O O0.0 O0.0 OH.O OO.HN O0.0 O0.0 OH HH
O O ON.OH O0.0N N0.0 ON.HN OO.H O0.0 OH OH
O O NO.¢¢ .O0.0N OH.O .O0.0H O0.0 ON.O OH O
O O O0.0 O0.0 O0.0 ON.ON OO.N O0.0 OH O
O O ON.OOH OO.HO O0.0 O0.0N Nm.H O0.0 OH O
Nm.c OH»O H¢.ON OO.NH OO.H O0.0H Nm.O OH.O OH O
O O NH.O ON.O O0.0 O0.0H O0.0 O0.0 OH O
Nm.c OH.O HOO.mN O0.0 vH.m OO.NN O0.0 OH.O OH O
O O ON.H O0.0 OH.O O0.0H O0.0 O0.0 OH O
O O O0.0 O0.0 O0.0 O0.0H O0.0 O0.0 OH N
O NO.v OO.H HN.O OO.HN OO.H O0.0 OH H
.>mo .cum 2mm: .>wa .ouw :mmz .>mo .Oum :mmz .>mo .Uum cum: 2 muHm
w~>EH©Hzmm mmumHocHnmm Heom‘mH£m< mm>mmH MHOHOEHHm wmmH OOIOOIO
muma

Q wacmmmd

 

 

APPENDIX E

APHID AND PREDATOR BIOLOGICAL OBSERVATIONS

In the following appendice "four tables are presented.
The first is the raw counts of terminals in several quadrats
of the tree (see also Results). The second are the results
from the examination of the 200 terminals, from 1976
sampling, for A. pgmi_eggs.' The data also shows the seasonal
growth total for each terminal. The third table provides
the aphid mean/terminal for each tree sampled in 1976 and
1977. The final table presents results of emergence cage

trapping of adult Aphidoletes aphidimyza.

 

89

Table E1.

Estimated for Lower and Upper Quadrat Terminal Numbers,

90

Taken at Graham Station 1977;

Tree

UlH

U1KOkD\)\IUI

Tree # ' Lower
2 107
3 68
4 68
6 37

10 73

12 78

15 66

17 47

29 93

33 54

34 47

if 67 . 09

S 20.75

N 11

Quadrat
NE
NE
SW
NE
SW
NE
SW

SW

 

 

‘Upper

116.

71

108

32

100

86

82

63

129

165

90

94.73

35.40

11
Canopy Level

Lower Upper

57 64
25 36
28 36
41 38
50 41
53 57
52 56
52 46

 

91

The following table presents the results from
-examining the terminals used in 1976 sampling for A, pomi
eggs. In all a total of 5 eggs were found, suggesting
limited oviposition occurred within the Red Delicious

block.

 

.1

J1.

 

92

Table E2.
Egg Deposition and shoot length on red delicious trees..
Samples taken in February 1977 prior to pruning and egg

hatch. Most terminals were those observed in 1976.

 

Terminal Length

Code In’ Status Eggs
0111 22 0
0121 26.5 0
0131 9.5 0
0141 16.5 0
0141 14 0
0112 35.5 0
0122 50 0
0132 31.5 0
0142 30.5 0
0152 37 O
0311 28 0
0321 35 0
0331 26 0
0341 13 0
0351 8 0
0312 33.0 New 0
0322 33 0
0332 35.5 0
0342 38.5 0
0352 25 0
1111 69 0
1121 5.5 O
1131 17 0
1141 17 0
1151 12 0
1112 16 New 0
1122 30 0
1132 18 0
1142 ll 0
1152 19.5 0

 

 

 

93

_ Table E2. (Cont'd)

 

Terminal Length

Code ‘ 'In’ Status
1311 26.5

1321 24.5

1331 20.5

1341 6 Broken
1351 31 New
1312 39

1322 58

1332 30.5

1342 35

1352 39

2111 22

2121 24

2131 11.5

2141 10

2151 9

2112 33

2122 31

2132 15.5

2142 22

2152 15

2311 23.5

2321 17

2331 20.5

2341 9

2351 19

2312 39.5

2322 32

2332 35 New
2342 4 Broken
2352 31

3111 38

3121 11.5

3131 5

3141 9.5

3151 7.5 New
3112 20

F5

OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOCOOOOO

 

Table E2.

