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Tomato pinworm, Keiferia lycopersicella (Walsingham)

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Title:
Tomato pinworm, Keiferia lycopersicella (Walsingham) population dynamics and assessment of plant injury in southern Florida
Added title page title:
Keiferia lycopersicella
Creator:
Peña, Jorge E., 1948-
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English
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xvii, 265 leaves : ill. ; 28 cm.

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Subjects / Keywords:
Crops ( jstor )
Eggs ( jstor )
Infestation ( jstor )
Insects ( jstor )
Larvae ( jstor )
Leaves ( jstor )
Pests ( jstor )
Planting ( jstor )
Tomatoes ( jstor )
Vegetation canopies ( jstor )
Tomato pinworm ( lcsh )
Tomatoes -- Diseases and pests -- Florida ( lcsh )
City of Gainesville ( local )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1983.
Bibliography:
Includes bibliographical references (leaves 230-242).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Jorge E. Peña.

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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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TOMATO PINWORM, KEIFERIA LYCOPERSICELLA (WALSINGHAM): POPULATION
DYNAMICS AND ASSESSMENT OF PLANT INJURY IN SOUTHERN FLORIDA








By

JORGE E. PENA


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLCRIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR
THE DEGREE OF DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


1983
















ACKNOWLEDGMENTS


I thank Dr. Van Waddill, my advisor and chairman, for his

encouragement, support and friendship, but most of all for his valuable

suggestions and allowing me freedom to conduct my research.

I would also like to express my appreciation to the following

people:

Dr. S.H. Kerr for his interest in teaching me the art of

communication and also for his help in solving administrative problems

during my studies.

Dr. J.L. Stimac for his help and constructive criticism, as

well as his ideas to improve the quality of this study.

Dr. K.H. Pohronezny for his constructive criticism, suggestions

and for reviewing this manuscript.

Dr. D.J. Schuster for supplying material for my research as

well as his interest in this study.

The Agricultural Research and Education Center, Homestead,

Florida, and to Dade Agricultural Council for providing the grantmanship

and scholarship to support my studies.

The staff of AREC, Homestead, for their cooperation, espe-

cially Jonnie Csterholdt, Carolyn Reitman, Susan Housley, Leslie

Sawyerlong, Rodney Chambers, Linda Douthit, and Wilbur Dankers for










their help during data collection. Mrs. Sheila Eldridge and Mrs.

Barbara Hollien for kindly typing this manuscript.

Drs. R.E. Litz and S.K. O'Hair for their friendship, encourage-

ment and support during the past years.

Mr. Ben Gregory for his honest friendship and willingness to

share ideas in research.

Ms. Annie Yao, Mr. A. Bustillo, Mr. W. Chongrattanameteekul,

Mr. K. Patel and Mr. C. Ho for their friendship and support.

I am indebted to my family for their love and encouragement,

the Litz family, Eleanor Merritt and Bunny Hendrix for their friend-

ship and support.


















TABLE OF CONTENTS


Page


ACKNOWLEDGMENTS . .


LIST OF TABLES . .

LIST OF FIGURES . .


ABSTRACT . .

INTRODUCTION . .


. ii


. vi

. xi
.... xv


CHAPTER I:


CHAPTER II:


LITERATURE REVIEW .

Family Gelechiidae .
Studies on Keiferia lycopersicella .
Tomato Plant Phenology and Measurement of TPW
Dispersion and Economic Damage .
Environmental Factors Affecting TPW Population


. 6
. 7


. 19
. 21


DESCRIPTION OF TOMATO PLANT PHENOLOGY AND EVALUATION
OF TOMATO PINWORM FOLIAR DAMAGE ASSESSMENT .


Introduction . .
Materials and Methods .
Results and Discussion .
Conclusions and General Discussion .

CHAPTER III: SPATIAL DISPERSION OF TOMATO PINWORM EGGS ON TOMATOES.


Introduction . .
Materials and Methods .
Results and Discussion .
Conclusions and General Discussion .


CHAPTER IV:


SPATIAL PATTERNS OF DISPERSION OF TOMATO PINWORM LARVAE
TN TOMATOES ... 89

Introduction ... 89
Materials and Methods. .. .90
Results and Discussion .. .91
Conclusions and General Discussion .. .117










CHAPTER V:


CHAPTER VI:


CHAPTER VII:


CHAPTER VIII:


CHAPTER IX:


TOMATO PINWORM ARTIFICIAL INFESTATION: EFFECT OF
FOLIAR AND FRUIT INJURY ON GROUND TOMATOES .

Introduction . .
Materials and Methods .
Results and Discussion .
Conclusions and General Discussion .


ADULT DISPERSION AND COLONIZATION OF TOMATO
FIELDS BY THE TOMATO PINWORM .

Introduction . .
Materials and Methods. .
Results and Discussion .
Conclusions and General Discussion .


. 119


. 154


EGG AND LARVAL PARASITISM OF TOMATO PINWORM IN
SOUTHERN FLORIDA . .. .171


Introduction . .
Materials and Methods. .
Results and Discussion .
Conclusions and General Discussion .

EFFECTS OF RAINFALL AND RELATIVE HUMIDITY ON
IMMATURE STAGES OF THE TOMATO PINWORM UNDER
GREENHOUSE AND FIELD CONDITIONS .

Introduction . .
Materials and Methods. .
Results and Discussion .
Conclusions and General Discussion .

INFLUENCE OF POST-HARVEST TOMATO FIELDS ON THE
POPULATION DYNAMICS OF THE TOMATO PINWORM. .

Introduction ... ...
Material and Methods .
Results and Discussion .


171
172
174
189


190

190
190
194
211


. 213


213
213
217


CONCLUSIONS AND GENERAL DISCUSSION .. .225


iREFERENCES . .


APPENDIX


. 230


EXPLANATORY TABLES FOR CHAPTERS II AND III .. .243


BIOGRAPHICAL SKETCH . .. 265

















LIST OF TABLES


Table Page

1 Larval parasites of Keiferia lycopersicella
reported from U.S.A. and South America until 17
1981 . .

2 Classification of tomato pinworm leaf
damage on 'Flora-Dade' tomatoes, based on
greenhouse and field observations. Homestead,
31
Florida, 1980 . .

3 Leaf area and reproductive plant structures in
tomatoes, cv Flora-Dade', planted on 5 dates
in Homestead, Dade County, Florida during 1980-
32
1981 . .

4 Stage of development description for tomato cv
Flora-Dade. Description is based on the average
of observations from tomato plants grown during
Fall 1980 through Winter 1981. Homestead,
42
Florida . .

5 Tomato leaf weight and leaf area consumed by
different larval instars of Keiferia lycopersicella
under greenhouse conditions; T 24+3C, 75+2% RH.

6 Percentage of tomato pinworm larval occurrence in
foliar injuries with different phenological charac-
52
teristics . .

7 Comparison of different sample sizes for tomato
pinworm eggs. Homestead, Dade County, Florida,
1980 . .

8 Mean number of tomato pinworm eggs per plant by
planting date for 8 tomato plantings in Homestead,
62
Florida, 1979-1981 .

9 Ovipositional preference of tomato pinworm for
upper and lower surfaces of tomato leaves from
plants grown under greenhouse and field
conditions .... .. *










Table Page

10 Mean number of tomato pinworm eggs in 2 plant
strata (upper and lower halves) per plant at
different sampling dates. Homestead, Dade
County, Florida, 1980 65

11 Mean number of tomato pinworm eggs per plant in
6 strata: upper, middle and lower external; upper,
middle and lower internal canopy of the tomato
plant. Homestead, Florida, 1981 67

12 Relationship between daily mean temperature ( C)
and TPW oviposition in 6 tomato plant strata.
Homestead, Florida, 1981 .. 71

13 Percentage distribution of TPW eggs for each stratum
of tomato plants in 5 tomat plantings. Homestead,
Dade County, Florida, 1980 ...... 75

14 TPW egg sample allocation for 6 plant strata during
3 different plant stages: second vegetative (TR ),
first reproductive (TR1), and second reproductive
stage (TR2) . .... 77

15 Mean tomato pinworm eggs on tomato leaves from
different strata of 45 day-old plants. Homestead,
Florida, 1980. ... 79

16 TPW egg sample allocation on tomato leaves numbered
from bottom to top. Plants 45 days old. ... 81

17 Relationship between frequency of occurrence of TPW
eggs per leaflet as dependent variable and distance
among eggs and leaflet area as independent vari-
ables ............. 82

18 TPW oviposition on tomato at different plant stages.
Homestead, Florida, 1981 .. 86

19 Sample size and relative net precision (RNP) for
sampling injuries at low and high population
densities. Homestead, Dade County, Florida, 1980. 93

20 Mean number of TPW foliar injuries and standard
error on different sampling units at specified
date. Crop planted in Nov., 1979. Homestead,
Florida . 96


vii









Table Page

21 Mean number of TPW foliar injuries and standard
error on different sampling units at specified
date. Crop planted in Jan., 1980. Homestead,
Florida ... 97

22 Sample size and relative net precision (RNP) for
sampling TPW larval injuries on upper and lower
plant canopy. Homestead, Florida, 1980 .. 100

23 Sample statistics: Mean tomato pinworm larval in-
juries per plant in 8 tomato plantings. Homestead,
Florida, 1979-81 ... 102

24 Mean tomato pinworm (TPW) foliar larval injuries
at 2 different plant levels for 3 different
plantings. Homestead, Dade County, Florida, 1980. 103

25 Mean tomato pinworm (TPW) larval injuries in 6
plant strata for 5 plantings. Homestead, Dade
County, Florida, 1981 ... .105

26 TPW larval injury sample allocation for 6 plant
strata at 3 different plant stages: second repro-
ductive (TR2), third reproductive (TR3) and
senescent (S1) . 113

27 Mean number and standard error of tomato pinworm
(TPW) injuries in 5 different plantings at speci-
fied date and plant growth stage ... 115

28 Tomato fruit damaged in the upper and lower plant
canopy, after a single artificial infestation with
K. lycopersicella larvae on ground tomatoes 124

29 Marketable value for tomato fruit damaged in the
lower and upper plant canopy after a single arti-
ficial larval infestation of K. lycopersicella
on ground tomatoes. ... 125

30 Tomato fruit damaged in the lower and upper plant
canopy after a double artificial infestation of
K. lycopersicella larvae on ground tomatoes ... .126

31 Marketable value for the tomato fruit damaged in
the lower and upper plant canopy after a double
infestation of K. lycopersicella on ground
tomatoes . ... .128


viii










Table Page

32 Effect of planting time on fruit injured by
K. lycopersicella larvae to ground tomatoes,
cv 'Flora-Dade' during 1981 149

33 Differences in cost and relative net precision
between sampling 6 plants per row and 1 random
150
plant per row ........ 150

34 Differences in mean fruit injured by K.
lycopersicella in pruned and not pruned
tomato plants 151

35 Effect of hedges and edgerows on tomato pinworm
field infestation at three fields in Home-
stead, Florida, 1981 . 166

36 Parasitism of the tomato pinworm larvae in
tomato fields in southern Florida, Dade
County, 1980 .. ...... 175

37 Parasitism of the tomato pinworm larvae in
tomato fields in southern Florida, Dade County,
1981 . 177

38 Keiferia lycopersicella eggs parasitism by 2
strains of Trichogramma pretiosum in the
laboratory. T25+10C; 75+2% RH 179

39 Number of Keiferia lycopersicella eggs col-
lected from two strata and percent of para-
sitism by Trichogramma pretiosum 1. 180

40 Distribution of normal and parasitized tomato
pinworm eggs in 2 tomato fields .. 181

41 Parasitism of tomato pinworm eggs by Tricho-
gramma pretiosum in 2 fields with different
host densities ......... 183

42 Plant water content in five tomato plantings
related to oviposition by the tomato pinworm 203

43 Effect of simulated rainfall on foliar larval
injuries caused by the tomato pinworm Keiferia
lycopersicella on plants grown under greenhouse
204
conditions .. ........... *

44 Mean percentage of tomato pinworm adults
emerged by day after pupal treatment with
different simulated rainfall regimes .. 208









Table Page

45 Effect of crop age of post-harvested tomato
plants on volunteer plants and number of tomato
pinworm larval injuries ... 220

46 General effect of cultural practices on
volunteer tomatoes and infestation by
tomato pinworm . ... 222

47 Effect of planting age and cultural practices
on volunteer tomato plants and number of TPW
injuries . ... 223

48 Tomato pinworm egg frequency distributions
determined on tomato plants during 1981 ...... 244

49 Tomato pinworm foliar injury frequency
distributions determined on tomato plants
during 1980 . ... .. 249

50 Tomato pinworm foliar injury frequency
distributions determined on tomato plants
during 1981 . ... .. 253

51 TPW egg allocation sample for 6 plant strata.
Planting 8, 1981. Age: 38 days. Stage of
development TV .................. 258

52 TPW egg allocation sample for 6 plant strata.
Planting 8, 1981. Age: 46 days. Stage of
development TR .................. 259

53 TPW egg allocation sample for 6 plant strata.
Planting 7, 1981. Age: 68 days. Stage of
development TR2 ... 260

54 TPW egg allocation sample for 6 plant strata.
Planting 4, 30 Oct. 1980. Age: 77 days. Stage
of development TR2 ................ 261

55 TPW larval injury sample allocation for 6 plant
strata. Planting 6, 1981. Age: 78 days.
Stage of development TR ............. 262

56 TPW larval injury sample allocation for 6
plant strata. Planting 5, 1981. Age: 108
days. Stage of development TR2 .......... 263

57 TPW larval injury sample allocation for 6
plant strata. Planting 4, 1981. Age: 120
days. Stage of development TR3 ... 264
3

















LIST OF FIGURES


Figure Page

1 Illustration of tomato cv Flora-Dade growth at 2 stages
of development. TV2=second vegetative stage; TR1=early
reproductive stage; a=primary leaf; bilateral develop-
ment . ... ....... .. .29

2 Influence of time on leaf area (dm2) expansion, flower-
ing and tomato fruit numbers of cv Flora-Dade grown
on 'Rockdale' soil under southern Florida conditions 39

3 Stages of development of tomato. TV1=early vegetative
stage; TV =late vegetative stage; TR, TR2, TR3=repro-
ductive stages; S =senescent stage .. 41

4 Linear relationship between tomato pinworm (Keiferia
lycopersicella) larval head capsule width (mm) and
foliar injury length, r2=0.47. ... 47

5 Linear relationship between tomato pinworm (Keiferia
lycopersicella) larval instars and visual leaf damage
scale, r =0.677 . 50

6 Average number of tomato pinworm eggs per plant
stratum during 6 different sampling dates in 2 to-
mato plantings at different growth stages.
A) Planting 7: Jan. 30, 1981. B) Planting 8: Feb.
28, 1981. TR =second reproductive stage of develop-
ment; TV2=sec8nd vegetative stage of development.
Plant strata: 1, 2, 3: upper, middle, lower external,
4, 5, 6: upper, middle, lower internal .... 73

7 Frequency of tomato pinworm eggs at different distances
(cm) between eggs when mean eggs were A) 2 eggs per leaf-
let and B) 5 eggs per leaflet. ... 85

8 Percentage of tomato pinworm (TPW) larval injuries in 2
sampling units from different plant portions, related to
number of injuries in the whole plant: 1) 1st planting,
Nov. 3, 1979; 2) 3rd planting, Jan. 8, 1980 ... 99










Figure

9 Percentage of tomato pinworm (TPW) foliar injuries found
at upper, medium and lower stratum in 4 tomato plantings:
1) Oct. 30, 1980; 2) Nov. 25, 1980; 3) Dec. 30, 1980; and
4) Jan. 30, 1981. Bars followed by different letters were
significantly different according to Duncan's Multiple
Range Test (P=0.05). Percentages were previously
transformed to arc sine. Percentages are expressed as
actual numbers before transformations .

10 Percentage of larval injuries at the external and
internal canopy evaluated from 5 tomato plantings.
Plantings 4, 5 and 6 planted in Oct., Nov., and Dec.,
1980; Plantings 7 and 8 planted in Jan. and Feb., 1981.
Homestead, Florida, 1980-81 .


11 Relationship between number of tomato pinworm
larvae per plant and number of injured fruits
and leaves in the lower plant canopy by a
single artificial infestation with TPW larvae.
Homestead, Florida, 1980-81 .

12 Relationship between number of tomato pinworm
larvae per plant and number of injured fruits
and leaves in the upper plant canopy by a single
infestation of TPW larvae. Homestead, Florida,
1980-81 . .

13 Relationship between number of leaves injured
in upper and lower canopy and number of fruits
injured in upper and lower canopy by a single
artificial infestation with TPW larvae. Home-
stead, Florida, 1980-81 .

14 Relationship between number of larvae per plant
and number of injured fruits and leaves (upper
and lower canopy) after a double artificial in-
festation with TPW larvae. Homestead, Florida,
1980-81 . .

15 Relationship between number of leaves injured
in upper and lower canopy and number of fruits
injured in upper and lower canopy by a double
artificial infestation with TPW larvae. Home-
stead, Florida, 1980-81 .


16 Regression of percent of yield reduction
against infestation densities per plant of
tomato pinworm larvae .


. 130






. 132






. 135






. 137


140


. 144


Page


108






111










Figure Page

17 Regression of percent yield reduction of to-
mato fruit against TPW number of foliar
injuries per plant ... 146

18 Abundance of tomato pinworm male moths at three
different field sites. A) Field 1, 1980; B)
Field 2, 1980; C) Field 3, 1981. Mean number
of moths at each site corresponds to the average
from 4 pheromone traps in north, south, west and
east directions. Homestead, Florida, 1980-81 .. 159

19 Mean number of tomato pinworm larval injuries
occurring in 4 commercial fields at 0, 30, and
120 m from the field border. Bars with the letter
"a" denote statistical differences at 0.05% dif-
ference level for a particular date ... 164

20 Field plan: Fields 1, 2, and 3, respectively,
each field was divided into 8-9 quadrats. The shaded
areas represent higher insect populations. Home-
stead, Florida, 1981 .. 168

21 A-B. Relationship of Keiferia lycopersicella
egg density to percent parasitism by Trichogramma
pretiosum in 2 fields . 185

22 A-C. Seasonal occurrence of Keiferia lycopersicella
eggs and parasitization by T. pretiosum in tomato
fields located at the (a) northern, (b) middle, and
(c) southern areas of Dade County, Florida. ... 188

23 Seasonal abundance of TPW eggs in experimental
fields, related to temperature and rainfall
regimes during A) 1980, and B) 1981, in Home-
stead, Florida .. 196

24 Seasonal abundance of TPW larvae in tomato fields,
related to temperature and rainfall regimes during
1980-81, in Homestead, Florida 200

25 Mean number of TPW injuries per plant during 9
days of simulated rainfall under greenhouse con-
ditions, avg. daily temperature 25+20C .. 206

26 Percentage of TPW adult emergence under greenhouse
conditions after treatment of pupae with 3 regimes
of artificial rainfall (200, 100, 50 and 0 ml water),
temperature 24+30C .... 210


xiii










Figure Page

27 Tomato field status following the main
harvest under S. Florida conditions. Home-
stead, Florida, 1980 ... .216

28 Number of tomato plants and TPW injuries
per m in 2 post-harvested tomato fields.
Homestead, Florida, 1980 ... .219


xiv
















Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

TOMATO PINWORM, KEIFERIA LYCOPERSICELLA (WALSINGHAM): POPULATION
DYNAMICS AND ASSESSMENT OF PLANT INJURY IN SOUTHERN FLORIDA



By

JORGE E. PENA

April 1983



Chairman: Dr. V.H. Waddill
Major Department: Entomology and Nematology

Experiments were conducted in Homestead, Florida, during 1979-1981

to describe tomato plant phenology, tomato pinworm (TPW), Keiferia lyco-

persicella (Walsingham), dispersion patterns, economic damage to tomato

and the effects of parasitoids, edgerows, rainfall and cultural practices

on TPW population dynamics.

Tomato cv. Flora-Dade phenology was described. Six stages were

designated based on the number of leaves, flowers, fruits and physiological

plant characteristics. This description can be of use in making pest

management decisions.

Based in the relative variation (RV) and sampling costs, sampling

units for TPW egg and larval stages were determined. Eggs were generally

(51%) found in the upper plant canopy, and larvae (50%) in the lower









plant canopy. Larger sampling units were allocated to the upper

and lower plant canopy for eggs and larvae, respectively. An

economic injury level was determined to be 1 larva per plant.

Yield can be reduced 10-40% when 1-12 larvae are attacking 45 day-old

plants. The results indicated a correlation between number of foliar

injuries in the lower plant canopy and fruit damage. In southern

Florida, higher TPW infestation occurred during March-May, 1980 and

March-April, 1981, compared with other months (Jan., Feb.). Trichogramma

pretiosum Riley caused 33-73% TPW egg mortality during May-July, 1981.

TPW larval parasitism fluctuated between 39-42% during 1980-1981. The

most abundant larval parasite was Apanteles spp., followed by Sympiesis

stigmatipennis and Temelucha spp.

TPW adult dispersion and effects of field edges on TPW dispersion

and field colonization were evaluated. Field areas surrounded by edge-

rows had higher TPW damage than areas surrounded by pastures.

The use of artificial rainfall demonstrated that when plant foliage

was irrigated there was a behavioral change in larval feeding which

resulted in 50% reduction of larval injuries compared to injuries on

soil-irrigated plants. TPW adult emergence was reduced 86% when high

levels of water were applied to pupae in the soil.

The effect of cultural practices on the TPW oversummering popu-

lation was evaluated. The mean number of injuries per m2 was 28 times

higher in crops planted later (December, 1980) than in crops planted

earlier (October-November, 1980). Lower numbers of injuries were found

in crops disced and mowed than in abandoned fields.










Parasitoids, cultural practices, and southern Florida climatological

patterns can have an impact on TPW population levels.


xvii
















INTRODUCTION


The tomato, Lycopersicon esculentum Mill., is one of the most popular

and important vegetables in the world (Purseglove 1968). Tomato produc-

tion in the U.S.A. is concentrated in California, Florida, Texas, New

York, New Jersey, Michigan and Virginia (Thompson and Kelly 1957).

Florida tomato acreage was 31360 ha during 1980-1981. Tomato produc-

tion is considered to comprise 28.31% of the total vegetable acreage

in Florida (Anonymous 1982). The tomato growing areas in Florida are

divided into 4 major districts: Palmetto-Ruskin, Pompano Beach-Fort Pierce,

Dade County and Immokalee-Naples (Anonymous 1981). Dade County has 18.3%

of the total state tomato production and supplies most of the winter (De-

cember through February) vegetables for the U.S.A.

The cultivation of fresh market tomatoes demands a high monetary in-

vestment from farmers. The cost of producing tomatoes in Dade County

during 1980 was $5123.25 per ha, which represents an increase of 1.44

times over the production cost of 1975 (Greene et al. 1980).

Expansion of tomato acreage in Florida resulted in changes of agro-

nomic practices to maximize tomato production (Geraldson 1975). Changes

in horticultural practices also established an agro-ecosystem with ento-

mological characteristics common to monocultures. From 1950-1975 insect

control in tomatoes was almost exclusively chemical. To help growers avoid

problems with insecticides such as insecticide resistance, secondary pest









outbreaks and objectionable pesticide residues, an integrated pest man-

agement program was established in Dade County on tomatoes (Pohronezny

et al. 1978). This program goal was to develop economically, technically

and ecologically sound systems of integrated pest management. This

approach had some constraints, however, such as the high crop value

which reduces the use of pest management tactics (Bottrell 1979). More-

over, the fruit quality standards for fresh tomatoes cause undue emphasis

on chemical control measures in order to prevent contamination of fruit

by insects and to prevent cosmetic damage to the fruit (Lange and

Bronson 1981).

Accordingly, insect pests in tomatoes can be categorized as direct

pests and indirect pests. Direct pests attack the product and directly

destroy a significant part of its value. Indirect pests attack plant parts

other than the saleable product but may reduce yield of the product (Ruesink

and Kogan 1975). Among direct pests of tomato in Florida are the corn ear-

worm, Heliothis zea Boddie, the southern armyworm, Spodoptera eridania

(Cramer), and the tomato pinworm, Keiferia lycopersicella (Walsingham).

Indirect pests are the serpentine leafminer complex, Liriomyza spp, the

tobacco hornworm, Manduca sexta (Joh.), and the granulate cutworm, Feltia

subterranea (Fab) (Poe 1972).

The tomato pinworm (TPW) can be either a direct or indirect pest

of tomatoes. The larva of this insect feeds in the mesophyll of the

leaves causing a serpentine-type mine during the first 2 larval instars.

In the latter instars the larvae can cause a blotch-type mine or they

tie leaflets together. The larvae also bore into fruit, providing an

entrance for plant pathogens which cause major damage to fruit.









The importance of the TPW as one of the most serious pests that

affect tomato production in semitropical areas of Florida has been docu-

mented by Poe (1974a) and Wolfenbarger et al. (1975). Tomato pinworm

incidence was noted in Florida as early as 1932 (Watson and Thompson

1932) with serious outbreaks occurring during 1942, and from 1970 through

1973. Several factors have been mentioned by Poe et al. (1975) as caus-

ing these outbreaks, i.e., type of insecticide used, change of tomato pro-

duction practices, and harmful effect of pesticides on natural enemies.

Other factors such as weather have been overlooked. Current practices

for TPW control in Florida have been almost exclusively chemical (Waddill

1975), although emphasis has also been given to breeding tomatoes for

TPW-resistance (Schuster 1977a) and less to TPW biological control

(V.H. Waddill, personal communication). The effects of several factors,

e.g., rainfall and cultural practices, that influence the life system of

the TPW are still not understood.

To develop effective integrated pest management for tomato, the

interrelationships among the crop (plant physiology, phenology), pests

(arthropods, weeds, pathogens) and environment (climate, natural enemies,

horticultural practices) must be carefully studied. It is necessary to

understand TPW ecology and basic biology by studying the role of several

factors that cause seasonal and annual changes in pest populations. The

ability to assess the presence and abundance of the pest by accurate

sampling techniques would permit a reliable study of TPW potential

for inflicting economic damage. By evaluating the role of extrinsic

factors, e.g., weather, natural enemies and agronomic practices, it may be

possible to reduce the TPW problem.









This study was initiated to answer these and related questions.

The specific objectives of research were:

1) to describe different stages of development of the tomato plant.

2) to evaluate techniques for tomato pinworm damage assessment.

3) to discuss sampling techniques for tomato pinworm immature
stages under southern Florida conditions and to describe TPW
spatial distribution.

4) to evaluate the importance and population dynamics of TPW
natural enemies.

5) to evaluate the effect of hedges and edgerows on TPW dispersion
and field colonization.

6) to determine a way to assess yield losses in ground tomatoes due
to TPW.

7) to determine the influence of rainfall on TPW population.

8) to study post-harvest field management practices that influence TPW
survival.

Therefore, the first chapter is a general literature review of

studies on K. lycopersicella and addresses the effects of biotic and

abiotic factors on the population dynamics of this insect. The

second chapter is a study of tomato plant phenology and also covers

the evaluation of foliar damage assessment techniques. In chapters III

and IV, I address sampling techniques and dispersion patterns of tomato

pinworm eggs and larvae. The fifth chapter deals with the effect of

tomato pinworm infestation on upper and lower parts of the plant. In the

same chapter I state the relationship between TPW population index and

yield losses. In chapter six I address the distribution of male moths

and larval stages in tomato fields, and the effect of edgerows in such

distribution. In chapter VII, I deal with the abundance of egg and

larval natural enemies of the tomato pinworm.






5


The interaction of rainfall and TPW is presented in chapter VIII.

Finally, I evaluated the data regarding horticultural practices

and the relationship between changes of tomato agroecosystem and

oversummering populations of tomato pinworm (chapter IX).
















CHAPTER I
LITERATURE REVIEW



Family Gelechiidae

The family Gelechiidae is one of the largest of the microlepidoptera

(about 580 North American species). Larvae vary in habits. Some are

leafminers, a few form leaf galls, many roll or tie leaves, and one

species, Sitotroga cerealella Olivier, is an important.pest of stored

grains (Borror et al. 1976). Studies on crop pests in this family have

been concentrated on pests of high economic importance, such as the pink

bollworm (Pectinophora gossypiella Saunders), the potato tuberworm

(Phthorimaea operculella Zeller), the angoumois grain moth (S.

cerealella), and Keiferia lycopersicella Walsingham, the tomato pinworm.

The pink bollworm and the potato tuberworm are generally considered

good colonizers with highly mobile behavior within and between fields

(Stern 1979, Van Steenwick et al. 1978); however, many experts considered

these moths weak fliers which move great distances by being carried pas-

sively by air currents (Kaae et al. 1977). They are capable of having

several generations per year, with the last generation showing a strong

dispersal tendency (Kaae et al. 1977). The potato tuberworm is perhaps

the most closely related to the tomato pinworm in patterns of behavior

and plant selection (Hofmaster 1949). Several authors (Shelton and

Wyman 1979, Meisner et al. 1974, Traynier 1975) have studied factors









influencing oviposition of potato tuberworm and the relationship between

populations of the pest and the host plant. Their studies were used as

a base in this research to compare with K. lycopersicella population

dynamics.



Studies on Keiferia lycopersicella (Walsingham)

The tomato pinworm (TPW), K. lycopersicella (Wals), is frequently

confused with other species (Povolny 1977), particularly with Scrobipal-

pula absolute (Meyr.) and Phthorimaea operculella (Zell.) (Doreste and

Nieves 1968), since they are also considered pests of potato and tomato

(Povolny 1973). K. lycopersicella and S. absolute are apparently iso-

lated from each other geographically and ecologically. K. lycopersicella

apparently avoids the cordillerian territory of the northern and southern

part of South America (Garcia et al. 1974, Mallea et al. 1972, Quiroz 1976).

The range of K. lycopersicella is in the eastern part of the American

continent and penetrates into Central America, Mexico (Povolny 1973) and

the U.S.A. (Elmore and Howland 1943). Phthorimaea operculella has been

reported on tomatoes in Venezuela, (Doreste and Nieves 1968), Bermuda

(Grooves 1974), and Egypt (Abdel-Salam et al. 1971).

In the U.S.A. K. lycopersicella is considered a key pest of tomatoes

in western California (Oatman 1970), Texas, Florida, Pennsylvania and

Hawaii (Swesey 1928, Thomas 1933). The tomato pinworm was first recog-

nized as a pest of tomatoes by Morrill (1925), and was later reported by

Elmore (1937) and Thomas (1933). In Florida, the TPW has been primarily

studied by Watson and Thompson (1932), Swank (1937), and recently by

Poe (1973). The seasonal history of the TPW was reported by Elmore









and Howland (1943) in California where it appears first during March

and April after overwintering in the pupal stage at or near the surface

of the soil. Later studies of the seasonal occurrence of TPW in Cali-

fornia showed that larval populations increased abruptly in September

and October (Oatman et al. 1979) and in April-June (Oatman 1970).

Batiste et al. (1970b) reported that there is no evidence for diapause in

this insect. Destruction of the tomato plants shortly after harvest may

prevent the insect from surviving the winter and infesting the crop

during the following season. Poe (1974a) reported that on the west coast

of Florida, severely infested fields occurred in the spring crop (February-

May) with less damage on plants during the autumn. Early infestations in

greenhouses also lead to heavy losses in the field.



Host Plants of Keiferia lycopersicella

Elmore and Howland (1943) reported that tomato and potato are pre-

ferred hosts of TPW. Several solanaceous plants, e.g. eggplant ISolanum

melongena (L)] and nightshade (Solanum nigrum L.), also are known hosts

for the TPW (Batiste et al. 1970b, Elmore and Howland 1943, Swank 1937,

Thomas 1933). Batiste and Olson (1973) demonstrated that K. lycopersicella

preferred tomato for oviposition over 12 other solanaceous plant species.

TPW could be reared on Solanum melongena L., S. dulcamara L., S. nigrum, and

S. elaegnifolium Cav. but not on S. nodiflorum Jacq., S. douglasi Dunal,

Datura meteloides A., D. stramonium L., D. ferox L., Nicotiana biglovii

(Torr.) and N. glauca Grah. The same author suggests that in California,

Solanum melongena, S. dulcamara and S. elaegnifolium may play a role in

the population dynamics and distribution of TPW.









Life Cycle of Keiferia lycopersicella

Accounts of the life history and behavior of K. lycopersicella have

been reported by Elmore and Howland (1943), Swank (1937), and Poe (1973).

Poe (1973) found that eggs are laid singly or in groups of two or

three on the host plant foliage. Elmore and Howland (1943) described the

egg as ellipsoid, 0.37 by 0.23 mm, light yellow when first deposited, grad-

ually darkening to a light orange before hatching. Eggs hatch 4-9 days

after deposition (Swank 1937) at 20.680C and after 4-4.5 days at 27-290C

(Elmore and Howland 1943). Weinberg and Lange (1980) determined that

eggs hatch in a range of 3.5 + 0 days at 350C and 7.8 + 0.2 days at

20C.

Keiferia lycopersicella has four larval instars (Elmore and Howland

1943, Swank 1937). Head capsule width of the larval instars are 1st in-

star 0.14-0.157 mm; 2nd instar 0.23-0.28 mm; 3rd instar 0.364 39 mm;

4th instar 0.52-0.61 mm (Elmore and Howland 1943). Newly hatched larvae

averaged 0.85 mm in length. The head capsule is dark brown and the re-

mainder of the body is a yellowish gray common to many newly hatched

lepidopterous larvae. The mature larvae are 5.8-7.9 mm in length and

characterized by an ash gray color with dark purple spots (Elmore and

Howland 1943). Larvae of K. lycopersicella characteristically possess

a pale prothoracic shield with conspicuous dark fuscous shading along

lateral and posterior margins (Capps 1946). Duration of the leaf mining

(1st-2nd instars) stage ranges between 4.7-5.8 days. The leaf folding

stage lasts between 5.6-16 days for a range of temperatures of 13-29 C

(Elmore and Howland 1943). Weinberg and Lange (1980) found that egg

hatching to pupation times range from 8 + 0.9 to 18 + 1.6 days when

reared at 350C.









The pupae are initially green, later turning to a brown typical

of lepidopterous pupae commonly found in the soil (Elmore and Howland,

1943). Before pupation the larvae form a loose pupal cell of sand grains

at a depth of 0.25-1.5 inches beneath the soil surface (Poe 1973). Wein-

berg and Lange (1980) recorded that pupation requires 11.3 + 0.5 at 20 C,

and 5.1 + 0.2 days at 350C. The length of the pupal stage was 38.7 and

11.4 days at temperatures of 12.65 and 26.4 C (Elmore and Howland 1943).

Swank (1936) obtained a range of 7-17 days with an average of 11 days

for the pupal stage at 260C.

Adults are characterized by an alar expanse of 9-12 mm. Labial

palpi have a short forrowed brush on the underside of the second joint,

a terminal joint somewhat thickened with scales, and are compressed

with the extreme tip pointed. The head and thorax are mottled with

dark brown. Forewings are elongate ovate, the hind wings have a

pointed apex, a strong pencil of hair scales, are dilated at tip of costa

in females, and dilated from base of costa in the males; the abdo-

men is dark fuscous above with basal joints slightly ochreous, the

underside is light ochreous sprinkled with dark fuscous spots. Adult

longevity is 7 days (24 + 20C) when they are fed on water and 8.5 days

at 240 + 20C when fed a 10% honey solution. At temperatures of 10 and

13C the respective longevities were 20.5 and 22.8 days (Elmore and

Howland 1943).



Insect Behavior

Elmore and Howland (1943) reported that copulation occurs within 24

to 48 hrs after moth emergence, and McLaughlin et al. (1979) stated that









sexual activity such as female calling was greatest during the 1st hr

of darkness. Very little copulation occurred after the 3rd hr. Males

ran or walked in their approach to calling females. Approach was gener-

ally from behind or at ca. 900 to the female and was accompanied by rapid

wing fanning. The copulatory strikes of the males were made laterally

beside the females. Moths remained in copula from 30 min to 2 hr.

Elmore and Howland (1943) and Poe (1973) described the behavior of

larval stages of K. lycopersicella. Newly closed larvae disperse briefly

from the hatched egg before entering the leaf. First instar larvae

spin a tent of silk over themselves and tunnel into the leaf. Further

feeding results in a blotch-like mine. The 3rd and 4th larval instars

feed from within tied leaves, folded portions of a leaf, or they

may enter stems or fruits. The 3rd instar appears to be the most

mobile and several types of behavior may occur (Poe 1973). This stage

larvae can draw 2 leaves together, may tunnel into stems or fruits at

the calyx, but usually the larvae form leaf folds on the upper leaf

surface. The four instars can cause injury to 3-6 leaves during develop-

ment (Poe 1973).

Elmore and Howland (1943) demonstrated that larvae that have mined

calyx lobes and nearby tissues enter the fruit instead of folding leaves.

Usually, the larvae enter the fruit beneath the calyx lobes or fruit

stems, but in heavily infested fields about 50% of the injured fruit may

be damaged in other places as well. The damaged areas caused by shallow

feeding just beneath the skin of the fruit appear as blotches. Larvae

that enter the fruit penetrate to a depth ranging from 0.9-1.9 cm.

Differences in the phenology of larval injuries were studied by

Batiste et al. (1970), who found that mines of the early stage larvae









superficially resembled the serpentine type mines produced by dipterous

leafminers of the genus Liriomyza. The mines could be distinguished

easily, because the dipterous leafminer leaves a trail of frass within

the mine, whereas the TPW larvae deposits nearly all the frass in a

single mass at the tunnel entrance.



Tomato Plant Resistance to TPW

Breeding for resistance work with tomatoes has largely been con-

cerned with pathogens, but currently there is a renewed emphasis on

insect resistance as part of integrated pest management (Lange and Bronson

1981). Resistance to many tomato insects does occur and includes resis-

tance to the fruitworm, Heliothis zea (Cosenza and Green 1979); leaf-

miners, Liriomyza spp. (Schuster et al. 1979); tomato pinworm, K. lyco-

persicella (Schuster 1977a); hornworms, Manduca spp (Kennedy and Henderson

1978), Colorado potato beetle, Leptinotarsa decemlineata Say (Schalk and

Stoner 1976; potato aphid, Macrosiphum euphorbiae (Thomas); flea beetles;

white flies (Aleyrodidae); spider mites (Acarina) and many others (Lange

and Bronson 1981). The mode of resistance in tomato is complex and may

involve many factors including antibiosis, preference, phenological devel-

opment (such as flowering time, time of fruiting, etc.), morphological

characteristics, presence or absence of foliage pigments, foliage vol-

atiles, and physiological incompatibility.

Resistance to the tomato pinworm has been studied by Schuster (1977a),

Schuster et al. (1979), and Kennedy and Yamamoto (1979). Schuster (1977a)

found that accessions of Lycopersicon sculentum Mill x L. pimpinelli

folium were more susceptible, while those of L. peruvianum (L) Mill, L.









peruvianum var. humifusum Mill., L. esculentum x L. peruvianum, L.

cheesmani f. minor (Hook F) Mull., and L. glandulosum Mull., were less

susceptible than the commercial cultivar 'Walter' (L. esculentum Mill.).

Selections of L. hirsutum Humb and L. hirsutum f. glabratum Mull. were

more resistant and had 25-50% and 50-75% less damage respectively

than 'Walter'. In laboratory studies the same author found that mine

lengths after 2 days were significantly shorter for PI numbers 129157

(L. hirsutum f glabratum) and 298933 (L. peruvianum). Schuster et al.

(1979) stated that levels of resistance to tomato pinworm and vegetable

leafminer appeared to be intermediate and the varieties PI 12930 and PI

1404403 of L. esculentum were found moderately resistant to both insects.

Kennedy and Yamamoto (1979) found an extractable toxic factor in the

foliage of L. hirsutum f. glabratum affecting Manduca sexta, H. zea, K.

lycopersicella, Aphis craccivora, A. gossypii, and Myzus persicae. Schuster

(1977b) reported that tomato varieties 'Pennorange E 160 A' and 'Pearson'

had less fruit damage by K. lycopersicella and armyworms, primarily

Spodoptera eridania (Cramer), than did the 'Walter' variety.



Chemical Control of TPW

Chemicals are widely used to control tomato pests. The need for

insecticides varies from year to year and from one area to another (Lange

and Bronson 1981). Chemical control of TPW was obtained in 1943 by Elmore

and Howland (1943) who recommended synthetic cryolite and talc dust

(50% sodium fluoaminate). In California, several insecticides were

evaluated by Middlekauff et al. (1963) and reevaluated by Batiste et al.

(1970a). The latter authors reported little or no control of larvae by









insecticides applied as soil treatments under greenhouse conditions.

These same authors stated that methyl parathion was the most effective

material in greenhouses, and also recommended parathion, methidathion,

phosphamidon, mexacarbamate and methamidophos. Spray deposits of para-

thion were found by the same authors to be significantly less effective

against eggs or early stage larva than was toxaphene-DDT.

Poe and Everett (1974) presented results of experiments to control

TPW in 2 locations in Florida. They reported that granular insecticides

in general did not perform as well as most spray materials for reduction

of the TPW mines and larvae in tomato transplants. They recommended

acephate, diazinon, endosulfan, and methomyl to keep seedlings nearly

mine free. Chlordimeform was co::isidered phytotoxic to seedlings but when

sprayed alone or combined with Bacillus thuringiensis Berliner on older

plants gave good control of TPW larvae without plant toxicity. Poe and

Everett (1974) recommended highly residual insecticides to maintain a crop

free of damaged fruit.

Waddill (1980) reported that certain insecticides used on demand

for tomato pinworm in Homestead, Florida, significantly reduced TPW

damage below that in the untreated check. Permethrin +

Bacillus thuringiensis were applied least often and were among the best

treatments. The author also showed that when used on demand a low rate

(0.225 Ibs ai) of methomyl resulted in significantly more damage than

the same rate plus 0.5 Ibs Bacillus thuringiensis.

Schuster (1977b) reported that when measured by the number of

damaged fruit, the degree of control of the TPW and southern armyworm

with Bacillus thuringiensis WP and chlordimeform was significantly depen-

dent on the tomato cultivar. The contact toxicity of 4 synthetic









pyrethroids and methomyl to some adult parasites of tomato pests indi-

cated that fenvalerate was generally the least toxic to parasites com-

pared to permethrin, burethrin, and NRD1C49 (+)-d-cyano-m phenoxybenzyl

(+) cis, trans-3-(2,2 dichlorovinyl)-2-dimethyl-cyclo-propanecarboxylate)

as well as methomyl (Waddill 1980). Fenvalerate was judged the most

promising candidate for use in a pest management program in tomatoes for

integrated control of the TPW and the vegetable leafminer. Lindquist

(1975) obtained the best control of TPW with synergized pyrethrins (MGK

pyrethrins) and endosulfan.

Emergence of K. lycopersicella and Apanteles spp from pupae and

soil treated with insect growth regulators (IGR's) resulted in 23%

suppression of pinworm adult emergence when applied directly to the TPW

pupae but was ineffective when applied to the soil. The IGR's caused a

reduced emergence of the parasite Apanteles spp from 61% to 0% (Poe

1974b). Prada and Gutierrez (1974) reported some results on microbial

insecticide control of Scrobipalpula absolute, the South American pinworm.

Seventy five to eighty percent control of the pest was obtained within

5-100 days after treatment at the rate of 500-200 Neoplectana carpo-

capsae Weiser nematodes per plant or with Bacillus thuringinesis (150-

500 g/ha). Schuster (1982) demonstrated that a mixture of B.

thuringiensis and Coax (454 g + 1.8 kg product/378 Its) when applied to

TPW infested tomato seedlings, increased TPW mortality up to 42.2%.




Cultural Practices for TPW Control

According to Lange and Bronson (1981), the mechanization of pro-

duction of processing tomatoes has not only revolutionized the industry

but has altered many control techniques and as a result, a few formerly









major pests have been reduced to a secondary position. Elmore and How-

land (1943) considered some cultural practices as undesirable because of

their adverse impact on TPW control. These include failure to destroy aban-

doned plantings, careless disposal of infested culled fruit, and use of

infested seedlings. In Florida, Swank (1937) recommended that all mate-

rial remaining in the field after the crop is harvested be carefully

plowed under. He suggested that the carelessly abandoned fields could

become a reservoir for infestation of a nearby or succeeding crop. Poe

(1973) stated that the best control for TPW is based on several cultural

practices: use of non-infested seedlings, destruction of plant debris,

use of light traps for adults in small areas, and destruction of plants

growing from seeds in compost heaps. Price and Poe (1977) reported that

staking and artificial mulching of tomato plants reduced damage caused by

K. lycopersicella and other pests.



Biological Control of TPW

Employment of biological control measures for insect and mite pests

of row crops has been limited, and the poor record probably relates

largely to the short-lived row crop environment, which presumably does

not permit establishment of the effective host-natural enemy relation-

ships which often characterize more stable environments (van den Bosch

et al. 1976). Modern-day biological control techniques have not been

fully exploited in tomato under field conditions (Lange and Bronson

1981). They have been widely accepted in European glasshouse tomato pro-

duction, however. Reports on parasitism of K. lycopersicella were

made by Elmore and Howland (1943), Swesey (1928), Thomas (1933), Oatman

et al. (1979) and Poe (1973) (Table 1).





























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Cardona and Oatman (1971) studied the biology of the larval para-

sitoid Apanteles dignus Muesebeck which is a solitary, primary, larval

endoparasite of K. lycopersicella and found that the total developmental

time from egg to adult was ca.18 days at 26.6 + 10C and 50 + 2% RH. Oat-

man (1970) stated also that the most common parasites at both Indio and

Escondido (California) during 1963-64 were A. scutellaris (Mues.) and

Parahormius pallidipes (Ashm.) followed by Sympiesis stigmatipennis

Girault. The biology of A. scutellaris (Mues.) was studied by Djamin

(1970). In California, A. dignus apparently occurs only along the coast

in the southern part of the state. Studies conducted by Oatman et al.

(1979) determined that in the south coast there was a range of larval

parasitization of 1.6-36.8% during 1972-73. Apanteles dignus was the most

abundant parasite reared from larvae followed by S. stigmatipennis Girault.

In Florida, Poe (1974b) reported that 50-70% of tomato pinworm larvae in

leaflets collected in the spring, 1973 were parasitized by Apanteles spp.



Behavioral Chemicals Used in Monitoring and Control of TPW

Pheromones. Sex pheromones of the adult TPW were obtained by

Antonio (1977) from extracts of the whole body of 2-day-old virgin fe-

males. A biological assay method was then devised to test males for

optimal response to the pheromone under varying conditions. Field evalu-

ation data by the same author indicate that the natural sex pheromones

were attractive to male tomato pinworm moths. McLaughlin et al. (1979)

found that males were more responsive when bioassayed with dim light

from both above and below an olfactometer than when illuminated only

from below. The effect of trap design and sex attractant release rates









on TPW catches was studied by Wyman (1979). It was determined that Zoe-

con Ic sticky traps were 6 times as effective in capturing TPW males as

were Delta sticky or mineral oil traps. An inverse relationship between

attractant release rate (fibres/dispenser) and trapping efficiency was

found. K. lycopersicella positively responded to a sex attractant of

unstated composition dispensed from rubber septa in traps in a tomato

field (Wyman 1979).

Deterrents. Beck and Schoonhoven (1980) stated that surface testing

of insects touching or piercing with the ovipositor or by biting and

probing with the mouth parts is in response to chemical factors that act

as incitants. If the stimuli received on initial testing indicate an

unacceptable plant, the behavior pattern is interrupted and the insect

abandons the plant. Such stimulants are deterrents.

Schuster (1980) reported that survival of TPW was reduced when larvae

fed on excised tomato leaflets dipped in solutions of cyhexatin, fentin

hydroxyde (triphenytin hydroxide) and guazatine (SN-513); N-n'''-(iminodi-

8,1-octanedilyl) bisguanidine. These compounds protected foliage and

fruit from insect damage when plants were sprayed in the field.



Tomato Plant Phenology and Measurement of TPW Dispersion and
Economic Damage

Little information is available that relates tomato plant phenology

to pest management or that concerns dispersion and economic damage of

the TPW. Plant phenology related to pest management tactics has been

determined already for different crops: alfalfa, cotton, potato, tobacco,

soybeans, etc. (Anonymous 1971, Reynolds et al. 1975, Ambrust and

Gyrisco 1975, Johnson 1979, Fehr et al. 1971). Tomato plant phenology









as related to pest management has been reported by Alvarez-Rodriguez

(1977) and Keularts (1980). Alvarez-Rodriguez (1977) evaluated pest

(pathogens and insects) damage as it is related to tomato life table

analysis and determined strategies for tomato production. Keularts

(1980) determined the effect of artificial defoliation in plants 30-100

days old. These studies should have been complemented by a phenological

description of the plant at different plant stages.

Studies of measurement and description of dispersion of TPW popu-

lations must involve a sampling program in which biological, statistical

and economical aspects of the program are evaluated (Southwood 1978).

This information is expected to result in 1) biological interpretation

of statistical parameters and 2) the use of this knowledge for measuring

of TPW control and for establishing a reliable scouting program.

Sampling designs for tomato pinworm larval stages have been studied by

Wolfenbarger et al. (1975) and Wellik et al. (1979). Wolfenbarger et

al. (1975) developed a sequential sampling program based on the detection

of larval feeding on the 3 top leaves per plant. Alternatively Wellik

et al. (1979) found that lower leaf and large fruit sampling methods

were best for detecting the presence of TPW. These opposing results

demand more detailed research in order to obtain more accurate TPW

density estimates.

The data concerning tomato pinworm damage range from estimations

of damage based on pesticide effectiveness (Batiste et al. 1970a, Poe

and Everett 1974, Waddill 1975, 1980), damage evaluation based on

effectiveness of parasitism of TPW larvae (Oatman 1970) to estimation









of economic injury levels (Wolfenbarger et al. 1975). Poe and Everett

(1974) determined the percentage of unmarketable fruit as 6.5 to 4%

when the plant was untreated. Waddill (1980) reported that plants

without chemical control may lose up to 75% of the fruit. Oatman (1970)

determined tomato fruit was infested up to 70% despite 68.9% larval

parasitization. Wolfenbarger et al. (1975) reported that an average of

0.3 TPW injuries per 3 top leaves caused 20% injured fruit.



Environmental Factors Affecting TPW Population

Characteristics of Agroecosystems

Agroecosystems vary widely in stability, continuity, complexity

and area. The kind of crops, agronomic practices, changes in land use

and weather are important elements affecting the degree of stability of

an agroecosystem (Stern et al. 1976). Since agroecosystems are

characterized by a short life (Loucks 19701, they are more susceptible

to pest damage and catastrophic outbreaks. This also occurs because of

a lack of diversity in plant species, insect species, and sudden alter-

ations imposed by man such as plowing, mowing and use of insecticides

(Luckman and Metcalf 1975, Pimentel 1961a, b, Smith 1970, van Emdem and

Williams 1974).