Terminal
Code

3122
3132
3142
3152
3311
3321
3331
3341
3351
3312
3322
3332
3342
3352
4111
4121
4131
4141
4151
4112
4122
4132
4142
4152
4311
4321
4331
4341
4351
4312
4322
4332
4342
4352
5211
5221
5231

(Cont'd)

Length
In’

17
16
23.5
14
29.5
11
14

6
7.5
26
28
25
24
21
94
42
9.5
9

15
39
38
23.5
27
28
18
38.5
31
18
15
48.5
37
38
23
33
28.0
28.5
20

Status

New

New

New

New

New
New

New

1%

 

Table E2.

Terminal
Code

5241
5251
5212
5222
5232
5242
5252
5411
5421
5431
5441
5451
5412
5422
5432
5442
5452
6211
6221
6231
6241
6251
6212
6222
6232
6242
6252
6411
6421
6431
6441
6451
6412
6422
6432
6442

 

(Cont'd)

Length
. In .

23.5
15
33.5
44
34

34.5
36.5
29
25
24
20.5
12
13
11
24
17
21.5
25

'Status

New

New

m
OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO ‘
U)

96

Table E2. (Cont'd)

 

11:53

Terminal Length
Code In Status
6452 23.5
7211 33
7221 11
7231 43
7241 18 New
7251 9.5
7212 19
7222 18
7232 28
7242 28.5
7252 30.5 New
7411 6.5
7421 8
7431 10.5
7441 11
7451 27.5
7412 18
7422 27
7432 27.5
7442 17
7452 11.5
8211 20
8221 32.5
8231 27.5
8241 12.5
8251 13.5
8212 47.5
8222 43.5
8232 23
8242 37
8252 35.25;
8411 15.25
8421 17.75
8431 21.5
8441 '21

8451 11.5

OOOOOOOOHOCOOOOOOOOOOOOOOOOOOOOOOOOO

 

97

Table E2. (Cont'd)

t1:
LO.
S
m

Terminal Length
Code In Status
8412 35
8422 4.8
8432 44
8442 30
8452 27
9211 37.5
9221 10
9231 3.5 Broken
9241 7.5
9251 4.5 - Broken
9212 32
9222 14.5
9232 27
9242 19.5
9252 8.75
9411 52
9421 25
9431 19
9441 12.5
9451 12
9412 20
9422 17
9432 33.5
9442 23.5
9452 34
200 Terminals
Mean Length = 23.93 S = 11.98

Total Length 4786.5 Inches
(398.88 Feet)

COOOOOOOOOOOOOOOOOOOOI—‘OI—‘N

 

98

O0.0V

 

 

NS.SS OO.nv ON.HO ON.OO H0.0N O0.0H O0.0 O0.0H OH.O «N.S SN.H.
N.OO OS0.0v N.ON OHv.N m.NH mOS.H n.O mNS.O N.v SS0.0
O.mO OSH.NO O.SS NvO.Sm m.NH NON.O 0.0N OSS.S O OO0.0
0.00 OON.OS O.NO OH0.0N 0.0H mNH.S 0.0N NOO.N S.O OON.H
.O.SS OHO.HS 0.0m NQ0.0 0.0N OHO.S 0.0N ONS.m S.S Nvm.O
m.NO ONO.SN m.O nnm.o 0.0H NO0.0 0.0S OOO.¢ m.O NvO.H
0.00_ OHS.NS n.SO OHO.HO O.NO mO0.0H S.O NON.O m.NH OOS.v m.O SOO.H
m.NS OSH.Om 0.00 SOO.Sn n.nO NON.OH N.ON NOO.N m.NH OO0.0 O.NH OOv.O
O.mh mNH.NS N.OO NOO.mm m.NO OOO.SH N.ON OOO.NN n.O ONS.O N.¢ OOH.O
n.Om mNH.vm N.vO OO¢.SN 0.00 OH0.0N 0.0 OO0.0 O.mm OOH.S N.v ONH.O
.S.Om Nvm.Nn 0.00 OO0.0S 0.00 SOO.Hv O.NH mOm.HH 0.0H OOH.N O OO0.0
fl C602 & 6mm: v flflwz w flmwz fl cmwz w CSO:

h m o

 