Tomato Agroecosystem

The tomato crop is a typical example of an agroecosystem with early

community succession (Price and Waldbauer 1975). In southern Florida 3

closely related tomato varieties are generally grown: MH1, 'Walter'

and 'Flora-Dade' (Volin and Bryan 1976). Horticultural practices are

characterized by direct-seeding in the field through mulched beds

that will aid in maintaining a regular amount of soil moisture, weed

control, and fertilization of the crop (Geraldson 1962, Davis et al.

1970, Bryan et al. 1967). In summary, the tomato crop is typical

of agricultural systems with high community energetic, small or low

community structure, rapid nutrient cycling, selection pressure

(r selected, many small progeny), and quantitative progeny production.

Also, the tomato agroecosystem is characterized by having a few major

key pests and secondary pests (Lange and Bronson 1981). Most of these

pests, e.g., lepidopterous larvae, stinkbugs, dipterous leafminers,

whiteflies, leafhoppers, aphids and some species of beetles, are con-

sidered as r selected species with rapid development, high maximal rate

of increase (rm), early reproduction, small body size, many small off-

spring and short length of life (Krebs 1978, Pianka 1978).



Biotic and Abiotic Factors Affecting Insect Population Dynamics

Biotic and abiotic factors exercise some influence on the fluctu-

ation in the number of insects in time and space. To reveal both char-

acteristics the inherent property of animals themselves and environmen-

tal conditions in their habitats must be studied (Shiyomi 1976). Among

the biotic factors, we should consider the habitat effect on insect









distribution. Effects of habitat have been studied by several research-

ers: Gossard and Jones 1977, Lyons 1964, Brazzell and Martin 1957, Yama-

moto et al. 1969, Wolfson 1980, Sparks and Cheatman 1970, Dethier 1959a,

Nishijima 1960. They demonstrated the effect of habitat on oviposition

and adult and larval dispersal. The effect of sheltered zones on

distribution of insects has been demonstrated by Lewis (1979) van Emdem

(1965) and Price (1976). They indicated the importance of crop edge

effect on colonization and dispersal of arthropods, especially for r

selected species, which show a "safety in numbers" strategy for progeny

reproduction and survival. Van Emdem (1965) considered that unculti-

vated land in regard to the insect fauna of a crop has 2 components:

1) Physical: shelter-survival in debris of woodland, 2) Biological:

plants of uncultivated land provide alternate food and breeding sites for

injurious insects, crop diseases or alternate hosts for predatory and

parasitic insects.

In most agricultural environments the principal pests are usually

controlled to a greater or lesser extent by natural enemies (Messenger

et al. 1976). The efficiency with which such natural enemies suppress

pest populations is influenced on the one hand by their own intrinsic pro-

erties and limitations and, on the other hand, by environmental factors

and conditions occurring in the agroecosystem under consideration

(Messenger et al. 1976).

Among the abiotic factors affecting insect populations, weather

and climate are commonly accepted by entomologists as dominant influ-

ences on the behavior, abundance and distribution of insects (Messenger









1959). Effects of climate on insect populations were studied by

Richards 1961, Nicholson 1958, Cloudsley-Thompson 1962, Andrewartha and

Birch 1974. Most authors agree that 2 of these factors, temperature and

RH possess a high degree of interaction and affect insect activity and

survival. As an example Chapman et al. (1960) and Hofmaster (1949)

have looked upon the effect of climate on survival of Gelechiidae.

Finally, pest control in an agroecosystem can be aided by proper use

of cultural practices. Two basic principles in the cultural control of

arthropod pests are 1) manipulation of the environment to make it less

favorable to the pest and 2) manipulation to make it more favorable for

their natural enemies (Stern et al. 1976). Cultural methods, however,

require a thorough knowledge of crop production and the biology and

ecology of the pest and its natural enemies in order to integrate the

techniques for pest control into proven agronomic procedures for crop

production.

















CHAPTER II
DESCRIPTION OF TOMATO PLANT PHENOLOGY AND EVALUATION OF
TOMATO PINWORM FOLIAR DAMAGE ASSESSMENT





Introduction

Description of tomato plant phenology and evaluation of tomato pin-

worm larval presence are major aspects of tomato pest management that need

to be determined. First, studies on tomato taxonomy, growth and develop-

ment, effects of fruit on vegetative growth, and relationship between

fresh weight and leaf area are well documented (Cooper 1964, Murneek

1924, Hurd et al. 1979, Romshe 1942, Thompson and Kelly 1957, Purseglove

1968). However, tomato crop phenology that divides the growing plant

into characteristic periods and shows the relative time in each period

needs to be studied. Second, description of TPW damage to the foliage

and TPW mine length correlation with plant resistance has been studied

by Elmore and Howland (1943), Batiste et al. (1970) and Schuster (1977).

Nevertheless, the evaluation of different techniques for TPW foliar

injury assessment is necessary to establish a relationship between larval

instars and amount of damage.

The objectives of this study were first, to define growth character-

istics of tomato plant during a typical southern Florida growing season,

second, to describe from a pest management point of view the phenology

of tomato, cv. Flora-Dade, and third, to determine constraints and prac-

tical use of TPW larval population indices.









Materials and Methods



The Tomato Crop

Tomatoes, cv Flora-Dade, were planted in 1979 (Nov. 3, Dec. 5),

1980 (Jan. 8, Oct. 30, Nov. 25, Dec. 30), and in 1981 (Jan. 30, Feb. 28)

at the University of Florida, Agricultural Research and Education Center,

in Homestead, Dade County, Florida. After metribuzin was incorporated

into the soil at a rate of 0.84 kg ai/ha, beds 45 m long were prepared

and fertilized with 7-14-14, at a rate of 2242 kg/ha. Immediately after

fumigation, drip tubing for irrigation was placed ca. 15 cm into the soil

and the beds were covered with plastic mulch. Tomato seeds were planted

with a seed drill 30 cm apart in the rows. One to two weeks after emer-

gence, the seedlings were thinned to one per hill. Plants were protected

from pests by application of fenvalerate 2.4 EC (.045 kg ai/ha), maneb and

tribasic copper sulfate (0.97 + 5.71 ai kg/ha) at weekly intervals.

Two to five plants were selected at random from each of the 8

plantings, and height, leaf area, number of leaves, suckers, flowers and

fruits were recorded every 8-15 days.



Description of Stages of Tomato Plant

The method used to describe tomato plant phenology was based on

the technique of Fehr et al. (1971) for soybeans, Glycine max (L).

Three developmental stages of tomatoes were defined: vegetative,

reproductive, and senescent. Number of leaves, plant height,










time of blooming and fruit formation were averaged from the 8 plantings.

Development of the plant was quantified by a nomenclature system, where

primary leaves were numbered from the bottom to the top; any secondary

growth, e.g., formation of primary laterals (Fig. 1), had the same node

number from which it originated, and it was distinguished with a let-

ter(s). The stem that originates from the bifurcation of the main stem

was called a secondary main stem (2M); laterals that develop from primary

laterals were considered secondary (2S).



Methods of Damage Assessment for TPW Larvae

A set of 3 tomato plants 40-50 days old grown in 20 cm pots was

introduced every 2 days into a cage (45 x 45 x 60 cm) for oviposition

by moths previously held at 24 + 30C and RH 75 + 2%. Plants were subse-

quently removed, and set aside for larval development. When each set of

plants was under attack by 1-4 larval instars, a total of 100 damaged

leaves was taken to the laboratory for inspection. A total of 20

individuals was studied per instar. Three methods of measurements were

used in separate experiments. First, a portion of leaf area mined by

the TPW larvae was separated from the leaf and then measured on a

LICOR model LI3100 area meter. For another set of damaged leaves,

leaf weight ingested by the larvae was determined by measuring the dif-

ferences in weight between the damaged leaflet and the juxtaposed leaf-

let. Larval instar in both experiments was determined by measurements

of the larval head capsule width. Second, a visual classification of dam-

age was made of leaf damage caused by TPW larval instars. A five class




















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scale of 0-4.5 (Table 2) was devised, based on personal observations and

the damage descriptions of Batiste et al. (1970a).

The leaf injury length (cm) was also measured and larval head capsule

width recorded. Finally, the presence or absence of TPW larvae in different

types of leaf injury was determined. A simple linear regression model was

used to examine the relationship between the head capsule width and injury

length, and between larval instar and the damage rating scale. The eval-

uation of the different methods of damage assessment was discussed with

regard to the practicality of their use for scouting programs.



Results and Discussion

The Tomato Crop

A summary of leaf area and number of flowers and fruits is shown in

Table 3. Tomatoes planted during October, 1980 began to flower 61 days

after plant emergence and fruit set occurred at 73 days. Maximum leaf

area was reached at 134 days. Tomatoes planted during November, 1980

started blooming at 54 days, and fruit set occurred at 68 days. Maximum

leaf area occurred 88-112 days after plant emergence. Tomatoes planted

during December, 1980 and January-February, 1981 had a shorter vegetative

period, with flowering at 42-62 days and fruiting at 49-62 days. Leaf

area reached a maximum at 63-89 days. The total leaf area during these

plantings was lower than that produced from fall plantings.

Under southern Florida conditions, average temperature changes

drastically from autumn to early spring (Mitchell and Ensign 1928). In

this area, the effect of planting date determines growth and tomato









Table 2. Classification of tomato pinworm leaf damage on 'Flora-Dade'
tomatoes, based on greenhouse and field observations. Home-
stead, Florida, 1980.


Degree of Damage Description


0 No damage

1-1.5 Mining of leaves, ca. 0.50 cm or less in

length; mine narrow and elongate; tissue

transparent; mine on any part of the leaf-

let; some leaves attacked by more than 1

larva; small larvae present.

2-2.5 Mining of leaves ca. 0.51-0.68 cm; 1/4 of the

mine is necrosed, but changing to a raised

area or oblong to ovoid blotch; frass accumu-

lation at the bottom of the injury.

3-3.5 Blotching of leaves; blotch necrosed over

60% of the injury; no holes indicating lar-

val exit; size 1-2 cm; epidermis of the leaf

opaque to chlorotic due to larval injury to

midvein; construction of silk tent in epi-

dermis.

4-4.5 Leaf folded; fold can occur at any lobe of

the leaflet. Necrosis extended to 75-80%

of the leaf; extensive frass accumulation

on blotch or fold; injury length 2-4 cm.



















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development. Tomato, cv Flora-Dade, was developed for production of

fresh market tomato fruit (Volin and Bryan 1976) during the months of

January-March. Consequently, planting before or after the autumn

months of October-December resulted in a high reduction of leaf area

and yield.



Developmental Stages of Tomato

Vegetative growth of the tomato plant passed through 3 distinct

phases (Fig. 2). In the first phase there was a steady increase in leaf

area, while in the second phase leaf area was relatively constant. The

third phase was characterized by reduction in rate of leaf expansion 130

days after plant emergence. The number of inflorescences rose rapidly to

a peak at 70 days and then steadily decreased, whereas fruit reached a

peak at 90 days post-planting and then steadily decreased. Flowering and

fruit formation were observed at 40 and 50 days, respectively. Three

major developmental stages were determined for tomatoes: vegetative,

reproductive and senescent (Fig. 3). Each stage was divided into sub-

stages. Each substage is explained in detail in Table 4.

The characteristics of tomato plant growth (Fig. 3) demonstrate the

relation between leaf area and crop age (days after emergence). The in-

crease in leaf area was observed until half of the second reproductive

stage (TR2). Leaf area is reduced during the third reproductive stage

(TR3) and senescent stage (S ). I consider that the TR2 stage can be

subdivided into another stage. This will allow a more detailed descrip-

tion of plant stages, as well as shorter time periods for better assess-

ment of pest management.





























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Table 4. Stage of development description for tomato cv Flora-Dade.
Description is based on the average of observations from
tomato plants grown during Fall 1980 through Winter 1981.
Homestead, Florida.


Plant Stage


Vegetative

TV1






TV
2





Reproductive

TR
1










TR
2



TR
3


Tomato Plant Description


Plants 1-15 days old. Complete formation of

2-3 primary leaves; loss of cotyledons; plant

height ca. 5-7 cm.

Plants 16-35 days old; plant erect (12-16 cm);

5-7 leaves, development of laterals; plant

with only 1 main stem.



Plant 35-40 days old; development of laterals

from nodes 1-5; at leaf 4-5 the stem bifurcates

producing another stem as vigorous as the first

main stem; production of floral clusters at

node 5 and second main stem; height 50 cm.

Plants 67-70 days old; fruit set; plant postrated;

yellowing of primary leaves.

Plant 109-135 days old; 90% fruit ripe; post-

harvest maturity; at least 60% of the primary

leaves necrosed, development of secondary laterals

at nodes 3-5; plant totally postrate; height ca.

32-57 cm.










Table 4--continued.

Senescence

S1


Plant 140-200 days old; dead leaves on main

stem and second main stem; regrowth of plant

from auxiliary buds at nodes 1,2 and produc-

tion of up to 3 floral clusters may occur;

possible fruit development.










The principal application of this nomenclature system is to deter-

mine the amount of yield reduction produced by damage inflicted at given

stages of plant development. As an example, if I use Keularts'(1980)

data from his experiment in tomato defoliation, 20% defoliation of lower

plant leaves at stages TV1 through TR2 did not alter mean yield

per plant. However, 20% defoliation of upper plant leaves at TR2 stage

caused yield reduction. The nomenclature system can apply to single plants

or entire crops. It would be worthwhile to apply this system to other

tomato cultivars.



Methods of Damage Assessment for TPW Larvae

Average leaf area and weight consumed by TPW larvae. The data from

this experiment demonstrated the complexity of measuring TPW foliar dam-

age. The average leaf weight (mg) and leaf area (cm2) consumed by larvae

of a determined instar are shown in Table 5. Average leaf area consumed

ranged from 0.5 to 1.57 cm2 for 1st to 4th instar. First and fourth

instar larvae consumed 5 and 13.42 mg of leaf, respectively. Variance of

leaf weight measurements was large suggesting that many uncontrolled

factors influence feeding of individual larvae in the field. Either

method might be used for laboratory and greenhouse experiments where the

researcher would have more control of the factors influencing variability

(e.g., leaflet size, leaf age).



Length of Foliar Injury and Use of Damage Scale

Length of foliar injury and TPW head capsule width were cor-

related (r=0.68; P=0.001, F=39.33) (Fig. 4). Furthermore, there was




















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a significant relationship between TPW (Fig. 5) larval instar and the

degree of damage observed (r=0.79). Both techniques suggest the possi-

bility of prediction of damage level in the tomato plant at stages

TV1 TR1. Such a prediction may be influenced by other factors such

as plant stages and larval density.

The use of larval instars to determine injury length has a reduced

bias compared to use of TPW damage degree scale. Foliar injury measure-

ment is only advisable for research experiments (e.g., plant resistance,

pesticide screening) in which the time frame available to determine the

dependent variable is not a constraint. Other aspects to be considered

for further study are larval preference for larger or smaller leaflets,

as well as presence of different larvae in the same leaflet.

The use of TPW damage scale is perhaps less precise than the tech-

nique mentioned above. Damage scale technique may introduce personal

error in measurement of larval instar in relation to degree of damage.

It is possible, however, to use this technique as an adjunct aid to the

population index (number of injuries per plant).

As an example, using the equation y=0.80 + 0.795x, where y = the

leaf injury damage scale and x = the tomato pinworm larval instar; if

the value of x equals 3, the average degree of damage in the plant will

be 3.18. This information will help to determine the effect of the

insect in economic terms, once the economic threshold is reached for

plant stages TV TR At this point there is no information available
1 1
for TPW EIL values for plants in these early stages. Therefore, further

studies will be necessary to indicate that the presence of a particular

larval instar is capable of producing a determined economic damage.


















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The ratio of percentage of larvae present to percentage of larvae

absent (Table 6) in the observed injuries was 4:1 for the folded necrosed

injuries, 31:1 for the folded with no necrosis, 1:3 for blotches with

necrosed tissue, and 3:1 for transparent blotches. Consequently, the

use of necrosed blotches will indicate that ca. 77% TPW larvae will be

absent from the observed blotches. If a high number of injuries per

plant falls in this category, the probability of not measuring

larval presence in each injury is increased. We can deduce that necrosed

tissue generally indicates that larvae are already attacking the fruit or

other leaves, or have left the canopy to pupate.

In a crop such as tomato where the margin of profit is great,

expensive methods of control are usually dictated. The use of a system

that will predict the damage level to the plant requires a high level of

accuracy. It is suggested that the method described here is advisable

for plants during stages TV1 to RV1.



Conclusions and General Discussion

Studies of tomato growth in different cropping seasons are useful

to determine effect of planting time on plant development. Tomatoes,

cv Flora-Dade, planted later in the winter have less (ca. 117 dm2)

leaf area than those planted early in the fall (ca. 253 dm2 leaf area).

Thus, those crops planted in October-November may be able to support

more damage than those planted in January-February. The proposed system

divides the plant stages into 2 vegetative stages (TV1 TV2), 3 repro-

ductive stages (TR1, TR2 and TR3), and a senescent stage (S ). The

description of the developmental stages of tomato can aid in using pest

management tactics. Definition of shorter developmental stages with

















Table 6. Percentage of tomato pinworm larval occurrence in foliar
injuries with different phenological characteristics.


Damage Description


Percentage of TPW Larval Occurrence
Present Absent


Transparent blotch 72.5 27.5

Necrosed blotch 23.3 76.6

Folded, no necrosed leaf 96.87 3.12

Folded, necrosed leaf 81.25 18.75










with more subdivisions would enhance phenological plant description.

This may allow better pest monitoring when plant development is in the

TR stage.
2
Results on leaf weight consumed (mg) and leaf area consumed (cm2

provided information on increments of those parameters for each larval

instar. Standard error and confidence intervals demonstrated a high

variability for both methods. Further research is necessary to deter-

mine if such variability is caused by larval behavior or by use of

different leaflet sizes and leaf area. I consider the leaf weight

method promising in such areas as plant resistance and behavioral

chemicals (deterrents) evaluation. Damage assessment based on the

leaf area mined by TPW is not considered appropriate for monitoring

TPW density because of inherent variability in insect behavior and

plant morphology.

Injury length has proven useful in evaluating plant resistance

(Schuster 1977a). The relationship between larval head capsule

and injury length was intermediate (r =0.47). The regression equation

developed in this study can be used by plant resistance evaluators to

determine feeding inhibition at a given instar. This technique has

to be carefully used, however, since it is dependent on the type of

leaflet consumed. Larvae that attack small leaflets might develop as

well as one in a large leaflet but the injury length will be smaller.

Data gathered from the visual damage classification proved to be

useful to evaluate damage inflicted by TPW. Since TPW instars have a

distinct behavior as leaf blotchers and leaf tiers, it will be easier

to develop knowledge in which the average larval instar will determine

the damage degree in a plant.






54


Scouts should use different techniques at the same time if

possible. A population index, degree damage scale and a survey deter-

mining the real presence of the larvae in the foliage will provide a

better estimate than a single technique. More research is needed to

evaluate these techniques together. Evaluation should be based on time

expended and reliability of the methods. Further study of the relation-

ship between several types of foliar damage and direct damage to the

tomato fruit is needed.

















CHAPTER III
SPATIAL DISPERSION OF TOMATO PINWORM EGGS ON TOMATOES



Introduction

Tomato pinworm (TPW) is one of the most important pests of tomato

Lycopersicon esculentum (Mill.) (Watson and Thompson 1932, Oatman 1970,

Poe et al. 1974). Little is known, however, about ovipositional pat-

terns of this pest on tomato plants under field conditions. There is

some indication that caged moths under laboratory conditions deposit

eggs indiscriminately on all parts of the plant including the upper

leaves (Elmore and Howland 1943). Wellik et al. (1979) indicated that

lower portions of the plant should be examined in the field for both

larvae and eggs of the TPW.

Studies of TPW egg dispersion are necessary because this knowledge

affects the sampling program as well as the method of analyzing the

data. Furthermore, dispersion patterns can be used to give a measure

of population size as well as to describe the factors that may affect

the condition of the population. This paper (1) describes the spatial

distribution of TPW eggs on field-grown tomato plants under varying

levels of TPW infestation, (2) presents an evaluation and discussion

of factors affecting this distribution and (3) discusses sampling

strategy.









Materials and Methods

Experimental Plots

To test for a possible relationship between oviposition of TPW and

different leaf strata of tomato cv Flora-Dade, 8 plantings (Oct. 3, 1979;

Dec. 5, 1979; Jan. 8, 1980; Oct. 30, 1980; Nov. 25, 1980; Dec. 30, 1980;

Jan. 30, 1981; Feb. 28, 1981) of non-staked tomatoes were evaluated at

the Agricultural Research and Education Center, University of Florida,

Homestead, Florida. Each planting (ca. 450-947 plants) was direct-seeded

in raised beds (3-5) (ca. 45 m long) of Rockdale soil, and mulched with

light colored plastic. The seedbed's midlines were 182 cm apart. Plants

were spaced 38 cm apart.



Sampling Methods

Sample size was selected by a preliminary random sampling of 50

plants on 2 dates. The method described by Elliott (1979) was adopted.

The relative variation (SE/x) x 100) was calculated to compare sam-

pling methods over a variety of sampling units (Hillhouse and

Pitre 1974, Ruesink 1980). Ten to twenty plants in each planting

were randomly selected on a weekly basis from February 7, 1980,

through May, 1980, and from Jan. 27, 1981, through May, 1981. Whole

leaves of the plant were first examined to determine differences in ovi-

position on lower and upper leaf surfaces (Plantings 1-3) and to detect

differences in oviposition in different plant strata (Plantings 1-8).

A plant was divided into upper half and lower half in the first 3

plantings (1979-80) and divided serially into 6 sections (upper, middle

and lower of each of the external and internal canopies) (1980-1981).










External canopy was defined as extending from the periphery to 5-15 cm

into the plant interior. The variance was stabilized by fitting the

number of eggs obtained to a suitable model (Poisson and negative

binomial) and transforming to logarithm (x+l) or x+0.5 (Elliott 1979)

depending upon the original frequency distribution of the counts.

The mean counts of eggs in upper and lower strata were compared by

student's t-test for plantings 1-3. Egg densities in the 6 strata for

plantings 4-8 were compared by analysis of variance (ANOVA). Means

were grouped by Duncan's Multiple Range Test (P=0.05). When tests

indicated significant differences in egg densities between strata of

plantings (4-8), optimum sample allocation among strata was determined

for each planting date (Cochran 1977).



Population Distribution Related to Leaf Position

To test differences in oviposition of TPW related to the vertical

distribution of the leaves with respect to the main axis, 17 randomly

sampled plants, each of which were 45 days old, were observed in a

commercial field. Leaves were numbered from bottom to top and the num-

ber of eggs recorded. Data were analyzed by ANOVA and means were separated

by use of Duncan's Multiple Range Test (P=0.05). When t-tests indicated

significant differences in egg densities between leaves, optimum sample

allocation among leaves was determined (Cochran 1977).



Distances Between Eggs and Effects on Distribution

To determine if TPW egg distribution pattern is influenced by leaf-

let size and egg density, the frequency of egg deposition on each










leaflet was recorded. Then, distance between eggs on each leaflet was

counted on 40 middle leaves collected from plants located in the same

field mentioned before. Several authors (Cottam and Curtis 1956)

proposed methods to evaluate randomness in spatial distribution of the

population by measurement of distances between individuals. In this

experiment, distance between eggs was checked by measuring the shortest

straight line between nearest neighbors with a metric ruler. Distance

accuracy was 0.05-0.25 cm. The frequency of occurrence of each dis-

tance was evaluated for egg densities. Also, simple linear regression

was applied to determine any relation between egg density and leaflet

size.



Oviposition Related to Plant Age

To determine if plant age affects oviposition, the number of eggs

on each plant was counted on 60-80 plants ranging in age from 2 to 21

weeks. Plants in this experiment were in the same field as previously

described tests (plantings 4-8). Plants were inspected weekly during

April and May, when the highest peaks of oviposition occurred. Data

were subjected to ANOVA, and means separated by use of Duncan's Multiple

Range Test (P=0.05).



Results and Discussion

Selection of Number of Sample Units

The main objective of planning a survey should be to obtain the

required information with a minimum amount of labor. To achieve this,

it is necessary to select a number of sample units that are in agreement










with the desired degree of precision and cost. This requirement is

difficult to meet in practice. First, an acceptable index of precision

(SEx100) is 25% (Barfield 1981). Secondly, the actual cost of sampling
x
tomatoes is 7 dollars per acre (Table 7).