 

 

OOOH

 

 

nouns onEmm

«HacHEuoa coummucH uo unmouom can HON u zv owns 20mm Hem memo: oHsma

.Sm wHQmB

mama:
mama

OH

“1th

O owns

 

99

 

 

 

 

 

NH.ON O0.0H NH.OH O0.0H H0.0v HS.NO OO.HO OH.HO
N.¢O OO0.0¢ O.NH OHv.O 0.0H OOH.OO O.NO OON.HO
O O 0.0H OOH.¢O 0.0N SOO.mm 0.0N NvO.m
O O m.O Nv0.0H O.Hv OH0.00 N.v Nv0.0
O.NH mmm.v m.O OOH.H O O 0.0N OO0.0m
m.O OOO.H O O 0.0N OH0.00 0.00 mmm.ON
N.O ON0.0 O.NH NON.O O.NH .ONH.O N.OO OO0.00
0.0m OOS.HS O.mN OHv.mH N.¢m NO0.0SH O.Hv mNS.OS
O.NH OH0.0H O.Hv OON.OO O.mm OON.NN O.NH OOO.v
0.0N OH0.00 0.0H OOO.mH 0.0N OO0.0H 0.0m ON0.00
0.0m NOO.N¢ O.NH ONH.O 0.00 OOH.NNH O.mm OO0.0N
w cme w cmmz O cmmz O com:
O m U

mama mHmEmm

 

wand:

mama

OH

O mmHB

mHmcHEume OmpmmmcH mo unmoumm cam mammz ©H5m<

Hw.ucouv

.Sm anma

 

 

100.

In 1977 emergence cages were distributed singly at the
base of ten trees. Cages were examined weekly and the number
of Syrphidae, Cecidomyiidae, and Chrysopidae adults counted.
These are the total catches each Week.

On every third sample date.the traps were moved in close
proximity to the prior site under the tree. This was to
capture Cecidomyiidae which had dropped to pupate in the soil

area around the trap, in the previous two weeks.

 

 

101

Hmumumousmzv mmeQOmmuno
AmumuOHOO mmOHHOeowHomo

HmumumHQO mMOHnmuwm

 

 

 

 

O ON S OHS
v SS ON SOS
S NS OH _ ONO
S SN O ONO
O OH N SHO
H OH O SOO
O Sm _ S SNS
O OH OH NNS
O HS OH OHS
H SN S OHS
O S O NOS
O OHH OS SNS
H SS ON SHS
O OH O HHO
O O OH OOO
mmoHQOmhunu wmcHHNEOUHowU mmcHnmuhm GHmQ

:ovaum Emnmuu um OOOH CH muoumwmum uHsod mo monoumu mmua mocmmumfim

.vm mHnt

 

APPENDIX F
Sample Size Estimation

IntroductiOn

 

The organization of the following appendices is to show
the steps involed in taking raw sample data through several
distributional analyses. The distribution coefficients
estimated were “ and B of the mean crowding-meanrelationship

(Iwao and Kuno 1968), k and kc of the negative binomial
(Bliss and Fisher 1953, Elliott 1971) and a and b for the
mean-variance relationship (Taylor 1961). Using the
appropriate formulas sample size estimates of several sample
designs are computed.

In the analysis which follow the results are shown
primarily for the 1977 data. Samples were not taken randomly
in 1976 and these initial attempts at defining the habitat.
sample unit and the sample universe are occasionally included
in an analysis as a general index only. Loss of randomness
makes interpretation of the 1976 data difficult and further
analysis unwarrented.

The sample universe chosen was the tree, with in which
the habitat sample unit, the apple terminal, was located.
Stratified sampling conducted in 1977 (see Methods, main
text) allowed several sample designs to be tested. Four

sample designs were selected and distribution analysis using

102

 

103

both A. pomi and §.'aphidimyza performed for designs

 

where sufficient data existed.
Throughout all analysis, the total tree sampling
scheme (N - 24) is presented for both A. pomi and A.

aphidimyza. Analysis is also presented for the lower and

 

upper canopy sampling quadrats. These were chosen for
accessibility and host growth-high aphid density charac-a
teristics, respectively. Finally, data from the four lower—
inner canopy positions (at the center of the lower canopy)
in analyzed and presented where sifficient information
existed.