Sample size was selected by a preliminary random sampling of 50

plants in 2 dates (Table 6). Three major criteria were followed to

select sample size. First, following the criteria outlined by Elliott

(1979), a suitable sample size was selected when the mean value ceased to

fluctuate. It is observed (Table 5) that with an increase in sample

size from 10 to 25 (at low egg density), the resultant mean (x)

fluctuates around 0.4-1.5 eggs/plant. Also, at higher egg density (2-7

eggs/plant) the number of selected sampling units is 20-25. Second,

the use of index of precision (SExlOO) over different sampling units is
x
a more adequate technique to select sample unit size.

Accordingly, the lower index of precision (Ip) was obtained when

the number of samples equals 50. Therefore, the percentage of the

standard error of the mean can be 34% if the TPW egg density per plant

is low (0.4-1.5 eggs/plant). This percentage is not good enough to make pest

management decisions. The index of precision can be 20% if the density is

higher (2-7 eggs/plant). Third, the sample number does not reconcile with

the actual budget per acre. Cost of sampling eggs is 1.4-77 dollars

(Table-5) more expensive than the actual sampling cost per acre.

The number of samples for a fixed level of precision (random sampling)

was calculated. A random egg distribution was assumed, n=(=-)2 where,
Ex

n=number of samples required, s=standard deviation, x=mean, and

E=predetermined standard error (e.g., 0.25). For instance, at























N0 Ckm'
rN m 00


H CNO CN k


0



aC
0


4
0
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4-1













02
a)

O



m







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ul




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a)

















04







0
-1





r
0



Ul
r-'-


04




0








-1
4-4i






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41

0- CO

COr








Ci
& o


COH
OHri-


o a' O> (oF W
r U)

m Ion c F






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* mIr
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01 M M01 0


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5 -I


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)
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n OD











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o a


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o
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o












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(a r


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01 0










04 0 (









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endemic levels of TPW egg population (0.4-1.33), the number of samples

to be taken, being S=2.79, E=0.25, x=1.15 will be 94, with a cost of

158 dollars per acre. If the TPW egg population is epidemic (2.16-7),

the number of samples to be taken will be 28, being s=3.82, x=2.9 and

E=0.25. The cost of sampling will be 47 dollars per acre.

Accordingly, under low TPW egg densities, increasing sample

precision as the sample size increases is not worth the work required

in taking larger samples. Consequently, I selected sample sizes of

10-20 which gave the best practical results per unit of work expended

($16.8-25.2 dollars per acre). It is considered that sampling TPW

eggs is not a practical method to make spray decisions.



Statistical Description of TPW Egg Spatial Distribution

The use of statistical methods, e.g., t-tests, analysis of vari-

ance, involves several conditions described by Snedecor and Cochran

(1967). One of them is that data must follow a normal distribution.

The distributions of density measurements on plant samples are sum-

marized for each planting in the Appendix. These statistics (Table 8)

support the hypothesis that TPW eggs are clustered on plants. This

clustering was more apparent when TPW egg densities on each plant

ranged from 0.302-1.3. As mean densities increased, variance also in-

creased except for planting 6. Values of the negative binomial parameter

(k) (Elliott 1979) range from 0.451-0.013 for my sampling. Thirty-two

percent of the weekly counts for each planting were fitted to the nega-

tive binomial distribution (see Appendix). For plantings with higher






















-E

-4



0




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C




4-i
a)

















-4








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04







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ar























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ct
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population densities (average 0.302-1.30), kurtosis and skewness

decreased as the mean increased. Skewness values were all positive.

This indicates that egg distribution tails off among higher counts.

This information in conjunction with the data indicating clumping can

aid in sampling design.



Distribution of Eggs on the Upper and Lower Surfaces of Leaves

Statistically significant differences (P=0.01) were found for egg

numbers on lower and upper leaf surfaces. Eighty-nine percent of

the total eggs found per plant were on the lower surface (Table 9).

These results and the results from the greenhouse contrast with those

found in caged plants by Elmore and Howland (1943), who detected 45%

of all egg deposition on the upper surface of the leaves. Insect pre-

ferences for oviposition on the underside might be correlated with dif-

ferences in pubescence of the 2 leaf surfaces. The average number of

trichomes on the underside was 1441 per leaflet as opposed to 469 on

the upper surface. This may also indicate preference to avoid egg

desiccation, or to avoid higher light intensities during oviposition

(Hinton 1981).



Distribution of Eggs on Upper and Lower Halves of the Plant

Statistically significant differences (P=0.05) were found in the

number of eggs deposited on the upper half of the plant vs the lower half

of the plant for the 3 sample dates in the first planting (Table 10). The

upper part of the plant had more eggs on 13 of the 15 sampling dates.

There were no significant differences between upper and lower halves

















Table 9. Ovipositional preference of tomato pinworm
for upper and lower surfaces of tomato
leaves from plants grown under greenhouse
and field conditions.


Mean Number of Eggs of TPW
Greenhouse
Leaf Side Fielda Caged Plantsb


Upper 0.857c 0.10c

Lower 7.3763 2.77


aMean based on counts from 80 plants.
b
Mean based on counts from 60 plants.
c
Numbers were significantly different at P=0.001.









65














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in the second planting (Dec., 1979). Analysis of the data from the third

planting (Jan., 1980) indicated significant differences in 6 of the 11

sampling dates. The upper half of the plant had more eggs except for

2 dates. In general, when numbers of eggs were higher in the lower

strata, this coincided with younger plant age (40-60 days after germin-

ation). Numbers of eggs were higher in the upper strata when plants

were in reproductive or older age (75-80 days after germination. These

data indicated that for 'Flora-Dade' ground tomatoes, ovipositional

preferences existed based on the level of the plant. Because of the low

economic threshold for TPW in tomatoes, it may be necessary to reduce

the sampling unit to detect major differences in internal and external

parts of the plant when populations are low. Consequently, smaller

sampling units were tested in subsequent experiments.



Distribution of TPW Eggs by Sampling Six Plant Strata

Statistically significant (P=0.05) (Table 11) differences were not

detected among the strata for the 4th (Oct., 1980) and 5th (Nov., 1980)

plantings possibly due to the relatively low mean egg numbers per plant.

However, the highest number of eggs oviposited was obtained in the upper

external canopy for planting 4 and in the middle internal canopy for

planting 5. There was an increase in eggs for the lower internal canopy

in planting 4 during January and February, when nocturnal temperatures

were lowest (20C), and an increase toward the upper external portion of

the plants when temperatures were fluctuating between 17-290C (April-May).








67















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r 4J o c o o o C

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SC C C





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4 -4 -r)


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Perhaps moths protect themselves from the cold temperatures by staying

close to the ground in the lower canopy. Despite these assumptions,

when number of eggs found per stratum was regressed (Table 12) against

temperature, there was no evidence of a relationship between the two

variables. Significant differences in numbers of eggs per stratum were

detected for the 6 (Dec., 1980), 7 (Jan., 1981) and 8 (Feb., 1981)

plantings. There was no significant variation among the six strata

during juvenile plant stages. Most of the significant differences were

observed (Fig. 6) during the mature stages (TR) of the plant. Concen-

trations of eggs in the upper external strata varied slightly among

plantings. In plantings 4 and 5, eggs generally occurred on the top and

middle external canopy during the last weeks of sampling (April and May),

and on all strata during the first weeks (juvenile stages) in March and

April. When mean numbers of eggs in the external and internal canopy

were added to reduce the strata to 3 (upper, middle and lower), no statis-

tical differences were observed despite the stratum reduction. This

agrees with the results expressed when 2 strata (upper and lower) were

sampled, indicating that differences in oviposition tend to be masked

if the units are widened. In general, the upper external stratum had the

highest number of eggs, followed by the middle external and internal strata,

during most of the sampling dates. At plantings 6 to 8, the TPW eggs

occurred in greatest abundance on the upper and middle external strata dur-

ing all growth stages. More eggs (44-68%) were deposited within the upper

external canopy of the plant than in any other stratum (Table 13). Four to

twenty eight percent of the eggs were laid in the next (middle external

stratum). The lower external stratum had the lowest range (1-11%); however,

















Table 12.


Relationship between daily mean temperature (C) and TPW
oviposition in 6 tomato plant strata. Homestead, Florida,
1981.


Dependent
Independent Variable
Variable No. Eggs/stratum ** ***
x y r r bo bl


Temperature upper internal 0.05 0.22 -0.007 0.005

upper external 0.13 0.36 0.46 0.04

middle internal 0.10 0.31 0.10 0.01

middle external 0.02 0.14 0.02 0.0056

lower internal 0.01 0.10 0.10 -0.02

lower external 0.13 0.36 -0.11 0.01


Coefficient of determination.
**
r
Correlation coefficient.
***
bo
Intercept of y axis.

Slope.

























4 *. O
CC OH

4- 0 0
H ** >





4t) r. 0)

-i > 0a




LO -4
S3 0) ( d






0 4--1 N
0) 4





3) cd-4
0)0 CC O

04 .) 4






H Cd c(0 3 0
CH H 0



4 -4 4J
O4)








-4 O4
Ha 0 0

n -4 0' 0











o o 40)
Ot QC( -4
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04 4 r4 a)
m 4 4-1 0













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4 *a 0)
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in 7 in
f0 W =3'










73






















w.

























L
U,

U I--





































m
I-





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74


































I--



















C= t c
r- -n %J
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mn o -S /so e o ua

unto Jt/s 3 B Q u oay4

















Table 13.


Percentage distribution of TPW eggs for each stratum of
tomato plants in 5 tomato plantings. Homestead, Dade
County, Florida, 1980.


Planting Date
Oct. 30 Nov. 25 Dec. 31 Jan. 30 Feb. 28
Stratum 1980 1980 1980 1981 1981


Upper external 68 32 44 46 51

Middle external 4 15 23 28 21

Lower external 1 3 10 11 10

Upper internal 0 3 7 0.6 5

Middle internal 9 43 11 10 9

Lower internal 16 0.8 4 1 2










growth of the plant upwards and outwards can mislead my interpretation

of actual ovipositional preference. The percentage of eggs found per

internal stratum ranged from 0-7% in the upper internal, 9-43% in the

middle internal, and 10-87% in the lower internal. TPW oviposits mainly

in the upper external canopy when egg populations range from 0.75-1.5

and when the plant was in its reproductive stage. A lower proportion of

eggs was found in all other strata.

Sampling 6 plant strata demonstrated that TPW tends to oviposit

in the middle and upper canopy. It is necessary to use sample allocation

(nh), as outlined by Cochran (1977) to minimize sampling cost or vari-

ance (s 2). I assumed equal sampling cost for each stratum. Sample

allocation was estimated on dates in which statistical differences in

oviposition were detected.

In general, more samples should be allocated to the upper and middle

external strata (Table 14). Because TPW eggs are clumped in the upper

and middle canopy, these strata had the highest variance (see Appendix)

(Tables 51-54). For a fixed total cost, n = (C-C ) NhSh where (C-C )=
o C-, o

L Nh Sh/ C
I C.n., nh= n W Sh n NnSh Therefore as S- increases so does nh.
ii hh= h. h
i=l WhSh ENhSh

The average number (n=20) for all planting dates was 6, 5 and 3 samples

from upper, middle and lower external canopy, and 1, 4 and 1 from upper,

middle and lower internal canopy. Allocation ranged from 5-10 samples

for the upper external canopy (see Appendix), and ranged from 2-8 samples

for the middle external canopy. I considered this sample allocation to

to be the best, because standard error (SE) of the sample mean was more


















0




o
(n







r4>
a4 r-
4J









cn
cy u




r- a)



*d U)

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C)


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0







0 -,
0 0


O-
-4 4






m

(0 -






(,
, -I




0C0)





r-4


O O m N -I 0 0 0 -I 0 0 0 N r-4










^ CO
in tn
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(N TI

























0 0 %
Lo n NON
(N In :T I 0 m CO in 'o 'r N In (N In









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(N ,-4 (N (N CN C4 (N (N (N cN (N (N (N 0)
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41




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en s -









constant through time (range: 0.20-0.66). There were exceptions for

these sample allocations. For instance, during the month of February

(planting 4), more numbers of samples were allocated to the lower inter-

nal stratum (see Appendix) (Table 54). Another aspect that requires more

understanding is the relation between phenological stages and sample

allocation. As an example, it was observed (see Appendix) that when

plants were in vegetative stage (TV), more samples (nh=18), should be

allocated for the upper external and middle internal canopy. When

plants are in first reproductive stage (TR ), more samples (n =61, are

allocated for upper external stratum. Finally, when plants reach the

second reproductive stage (TR2), all strata had similar sample allo-

cations except for lower internal canopy (nh=0).




Egg Distribution Influenced by Leaf Position

During heavy oviposition (avg 21.94 eggs per plant) on 45 day-old

tomato plants, the highest number of eggs was observed on leaf number 4

(Table 15). The numbers of eggs on leaves 3 and 5 were statistically

equal to those found on leaf 4. The number of eggs decreased sharply on

leaves adjacent to the apical point toward the bottom of the plant

(leaves 1-2). Tjese results indicated that middle leaves of 45 day-old

plants under conditions of high egg oviposition (1-5.5 eggs per leaf)

have 65% of the total egg population. These data differ from those

obtained in experiment 1. The higher number of eggs per plant indicates

that the insect tends to oviposit in the upper-middle canopy, avoiding

the 2 top and bottom leaves of the plant. Several factors may influence

the ovipositional pattern. First, these results agree with Hinton (1981),

















Table 15. Mean tomato pinworm eggs on tomato leaves from different
strata of 45 day-old plants. Homestead, Florida, 1980.


Mean No. No.
Leaf Position Eggs/Leaf Leaflets/Leaf Eggs/Leaflet


1 bottom 2.20b* 7 0.31

2 3.20a 8 0.40

3 middle 5.40a 11 0.49

4 5.50a 11 0.50

5 5.00a 11 0.45

6 top 2.00b 8 0.25

7 1.00b 7 0.14

*
Numbers followed by different letters were significantly differ-
ent statistically at P=0.05 according to Duncan's Multiple Range
Test.









who stated that species that lay eggs on plants have a marked preference

for laying a certain height above the ground. Secondly, the insect may

be avoiding overcrowding in the smaller top leaves and competition of

foliar consumption by TPW larvae on the lower leaves. The highest

sample (n=17) allocation was for leaves in the middle canopy (Table 16).

Higher variance (s2=44.4) was found for eggs deposited on those leaves,

as was a high mean (x=6.6). This is caused by egg clumping in the canopy.

The fourth leaf had the highest allocation sample (nh=5), followed by the

fifth leaf (nh=4). The lowest allocation was for the bottom leaf (nh=l).

The standard error of the mean sample was lowest (SE/x=0.21), for the

third leaf and slightly higher (SE/x=0.24) for the fourth leaf. There-

fore, when higher density and large variance are found, the leaves

selected should be the middle ones. Sample allocation was reduced for

bottom and top leaves. These leaves had smaller variance and smaller

density than the middle ones.



Distances Between Eggs per Leaflet and Effect on Distribution

In the present study, the results indicated that TPW egg density

was not related to leaflet area (Table 17). The coefficient of deter-

mination (r2=0.026) indicated that females tend to oviposit different

egg densities disregarding leaflet size. Therefore, any leaflet can be

selected as the sampling unit. Frequency of egg occurrence per leaflet

was not related to distance between eggs. Low coefficients of deter-

mination (r2=0.19-0.23) between frequency of occurrence at different egg

densities (2, 5 and 10 eggs/leaflet) and egg distances indicate lack of

linear relationship between these variables. The slope (bl) obtained
























U)








4-i
4-







4-)
0
0









w
4





O)














0



0



4-)
0
(d





-40
0)
-!

C)
04








aU


-.~ o in o
o 10 oN 0

N in o N o
oco um


N w I I) c
IX En e \ .c
l
0n


n H n H
m m 0
H D N.

















co o ON o






0 0 N
I m ON oN





m 0 m
C 0 N 0
in o m o
i-i~ r


^r m o
*q o TI (



c o m o


II


0
0
C






.1



II
i ^

















Table 17.


Relationship between frequency of occurrence of TPW eggs per
leaflet as dependent variable and distance among eggs and
leaflet area as independent variables.


2* ** ** t
Dependent Independent r r b b1
Variable Variable
y x


No eggs Leaflet area 0.026 0.16 1.99a 0.04

2 TPW eggs Distance among 0.23 0.48 1.75a -0.15b
eggs

5 TPW eggs Distance among 0.19 0.44 1.47a -0.12b
eggs

10 TPW eggs Distance among 0.23 0.48 1.38a -0.22b
eggs


* 2
r =coefficient


of determination.


r=correlation coefficient.
k*
bo=intercept of y asis.
bl,=slope.

numbers were highly statistically significant (P=0.01).
numbers were statistically significant (P=0.05).










for any egg density was negative and highly significant (P=0.001). This

can be explained in Fig. 7, where the frequency of egg occurrence at dis-

tances higher than 3 cm was as low as 5%. The average distance between

eggs was 0.5-0.75 cm. The average number of eggs found on each leaflet

was 2-3. These results agree with those expressed by Poe (1973); in the

present study the number of eggs on each leaflet was as high as 11.

Eggs tended to be laid more uniformly in some parts of the leaflet. Per-

haps the female lays 2 eggs successively on a certain part of the leaf-

let, but is likely to move away after oviposition. The arrangement of

eggs may also be a reflection of heterogeneity of conditions among parts

of a leaflet such as pubescence and leaf venation. From the practical

standpoint, these results can be used to determine use of single leaflets

as less variable sampling units compared to the whole plant. A more

detailed study of female behavior is necessary to determine the role of

leaf factors (e.g., pubescence) affecting oviposition.



Differences in Oviposition Related to Plant Age

The relationship between oviposition and stage of plant development

was determined during the study of plantings 4-8. Statistical differ-

ences were detected among these plantings (Table 18), when plantings

were 19, 15, 11, 7 and 3 weeks old (stages S1, TR2, TR TV2, TV1 respec-

tively. The largest number of eggs was detected in planting 7, when

this planting was in the TR TR2 stages. At the same time, egg num-

bers decreased for planting 6 after the 10th week of plant growth. The

mean number of eggs in planting 8 increased slightly from week 3 (TV2),

through 7 (TR ). These data indicate that there may be several factors,




Full Text
TOMATO PINWCRM, KEIFERIA LYCOPERSICELLA (WALSINGKAM): POPULATION
DYNAMICS AND ASSESSMENT OF PLANT INJURY IN SOUTHERN FLORIDA
By
JORGE S. PENA
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR
THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1933

ACKNOWLEDGMENTS
I thank Dr. Van Waddill, my advisor and chairman, for his
encouragement, support and friendship, but most of all for his valuable
suggestions and allowing me freedom to conduct my research.
I would also like to express my appreciation to the following
people:
Dr. S.K. Karr for his interest in teaching me the art of
communication and also for his help in solving administrative problems
during my studies.
Dr. J.L. Stimac for his help and constructive criticism, as
well as his ideas to improve the quality of this study.
Dr. K.H. Pohronezny for his constructive criticism, suggestions
and for reviewing this manuscript.
Dr. D.J. Schuster for supplying material for my research as
well as his interest in this study.
The Agricultural Research and Education Center, Homestead,
Florida, and to Dade Agricultural Council for providing the grantmanship
and scholarship to support my studies.
The staff of AREC, Homestead, for their cooperation, espe¬
cially Connie Csterholdt, Carolyn Reitman, Susan Housley, Leslie
Sawyerlcng, Rodney Chambers, Linda Douthit, and Wilbur Dankers for
n

their help during data collection. Mrs. Sheila Eldridge and Mrs.
Barbara Hollien for kindly typing this manuscript.
Drs. R.E. Litz and S.K. O'Hair for their friendship, encourage¬
ment and support during the past years.
Mr. Ben Gregory for his honest friendship and willingness to
share ideas in research.
Ms. Annie Yao, Mr. A. Bustillo, Mr. W. Chongrattanameteekul,
Mr. K. Patel and Mr. C. Ho for their friendship and support.
I am indebted to my family for their love and encouragement,
the Litz family, Eleanor Merritt and Bunny Hendrix for their friend¬
ship and support.
lu

TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS ii
LIST OF TABLES vi
LIST OF FIGURES xi
ABSTRACT xv
INTRODUCTION 1
CHAPTER I: LITERATURE REVIEW 6
Family Gelechiidae 6
Studies on Keiferia lvcopersicella 7
Tomato Plant Phenology and Measurement of TPW
Dispersion and Economic Damage 19
Environmental Factors Affecting TPW Population .... 21
CHAPTER II: DESCRIPTION OF TOMATO PLANT PHENOLOGY AND EVALUATION
OF TOMATO PINWORM FOLIAR DAMAGE ASSESSMENT 25
Introduction 25
Materials and Methods 26
Results and Discussion 30
Conclusions and General Discussion 51
CHAPTER III: SPATIAL DISPERSION OF TOMATO PINWORM EGGS ON TOMATOES. 55
Introduction 55
Materials and Methods 56
Results and Discussion 58
Conclusions and General Discussion 87
CHAPTER IV: SPATIAL PATTERNS OF DISPERSION OF TOMATO PINWORM LARVAE
IN TOMATOES 89
Introduction 89
Materials and Methods 90
Results and Discussion 91
Conclusions and General Discussion 117
IV

CHAPTER V: TOMATO PINWORM ARTIFICIAL INFESTATION: EFFECT OF
FOLIAR AND FRUIT INJURY ON GROUND TOMATOES 119
Introduction 119
Materials and Methods 120
Results and Discussion 122
Conclusions and General Discussion 152
CHAPTER VI: ADULT DISPERSION AND COLONIZATION OF TOMATO
FIELDS BY THE TOMATO PINWORM 154
Introduction 154
Materials and Methods 155
Results and Discussion 157
Conclusions and General Discussion 169
CHAPTER VII: EGG AND LARVAL PARASITISM OF TOMATO PINWORM IN
SOUTHERN FLORIDA 171
Introduction 171
Materials and Methods 172
Results and Discussion 174
Conclusions and General Discussion 189
CHAPTER VIII: EFFECTS OF RAINFALL AND RELATIVE HUMIDITY ON
IMMATURE STAGES OF THE TOMATO PINWORM UNDER
GREENHOUSE AND FIELD CONDITIONS 190
Introduction 190
Materials and Methods 190
Results and Discussion 194
Conclusions and General Discussion 211
CHAPTER IX: INFLUENCE OF POST-HARVEST TOMATO FIELDS ON THE
POPULATION DYNAMICS OF THE TOMATO PINWORM 213
Introduction 213
Material and Methods 213
Results and Discussion 217
CONCLUSIONS AND GENERAL DISCUSSION 225
REFERENCES 230
APPENDIX EXPLANATORY TABLES FOR CHAPTERS II AND III 243
BIOGRAPHICAL SKETCH 265
v

LIST CF TABLES
Table Page
1 Larval parasites of Keiferia lycopersicella
reported from U.S.A. and South America until
1981 1
2
3
4
Classification of tomato pinworm leaf
damage on 'Flora-Dade' tomatoes, based on
greenhouse and field observations. Homestead,
Florida, 1980
Leaf area and reproductive plant structures in
tomatoes, cv Flora-Dade', planted on 5 dates
in Homestead, Dade County, Florida during 1980-
1931
Stage of development description for tomato cv
Flora-Dade. Description is based on the average
of observations from tomato plants grown during
Fall 1980 through Winter 1981. Homestead,
Florida
5 Tomato leaf weight and leaf area consumed by
different larval instars of Keiferia lycopersicella
under greenhouse conditions; T 24+_3°C, 75+2% RH. . .
6 Percentage of tomato pinworm larval occurrence in
foliar injuries with different phenological charac¬
teristics
7
8
o
Comparison of different sample sizes for tomato
pinworm eggs. Homestead, Dade County, Florida,
1980
Mean number of tomato pinworm eggs per plant by
planting date for 8 tomato plantings in Homestead,
Florida, 1979-1981
Ovipositional preference of tomato pinworm for
upper and lower surfaces of tomato leaves from
plants grown under greenhouse and field
conditions
60
62
64
vi

Table
Page
10 Mean number of tomato pinworm eggs in 2 plant
strata (upper and lower halves) per plant at
different sampling dates. Homestead, Dade
County, Florida, 1980 65
11 Mean number of tomato pinworm eggs per plant in
6 strata: upper, middle and lower external; upper,
middle and lower internal canopy of the tomato
plant. Homestead, Florida, 1981 67
12 Relationship between daily mean temperature (°C)
and TPW oviposition in 6 tomato plant strata.
Homestead, Florida, 1981 71
13 Percentage distribution of TPW eggs for each stratum
of tomato plants in 5 tomat plantings. Homestead,
Dade County, Florida, 1980 75
14 TPW egg sample allocation for 6 plant strata during
3 different plant stages: second vegetative (TR^),
first reproductive (TR^), and second reproductive
stage 77
15 Mean tomato pinworm eggs on tomato leaves from
different strata of 45 day-old plants. Homestead,
Florida, 1980 79
16 TPW egg sample allocation on tomato leaves numbered
from bottom to top. Plants 45 days old 81
17 Relationship between frequency of occurrence of TPW
eggs per leaflet as dependent variable and distance
among eggs and leaflet area as independent vari¬
ables 82
18 TPW oviposition on tomato at different plant stages.
Homestead, Florida, 1981 86
19 Sample size and relative net precision (RNP) for
sampling injuries at low and high population
densities. Homestead, Dade County, Florida, 1980. . 93
20 Mean number of TPW foliar injuries and standard
error on different sampling units at specified
date. Crop planted in Nov., 1979. Homestead,
Florida 96
vii

22
23
24
25
26
27
28
29
30
31
Page
Mean number of TPW foliar injuries and standard
error on different sampling units at specified
date. Crop planted in Jan., 1980. Homestead,
Florida 97
Sample size and relative net precision (RNP) for
sampling TPW larval injuries on upper and lower
plant canopy. Homestead, Florida, 1980 100
Sample statistics: Mean tomato pinworm larval in¬
juries per plant in 8 tomato plantings. Homestead,
Florida, 1979-81 102
Mean tomato pinworm (TPW) foliar larval injuries
at 2 different plant levels for 3 different
plantings. Homestead, Dade County, Florida, 1980. . . 103
Mean tomato pinworm (TPW) larval injuries in 6
plant strata for 5 plantings. Homestead, Dade
County, Florida, 1981 105
TPW larval injury sample allocation for 6 plane
strata at 3 different plant stages: second repro¬
ductive (TR2), third reproductive (TR^) and
senescent (S^) 113
Mean number and standard error of tomato pinworm
(TPW) injuries in 5 different plantings at speci¬
fied date and plant growth stage 115
Tomato fruit damaged in the upper and lower plant
canopy, after a single artificial infestation with
K. lvcopersicella larvae on ground tomatoes .... 124
Marketable value for tomato fruit damaged in the
lower and upper plant canopy after a single arti¬
ficial larval infestation of K. lycopersicella
on ground tomatoes 125
Tomato fruit damaged in the lower and upper plant
canopy after a double artificial infestation of
K. lycopersicella larvae on ground tomatoes 126
Marketable value for the tomato fruit damaged in
the lower and upper plant canopy after a double
infestation of K. lycopersicella on ground
tomatoes
viii
128

Table
Page
32
33
34
35
36
37
38
39
40
41
42
43
44
Effect of planting time on fruit injured by
K. lycopersicella larvae to ground tomatoes,
cv 'Flora-Dade' during 1981
Differences in cost and relative net precision
between sampling 6 plants per row and 1 random
plant per row
Differences in mean fruit injured by K.
lycopersicella in pruned and not pruned
tomato plants
Effect of hedges and edgerows on tomato pinworm
field infestation at three fields in Home¬
stead, Florida, 1981
Parasitism of the tomato pinworm larvae in
tomato fields in southern Florida, Dade
County, 1980
Parasitism of the tomato pinworm larvae in
tomato fields in southern Florida, Dade County,
1981
Keiferia lycopersicella eggs parasitism by 2
strains of Trichogramma pretiosum in the
laboratory. T25+1°C; 75+2% RH
Number of Keiferia lycopersicella eggs col¬
lected from two strata and percent of para¬
sitism by Trichogramma pretiosum
Distribution of normal and parasitized tomato
pinworm eggs in 2 tomato fields
Parasitism of tomato pinworm eggs by Tricho¬
gramma pretiosum in 2 fields with different
host densities
149
150
151
166
175
177
179
180
181
183
Plant water content in five tomato plantings
related to oviposition by the tomato pinworm
Effect of simulated rainfall on foliar larval
injuries caused by the tomato pinworm Keiferia
lycopersicella on plants grown under greenhouse
conditions
Mean percentage of tomato pinworm adults
emerged by day after pupal treatment with
different simulated rainfall regimes
203
204
208
ix

45
46
47
43
49
50
51
52
53
54
55
56
57
Page
Effect of crop age of post-harvested tomato
plants on volunteer plants and number of tomato
pinworm larval injuries
General effect of cultural practices on
volunteer tomatoes and infestation by
tomato pinworm
Effect of planting age and cultural practices
on volunteer tomato plants and number of TPW
injuries
Tomato pinworm egg frequency distributions
determined on tomato plants during 1981 . . . .
Tomato pinworm foliar injury frequency
distributions determined on tomato plants
during 1980
Tomato pinworm foliar injury frequency
distributions determined on tomato plants
during 1981
TPW egg allocation sample for 6 plant strata.
Planting 8, 1981. Age: 38 days. Stage of
development TV„
TPW egg allocation sample for 6 plant strata.
Planting 8, 1981. Age: 46 days. Stage of
development TR^
TPW egg allocation sample for 6 plant strata.
Planting 7, 1981. Age: 68 days. Stage of
development TR^
TPW egg allocation sample for 6 plant strata.
Planting 4, 30 Oct. 1980. Age: 77 days. Stage
of development TR^
TPW larval injury sample allocation for 6 plant
strata. Planting 6, 1981. Age: 78 days.
Stage of development TR^
TPW larval injury sample allocation for 6
plant strata. Planting 5, 1981. Age: 108
days. Stage of development TR^
TPW larval injury sample allocation for 6
plant strata. Planting 4, 1981. Age: 120
days. Stage of development TR
220
222
223
244
249
253
258
259
260
261
262
263
264
x

LIST OF FIGURES
Figure
Page
1 Illustration of tomato cv Flora-Dade growth at 2 stages
of development. TV2=second vegetative stage; TR^=early
reproductive stage; a=primary leaf; b=lateral develop¬
ment
2
2 Influence of time on leaf area (dm ) expansion, flower¬
ing and tomato fruit numbers of cv Flora-Dade grown
on 'Rockdale' soil under southern Florida conditions . . . .
3 Stages of development of tomato. TV^=early vegetative
stage; TV =late vegetative stage; TR^, TR^, TR3=rePro-
ductive stages; S^=senescent stage
4 Linear relationship between tomato pinworm (Keiferia
lycopersicella) larval head capsule width (mm) and
foliar injury length, r2=0.47
5Linear relationship between tomato pinworm (Keiferia
lycopersicella) larval instars and visual leaf damage
scaled r2=0.677
6 Average number of tomato pinworm eggs per plant
stratum during 6 different sampling dates in 2 to¬
mato plantings at different growth stages.
A) Planting 7: Jan. 30, 1981. B) Planting 8: Feb.
28, 1981. TR =second reproductive stage of develop¬
ment; TV9=sec¿nd vegetative stage of development.
Plant strata: 1, 2, 3: upper, middle, lower external,
4, 5, 6: upper, middle, lower internal
7 Frequency of tomato pinworm eggs at different distances
(cm) between eggs when mean eggs were A) 2 eggs per leaf¬
let and B) 5 eggs per leaflet
8 Percentage of tomato pinworm (TPW) larval injuries in 2
sampling units from different plant portions, related to
number of injuries in the whole plant: 1) 1st planting,
Nov. 3, 1979; 2) 3rd planting, Jan. 8, 1980
xi

Fig
9
10
11
12
13
14
15
16
Page
Percentage of tomato pinworm (TPW) foliar injuries found
at upper, medium and lower stratum in 4 tomato plantings:
1) Oct. 30, 1980; 2) Nov. 25, 1980; 3) Dec. 30, 1980; and
4) Jan. 30, 1981. Bars followed by different letters were
significantly different according to Duncan’s Multiple
Range Test (P=0.05). Percentages were previously
transformed to arc sine. Percentages are expressed as
actual numbers before transformations 108
Percentage of larval injuries at the external and
internal canopy evaluated from 5 tomato plantings.
Plantings 4, 5 and 6 planted in Oct., Nov., and Dec.,
1980; Plantings 7 and 8 planted in Jan. and Feb., 1981.
Homestead, Florida, 1980-81 Ill
Relationship between number of tomato pinworm
larvae per plant and number of injured fruits
and leaves in the lower plant canopy by a
single artificial infestation with TPW larvae.
Homestead, Florida, 1980-81 130
Relationship between number of tomato pinworm
larvae per plant and number of injured fruits
and leaves in the upper plant canopy by a single
infestation of TPW larvae. Homestead, Florida,
1980-81 132
Relationship between number of leaves injured
in upper and lower canopy and number of fruits
injured in upper and lower canopy by a single
artificial infestation with TPW larvae. Home¬
stead, Florida, 1980-81 135
Relationship between number of larvae per plant
and number of injured fruits and leaves (upper
and lower canopy) after a double artificial in¬
festation with TPW larvae. Homestead, Florida,
1980-81 137
Relationship between number of leaves injured
in upper and lower canopy and number of fruits
injured in upper and lower canopy by a double
artificial infestation with TPW larvae. Home¬
stead, Florida, 1980-81 140
Regression of percent of yield reduction
against infestation densities per plant of
tomato pinworm larvae 144
xii

Figure
17 Regression of percent yield reduction of to¬
mato fruit against TPW number of foliar
injuries per plant
18 Abundance of tomato pinworm male moths at three
different field sites. A) Field 1, 1980; B)
Field 2, 1980; C) Field 3, 1981. Mean number
of moths at each site corresponds to the average
from 4 pheromone traps in north, south, west and
east directions. Homestead, Florida, 1980-81 . . . .
19 Mean number of tomato pinworm larval injuries
occurring in 4 commercial fields at 0, 30, and
120 m from the field border. Bars with the letter
"a" denote statistical differences at 0.05% dif¬
ference level for a particular date
20 Field plan: Fields 1, 2, and 3, respectively,
each field was divided into 8-9 quadrats. The shaded
areas represent higher insect populations. Home¬
stead, Florida, 1981
21 A-3. Relationship of Keiferia lycopersicella
egg density to percent parasitism by Trichogramma
pretiosum in 2 fields
22 A-C. Seasonal occurrence of Keiferia lycopersicella
eggs and parasitization by T^. pretiosum in tomato
fields located at the (a) northern, (b) middle, and
(c) southern areas of Dade County, Florida
23 Seasonal abundance of TPW eggs in experimental
fields, related to temperature and rainfall
regimes during A) 1980, and B) 1981, in Home¬
stead, Florida
24 Seasonal abundance of TPW larvae in tomato fields,
related to temperature and rainfall regimes during
1980-81, in Homestead, Florida
25 Mean number of TPW injuries per plant during 9
days of simulated rainfall under greenhouse con¬
ditions, avg. daily temperature 25+_2°C
26 Percentage of TPW adult emergence under greenhouse
conditions after treatment of pupae with 3 regimes
of artificial rainfall (200, 100, 50 and 0 ml water),
temperature 24+3°C
Page
146
159
164
168
185
188
196
200
206
xiii
210

Figure Page
27 Tomato field status following the main
harvest under S. Florida conditions. Home¬
stead, Florida, 1980 216
28 Number of tomato plants and TPW injuries
per m in 2 post-harvested tomato fields.
Homestead, Florida, 1980 219
xiv

Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
TOMATO PINWORM, KEIFERIA LYCOPERSICELLA (WALSINGHAM): POPULATION
DYNAMICS AND ASSESSMENT OF PLANT INJURY IN SOUTHERN FLORIDA
By
JORGE E. PENA
April 1983
Chairman: Dr. V.H. Waddill
Major Department: Entomology and Nematology
Experiments were conducted in Homestead, Florida, during 1979-1981
to describe tomato plant phenology, tomato pinworm (TPW), Keiferia lyco-
persicella (Walsingham), dispersion patterns, economic damage to tomato
and the effects of parasitoids, edgerows, rainfall and cultural practices
on TPW population dynamics.
Tomato cv. Flora-Dade phenology was described. Six stages were
designated based on the number of leaves, flowers, fruits and physiological
plant characteristics. This description can be of use in making pest
management decisions.
Based in the relative variation (RV) and sampling costs, sampling
units for TPW egg and larval stages were determined. Eggs were generally
(51%) found in the upper plant canopy, and larvae (50%) in the lower
:v

plant canopy. Larger sampling units were allocated to the upper
and lower plant canopy for eggs and larvae, respectively. An
economic injury level was determined to be 1 larva per plant.
Yield can be reduced 10-40% when 1-12 larvae are attacking 45 day-old
plants. The results indicated a correlation between number of foliar
injuries in the lower plant canopy and fruit damage. In southern
Florida, higher TPW infestation occurred during March-May, 1980 and
March-April, 1981, compared with other months (Jan., Feb.). Trichogramma
pretiosum Riley caused 33-73% TPW egg mortality during May-July, 1981.
TPW larval parasitism fluctuated between 39-42% during 1980-1981. The
most abundant larval parasite was Apanteies spp., followed by Sympiesis
stigmatipennis and Teme.lucha spp.
TPW adult dispersion and effects of field edges on TPW dispersion
and field colonization were evaluated. Field areas surrounded by edge-
rows had higher TPW damage than areas surrounded by pastures.
The use of artificial rainfall demonstrated that when plant foliage
was irrigated there was a behavioral change in larval feeding which
resulted in 50% reduction of larval injuries compared to injuries on
soil-irrigated plants. TPW adult emergence was reduced 86% when high
levels of water were applied to pupae in the soil.
The effect of cultural practices on the TPW oversummering popu-
2
lation was evaluated. The mean number of injuries per m was 28 times
higher in crops planted later (December, 1980). than in crops planted
earlier (October-November, 1980). Lower numbers of injuries were found
in crops disced and mowed than in abandoned fields.
xvi

Parasitoids,
patterns can have
cultural practices, and southern Florida climatological
an impact on TPW population levels.
xvi i

INTRODUCTION
The tomato, Lycopersiccn esculentum Mill., is one of the most popular
and important vegetables in the world (Purseglove 1968). Tomato produc¬
tion in the U.S.A. is concentrated in California, Florida, Texas, New
York, New Jersey, Michigan and Virginia (Thompson and Kelly 1957).
Florida tomato acreage was 31360 ha during 1980-1981. Tomato produc¬
tion is considered to comprise 28.31% of the total vegetable acreage
in Florida (Anonymous 1982). The tomato growing areas in Florida are
divided into 4 major districts: Palmetto-Ruskin, Pompano Beach-Fort Pierce,
Dade County and Immokalee-Naples (Anonymous 1981). Dade County has 18.3%
of the total state tomato production and supplies most of the winter (De¬
cember through February) vegetables for the U.S.A.
The cultivation of fresh market tomatoes demands a high monetary in¬
vestment from farmers. The cost of producing tomatoes in Dade County
during 1980 was $5123.25 per ha, which represents an increase of 1.44
times over the production cost of 1975 (Greene et al. 198C).
Expansion of tomato acreage in Florida resulted in changes of agro¬
nomic practices to maximize tomato production (Geraidson 1975). Changes
in horticultural practices also established an agro-ecosystem with ento¬
mological characteristics commcn to monocultures. From 1950-1975 insect
control in tomatoes was almost exclusively chemical. To help growers avoid
problems with insecticides such as insecticide resistance, secondary pest
1

2
outbreaks and objectionable pesticide residues, an integrated pest man¬
agement program was established in Dade County on tomatoes (Pohronezny
et al. 1978). This program goal was to develop economically, technically
and ecologically sound systems of integrated pest management. This
approach had some constraints, however, such as the high crop value
which reduces the use of pest management tactics (Bottrell 1979) . More¬
over, the fruit quality standards for fresh tomatoes cause undue emphasis
on chemical control measures in order to prevent contamination of fruit
by insects and to prevent cosmetic damage to the fruit (Lange and
Bronson 1981).
Accordingly, insect pests in tomatoes can be categorized as direct
pests and indirect pests. Direct pests attack the product and directly
destroy a significant part of its value. Indirect pests attack plant parts
other than the saleable product but may reduce yield of the product (Ruesink
and Kogan 1975). Among direct pests of tomato in Florida are the corn ear-
worm, Heliothis zea Boddie, the southern armyworm, Spodoptera eridania
(Cramer), and the tomato pinworm, Keiferia lycopersicella (Walsingham).
Indirect pests are the serpentine leafminer complex, Liriomyza spp, the
tobacco hornworm, Manduca sexta (Joh.), and the granulate cutworm, Feltia
subterránea (Fab) (Poe 1972).
The tomato pinworm (TPW) can be either a direct or indirect pest
of tomatoes. The larva of this insect feeds in the mesophyll of the
leaves causing a serpentine-type mine during the first 2 larval instars.
In the latter instars the larvae can cause a blotch-type mine or they
tie leaflets together. The larvae also bore into fruit, providing an
entrance for plant pathogens which cause major damage to fruit.

3
The importance of the TPW as one of the most serious pests that
affect tomato production in semitropical areas of Florida has been docu¬
mented by Poe (1974a) and Wolfenbarger et al. (1975). Tomato pinworm
incidence was noted in Florida as early as 1932 (Watson and Thompson
1932) with serious outbreaks occurring during 1942, and from 1970 through
1973. Several factors have been mentioned by Poe et al. (1975) as caus¬
ing these outbreaks, i.e., type of insecticide used, change of tomato pro¬
duction practices, and harmful effect of pesticides on natural enemies.
Other factors such as weather have been overlooked. Current practices
for TPW control in Florida have been almost exclusively chemical (Waddill
1975), although emphasis has also been given to breeding tomatoes for
TPW-resistance (Schuster 1977a) and less to TPW biological control
(V.K. Waddill, personal communication). The effects of several factors,
e.g., rainfall and cultural practices, that influence the life system of
the TPW are still not understood.
To develop effective integrated pest management for tomato, the
interrelationships among the crop (plant physiology, phenology), pests
(arthropods, weeds, pathogens) and environment (climate, natural enemies,
horticultural practices) must be carefully studied. It is necessary to
understand TPW ecology and basic biology by studying the role of several
factors that cause seasonal and annual changes in pest populations. The
ability to assess the presence and abundance of the pest by accurate
sampling techniques would permit a reliable study of TPW potential
for inflicting economic damage. By evaluating the role of extrinsic
factors, e.g., weather, natural enemies and agronomic practices, it may be
possible to reduce the TPW problem.

4
This study was initiated to answer these and related questions.
The specific objectives of research were:
1) to describe different stages of development of the tomato plant.
2) to evaluate techniques for tomato pinworm damage assessment.
3) to discuss sampling techniques for tomato pinworm immature
stages under southern Florida conditions and to describe TPW
spatial distribution.
4) to evaluate the importance and population dynamics of TPW
natural enemies.
5) to evaluate the effect of hedges and edgerows on TPW dispersion
and field colonization.
6) to determine a way to assess yield losses in ground tomatoes due
to TPW.
7) to determine the influence of rainfall on TPW population.
8) to study post-harvest field managment practices that influence TPW
survival.
Therefore, the first chapter is a general literature review of
studies on K. lvcopersicella and addresses the effects of biotic and
abiotic factors on the population dynamics of this insect. The
second chapter is a study of tomato plant phenology and also covers
the evaluation of foliar damage assessment techniques. In chapters III
and IV, I address sampling techniques and dispersion patterns of tomato
pinworm eggs and larvae. The fifth chapter deals with the effect of
tomato pinworm infestation on upper and lower parts of the plant. In the
same chapter I state the relationship between TPW population index and
yield losses. In chapter six I address the distribution of male moths
and larval stages in tomato fields, and the effect of edgerows in such
distribution. In chapter VII, I deal with the abundance of egg and
larval natural enemies of the tomato pinworm.

5
The interaction of rainfall and TPW is presented in chapter VIII.
Finally, I evaluated the data regarding horticultural practices
and the relationship between changes of tomato agroecosystem and
oversummering populations of tomato pinworm (chapter IX).

CHAPTER I
LITERATURE REVIEW
Family Gelechiidae
The family Gelechiidae is one of the largest of the microlepidoptera
(about 580 North American species). Larvae vary in habits. Some are
leafminers, a few form leaf galls, many roll or tie leaves, and one
species, Sitotroga cerealella Olivier, is an important pest of stored
grains (Borror et al. 1976). Studies on crop pests in this family have
been concentrated on pests of high economic importance, such as the pink
bollworm (Pectinophora gossypiella Saunders), the potato tuberworm
(Phthorimaea operculella Zeller) , the angoumois grain moth (S_.
cerealella), and Keiferia lycopersicella Walsingham, the tomato pinworm.
The pink bollworm and the potato tuberworm are generally considered
good colonizers with highly mobile behavior within and between fields
(Stern 1979, Van Steenwick et al. 1978); however, many experts considered
these moths weak fliers which move great distances by being carried pas¬
sively by air currents (Kaae et al. 1977) . They are capable of having
several generations per year, with the last generation showing a strong
dispersal tendency (Kaae et al. 1977) . The potato tuberworm is perhaps
the most closely related to the tomato pinworm in patterns of behavior
and plant selection (Hofmaster 1949). Several authors (Shelton and
Wyman 1979, Meisner et al. 1974, Traynier 1975) have studied factors

influencing oviposition of potato tuberworm and the relationship between
populations of the pest and the host plant. Their studies were used as
a base in this research to compare with K. lycopersicella population
dynamics.
Studies on Keiferia lvcopersicella (Walsingham)
The tomato pinworm (TPW), K. lycopersicella (Wals), is frequently
confused with other species (Povolny 1977) , particularly with Scrobipal-
pula absoluta (Meyr.) and Phthorimaea operculella (Zell.) (Doreste and
Nieves 1963), since they are also considered pests of potato and tomato
(Povolny 1973) . K. lycopersicella and S_. absoluta are apparently iso¬
lated from each ether geographically and ecologically. K. lycopersicella
apparently avoids the cordillerian territory of the northern and southern
part of South America (Garcia et al. 1974, Mallea et al. 1972, Quiroz 1976).
The range of K. lycopersicella is in the eastern part of the American
continent and penetrates into Central America, Mexico (Povolny 1973) and
the Ü.S.A. (Elmore and Howland 1943). Phthorimaea operculella has been
reported on tomatoes in Venezuela, (Doreste and Nieves 1968), Bermuda
(Grooves 1974), and Egypt (Abdel-Salam et al. 1971).
In the U.S.A. K. lycopersicella is considered a key pest of tomatoes
in western California (Oatman 1970), Texas, Florida, Pennsylvania and
Hawaii (Swesey 1928, Thomas 1933) . The tomato pinworm was first recog¬
nized as a pest of tomatoes by Morrill (1925) , and was later reported by
Elmore (1937) and Thomas (1933) . In Florida, the TPW has been primarily
studied by Watson and Thompson (1932) , Swank (1937) , and recently by
Poe (1973). The seasonal history of the TPW was reported by Elmore

8
and Howland (1943) in California where it appears first during March
and April after overwintering in the pupal stage at or near the surface
of the soil. Later studies of the seasonal occurrence of TPW in Cali¬
fornia showed that larval populations increased abruptly in September
and October (Oatman et al. 1979) and in April-June (Oatman 1970) .
Batiste et al. (1970b) reported that there is no evidence for diapause in
this insect. Destruction of the tomato plants shortly after harvest may
prevent the insect from surviving the winter and infesting the crop
during the following season. Poe (1974a) reported that on the west coast
of Florida, severely infested fields occurred in the spring crop (February-
May) with less damage on plants during the autumn. Early infestations in
greenhouses also lead to heavy losses in the field.
Host Plants of Keiferia lycopersicella
Elmore and Howland (1943) reported that tomato and potato are pre¬
ferred hosts of TPW. Several solanaceous plants, e.g. eggplant iSolanum
melongena (L)] and nightshade (Solanum nigrum L.), also are known hosts
for the TPW (Batiste et al. 1970b, Elmore and Howland 1943, Swank 1937,
Thomas 1933). Batiste and Olson (1973) demonstrated that K. lycopersicella
preferred tomato for oviposition over 12 other solanaceous plant species.
TPW could be reared on Solanum melongena L., S_. dulcamara L., S_. nigrum, and
S_. elaegnifolium Cav. but not on S_. nodiflorum Jacq., S_. douglasi Dunal,
Datura me telo ides A. , D_. stramonium L., D_. ferox L., Nicotiana biglovii
(Torr.) and N. glauca Grah. The same author suggests that in California,
Solanum melongena, S. dulcamara and S_. elaegnifolium may play a role in
the population dynamics and distribution of TPW.

9
Life Cycle of Keiferia lycopersicella
Accounts of the life history and behavior of K. lycopersicella have
been reported by Elmore and Howland (1943) , Swank (1937) , and Poe (1973).
Poe (1973) found that eggs are laid singly or in groups of two or
three on the host plant foliage. Elmore and Howland (1943) described the
egg as ellipsoid, 0.37 by 0.23 mm, light yellow when first deposited, grad¬
ually darkening to a light orange before hatching. Eggs hatch 4-9 days
after deposition (Swank 1937) at 20.68°C and after 4-4.5 days at 27-29°C
(Elmore and Howland 1943) . Weinberg and Lange (1980) determined that
eggs hatch in a range of 3.5 +_ 0 days at 35°C and 7.8 +_ 0.2 days at
20°C.
Keiferia lycopersicella has four larval instars (Elmore and Howland
1943, Swank 1937). Head capsule width of the larval instars are 1st in¬
star 0.14-0.157 mm; 2nd instar 0.23-0.28 mm; 3rd instar 0.36-> 5 39 mm;
4th instar 0.52-0.61 mm (Elmore and Howland 1943). Newly hatched larvae
averaged 0.85 mm in length. The head capsule is dark brown and the re¬
mainder of the body is a yellowish gray common to many newly hatched
lepidopterous larvae. The mature larvae are 5.8-7.9 mm in length and
characterized by an ash gray color with dark purple spots (Elmore and
Howland 1943). Larvae of K. lycopersicella characteristically possess
a pale prothoracic shield with conspicuous dark fuscous shading along
lateral and posterior margins (Capps 1946) . Duration of the leaf mining
(lst-2nd instars) stage ranges between 4.7-5.8 days. The leaf folding
stage lasts between 5.6-16 days for a range of temperatures of 13-29°C
(Elmore and Howland 1943). Weinberg and Lange (1980) found that egg
hatching to pupation times range from 8 +_ 0.9 to 18 +_ 1.6 days when
reared at 35°C.