The analysis were preformed using the estimates per
tree from each sample date. These were also used as
estimates when combining all dates. Thus, results presented
include each sample date and the combination of all dates

for a sample design.

Mean Crowding and the Mean

 

Calculation of Iwao and Kuno's (1968)a:and 8
required obtaining estimates of mean and m*, a mean crowding
index proposed by Llyod (1967). Using the estimates, linear
least squares regression was performed to obtain a and 8
for each sample date and for the total season. Tables F1 and
F2 present the 1976 and 1977 values of a, B and r2 for A. pomi

on each sample date and for the total season.

 

 

104

Using the total season values (1977) sample size
-estimates in the three sample design for A. pomi are
presented in Table F3. The formula used in their

computation was

 

q = t2/D2 («+1 +3-1)
32
where q = sample size, corrected for error relative
to the mean
t = students modified t statistics

D = error relative to the mean

O and B: Iwao and Kuno's regression
coefficients

i = desired mean value
Table F4 shows the regression coefficients for A.

aphidimyza on each sample date and for the combined

 

sample date. Table F5 provides the total tree sample
size estimates for the predatOr using the combined

seasonal estimates of‘xandB .

1‘)

 

105

ucmHonmsmcH on an: I mucHom N no

HO¢.OS

«O0.0

OSS. SS.H NS.Nv

«SO. O0.0H OS.H
OOO. ON.OH O0.0:
OOO. ON.OH HN.O:
SOO. O0.0H O0.0H:
SOS. SO.N OS.SH:
OHO. NH.N ON.Nmu
OOO. OO.H NN.HO
SHO. OO.N O0.0H
OOO. OO.N NH.O
H O a
wth. HMHHOH.

 

N

 

mHmNHma< Mom

H NHCO :Hmucou H O O mmuma umucmso HmBOH mpoz

 

 

 

SO0.0
SNH.v
OOO. ON.H Nv.SS
NOO. OO.O N0.0-
HOO. O0.0 O0.0!
OOO. O0.0 O0.0t
OOO.H HH.O O0.0:
NSO. OS.H O0.0N
ONO. O0.0 HO.NOH
HOO. SH.H NH.SSI
NOO. OS.H N0.0N
ONO. SS.N O0.0
NH _O a
umuomsa Momma

HON.OH HIO mocmHHm>
OOO.O HIO uom msmmz
OOO. OO.N OO.NH
mmumo HHS
OO.H O0.0 OO.O H
OOO. OO.OH OO.N: m
OO.H OO.O OO.O o
HOO. O0.0H Hm.O O
OHO. OO.H NO.NO m
ONO. NO.O OO.HON: o
OOO. OO.H OO.HO o
HOO. NO.N OO.ON m
HOO. OO.N OO.OH .4
NH O a OOOO

 

umncmso umBOH

.mucsoo cOHHMHsmom OHSQO Mom mpHSmmm :onmmummm Sadx Sam omzH OOOH .Hm mHQMB

     

   

NE...

  

-.....“ ...»:ch “O..—

suntan sob.

H

..... .. :

 

106

 

 

O0.0H

«HH.S
ONO. SOO.S OOS.SS
SSH. OOH.¢ SHO.vm
SSS. HNS.v SS0.0S
SOS. OSS.S SOS.Sm
SSS. SON.S «ON.OH!
OOO. OOO.H HS0.0m
SSO. OSS.m SSN.SHI
SSS. NOS.VH SSS.¢HI
NNO. OSS.O «SO.N
SOS. OSS.HH SN0.0
NH m 8

OOHB HM#OB

 

HNS.S

OSS.v
OHS. OOH.S OSv.SH
SSS. SSS.S NON.mN
SHS. vvm.S SSS.HS
OSS. SHS.S th.OH
HSN. HSv.S SOS.¢HI
HHS. OSv.H HNS.SS
SNS. SmH.v SSm.HHI
OSS. OO0.0H SSS.vl
SSS. OHS.S NOS.SI
SSS. SSS.S NNS.H

H 8

N O

 

 

umummao Hmaaa

 

OO0.0 Nm
Omm.O M.
HOO. HO0.0 OOO.NH
mmuma HH<
OOO. OO0.0 OO0.0 5
OOO. NN0.0 OOO.SN H
OOO. OON.O OON.ON 0
OOO. OOO.N OOO.OI a
OOO. OOO.N OO0.0N m
ONO. OOH.m OOO.N a
OOO. ON0.0 OOO.v| 0
OOO. OO0.0 NOO.H: a
OOO. ON0.0H OON.OI <
NH O 8 mama
pmummso Hm3oa

mumaou :oHumHmaoa OHaa< How muHsmmm GOHmmmHOmm ocsx mam om3H

OOOH .Nm mHQmB

 

107

 

HO.S OS.SS

S 8
SS SNH SSS
SS OSH SOS
HS OSH HOS
Sv NHH NOS
mv SSH OSS
Sv OOH HOOH
Sm SOH SSHH
Sm SSH SONH
NS SON SSSH
SO HON OOSH
SS OHS vSHN
OOH NSS OvSv
SHSN NSOOH SSSSSH

S. N. H.
mmHB Hmvoa

mmmmm mcoHumHsaoa Han .< ScHumEHumm Mom Humafiaz mmHBO mmNHm mHaEmm

SN
SN
OS
HS
NS
SS
SS
OS
OS
mv
vm
OS

.SOOH

m.

SH.S

OS
SS
SS
NOH
OOH
SOH
OHH
SNH
NSH
SSH
SSH
SOS
OSSv
N.

SS.SH

SNS
SSS
OSS
SOS
OOO
OSO
OSO
SNS
SSS
OHOH
SONH
SSNN

SHOSO

H.

 

umnmmso Hmaaa

OH
OH
HO
Nv
SS
«V
OH
OH
om
Sm
NS
SS
SNS
S.

O0.0

HSH
SSH
OSH
SSH
OOH
SOH
HSH
SSH
«SH
SOH
.SON
SHS
HSSS
N.

SO.NH

OOS
HSS
OOS
SHS
SSS
SSS
HHOH
HOOH
SHHH
SHNH
SOvH
SSSN

NHOOS

H.

 

umHUmaO Hm30H

.monsz oasm mam szH mconH>Ha ancmo mmune :0

ONH
OOH

SO
OS
Om
OS
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ON
ON
OH
OH

2mm:

.Sm mHnt

 

 

108

Table F4. 1977 Predator Linear Regression Values From Iwao

and Kuno Analysis

 

Date _. _ a. V 3. .5 . ._ r2
6-22-77 -.259 19.955 .982
6-29-77 .311 . 2.832 .144
7-06-77 -o.088 6.552 .972
7-13-77 .344 6.895 .446
*7-20-77 2.549 3.361 .626
7-27-77 -0.869 13.923 .929
8-03-77 -o.022 7.184 .966
Combined .222 7.014 .637

*
Does not include lower canopy level of last five trees

sampled.

109

Table F5. 1977 Predator Sample Sizes (Trees) Estimated.
Using Iwao and Kuno's Method

Error Relative to Mean““

 

 

7.014

Mean .1 ".2 ".3
1.0 28847 1714 397
1.5 14556 1052 260
2.0 5649 589 160
2.5 4522 499 140
3.0 3555 431 122
3.5 3198 400 115
4.0 2872 371 107
4.5 2704 356 104
5.0 2541 241 100
6.0 2347 322 95
7.0 2222 310 92
8.0 2133 301 90
. 9.0 2066 294 88
10.0 2015 289 86
11.0 1976 284 85
12.0 1942 281 84
13.0 1916 278 84
14.0 1892 276 83
15.0 1873 274 82
20.0 1807 267 81
a = 0.222

 

 

110

Negative‘BinOmial

 

The estimates of the negative binomial k obtain
was derived with formulas presented by Elliott (1971).
Difficulties in estimation of the negative binomial coefficient
by other methods (e.g. Bliss and Fisher 1953) limits the
_study to perhaps a bit less regorous methodology.