10
The pupae are initially green, later turning to a brown typical
of lepidopterous pupae commonly found in the soil (Elmore and Howland,
1943). Before pupation the larvae form a loose pupal cell of sand grains
at a depth of 0.25-1.5 inches beneath the soil surface (Poe 1973). Wein¬
berg and Lange (1980) recorded that pupation requires 11.3 _+ 0.5 at 20°C,
and 5.1 +_ 0.2 days at 35°C. The length of the pupal stage was 38.7 and
11.4 days at temperatures of 12.65 and 26.4°C (Elmore and Howland 1943).
Swank (1936) obtained a range of 7-17 days with an average of 11 days
for the pupal stage at 26°C.
Adults are characterized by an alar expanse of 9-12 mm. Labial
palpi have a short forrowed brush on the underside of the second joint,
a terminal joint somewhat thickened with scales, and are compressed
with the extreme tip pointed. The head and thorax are mottled with
dark brown. Forewings are elongate ovate, the hind wings have a
pointed apex, a strong pencil of hair scales, are dilated at tip of costa
in females, and dilated from base of costa in the males; the abdo¬
men is dark fuscous above with basal joints slightly ochreous, the
underside is light ochreous sprinkled with dark fuscous spots. Adult
longevity is 7 days (24 +_ 2°C) when they are fed on water and 8.5 days
at 24° + 2°C when fed a 10% honey solution. At temperatures of 10 and
13°C the respective longevities were 20.5 and 22.8 days (Elmore and
Howland 1943).
Insect Behavior
Elmore and Howland (19431 reported that copulation occurs within 24
to 48 hrs after moth emergence, and McLaughlin et al. (1979) stated that

11
sexual activity such as female calling was greatest during the 1st hr
of darkness. Very little copulation occurred after the 3rd hr. Males
ran or walked in their approach to calling females. Approach was gener-
o
ally from behind or at ca. 90 to the female and was accompanied by rapid
wing fanning. The copulatory strikes of the males were made laterally
beside the females. Moths remained in copula from 30 min to 2 hr.
Elmore and Howland (1943) and Poe (1973) described the behavior of
larval stages of K. lycopersicella. Newly eclosed larvae disperse briefly
from the hatched egg before entering the leaf. First instar larvae
spin a tent of silk over themselves and tunnel into the leaf. Further
feeding results in a blotch-like mine. The 3rd and 4th larval instars
feed from within tied leaves, folded portions of a leaf, or they
may enter stems or fruits. The 3rd instar appears to be the most
mobile and several types of behavior may occur (Poe 1973),. This stage
larvae can draw 2 leaves together, may tunnel into stems or fruits at
the calyx, but usually the larvae form leaf folds on the upper leaf
surface. The four instars can cause injury to 3-6 leaves during develop¬
ment (Poe 1973).
Elmore and Howland (1943) demonstrated that larvae that have mined
calyx lobes and nearby tissues enter the fruit instead of folding leaves.
Usually, the larvae enter the fruit beneath the calyx lobes or fruit
stems, but in heavily infested fields about 50% of the injured fruit may
be damaged in other places as well. The damaged areas caused by shallow
feeding just beneath the skin of the fruit appear as blotches. Larvae
that enter the fruit penetrate to a depth ranging from 0.9-1.9 cm.
Differences in the phenology of larval injuries were studied by
Batiste et al. (1970), who found that mines of the early stage larvae

12
superficially resembled the serpentine type mines produced by dipterous
leafminers of the genus Liriomyza. The mines could be distinguished
easily, because the dipterous leafminer leaves a trail of frass within
the mine, whereas the TPW larvae deposits nearly all the frass in a
single mass at the tunnel entrance.
Tomato Plant Resistance to TPW
Breeding for resistance work with tomatoes has largely been con¬
cerned with pathogens, but currently there is a renewed emphasis on
insect resistance as part of integrated pest management (Lange and Bronson
1981) . Resistance to many tomato insects does occur and includes resis¬
tance to the fruitworm, Heliothis zea (Cosenza and Green 1979); leaf-
miners, Liriomyza spp. (Schuster et al. 1979); tomato pinworm, K. lyco-
persicella (Schuster 1977a); hornworms, Manduca spp (Kennedy and Henderson
1978), Colorado potato beetle, Leptinotarsa decemlineata Say (Schalk and
Stoner 1976; potato aphid, Macrosiphum euphorbiae (Thomas); flea beetles;
white flies (Aleyrodidae); spider mites (Acariña) and many others (Lange
and Bronson 1981). The mode of resistance in tomato is complex and may
involve many factors including antibiosis, preference, phenological devel¬
opment (such as flowering time, time of fruiting, etc.), morphological
characteristics, presence or absence of foliage pigments, foliage vol¬
atiles, and physiological incompatibility.
Resistance to the tomato pinworm has been studied by Schuster (1977a),,
Schuster et al. (1979) , and Kennedy and Yamamoto (1979) . Schuster (1977a)
found that accessions of Ly copers icon sculentum Mill x Ij. pimpinelli
folium were more susceptible, while those of L. peruvianum (L) Mill, L.

13
peruvianum var. humifusum Mill., L. esculentum x L. peruvianum, L.
cheesmani f. minor (Hook F) Mull., and L. glandulosum Mull., were less
susceptible than the commercial cultivar 'Walter' (L. esculentum Mill.).
Selections of L. hirsutum Humb and L. hirsutum f. glabratum Mull, were
more resistant and had 25-50% and 50-75% less damage respectively
than 'Walter1. In laboratory studies the same author found that mine
lengths after 2 days were significantly shorterâ–  for PI numbers 129157
(L. hirsutum f glabratum) and 298933 (L. peruvianum). Schuster et al.
(1979) stated that levels of resistance to tomato pinworm and vegetable
leafminer appeared to be intermediate and the varieties PI 12930 and PI
1404403 of L. esculentum were found moderately resistant to both insects.
Kennedy and Yamamoto (1979) found an extractable toxic factor in the
foliage of L. hirsutum f. glabratum affecting Manduca sexta, H_. zea, K.
lycopersicella, Aphis craccivora, A. gossypii, and Myzus persicae. Schuster
(1977b) reported that tomato varieties 'Pennorange E 160 A' and 'Pearson'
had less fruit damage by K. lycopersicella and armyworms, primarily
Spodoptera eridania (Cramer), than did the 'Walter' variety.
Chemical Control of TPW
Chemicals are widely used to control tomato pests. The need for
insecticides varies from year to year and from one area to another (Lange
and Bronson 1981). Chemical control of TPW was obtained in 1943 by Elmore
and Howland (1943) who recommended synthetic cryolite and talc dust
(50% sodium fluoaminate). In California, several insecticides were
evaluated by Middlekauff et al. (1963) and reevaluated by Batiste et al.
(1970a). The latter authors reported little or no control of larvae by

14
insecticides applied as soil treatments under greenhouse conditions.
These same authors stated that methyl parathion was the most effective
material in greenhouses, and also recommended parathion, methidathion,
phosphamidon, mexacarbamate and methamidophos. Spray deposits of para¬
thion were found by the same authors to be significantly less effective
against eggs or early stage larva than was toxaphene-DDT.
Poe and Everett (1974) presented results of experiments to control
TPW in 2 locations in Florida. They reported that granular insecticides
in general did not perform as well as most spray materials for reduction
of the TPW mines and larvae in tomato transplants. They recommended
acephate, diazinon, endosulfan, and methomyl to keep seedlings nearly
mine free. Chlordimeform was considered phytotoxic to seedlings but when
sprayed alone or combined with Bacillus thuringiensis Berliner on older
plants gave good control of TPW larvae without plant toxicity. Poe and
Everett (1974) recommended highly residual insecticides to maintain a crop
free of damaged fruit.
Waddill (1980) reported that certain insecticides used on demand
for tomato pinworm in Homestead, Florida, significantly reduced TPW
damage below that in the untreated check. Permethrin +
Bacillus thuringiensis were applied least often and were among the best
treatments. The author also showed that when used on demand a low rate
(0.225 lbs ai) of methomyl resulted in significantly more damage than
the same rate plus 0.5 lbs Bacillus thuringiensis.
Schuster (1977b) reported that when measured by the number of
damaged fruit, the degree of control of the TPW and southern armyworm
with Bacillus thuringiensis WP and chlordimeform was significantly depen¬
dent on the tomato cultivar. The contact toxicity of 4 synthetic

15
pyrethroids and methomyl to some adult parasites of tomato pests indi¬
cated that fenvalerate was generally the least toxic to parasites com¬
pared to permethrin, burethrin, and NRD1C49 (+) -d-cyano-m phenoxybenzyl
( + ) cis, trans-3-(2,2 dichlorovinyl)-2-dimethyl-cyclo-propanecarboxylate)
as well as methomyl (Waddill 1980) . Fenvalerate was judged the most
promising candidate for use in a pest management program in tomatoes for
integrated control of the TPW and the vegetable leafminer. Lindquist
(1975) obtained the best control of TPW with synergized pyrethrins (MGK
pyrethrins) and endosulfan.
Emergence of K. lycopersicella and Apanteles spp from pupae and
soil treated with insect growth regulators (IGR's) resulted in 23%
suppression of pinworm adult emergence when applied directly to the TPW
pupae but was ineffective when applied to the soil. The IGR's caused a
reduced emergence of the parasite Apanteles spp from 61% to 0% (Poe
1974b). Prada and Gutierrez (1974) reported some results on microbial
insecticide control of Scrobipalpula absoluta, the South American pinworm.
Seventy five to eighty percent control of the pest was obtained within
5-100 days after treatment at the rate of 500-200 Neoplectana carpo-
capsae Weiser nematodes per plant or with Bacillus thuringinesis (150-
500 g/ha). Schuster (1982) demonstrated that a mixture of B.
thuringiensis and Coax® (454 g + 1.8 kg product/378 Its) when applied to
TPW infested tomato seedlings, increased TPW mortality up to 42.2%.
Cultural Practices for TPW Control
According to Lange and Bronson (1981) , the mechanization of pro¬
duction of processing tomatoes has not only revolutionized the industry
but has altered many control techniques and as a result, a few formerly

16
major pests have been reduced to a secondary position. Elmore and How¬
land (1943) considered some cultural practices as undesirable because of
their adverse impact on TPW control. These include failure to destroy aban¬
doned plantings, careless disposal of infested culled fruit, and use of
infested seedlings. In Florida, Swank (1937) recommended that all mate¬
rial remaining in the field after the crop is harvested be carefully
plowed under. He suggested that the carelessly abandoned fields could
become a reservoir for infestation of a nearby or succeeding crop. Poe
(1973) stated that the best control for TPW is based on several cultural
practices: use of non-infested seedlings, destruction of plant debris,
use of light traps for adults in small areas, and destruction of plants
growing from seeds in compost heaps. Price and Poe (1977) reported that
staking and artificial mulching of tomato plants reduced damage caused by
K. lycopersicella and other pests.
Biological Control of TPW
Employment of biological control measures for insect and mite pests
of row crops has been limited, and the poor record probably relates
largely to the short-lived row crop environment, which presumably does
not permit establishment of the effective host-natural enemy relation¬
ships which often characterize more stable environments (van den Bosch
et al. 1976). Modern-day biological control techniques have not been
fully exploited in tomato under field conditions (Lange and Bronson
1981). They have been widely accepted in European glasshouse tomato pro¬
duction, however. Reports on parasitism of K. lycopersicella were
made by Elmore and Howland (1943) , Swesey (1928) , Thomas (193 3) , Oatman
et al. (1979) and Poe (1973) (Table 1) .

Table 1. Larval parasites of Keiferia lycopersicella reported from U.S.A. and South America until 1981.
Scientific Name
Family
Place
Reference
Angitia blackburni Cam
Ichneumonidae
Hawaii
Swesey, 1928
Angitia ferrugipeneipes (Ashm.)
Ichneumonidae
Pennsylvania,
California
Thomas, 1933
Apanteles epinotiae Vier.
Braconidae
California
Thomas, 1933
Apanteles scutellaris Mues.
Braconidae
California
Elmore and Howland, 1943
Apanteles dignus Mues.
Braconidae
California,
Florida
Elmore and Howland (1943),
Krombreii et al. (1979)
Apanteles gelechidivorus sp. n.
Braconidae
Colombia,a
California
March, 1975
Catolaccus aeneoviridis (Gir)
Pteromalidae
California
Elmore and Howland, 1943
Chelonus phthorimae Gahan
Braconidae
California
Elmore and Howland, 1943
h-*
Campoplex phthorimaeae (Cus)
Ichneumonidae
California
Elmore and Howland, 1943
'-l
Chrysocharis sp Foerster
Eulophidae
California
Elmore and Howland, 1943
Horismenus spp
Eulophidae
Florida
Krombreii et al. 1979
Hormius pallidipes Ashm.
Braconidae
California
Elmore and Howland, 1943
Microbracon junicola (Ashm.)
Braconidae
Pennsylvania,
California
Thomas, 1933
Parahormius pallidipes (Ashm.)
Braconidae
California
Oatman et al., 1979
Sympiesis stigmatipennis Girault
Spilochalcis hirtifemora Ashmead^*
Eulophidae
California,
Florida
Cuba
Elmore and Howland, 1943,
Krombreii et al., 1979
Castineira and Hernandez, 1980
Tetrastichus sp Holiday
Eulophidae
California
Elmore and Howland, 1943
Zagrammosoma multilineatum
(Ashmead)
Eulophidae
Florida
Waddill, 1980
Zatropis sp Crawford
Pteromalidae
California
Elmore and Howland, 1943
Parasite of the South American pinworm £>. absoluta.
Hyperparasite of A. dignus.

18
Cardona and Oatman (1971) studied the biology of the larval para-
sitoid Apanteles dignus Muesebeck which is a solitary, primary, larval
endonarasite of K. lycopersicella and found that the total developmental
time from egg to adult was ca.18 days at 26.6 + 1°C and 50 _+ 2% RH. Oat¬
man (1970) stated also that the most common parasites at both Indio and
Escondido (California) during 1963-64 were A, scutellaris (Mues.) and
Parahormius pallidipes (Ashm.) followed by Sympiesis stigmatipennis
Girault. The biology of A. scutellaris (Mues.) was studied by Djamin
(1970). In California, A. dignus apparently occurs only along the coast
in the southern part of the state. Studies conducted by Oatman et al.
(1979) determined that in the south coast there was a range of larval
parasitization of 1.6-36.8% during 1972-73. Apanteles dignus was the most
abundant parasite reared from larvae followed by _S. stigma tipennis Girault.
In Florida, Poe (1974b) reported that 50-70% of tomato pinworm larvae in
leaflets collected in the spring, 1973 were parasitized by Apanteles spp.
Behavioral Chemicals Used in Monitoring and Control of TPW
Pheromones. Sex pheromones of the adult TPW were obtained by
Antonio (1977) from extracts of the whole body of 2-day-old virgin fe¬
males. A biological assay method was then devised to test males for
optimal response to the pheromone under varying conditions. Field evalu¬
ation data by the same author indicate that the natural sex pheromones
were attractive to male tomato pinworm moths. McLaughlin et al. (1979)
found that males were more responsive when bioassayed with dim light
from both above and below an olfactometer than when illuminated only
from below. The effect of trap design and sex attractant release rates

19
on TPW catches was studied by Wyman (1979). It was determined that Zoe-
con lc® sticky traps were 6 times as effective in capturing TPW males as
were Delta® sticky or mineral oil traps. An inverse relationship between
attractant release rate (fibres/dispenser) and trapping efficiency was
found. K. lycopersicella positively responded to a sex attractant of
unstated composition dispensed from rubber septa in traps in a tomato
field (Wyman 1979).
Deterrents. Beck and Schoonhoven (1980) stated that surface testing
of insects touching or piercing with the ovipositor or by biting and
probing with the mouth parts is in response to chemical factors that act
as incitants. If the stimuli received on initial testing indicate an
unacceptable plant, the behavior pattern is interrupted and the insect
abandons the plant. Such stimulants are deterrents.
Schuster (1980) reported that survival of TPW was reduced when larvae
fed on excised tomato leaflets dipped in solutions of cyhexatin, fentin
hydroxyde (triphenytin hydroxide) and guaratine (SN-513); N-n1''-(iminodi-
8,1-octanedilyl) bisguanidine. These compounds protected foliage and
fruit from insect damage when plants were sprayed in the field.
Tomato Plant Phenology and Measurement of TPW Dispersion and
Economic Damage
Little information is available that relates tomato plant phenology
to pest management or that concerns dispersion and economic damage of
the TPW. Plant phenology related to pest management tactics has been
determined already for different crops: alfalfa, cotton, potato, tobacco,
soybeans, etc. (Anonymous 1971, Reynolds et al. 1975, Ambrust and
Gyrisco 1975, Johnson 1979, Fehr et al. 1971) . Tomato plant phenology

20
as related to pest management has been reported by Alvarez-Rodriguez
(1977) and Keularts (1980). Alvarez-Rodriguez (1977) evaluated pest
(pathogens and insects) damage as it is related to tomato life table
analysis and determined strategies for tomato production. Keularts
(1980) determined the effect of artificial defoliation in plants 30-100
days old. These studies should have been complemented by a phenological
description of the plant at different plant stages.
Studies of measurement and description of dispersion of TPW popu¬
lations must involve a sampling program in which biological, statistical
and economical aspects of the program are evaluated (Southwood 1978).
This information is expected to result in 1) biological interpretation
of statistical parameters and 2) the use of this knowledge for measuring
of TPW control and for establishing a reliable scouting program.
Sampling designs for tomato pinworm larval stages have been studied by
Wolfenbarger et al. (1975) and Wellik et al. (1979). Wolfenbarger et
al. (1975) developed a sequential sampling program based on the detection
of larval feeding on the 3 top leaves per plant. Alternatively Wellik
et al. (1979) found that lower leaf and large fruir sampling methods
were best for detecting the presence of TPW. These opposing results
demand more detailed research in order to obtain more accurate TPW
density estimates.
The data concerning tomato pinworm damage range from estimations
of damage based on pesticide effectiveness (Batiste et al. 1970a, Poe
and Everett 1974, Waddill 1975, 1980), damage evaluation based on
effectiveness of parasitism of TPW larvae (Oatman 1970) to estimation

21
of economic injury levels (Wolfenbarger et al. 1975). Poe and Everett
(1974) determined the percentage of unmarketable fruit as 6.5 to 4%
when the plant was untreated. Waddill (1980) reported that plants
without chemical control may lose up to 75% of the fruit. Oatman (1970)
determined tomato fruit was infested up to 70% despite 68.9% larval
parasitization. Wolfenbarger et al. (1975) reported that an average of
0.3 TPW injuries per 3 top leaves caused 20% injured fruit.
Environmental Factors Affecting TPW Population
Characteristics of Agroecosystems
Agroecosystems vary widely in stability, continuity, complexity
and area. The kind of crops, agronomic practices, changes in land use
and weather are important elements affecting the degree of stability of
an agroecosystem (Stern et al. 1976). Since agroecosystems are
characterized by a short life (Loucks 19701, they are more susceptible
to pest damage and catastrophic outbreaks. This also occurs because of
a lack of diversity in plant species, insect species, and sudden alter¬
ations imposed by man such as plowing, mowing and use of insecticides
(Luckman and Metcalf 1975, Pimentel 1961a, b, Smith 1970, van Emdem and
Williams 1974).

22
Tomato Agroecosystem
The tomato crop is a typical example of an agroecosystem with early
community succession (Price and Waldbauer 1975) . In southern Florida 3
closely related tomato varieties are generally grown: MHl, 'Walter'
and 'Flora-Dade' (Volin and Bryan 1976). Horticultural practices are
characterized by direct-seeding in the field through mulched beds
that will aid in maintaining a regular amount of soil moisture, weed
control, and fertilization of the crop (Geraldson 1962, Davis et al.
1970, Bryan et al. 1967). In summary, the tomato crop is typical
of agricultural systems with high community energetics, small or low
community structure, rapid nutrient cycling, selection pressure
(r - selected, many small progeny), and quantitative progeny production.
Also, the tomato agroecosystem is characterized by having a few major
key pests and secondary pests (Lange and Bronson 1981) . Most of these
pests, e.g., lepidopterous larvae, stinkbugs, dipterous leafminers,
whiteflies, leafhoppers, aphids and some species of beetles, are con¬
sidered as r selected species with rapid development, high maximal rate
of increase (rm) , early reproduction, small body size, many small off¬
spring and short length of life (Krebs 1978, Pianka 1978) .
Biotic and Abiotic Factors Affecting Insect Population Dynamics
Biotic and abiotic factors exercise some influence on the fluctu¬
ation in the number of insects in time and space. To reveal both char¬
acteristics the inherent property of animals themselves and environmen¬
tal conditions in their habitats must be studied (Shiyomi 1976) . Among
the biotic factors, we should consider the habitat effect on insect

23
distribution. Effects of habitat have been studied by several research¬
ers: Gossard and Jones 1977, Lyons 1964, Brazzell and Martin 1957, Yama¬
moto et al. 1969, Wolfson 1980, Sparks and Cheatman 1970, Dethier 1959a,
Nishijima 1960. They demonstrated the effect of habitat on oviposition
and adult and larval dispersal. The effect of sheltered zones on
distribution of insects has been demonstrated by Lewis (1979) van Emdem
(1965) and Price (1976). They indicated the importance of crop edge
effect on colonization and dispersal of arthropods, especially for r
selected species, which show a "safety in numbers" strategy for progeny
reproduction and survival. Van Emdem (1965) considered that unculti¬
vated land in regard to the insect fauna of a crop has 2 components:
1) Physical: shelter-survival in debris of woodland, 2) Biological:
plants of uncultivated land provide alternate food and breeding sites for
injurious insects, crop diseases or alternate hosts for predatory and
parasitic insects.
In most agricultural environments the principal pests are usually
controlled to a greater or lesser extent by natural enemies (Messenger
et al. 1976). The efficiency with which such natural enemies suppress
pest populations is influenced on the one hand by their own intrinsic pro-
erties and limitations and, on the other hand, by environmental factors
and conditions occurring in the agroecosystem under consideration
(Messenger et al. 1976).
Among the abiotic factors affecting insect populations, weather
and climate are commonly accepted by entomologists as dominant influ¬
ences on the behavior, abundance and distribution of insects (Messenger

24
1959). Effects of climate on insect populations were studied by
Richards 1961, Nicholson 1958, Cloudsley-Thompson 1962, Andrewartha and
Birch 1974. Most authors agree that 2 of these factors, temperature and
RH possess a high degree of interaction and affect insect activity and
survival. As an example Chapman et al. (1960) and Hofmaster (1949)
have looked upon the effect of climate on survival of Gelechiidae.
Finally, pest control in an agroecosystem can be aided by proper use
of cultural practices. Two basic principles in the cultural control of
arthropod pests are 1) manipulation of the environment to make it less
favorable to the pest and 2) manipulation to make it more favorable for
their natural enemies (Stern et al. 1976). Cultural methods, however,
require a thorough knowledge of crop production and the biology and
ecology of the pest and its natural enemies in order to integrate the
techniques for pest control into proven agronomic procedures for crop
production.