Tables F6-F8 present the eStimates used to obtain.k
values on each sample date for lower, upper and total tree
sample designs. To determine if computation of c common
K (Kc) was applicable, linear regression of l/k and i was
performed (see lower portion of each table). If the computed
slopes were not significantly different from a horizontal
line, the common K was computed.

Using the formula

11 = t2/132 (1/2’ + l/Kc)
tree sample number estimates were computed (Elliott 1971).

These are presented in Table F9 for A. pOmi.

 

Similarly the,approaches used in calculation of K values

for A. aphidimyza are presented in Table F 10. The sample

 

size estimates (trees) for the total tree sample design are

presented in Table F10.

 

111

.acmvcmamSGH ma oa mESmmm
SmE maaHoa mna mcHH HmaGONHao: m ana acmamaaHHO OHammaS a0: mH maon mm .

O " H
OH N

HMO va.OI + HHS.SN H S

:OHamaSm GOHmmmaSmm

X m> M\H

mo aoHa “moamwcmamwcH How Homau

 

o
mSH. H M
u OO0.0 n ox\H
mS0.0 SSS.SH OOO.SSvH SNS.SS ONH SS.vaH SN.OH
mm0.0 SSH.SH OOS.OSSS SNS.SNS ONH OS.SSSS OS.SH
SOH.O SSN.S OOH.SmSmH NOO.SmSH ONH NS.SSSSH SN.NO
OV0.0 SON.HN OSS.SSSS OSS.OSH ONH O0.0HOO SS.VH
OSm.O SNO.H OSS.SNOS OHN.mSSH OS SO.SSOS ON.S¢
SHS.O NvH.S OSH.OSSm NOO.HOSH ONH SS.SNSS SS.SV
SSH. SOS.S OSS.SSNH SHv.va . ONH OS.NHSH SS.SH
OSO. SHH.SS . SS.OSSH SSS.SO ONH OS.SSSH SS.O
SOH. OmS.S OSN.¢VH OSO.HN ONH SS.SwH NO.v
ONO. vSm.Hv HNO.N SO0.0 ONH SN.N 7 SN.O
>ao:mu.am3oa .
o
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h H
Hx\H H\M
Wm .... NW I. .
:\NO I NM u _x
..Eom.am HOS M :OEEOU mo COHamHSOHmU OOOH

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SclS

mama
mHaEmm

.Sm mHQmB

 

mHamoHHaam & aoafioo HO.H.O maonO moamcamamm oz

112

ONS. u Na HMO SOH.O + NSO.OS n O :OHamaUm COHmmmaSmm
m m> M\H mo aOHm "mocmoamamvcH a0m_xomau
o
OON.O n M

o
OOO.O + O\H
OHH.O NS0.0 OS0.000H OON.OOO ONH ON.ONOv NS.SN SOIO
NO0.0 NHH.SH OOS.ONNO OHS.NSN ONH OO.HONO ON.OH ONIO
OOS.O OOO.N ONO.HOHON OO0.0000 ONH SH.NONON NH.HO ONIO
HOH.O OSS.O OON.OHONH OOO.HvHN ONH SS.SSONH .Sv.Ov SHIO
ONS.O SSO.S OOH.OONO OO0.0SOH OS ON.HNSO . OO.NO SOIO
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SN0.0 OOO.Nv OOS.OSO «SS.NH ONH O0.0SO OH.O OHIO
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ON0.0 OON.Ov OOO.NHH «O0.0 ONH OO.HHH SH.N mOIS
O OxH .O x c m m mama
. N mHaEmm

 

Smocmo amaaa

Han am How M £05800 ao.aoHamH50HmU;.OOOH .Om mHame

 

 

113

mHamoHHaad M GOEEOU "

SOH. u NH HMOSO.OI SO.SN n O

COHamSSm GOHmmmaSmM
M m> M\H

So aOHa "mocmccmamSGH How Momau

0
OON.O H M

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NO0.0 HOO.N OH0.00HO HHO.mmON OHN O0.0SNO O0.0H ONO
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M O\H _O .x 2 NO m 11:11:.

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mmaa HmaOB mHaEmm

.som am How O Oossoo mo OoHamHsono OOOH .OO_6HQOO

114

H0M\H + M\HO Na\Na u a mHSEHom

 

 

 

o o o
OON. u M OON. u M OOH. n M

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OO HON «NON 7 NS SHN OOOH OOH HHS NOON O

OHm OOOH NOOmN OON HHHH NOOOH OOO. NOOH OHHON H
m. N. H. m. N. H. m. N. H. cmmz

mmaanHmaoa Smocmo Hmmaa maoamo amBoa

:mmz maa oa m>HamHmM maoaam mcHOam> am Hmmmaa mo amafiscv
M :0 Ummma OCOHamHsaom Han .¢.aom mmamEHamm mNHm mHaEmm OOOH .Om_mHamB

 

O

 

115

NH.OH
Hm.O
O0.0H
ON.OH
O0.0
O0.00N

MammO M\H

NNOHm.O
MNNO0.0
NOOH0.0
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munsoo mNmEHwOnm< mm#mHowH:mm.OOOH Mom .M :08800 mo COHumusmEoo .OHm.mHQme

 

NHH

116

Table F11. 1977 Sample Size Estimates for g,
_ Aphidimyza Using Kc (number of Trees)

 

 

"‘Total'Tree

Mean .1 .2 .3
1.0 45294 2691 626
1.5 23509 1698 420
2.0 9261 . 966 262
2.5 7483 '826 231
3.0 5921 757 203
3.5 5352 -670 192
4.0 4823 623 180
4.5 4553 .599 174
5.0 4288 575 168
6.0 3975 545 161
7.0 3772 525 156
8.0 3628 512 152
9.0 3519 501 149
0.0 3435 492 147
5.0 3205 469 141
0.0 3098 457 138
Kc - .09651
n = t2 ( 1 1

“'2' 27+?

I) X c

 

117

Mean Variance Relationship
The computation of Taylor's (1961) coefficients from
the mean-variance relationship involved obtaining individual

tree estimates for‘g.‘pomi and §.@aphidimyza in three sample

 

designs. The results, from log-log regression, for total tree,
lower canopy and lower inner canopy sampling designs of'g.

pomi and g. aphidimyza populations are presented in Table

 

F11.
The formula for sample size estimation using Taylor's

coefficients was presented by Croft, Welch and Dover (1975).

 

It is:
n='_1_:_2_ (H1 + (m+'-]%—)‘V(5<') +l/k)
D2 .9X f2
where

n = number of samples

t = students modified t statistic (<r+ .05)

D = error relative to the mean

a = sampling efficiency (a = 0.9)

_X = desired mean

M = number of secondary sample units (terminals)

K = negative binomial coefficient

V(x) = Variance estimate using Taylor's coefficients

 

r

118

Tables F12 and F13 show the eStimated variances for several

means of'g.‘pomi and §.'aphidimyza, respectively. Also

 

shown are the Kc values used for each sample design for both
species. Table F14 presents the student t values used in
the formula. Table F15. and Table F16 present the terminal
sample number and the tree sample number at varying densities
‘and at three levels of error relative to the mean for the

él‘pomi and é.‘aphidimyza sampling designs.

 

   
 

Table F12.

 

Intercept

SlOpe

2
r

K
C

119

Results For 1977 Taylor's Mean Variance
Relationship (per tree bases)

...... . . Samp.l.e. De.sign . .

Total Lower Lower
Tree ' Canopy Inside
19.82 7.71 3.28
1.4956! 1.771 1.754
.760 .965 .957
.209 .165 .209

a. aphidimyza

 

Intercept

SlOpe

2
r

K
C

Sample Design

Total Lower Lower

Tree Canopy Inside

7.32 7.348 3.45

1.658 1.692 1.495
3943 .841 .715
.097

   

120

Table F13. Computed Variances xvi) From Taylor's Mean

UI+H NI

10
15
20
25
40
50
60
75
100

Variance Relationship’§.'pomi 1977.