CHAPTER II
DESCRIPTION OF TOMATO PLANT PHENOLOGY AND EVALUATION OF
TOMATO PINWORM FOLIAR DAMAGE ASSESSMENT
Introduction
Description of tomato plant phenology and evaluation of tomato pin-
worm larval presence are major aspects of tomato pest management that need
to be determined. First, studies on tomato taxonomy, growth and develop¬
ment, effects of fruit on vegetative growth, and relationship between
fresh weight and leaf area are well documented (Cooper 1964, Murneek
1924, Hurd et al. 1979, Romshe 1942, Thompson and Kelly 1957, Purseglove
1968). However, tomato crop phenology that divides the growing plant
into characteristic periods and shows the relative time in each period
needs to be studied. Second, description of TPW damage to the foliage
and TPW mine length correlation with plant resistance has been studied
by Elmore and Howland (1943), Batiste et al. (1970) and Schuster (1977).
Nevertheless, the evaluation of different techniques for TPW foliar
injury assessment is necessary' to establish a relationship between larval
instars and amount of damage.
The objectives of this study were first, to define growth character¬
istics of tomato plant during a typical southern Florida growing season,
second, to describe from a pest management point of view the phenology
of tomato, cv. Flora-Dade, and third, to determine constraints and prac¬
tical use of TPW larval population indices.
25

26
Materials and Methods
The Tomato Crop
Tomatoes, cv Flora-Dade, were planted in 1979 (Nov. 3, Dec. 5),
1980 (Jan. 8, Oct. 30, Nov. 25, Dec. 30), and in 1981 (Jan. 30, Feb. 28)
at the University of Florida, Agricultural Research and Education Center,
in Homestead, Dade County, Florida. After metribuzin was incorporated
into the soil at a rate of 0.84 kg ai/ha, beds 45 m long were prepared
and fertilized with 7-14-14, at a rata of 2242 kg/ha. Immediately after
fumigation, drip tubing for irrigation was placed ca. 15 cm into the soil
and the beds were covered with plastic mulch. Tomato seeds were planted
with a seed drill 30 cm apart in the rows. One to two weeks after emer¬
gence, the seedlings were thinned to one per hill. Plants were protected
from pests by application of fenvalerate 2.4 EC (.045 kg ai/ha), maneb and
tribasic copper sulfate (0.97 + 5.71 ai kg/ha) at weekly intervals.
Two to five plants were selected at random from each of the 8
plantings, and height, leaf area, number of leaves, suckers, flowers and
fruits were recorded every 8-15 days.
Description of Stages of Tomato Plant
The method used to describe tomato plant phenology was based on
the technique of Fehr et al. (1971) for soybeans, Glycine max (L).
Three developmental stages of tomatoes were defined: vegetative,
reproductive, and senescent. Number of leaves, plant height,

27
time of blooming and fruit formation were averaged from the 8 plantings.
Development of the plant was quantified by a nomenclature system, where
primary leaves were numbered from the bottom to the top; any secondary
growth, e.g., formation of primary laterals (Fig. 1), had the same node
number from which it originated, and it was distinguished with a let-
ter(s). The stem that originates from the bifurcation of the main stem
was called a secondary main stem (2M); laterals that develop from primary
laterals were considered secondary (2S).
Methods of Damage Assessment for TPW Larvae
A set of 3 tomato plants 40-50 days old grown in 20 cm pots was
introduced every 2 days into a cage (45 x 45 x 60 cm) for oviposition
by moths previously held at 24 + 3°C and RH 75 _+ 2%. Plants were subse¬
quently removed, and set aside for larval development. When each set of
plants was under attack by 1-4 larval instars, a total of 100 damaged
leaves was taken to the laboratory for inspection. A total of 20
individuals was studied per instar. Three methods of measurements were
used in separate experiments. First, a portion of leaf area mined by
the TPW larvae was separated from the leaf and then measured on a
LICOR® model LI3100 area meter. For another set of damaged leaves,
leaf weight ingested by the larvae was determined by measuring the dif¬
ferences in weight between the damaged leaflet and the juxtaposed leaf¬
let. Larval instar in both experiments was determined by measurements
of the larval head capsule width. Second, a visual classification of dam¬
age was made of leaf damage caused by TPW larval instars. A five class

Figure 1.
vegetative
Illustration of tomato cv Flora-Dade
stage; TR =early reproductive stage;
growth at 2 stages of development. TV^-second
a=primary leaf; b=lateral development.

to
Tomato Stages of Development

30
scale of 0-4.5 (Table 2) was devised, based on personal observations and
the damage descriptions of Batiste et al. (1970a).
The leaf injury length (cm) was also measured and larval head capsule
width recorded. Finally, the presence or absence of TPW larvae in different
types of leaf injury was determined. A simple linear regression model was
used to examine the relationship between the head capsule width and injury
length, and between larval instar and the damage rating scale. The eval¬
uation of the different methods of damage assessment was discussed with
regard to the practicality of their use for scouting programs.
Results and Discussion
The Tomato Crop
A summary of leaf area and number of flowers and fruits is shown in
Table 3. Tomatoes planted during October, 1980 began to flower 61 days
after plant emergence and fruit set occurred at 73 days. Maximum leaf
area was reached at 134 days. Tomatoes planted during November, 1980
started blooming at 54 days, and fruit set occurred at 68 days. Maximum
leaf area occurred 88-112 days after plant emergence. Tomatoes planted
during December, 1980 and January-February, 1981 had a shorter vegetative
period, with flowering at 42-62 days and fruiting at 49-62 days. Leaf
area reached a maximum at 63-89 days. The total leaf area during these
plantings was lower than that produced from fall plantings.
Under southern Florida conditions, average temperature changes
drastically from autumn to early spring (Mitchell and Ensign 1928) . In
this area, the effect of planting date determines growth and tomato

31
Table 2. Classification of tomato pinworm leaf damage on 'Flora-Dade'
tomatoes, based on greenhouse and field observations. Home-
stead, Florida,
1980.
Degree of Damage
Description
0
No damage
1-1.5
Mining of leaves, ca. 0.50 cm or less in
length; mine narrow and elongate; tissue
transparent; mine on any part of the leaf¬
let; some leaves attacked by more than 1
larva; small larvae present.
2-2.5
Mining of leaves ca. 0.51-0.68 cm; 1/4 of the
mine is necrosed, but changing to a raised
area or oblong to ovoid blotch; frass accumu¬
lation at the bottom of the injury.
3-3.5
Blotching of leaves; blotch necrosed over
60% of the injury; no holes indicating lar¬
val exit; size 1-2 cm; epidermis of the leaf
opaque to chlorotic due to larval injury to
midvein; construction of silk tent in epi¬
dermis .
4-4.5
Leaf folded; fold can occur at any lobe of
the leaflet. Necrosis extended to 75-30%
of the leaf; extensive frass accumulation
on blotch or fold; injury length 2-4 cm.

Table 3. Leaf area and reproductive
in Homestead, Dade County,
plant structures in
Florida during 1980-
tomatoes, cv
â– 1981.
Flora-Dade, planted
on 5 dates
Leaf Areaa
No. Flowers
No. Fruits
Height
Age:Days
Planting Date After Germination
x + SE
x + SE
x + SE
(cm)
Oct. 30, 1980 21
1.60+0.32
0
0
15
35
9.20+1.34
0
0
22
61
31.20+2.74
7.8+0.46
0
17
67
22.00+1.06
5.8+1.00
0
49.45
73
9.80+0.95
14.8+1.77
3.75+0.83
50.00
79
11.60+1.36
18.6+1.98
7.4 +0.86
b
86
23.40+2.67
13.5+1.73
7.4 +1.00
99
27.4 +2.54
19.4+2.10
11.00+1.50
109
20.00+2.40
15.2+1.79
17.20+1.95
116
10.6 +0.54
8.2+1.22
20.2 +2.08
123
23.80+2.72
2.8+0.93
25.80+2.36
134
37.20+2.90
0
23.2 +2.32
151
17.80+1.82
0
11.60+1.46
159
2.80+0.87
0
0
175
3.60+0.32
0
0

Tab Le 3--continued
1980
32
3.4
+ 0.64
0
0
16
39
5.6
+ 0.51
0
0
14.5
45
4.2
+ 0.87
0
0
20.00
54
8.2
+ 1.43
1.40+0.51
0
20.00
62
15.0
+ 1.85
8.00+0.69
0
41.00
68
20.2
+ 2.13
7.4 +1.23
0.8+0.46
57.6
74
26.0
+ 2.72
9.6 +1.37
5.8+0.85
67.2
81
30.0
+ 2.62
9.0 +1.55
6.4+1.29
52.0
88
41.0
+ 3.08
9.6 +1.44
11.4+1.66
64.0
94
24.8
+ 2.43
7.2 +1.11
6.2+0.97
b
106
29.4
+ 2.54
0
10.0+1.51
112
32.8
+ 2.82
0
15.0+1.93
132
2.2
+ 0.57
0
0
139
2.0
+ 0.10
0
0
151
2.4
+ 0.54
0
0
156
13.4
+13.10
0
0

Table 3—continued
Dec. 30, 1980
1.80+0.41
2.80+0.51
2.40+0.57
6.00+0.96
6.8C+0.97
16.60+1.91
8.00+1.28
12.3 +1.73
7.4 +1.00
9.2 +1.31
4.6 +1.03
9.8 +1.53
3.4 +0.32
8.4 +0.32
18
24
34
42
49
59
66
80
87
95
107
112
119
125
i
0
0
7
0
0
14.6
0
0
21.6
0.40+0.42
0
15.0
8.40+0.32
1.0+0.02
21.0
9.6 +1.49
7.4+1.01
37.0
6.2 +0.95
5.2+0.46
48.0
1.2 +0.57
0.6+0.33
36.0
0
7.0+1.00
b
0
0
0
0
0
0
0
0
0
0

Table 3—continued.
Jan. 30, 1981
Feb. 28, 1981
1.3 +0.54
6.6+1.23
6.6+0.91
14.6 +3.15
12.6 +1.57
23.4 +2.28
8.2 +1.04
5.2 +1.15
6.4 +0.66
7.00+0.73
8.20+1.26
1.40+0.32
9.40+0.32
22.00+2.26
25.00+2.51
27
34
62
75
81
89
100
106
113
119
25
34
48
55
63
0
0
9.0
0
0
4.0
5.20+0.17
1.20+0.57
15.0
7.8 +0.57
6.0 +0.73
39.0
2.0 +0.0
3.6 +0.81
15.0
7.8 +0.46
0
37.0
0
0
b
0
0
0
0
0
0
0
0
13.00
0
0
13.00
3.00+0.00
0
29.00
4.20+0.94
3.80+1.11
43b
5.20+1.03
5.60+1.08

Table ^--continued
73
25.00+2.29
8.20+1.13
8.60+1.26
79
13.00+1.72
4.20+0.87
5.80+1.15
84
7.40+0.89
0
0
90
5.60+1.03
0
0
aLeaf area measured in square decimeters,
b
Plant becomes póstrate at this time, changing height patterns.

37
development. Tomato, cv Flora-Dade, was developed for production of
fresh market tomato fruit (Volin and Bryan 1976) during the months of
January-Marc'n. Consequently, planting before or after the autumn
months of October-December resulted in a high reduction of leaf area
and yield.
Developmental Stages of Tomato
Vegetative growth of the tomato plant passed through 3 distinct
phases (Fig. 2). In the first phase there was a steady increase in leaf
area, while in the second phase leaf area was relatively constant. The
third phase was characterized by reduction in rate of leaf expansion 130
days after plant emergence. The number of inflorescences rose rapidly to
a peak at 70 days and then steadily decreased, whereas fruit reached a
peak at 90 days post-planting and then steadily decreased. Flowering and
fruit formation were observed at 40 and 50 days, respectively. Three
major developmental stages were determined for tomatoes: vegetative,
reproductive and senescent (Fig. 3). Each stage was divided into sub¬
stages. Each substage is explained in detail in Table 4.
The characteristics of tomato plant growth (Fig. 3) demonstrate the
relation between leaf area and crop age (days after emergence). The in¬
crease in leaf area was observed until half of the second reproductive
stage (TR^) • Leaf area is reduced during the third reproductive stage
(TR^) and senescent stage (S^). I consider that the TR^ stage can be
subdivided into another stage. This will allow a more detailed descrip¬
tion of plant stages, as well as shorter time periods for better assess¬
ment of pest management.

Figure 2. Influence of time on leaf area (dm*”) expansion, flowering and tomato fruit
numbers of cv Flora-Dade grown on 'Rockdale' soil under southern Florida conditions.

<1
<
u-
<
80-
6 0_
4 0-
20 _
u>
kO
PLANT AGE (Days)
No. FLOWERS

Figure 3. Stages of development of tomato. TV^early vegetative stage; TV =late vegetative
stage; TR^, TR2, TR3=reproductive stages; S^=senescent stages.

Leaf
Crop Age( Days after Emergence)

42
Table 4. Stage of development description for tomato cv Flora-Dade.
Description is based on the average of observations from
tomato plants grown during Fall 1980 through Winter 1981.
Homestead, Florida.
Plant Stage
Tomato Plant Description
Vegetative
TV
1
TV
2
Reproductive
TR
1
TR
2
TR
3
Plants 1-15 days old. Complete formation of
2-3 primary leaves; loss of cotyledons; plant
height ca. 5-7 cm.
Plants 16-35 days old; plant erect (12-16 cm);
5-7 leaves, development of laterals; plant
with only 1 main stem.
Plant 35-40 days old; development of laterals
from nodes 1-5; at leaf 4-5 the stem bifurcates
producing another stem as vigorous as the first
main stem; production of floral clusters at
node 5 and second main stem; height 50 cm.
Plants 67-70 days old; fruit set; plant postrated;
yellowing of primary leaves.
Plant 109-135 days old; 90% fruit ripe; post¬
harvest maturity; at least 60% of the primary
leaves necrosed, development of secondary laterals
at nodes 3-5; plant totally póstrate; height ca.
32-57 cm.

43
Table 4—continued.
Senescence
stem and second main stem; regrowth of plant
from auxiliary buds at nodes 1,2 and produc¬
tion of up to 3 floral clusters may occur;
possible fruit development.

44
The principal application of this nomenclature system is to deter¬
mine the amount of yield reduction produced by damage inflicted at given
stages of plant development. As an example, if I use Keularts1(1980)
data from his experiment in tomato defoliation, 20% defoliation of lower
plant leaves at stages TV^ through TR2 did not alter mean yield
per plant. However, 20% defoliation of upper plant leaves at TR^ stage
caused yield reduction. The nomenclature system can apply to single plants
or entire crops. It would be worthwhile to apply this system to other
tomato cultivars.
Methods of Damage Assessment for TPW Larvae
Average leaf area and weight consumed by TPW larvae. The data from
this experiment demonstrated the complexity of measuring TPW foliar dam-
2
age. The average leaf weight (mg) and leaf area (cm ) consumed by larvae
of a determined instar are shown in Table 5. Average leaf area consumed
2
ranged from 0.5 to 1.57 cm for 1st to 4th instar. First and fourth
instar larvae consumed 5 and 13.42 mg of leaf, respectively. Variance of
leaf weight measurements was large suggesting that many uncontrolled
factors influence feeding of individual larvae in the field. Either
method might be used for laboratory and greenhouse experiments where the
researcher would have more control of the factors influencing variability
(e.g., leaflet size, leaf age).
Length of Foliar Injury and Use of Damage Scale
Length of foliar injury and TPW head capsule width were cor¬
related (r=0.63; F=0.001, F=39.33) (Fig. 4). Furthermore, there was

Table 5. Tomato leaf weight and leaf area consumed by different larval instars of
Keiferia lycopersice11a under greenhouse conditions; T 24+3°C; 75+2% RH.
Head Capsule
Width9
0.14 -
0.16
0.23 - 0.30
0.36 - 0.40
0.525 - 0.61
x +
SE
x + SE
x + SE
x + SE
Leaf Weight
Consumed (mg)
5.0 +
2.86 ,
6.42 +
1.31
12.7 + 2.99
13.42 +
6.75
(11.46,-
1.46)b
(10.04,
2.8 )
(21.05, 4.42)
(20.17,
6.67)
Leaf Area
Consumed (cm )
0.5 +
0.16
1.32 +
0.23
0.85 + 0.11
1.57 +
1.22
(0.96,
0.14)
(1.82,
0.72)
(0.96, 0.74)
(4.54,
0.35)
a
Head capsule in
mm; each
width
range corresponds to
1-4 instar.
b
Confidence limits expressed at 0.05% significance level.
Ln

Figure 4.
width (mm)
Linear relationship between tomato pinworm (Keiferia lycopersicella) larval head capsule
and foliar injury length, r =0.47.

Head Capsule TPW Larvae (mm)

48
a significant relationship between TPW (Fig. 5) larval instar and the
degree of damage observed (r=0.79). Both techniques suggest the possi¬
bility of prediction of damage level in the tomato plant at stages
TV-^ - TR^. Such a prediction may be influenced by other factors such
as plant stages and larval density.
The use of larval instars to determine injury length has a reduced
bias compared to use of TPW damage degree scale. Foliar injury measure¬
ment is only advisable for research experiments (e.g., plant resistance,
pesticide screening) in which the time frame available to determine the
dependent variable is not a constraint. Other aspects to be considered
for further study are larval preference for larger or smaller leaflets,
as well as presence of different larvae in the same leaflet.
The use of TPW damage scale is perhaps less precise than the tech¬
nique mentioned above. Damage scale technique may introduce personal
error in measurement of larval instar in relation to degree of damage.
It is possible, however, to use this technique as an adjunct aid to the
population index (number of injuries per plant).
As an example, using the equation y=0.80 + 0.795x, where y = the
leaf injury damage scale and x = the tomato pinworm larval instar; if
the value of x equals 3, the average degree of damage in the plant will
be 3.18. This information will help to determine the effect of the
insect in economic terms, once the economic threshold is reached for
plant stages TV^ - TR^. At this point there is no information available
for TPW EIL values for plants in these early stages. Therefore, further
studies will be necessary to indicate that the presence of a particular
larval instar is capable of producing a determined economic damage.

Figure 5. Linear relationship between tomato pinworm (Keiferia lycopersicella) larval instars
O 1
and visual leaf damage scale, r =0.677.

5
Tomato Pinworm Larval
Instar
Y: 0.80 + O .795 X
Ln
O

51
The ratio of percentage of larvae present to percentage of larvae
absent (Table 6) in the observed injuries was 4:1 for the folded necrosed
injuries, 31:1 for the folded with no necrosis, 1:3 for blotches with
necrosed tissue, and 3:1 for transparent blotches. Consequently, the
use of necrosed blotches will indicate that ca. 77% TPW larvae will be
absent from the observed blotches. If a high number of injuries per
plant falls in this category, the probability of not measuring
larval presence in each injury is increased. We can deduce that necrosed
tissue generally indicates that larvae are already attacking the fruit or
other leaves, or have left the canopy to pupate.
In a crop such as tomato where the margin of profit is great,
expensive methods of control are usually dictated. The use of a system
that will predict the damage level to the plant requires a high level of
accuracy. It is suggested that the method described here is advisable
for plants during stages TV to RV^.
Conclusions and General Discussion
Studies of tomato growth in different cropping seasons are useful
to determine effect of planting time on plant development. Tomatoes,
cv Flora-Dade, planted later in the winter have less (ca. 117 dm )
leaf area than those planted early in the fall (ca. 253 dm leaf area).
Thus, those crops planted in October-November may be able to support
more damage than those planted in January-February. The proposed system
divides the plant stages inco 2 vegetative stages (TV^ - TV2), 3 repro¬
ductive stages (TR-, , TR and TR_) , and a senescent stage (S ) . The
2. 1
description of the developmental stages of tomato can aid in using pest
management tactics. Definition of shorter developmental stages with

52
Table 6. Percentage of tomato pinworm larval occurrence in foliar
injuries with different phenological characteristics.
Percentage of TPW Larval Occurrence
Damage Description Present Absent
Transparent blotch
72.5
27.5
Necrosed blotch
23.3
76.6
Folded, no necrosed leaf
96.87
3.12
Folded, necrosed leaf
81.25
18.75

53
with more subdivisions would enhance phenological plant description.
This may allow better pest monitoring when plant development is in the
TR stage.
2
2
Results on leaf weight consumed (mg) and leaf area consumed (cm )
provided information on increments of those parameters for each larval
instar. Standard error and confidence intervals demonstrated a high
variability for both methods. Further research is necessary to deter¬
mine if such variability is caused by larval behavior or by use of
different leaflet sizes and leaf area. I consider the leaf weight
method promising in such areas as plant resistance and behavioral
chemicals (deterrents) evaluation. Damage assessment based on the
leaf area mined by TPW is not considered appropriate for monitoring
TPW density because of inherent variability in insect behavior and
plant morphology.
Injury length has proven useful in evaluating plant resistance
(Schuster 1977a). The relationship between larval head capsule
2
and injury length was intermediate (r =0.47). The regression equation
developed in this study can be used by plant resistance evaluators to
determine feeding inhibition at a given instar. This technique has
to be carefully used, however, since it is dependent on the type of
leaflet consumed. Larvae that attack small leaflets might develop as
well as one in a large leaflet but the injury length will be smaller.
Data gathered from the visual damage classification proved to be
useful to evaluate damage inflicted by TPW. Since TPW instars have a
distinct behavior as leaf blotchers and leaf tiers, it will be easier
to develop knowledge in which the average larval instar will determine
the damage degree in a plant.

54
Scouts should use different techniques at the same time if
possible. A population index, degree damage scale and a survey deter¬
mining the real presence of the larvae in the foliage will provide a
better estimate than a single technique. More research is needed to
evaluate these techniques together. Evaluation should be based on time
expended and reliability of the methods. Further study of the relation¬
ship between several types of foliar damage and direct damage to the
tomato fruit is needed.

CHAPTER III
SPATIAL DISPERSION OF TOMATO PINWORM EGGS ON TOMATOES
Introduction
Tomato pinworm (TPW) is one of the most important pests of tomato
Lycopersicon esculentum (Mill.) (Watson and Thompson 1932, Oatman 1970,
Poe et al. 1974). Little is known, however, about ovipositional pat¬
terns of this pest on tomato plants under field conditions. There is
some indication that caged moths under laboratory conditions deposit
eggs indiscriminately on all parts of the plant including the upper
leaves (Elmore and Howland 1943). Wellik et al. (1979) indicated that
lower portions of the plant should be examined in the field for both
larvae and eggs of the TPW.
Studies of TPW egg dispersion are necessary because this knowledge
affects the sampling program as well as the method of analyzing the
data. Furthermore, dispersion patterns can be used to give a measure
of population size as well as to describe the factors that may affect
the condition of the population. This paper (1) describes the spatial
distribution of TPW eggs on field-grown tomato plants under varying
levels of TPW infestation, (2) presents an evaluation and discussion
of factors affecting this distribution and (3) discusses sampling
strategy.
55

56
Materials and Methods
Experimental Plots
To test for a possible relationship between oviposition of TPW and
different leaf strata of tomato cv Flora-Dade, 8 plantings (Oct. 3, 1979;
Dec. 5, 1979; Jan. 8, 1980; Oct. 30, 1980; Nov. 25, 1980; Dec. 30, 1980;
Jan. 30, 1981; Feb. 28, 1981) of non-staked tomatoes were evaluated at
the Agricultural Research and Education Center, University of Florida,
Homestead, Florida. Each planting (ca. 450-947 plants) was direct-seeded
in raised beds (3-5) (ca. 45 m long) of Rockdale soil, and mulched with
light colored plastic. The seedbed's midlines were 182 cm apart. Plants
were spaced 38 cm apart.
Sampling Methods
Sample size was selected by a preliminary random sampling of 50
plants on 2 dates. The method described by Elliott (1979) was adopted.
The relative variation (SE/x) x 100) was calculated to compare sam¬
pling methods over a variety of sampling units (Hillhouse and
Pitre 1974, Ruesink 1980). Ten to twenty plants in each planting
were randomly selected on a weekly basis from February 7, 1980,
through May, 1980, and from Jan. 27, 1981, through May, 1981. Whole
leaves of the plant were first examined to determine differences in ovi¬
position on lower and upper leaf surfaces (Plantings 1-3) and to detect
differences in oviposition in different plant strata (Plantings 1-8).
A plant was divided into upper half and lower half in the first 3
plantings (1979-SO) and divided serially into 6 sections (upper, middle
and lower of each of the external and internal canopies) (1980-1981) .

57
External canopy was defined as extending from the periphery to 5-15 cm
into the plant interior. The variance was stabilized by fitting the
number of eggs obtained to a suitable model (Poisson and negative
binomial) and transforming to logarithm (x+1) or x+0.5 (Elliott 1979)
depending upon the original frequency distribution of the counts.
The mean counts of eggs in upper and lower strata were compared by
student's t-test for plantings 1-3. Egg densities in the 6 strata for
plantings 4-8 were compared by analysis of variance (ANOVA). Means
were grouped by Duncan's Multiple Range Test (P=0.05). When tests
indicated significant differences in egg densities between strata of
plantings (4-8), optimum sample allocation among strata was determined
for each planting date (Cochran 1977) .
Population Distribution Related to Leaf Position
To test differences in oviposition of TPW related to the vertical
distribution of the leaves with respect to the main axis, 17 randomly
sampled plants, each of which were 45 days old, were observed in a
commercial field. Leaves were numbered from bottom to top and the num¬
ber of eggs recorded. Data were analyzed by ANOVA and means were separated
by use of Duncan's Multiple Range Test (P=0.05). When t-tests indicated
significant differences in egg densities between leaves, optimum sample
allocation among leaves was determined (Cochran 1977).
Distances Between Eggs and Effects on Distribution
To determine if TPW egg distribution pattern is influenced by leaf¬
let size and egg density, the frequency of egg deposition on each

58
leaflet was recorded. Then, distance between eggs on each leaflet was
counted on 40 middle leaves collected from plants located in the same
field mentioned before. Several authors (Cottam and Curtis 1956)
proposed methods to evaluate randomness in spatial distribution of the
population by measurement of distances between individuals. In this
experiment, distance between eggs was checked by measuring the shortest
straight line between nearest neighbors with a metric ruler. Distance
accuracy was 0.05-0.25 cm. The frequency of occurrence of each dis¬
tance was evaluated for egg densities. Also, simple linear regression
was applied to determine any relation between egg density and leaflet
size.
Oviposition Related to Plant Age
To determine if plant age affects oviposition, the number of eggs
on each plant was counted on 60-80 plants ranging in age from 2 to 21
weeks. Plants in this experiment were in the same field as previously
described tests (plantings 4-8). Plants were inspected weekly during
April and May, when the highest peaks of oviposition occurred. Data
were subjected to ANOVA, and means separated by use of Duncan's Multiple
Range Test (P=0.05).
Results and Discussion
Selection of Number of Sample Units
The main objective of planning a survey should be to obtain the
required information with a minimum amount of labor. To achieve this,
it is necessary to select a number of sample units that are in agreement

59
with the desired degree of precision and cost. This requirement is
difficult to meet in practice. First, an acceptable index of precision
(SExlOO) is 25% (Barfield 1981). Secondly, the actual cost of sampling
x
tomatoes is 7 dollars per acre (Table 7).
Sample size was selected by a preliminary random sampling of 50
plants in 2 dates (Table 6). Three major criteria were followed to
select sample size. First, following the criteria outlined by Elliott
(1979), a suitable sample size was selected when the mean value ceased to
fluctuate. It is observed (Table 5) that with an increase in sample
size from 10 to 25 (at low egg density), the resultant mean (x)
fluctuates around 0.4-1.5 eggs/plant. Also, at higher egg density (2-7
eggs/plant) the number of selected sampling units is 20-25. Second,
the use of index of precision (SExlOO) over different sampling units is
a more adequate technique to select sample unit size.
Accordingly, the lower index of precision (Ip) was obtained when
the number of samples equals 50. Therefore, the percentage of the
standard error of the mean can be 34% if the TPW egg density per plant
is low (0.4-1.5 eggs/plant). This percentage is not good enough to make pest
management decisions. The index of precision can be 20% if the density is
higher (2-7 eggs/plant). Third, the sample number does not reconcile with
the actual budget per acre. Cost of sampling eggs is 1.4-77 dollars
(Table- 5) more expensive than the actual sampling cost per acre.
The number of samples for a fixed level of precision (random sampling)
s 2
was calculated. A random eag distribution was assumed, n= (—=-)" where,
Ex
n=number of samples required, s=standard deviation, x=mean, and
E=predetermined standard error (e.g., 0.25). For instance, at

Table 7. Comparison of different sample sizes for tomato pinworm eggs. Homestead, Dade County,
Florida, 1980.
Egg Density
No. Plants
Sampled
9
Mean Eggs/Plant S
SE
(SE/x)xlOO
Cost of
Sampling/Acrea
T b
IiOW
5
0.4
0.8
0.4
200
8.4
10
1.5
14.5
1.18
78
16.8
15
1.33
9.95
0.814
61
25.2
20
1.15
7.81
0.624
54
33.6
25
1.08
8.26
0.574
53
42
50
0.88
4.59
0.302
34
84
High
5
7
23.5
2.16
31
8.4
10
3.9
19.78
1.4
36
16.8
15
3.4
17.66
1.08
31
25.2
20
2.9
14.62
0.854
29
33.6
25
2.96
15.29
0.874
29
42
50
2.16
9.28
0.43
20
84
a
Cost of sampling was calculated on the basis of $7.00/man hr for scouting tomatoes. Average
time spent per plant was 14.4 min.
Egg density was considered low when mean eggs/plant ranged 0.4-0.88; egg density was considered
high when mean eggs/plant ranged 2.16-7.