Vx
Tree

7.320
105.320
333.051
‘652.335
1051.039
2058.629
3316.855
4801.792
6496.596
9405.084

15153.435

.209

Vx
Lower

7.348
111.899
361-551
717.985

1168.186
2319.841
3774.458
5505.876
7495.508
10933.842
17789.721

.165

Vx
Inner

3.450
38.263
107.850
197.731
303.990.
557.334
856.838
1196.132
1570.924
2197.985
3371.468

.209

 

 

121

Table F14. Computed Variances Xvi) From Taylor's Mean
Variance Relationships é.‘aphidimyza 1977.

 

 

— W _‘E— _Vj;

X Tree Lower Inner
1 19.820 7-710 3.008
1.5 35.965 15.809 6.037
2. 54.889 26.313 9.896
. 76.190 39.067 14.519
3.0 99.601 53.956 19.860
3.5 124.924 70.892 25.881
4.0 152.008 89.805 32.555
4.5 180.733 110.635 39.856
5.0 211.000 133.331 47.765
6.0 275.831 184.145 65.334
7.0 345.960 241.949 85.144
8.0 420.968 ~306.497 107.099
9.0 500.519 377.508 131.119
10.0 584.337 445.045 157.137
15.0 1060.328 933.065 315.357
20.0 1618.249 1553.025 516.949

K = 0.0 9651
c

 

 

122

Table F15. t - Values Used In Computation of Sample Size

 

Estimates

g. pomi. é. aphidimyza

Mean t = 0.05 Mean t = 0.05
1 6.314 1.0 6.314
5 2.015 1.5 4.617
10 1.812 ‘ 2.0 2.920
15 1.753 2.5 2.637
20 1.725 ' 3.0 2.353
25 1.708 3.5 2.242
30 1.697 4.0 2.132
40 1.684 4.5 2.074
50 1.678 5.0 2.015
60 1.671 6.0 1.943
75 1.668 7.0 1.895
100 1.662 8.0 1.860
9.0 1.833
10.0 1.812
15.0 1.753
20.0 1.725

Error Relative to the mean

D = 0.1
0.3
0.5

 

 

 

123

 

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124

 

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APPENDIX F 3
Pertinent Literature

Bliss, C. I. and R. A. Fisher, 1953. Fitting the negative
bionomial distribution to biological data. Biometrics
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Croft, B. A., Welch S. M. and M. Dover. 1976 Supresseion
statistics and sample size estimates for populations
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fallocis on apple. Env. Ent. 5(2): 237-234.

 

 

Elliott, J. M. 1971. Some methods for the statistical
analysis of samples of benthic invertebrates,
Freshwater Bio.Ans. Sci. Pub. 25

Iwao, S. 1968. A new regression method for analyzing the
aggregation pattern of arrival populations. Res. Pop.
Ecol. 10: 1-20.

Iwao, S. 1975. A new method of sequential sampling to
calssify populations relative to a critical density.
Res. Pop. Ecol. 16: 281-288.

Iwao, S. and E. Kuno, 1968. Use of the regression of mean
crowding on mean density for estimating sample size
and the transformation of data for the analysis of
variance. Res. Pop. Ecol. 10: 210-214.

Karandinos, M. G. 1976. Optimum sample size and comments
on some published formula. Bull. Ent. Soc. of Amer.
Vol. 1. 417-421.

Kuno, E. 1969. A new method of sequential sampling to
obtain the population estimates with a fixed level
of precision. Res. Pop. Ecol. 11: 127-136.

Kuno, E. 1976. Multi-stage Sampling for population
estimation. Res. Pop. Ecol. 18: 39-56.

Lloyd, M. 1967. Mean Crowding. J. Animal Ecol. 36: 1-30.
Iwao, A. 1976. A note on the related concepts 'Mean

Crowding' and 'mean concentration' Res. Pop. Ecol.
17: 240-242

125

 

126

Hagihara, A. 1976. The time sequence of the relation
between mean crowding and mean density with some
population processes. Res. Pop. Ecol. 17: 244-239.

Sekita, N. and.M. Tamosa 1973. Applicability of a new
sequential sampling method in the field population
surveys. Appl. Ent. 2001. 8(1): 8-17.

 

 

LITERATURE CITATIONS

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Baker, A. C. and W. F. Turner. 1916. Morphology and
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127

 

128

Dixon, A. F. G. 1971a. The role of aphids in wood formation.
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129

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