61
endemic levels of TPW egg population (0.4-1.33), the number of samples
to be taken, being S=2.79, E=0.25, x=1.15 will be 94, with a cost of
158 dollars per acre. If the TPW egg population is epidemic (2.16-7),
the number of samples to be taken will be 28, being s=3.82, x=2.9 and
E=0.25. The cost of sampling will be 47 dollars per acre.
Accordingly, under low TPW egg densities, increasing sample
precision as the sample size increases is not worth the work required
in taking larger samples. Consequently, I selected sample sizes of
10-20 which gave the best practical results per unit of work expended
($16.8-25.2 dollars per acre). It is considered that sampling 'TPW
eggs is not a practical method to make spray decisions.
Statistical Description of TPW Egg Spatial Distribution
The use of statistical methods, e.g., t-tests, analysis of vari¬
ance, involves several conditions described by Snedecor and Cochran
(1967). One of them is that data must follow a normal distribution.
The distributions of density measurements on plant samples are sum¬
marized for each planting in the Appendix. These statistics (Table 8)
support the hypothesis that TPW eggs are clustered on plants. This
clustering was more apparent when TPW egg densities on each plant
ranged from 0.302-1.3. As mean densities increased, variance also in¬
creased except for planting 6. Values of the negative binomial parameter
(k) (Elliott 1979) range from 0.451-0.013 for my sampling. Thirty-two
percent of the weekly counts for each planting were fitted to the nega¬
tive binomial distribution (see Appendix). For plantings with higher

Table
8. Mean number of tomato pinworm eggs
in Homestead, Florida, 1979-1981.
per plant by planting
date for
8 tomato
plantings
No.
Planting
Date
Mean
Variance
Skewness
Kurtosis
CV
ka
Ib
1
Nov.
3,
1979
1.30
5.04
2.96
13.74
172.41
0.451
4.17
2
Dec.
5,
1979
0.736
2.44
2.78
8.18
212.24
0.317
3.05
3
Jan.
8,
1980
0.345
0.97
4.332
23.97
285.73
0.190
2.15
4
Oct.
30,
1980
0.037
0.139
18.28
405.40
993.60
0.013
2.79
5
Nov.
25,
1980
0.135
1.34
16.18
314.24
858.16
0.015
9.06
6
Dec.
30,
1980
0.532
0.315
5.44
33.707
423.38
-1
0.124
7
Jan.
30,
1981
0.221
0.52
4.91
31.04
328.65
0.1633
1.57
8
Feb.
28,
1981
0.302
0.74
4.34
24.35
286.19
0.208
1.75
al
k= x
2
S - 1
x
b 2 , 2
1= S + (x) - 1
X

63
population densities (average 0.302-1.30), kurtosis and skewness
decreased as the mean increased. Skewness values were all positive.
This indicates that egg distribution tails off among higher counts.
This information in conjunction with the data indicating clumping can
aid in sampling design.
Distribution of Eggs on the Upper and Lower Surfaces of Leaves
Statistically significant differences (P=0.01) were found for egg
numbers on lower and upper leaf surfaces. Eighty-nine percent of
the total eggs found per plant were on the lower surface (Table 9).
These results and the results from the greenhouse contrast with those
found in caged plants by Elmore and Howland (1943), who detected 45%
of all egg deposition on the upper surface of the leaves. Insect pre¬
ferences for oviposition on the underside might be correlated with dif¬
ferences in pubescence of the 2 leaf surfaces. The average number of
trichomes on the underside was 1441 per leaflet as opposed to 469 on
the upper surface. This may also indicate preference to avoid egg
desiccation, or to avoid higher light intensities during oviposition
(Hinton 1981).
Distribution of Eggs on Upper and Lower Halves of the Plant
Statistically significant differences (P=0.05) were found in the
number of eggs deposited on the upper half of the plant vs the lower half
of the plant for the 3 sample dates in the first planting (Table 10). The
upper part of the plant had more eggs on 13 of the 15 sampling dates.
There were no significant differences between upper and lower halves

64
Table 9. Ovipositional preference of tomato pinworm
for upper and lower surfaces of tomato
leaves from plants grown under greenhouse
and field conditions.
Mean Number
of Eggs of TPW
Greenhouse
Leaf Side
Fielda
Caged Plants0
Upper
0.857C
o.ioc
Lower
7.3763
2.77
aMean based on counts from 80 plants.
Mean based on counts from 60 plants,
c
Numbers were significantly different at P=0.001.

Table 10. Mean number of tomato pinworm eggs in 2 plant strata (upper and lower halves)
per plant at different sampling dates. Homestead, Dade County, Florida, 1980.
Planting
Stra ta
Mean Egg Density* of
Tl’W E 3 Per S
¡tratum On Specified Dates
2/7
2/12
2/21
3/7
3/14
3/21
3/27
4/5
4/11
4/18
4/24
5/2
5/10
i
Upper half
0.05**
0.05
0.50a
0.35
0.55
0.35
1.45
1.80
2.30a
2.70
1.25a
Lower ha]f
0. f>0
0.00
0.10b
0.20
0.65
0.15
1.25
1.20
1.20b
2.55
0.75b
2
Upper half
0.00
0.07
0.07
0.15
1.04a
0.18
0.39
0.18
-
0.311
Lower half
0.00
0.07
0.07
0.12
0.45b
0.12
0.23
0.04
-
0.139
3
Upper half
0.00
0.00
0.15
0.25a
0.15
0.25
1,7a
1.15
0.10
1 .05°
l.la
Lower half
0.00
0.15
0.00
0.15b
0.10
0.30
0.6b
0.25
0.00
0.35b
0.2b
*
Data transformed back to the original units after statistical analysis cf transformed data.
* *
Means followed by different letters, in the same planting and date, are significantly different at p=0.05 according
to t-test.

66
in the second planting (Dec., 1979). Analysis of the data from the third
planting (Jan., 1980) indicated significant differences in 6 of the 11
sampling dates. The upper half of the plant had more eggs except for
2 dates. In general, when numbers of eggs were higher in the lower
strata, this coincided with younger plant age (40-60 days after germin¬
ation) . Numbers of eggs were higher in the upper strata when plants
were in reproductive or older age (75-80 days after germination. These
data indicated that for 'Flora-Dade' ground tomatoes, ovipositional
preferences existed based on the level of the plant. Because of the low
economic threshold for TPW in tomatoes, it may be necessary to reduce
the sampling unit to detect major differences in internal and external
parts of the plant when populations are low. Consequently, smaller
sampling units were tested in subsequent experiments.
Distribution of TPW Eggs by Sampling Six Plant Strata
Statistically significant (P=0.05) (Table 11) differences were not
detected among the strata for the 4th (Oct., 1980) and 5th (Nov., 1980)
plantings possibly due to the relatively low mean egg numbers per plant.
However, the highest number of eggs oviposited was obtained in the upper
external canopy for planting 4 and in the middle internal canopy for
planting 5. There was an increase in eggs for the lower internal canopy
in planting 4 during January and February, when nocturnal temperatures
were lowest (2°C), and an increase toward the upper external portion of
the plants when temperatures were fluctuating between 17-29°C (April-May).

Table 11. Mean number of tomato pinworm eggs per plant in 6 strata; upper, middle and
lower external; upper, middle and lower internal canopy of the tomato plant.
Homestead, Florida, 1981.
Mean Kc
jg Density* of
TPW Eggs per
Str.it tun
on Spe<
•i £ ied
Dates
P1 a n t i ng
Strata
1/27
2/4
2/10
2/17
2/25
3/18
•1/1
4/8
4/16
4/24
5/1
5/8
5/14
4
Upper external
A A
0.00
0.05
0.00
0.00
0.05
0.15
0.00
0.30
0.20
0.20
1.3
Middle external
0.05
0.00
0.00
0.20
0.00
0.00
0.00
0.00
0.00
0.10
0.00
Lower external
0.00
0.05
0.00
0.15
0.00
0.00
0.00
0.00
0.00
0.00
0.00
Upper, internal
0.00
0.00
0.00
0.00
0.00
0.00
. 0.00
0.00
0.00
0.00
0.00
Middle internal
0.00
0.10
0.00
0.05
0.05
0.05
0.00
0.00
0.00
o. or»
0.00
I.ower internal
0.05
0.40
0.05
0.00
0.40
0.00
0.05
0.00
0.00
0.00
0.00
5
Upper external
0.00
0.00
0.60
0.05
0.40
0.10
0.60
0.10
Middle external
0.00
0.00
0.10
0.10
0.00
0.00
0.10
0.10
I/jwer external
0.00
0.00
0.00
0.10
0.00
0.00
0.00
0.10
Upper internal
0.00
0.00
0.20
0.00
0.00
0.00
0.00
0.00
Middle internal
0.00
0.05
0.40
0.00
2.05
0.00
0.00
0.00
Lower internal
0.00
0.05
0.00
0.00
0.00
0.00
0.00
0.00

Table 11—continued
6 Upper external 0.00
Middle external 0.00
Lower external 0.00
Upper internal 0.00
Middle internal 0.00
Lower internal 0.00
7 Upper external
Middle external
Lower external
Upper internal
Middle internal
Lower internal
0.70
0.30
0.25
0.10
a
0.30
0.90a
0.20
0.40
0.20
0.15
0.20
0.00
0 . i ob
0.40
0.20
0.30
0.00
0.00
0.15ab
o.oob
0.00
0.10
0.10
0.25
0.00
o.oob
o.oob
0.00
0.30
0.30
0.10
0.00
o.oob
0.00
'0.00
0.20
0.00
0.00
0.00
0.05
0.00
0.00
0.70
0.60ab
1.00a
0.75a
0.50a
0.30a
0.00
0.80a
0.4 5b
0.55a
0.40ab
o.io1
0.40
o.iobc
0.05b
0.05b
0.35b
0.00
0.00
0.00
0.05b
b
0.00
0.00b
0.00
0.40
0.00
0.30b
0.20b
0.00
0.00
0.10
0.00
0.0 5b
0.00
0.00
0.00

Table 11—continued
Upper external
0.10
0.40
0.55
1.20“b
0.55“
1 .B0a
Middle external
0.30
0.20
0.25
0.85b
o.ioab
0.25jb
Lower external
0.00
0.40
0.05
0.40b
ab
0.25
0.20ab
Upper internal
0.00
0.00
0.00
o.iob
0.25Sb
0.10b
Middle internal
0.00
0.00
0.25
0.15b
0.25“b
0.20ab
Lower internal
0.00
0.00
0.00
0.15b
0.00
0.10b
Data retransformed to the original units after statistical analysis.
Numbers followed by different letters, in the same date and planting are significantly different at P-0.05 according
to Duncan's Multiple Range Test.

70
Perhaps moths protect themselves from the cold temperatures by staying
close to the ground in the lower canopy. Despite these assumptions,
when number of eggs found per stratum was regressed (Table 12) against
temperature, there was no evidence of a relationship between the two
variables. Significant differences in numbers of eggs per stratum were
detected for the 6 (Dec., 1980), 7 (Jan., 1981) and 8 (Feb., 1981)
plantings. There was no significant variation among the six strata
during juvenile plant stages. Most of the significant differences were
observed (Fig. 6) during the mature stages (TR) of the plant. Concen¬
trations of eggs in the upper external strata varied slightly among
plantings. In plantings 4 and 5, eggs generally occurred on the top and
middle external canopy during the last weeks of sampling (April and May),
and on all strata during the first weeks (juvenile stages) in March and
April. When mean numbers of eggs in the external and internal canopy
were added to reduce the strata to 3 (upper, middle and lower), no statis¬
tical differences were observed despite the stratum reduction. This
agrees with the results expressed when 2 strata (upper and lower) were
sampled, indicating that differences in oviposition tend to be masked
if the units are widened. In general, the upper external stratum had the
highest number of eggs, followed by the middle external and internal strata,
during most of the sampling dates. At plantings 6 to 8, the TPW eggs
occurred in greatest abundance on the upper and middle external strata dur¬
ing all growth stages. More eggs (44-68%) were deposited within the upper
external canopy of the plant than in any other stratum (Table 13). Four to
twenty eight percent of the eggs were laid in the next (middle external
stratum). The lower external stratum had the lowest range (1-11%); however.

71
Table 12. Relationship between daily mean temperature (°C) and TPW
oviposition in 6 tomato plant strata. Homestead, Florida,
1981.
Independent
Variable
X
Dependent
Variable
No. Eggs/stratum
y
2*
r
* *
r
* * *
bo
bl^
Temperature
upper internal
0.05
0.22
-0.007
0.005
upper external
0.13
0.36
0.46
0.04
middle internal
0.10
0.31
0.10
0.01
middle external
0.02
0.14
0.02
0.0056
lower internal
0.01
0.10
0.10
-0.02
lower external
0.13
0.36
-0.11
0.01
*
r^
Coefficient of determination.
★ *
r . . .
Correlation coefficient.
* * *
bo
Intercept of y axis.
T
1 Slope.

Figure 6. Average number of tomato pinworm eggs per plant stratum during 6 different
sampling dates in 2 tomato plantings at different growth stages. A) Planting 7: Jan.
30, 1981. B) Planting 8: Feb. 28, 1981. TR2=second reproductive stage of development;
TV2--second vegetative stage of development. Plant strata; 1, 2, 3: upper, middle, lowe
external, 4, 5, 6: upper, middle, lower internal.

Mean Eggs /Stratu
A
PLANT STRAT A

VIVI X S INVld
n

75
Table 13. Percentage distribution of TPW eggs for each stratum of
tomato
County
plants in
, Florida,
5 tomato
1980.
plantings.
Homestead,
Dade
Stratum
Planting Date
Oct. 30
1980
Nov. 25
1980
Dec. 31
1980
Jan. 30
1981
Feb. 28
1981
Upper external
68
32
44
46
51
Middle external
4
15
23
28
21
Lower external
1
3
10
11
10
Upper internal
0
3
7
0.6
5
Middle internal
9
43
11
10
9
Lower internal
16
0.8
4
1
2

76
growth of the plant upwards and outwards can mislead my interpretation
of actual ovipositional preference. The percentage of eggs found per
internal stratum ranged from 0-7% in the upper internal, 9-43% in the
middle internal, and 10-87% in the lower internal. TPW oviposits mainly
in the upper external canopy when egg populations range from 0.75-1.5
and when the plant was in its reproductive stage. A lower proportion of
eggs was found in all other strata.
Sampling 6 plant strata demonstrated that TPW tends to oviposit
in the middle and upper canopy. It is necessary to use sample allocation
(n ), as outlined bv Cochran (1977) to minimize sampling cost or vari-
h
2
anee (s ). I assumed equal sampling cost for each stratum. Sample
allocation was estimated on dates in which statistical differences in
oviposition were detected.
In general, more samples should be allocated to the upper and middle
external strata (Table 14). Because TPW eggs are clumped in the upper
and middle canopy, these strata had the highest variance (see Appendix)
N S
(Tables 51-54). For a fixed total cost, n = (C-C )E h h where (C-C )=
o —o
h_
L ZNhSh/ Ch
E C.n., nh= n W S n NnS Therefore as S,_ increases so does nh.
i i h h = h. h
i=l EW.S EN s
h h h h
The average number (n=20) for all planting dates was 6, 5 and 3 samples
from upper, middle and lower external canopy, and 1, 4 and 1 from upper,
middle and lower internal canopy. Allocation ranged from 5-10 samples
for the upper external canopy (see Appendix), and ranged from 2-8 samples
for the middle external canopy. I considered this sample allocation to
to be the best, because standard error (SE) of the sample mean was more

Table 14.
TPW egg sample
vegetative (TV2
allocation for 6 plant
), first reproductive
strata during 3 different plant stages:
(TR^) , and second reproductive stage (TR2
second
) .
External
Stratum
Internal
Plant
Stage
Upper
Middle
Lower
Upper
Middle
Lower
TV0
8.59*
2.10
0
2.6
8.59
0
TR^
5.70
5.75
8.24
0
0
0
tr2
0
6.47
3.44
0
1.58
8.49
tr2
5.96
5
2
1
4
2
Tr2
8.76
0
4
0
6
1
tr2
4
3
6
2
4
0
tr2
5
8
6
8
0
0
tr2
5
5
0
1
2
0
tr2
6
6
1
0
4
1
TR
2
9
4
2
0
5
0
TR„
2
5
7
2
5
5
0
tr2
5
5
2
0
3
0
tr2
10
2
2
1
3
2
Average
6
5
3
2
4
1
•k
n =(N S, )n; proportional allocation, assumes cost equal for sampling in each stratum,
h h h

78
constant through time (range: 0.20-0.66). There were exceptions for
these sample allocations. For instance, during the month of February
(planting 4), more numbers of samples were allocated to the lower inter¬
nal stratum (see Appendix) (Table 54). Another aspect that requires more
understanding is the relation between phenologic'al stages and sample
allocation. As an example, it was observed (see Appendix) that when
plants were in vegetative stage (TV), more samples (n^=18), should be
allocated for the upper external and middle internal canopy. When
plants are in first reproductive stage (TR^), more samples (n^=6), are
allocated for upper external stratum. Finally, when plants reach the
second reproductive stage (TR0), all strata had similar sample allo¬
cations except for lower internal canopy (n, =0).
Egg Distribution Influenced by Leaf Position
During heavy oviposition (avg 21.94 eggs per plant) on 45 day-old
tomato plants, the highest number of eggs was observed on leaf number 4
(Table 15). The numbers of eggs on leaves 3 and 5 were statistically
equal to those found on leaf 4. The number of eggs decreased sharply on
leaves adjacent to the apical point toward the bottom of the plant
(leaves 1-2). Tjese results indicated that middle leaves of 45 day-old
plants under conditions of high egg oviposition (1-5.5 eggs per leaf)
have 65% of the total egg population. These data differ from those
obtained in experiment 1. The higher number of eggs per plant indicates
that the insect tends to oviposit in the upper-middle canopy, avoiding
the 2 top and bottom leaves of the plant. Several factors may influence
the ovipositional pattern. First, these results agree with Hinton (1981) ,

79
Table 15. Mean tomato pinworm eggs on tomato leaves from different
strata of 45 day-old plants. Homestead, Florida, 1980.
Leaf Position
Mean No.
Eggs/Leaf
No.
Leaflets/Leaf
Eggs/Leaflet
1 bottom
2.20b*
7
0.31
2
3.20a13
8
0.40
3 middle
5.40a
11
0.49
4
5.50a
11
0.50
5
5.00a
11
0.45
6 top
2.00b
8
0.25
7
i.oob
7
0.14
*
Numbers followed by different letters were significantly differ¬
ent statistically at P=0.05 according to Duncan's Multiple Range
Test.

80
who stated that species that lay eggs on plants have a marked preference
for laying a certain height above the ground. Secondly, the insect may
be avoiding overcrowding in the smaller top leaves and competition of
foliar consumption by TPW larvae on the lower leaves. The highest
sample (n=17) allocation was for leaves in the middle canopy (Table 16).
Higher variance (s =44.4) was found for eggs deposited on those leaves,
as was a high mean (x=6.6). This is caused by egg clumping in the canopy.
The fourth leaf had the highest allocation sample (n^=5), followed by the
fifth leaf (n^=4). The lowest allocation was for the bottom leaf
The standard error of the mean sample was lowest (SE/x=0.21), for the
third leaf and slightly higher (SE/x=0.24) for the fourth leaf. There¬
fore, when higher density and large variance are found, the leaves
selected should be the middle ones. Sample allocation was reduced for
bottom and top leaves. These leaves had smaller variance and smaller
density than the middle ones.
Distances Between Eggs per Leaflet and Effect on Distribution
In the present study, the results indicated that TPW egg density
was not related to leaflet area (Table 17). The coefficient of deter-
mination (r =0.026) indicated that females tend to oviposit different
egg densities disregarding leaflet size. Therefore, any leaflet can be
selected as the sampling unit. Frequency of egg occurrence per leaflet
was not related to distance between eggs. Low coefficients of deter-
mination (r^=0.19-0.23) between frequency of occurrence at different egg
densities (2, 5 and 10 eggs/leaflet) and egg distances indicate lack of
linear relationship between these variables. The slope (bl) obtained

Table 16.
TPW egg sample
45 days old.
allocation
on tomato
leaves
numbered from
bottom to
top. Plants
Parameter
Leaf Number
1
2
3
4
5
6
7
X
2
3.50
4.58
6.58
4.70
1.91
1.25
s2
5.07
8.44
15.88
44.38
35.22
7.17
12.5
SE
0.60
0.83
0.96
1.61
1.43
0.77
1.25
Sh
2.25
2.90
3.98
6.66
5.93
2.67
3.53
SE/x
0.30
0.23
0.21
0.24
0.30
0.40
1.00
*
n,
1
2
3
5
4
4
2
h
•k
nh--(NhSh)n, Nh=200, n=17.
ENhSh

82
Table 17. Relationship between frequency of occurrence of TPW eggs per
leaflet as dependent variable and distance among eggs and
leaflet area as independent variables.
Dependent
Variable
y
Independent
Variable
X
2*
r
â– k k
r
k k k
b
o
bl
No eggs
Leaflet area
0.026
0.16
1.99a
0.04
2 TPW eggs
Distance among
eggs
0.23
0.48
1.75a
b
-0.15
5 TPW eggs
Distance among
eggs
0.19
0.44
1.47a
-0.12b
10 TPW eggs
Distance among
eggs
0.23
0.48
1.38a
-0.22b
r coefficient of determination.
r=correlation coefficient.
k k k
bQ=intercept of y asis.
tb^=slope.
numbers were highly statistically significant (P=0.01).
T_
numbers were statistically significant (P=0.05).

83
for any egg density was negative and highly significant (P=0.001). This
can be explained in Fig. 7, where the frequency of egg occurrence at dis¬
tances higher than 3 cm was as low as 5%. The average distance between
eggs was 0.5-0.75 cm. The average number of eggs found on each leaflet
was 2-3. These results agree with those expressed by Poe (1973) ; in the
present study the number of eggs on each leaflet was as high as 11.
Eggs tended to be laid more uniformly in some parts of the leaflet. Per¬
haps the female lays 2 eggs successively on a certain part of the leaf¬
let, but is likely to move away after oviposition. The arrangement of
eggs may also be a reflection of heterogeneity of conditions among parts
of a leaflet such as pubescence and leaf venation. From the practical
standpoint, these results can be used to determine use of single leaflets
as less variable sampling units compared to the whole plant. A more
detailed study of female behavior is necessary to determine the role of
leaf factors (e.g., pubescence) affecting oviposition.
Differences in Oviposition Related to Plant Age
The relationship between oviposition and stage of plant development
was determined during the study of plantings 4-8. Statistical differ¬
ences were detected among these plantings (Table 18), when plantings
were 19, 15, 11, 7 and 3 weeks old (stages TR0, TR , TV2, TV1 respec¬
tively. The largest number of eggs was detected in planting 7, when
this planting was in the TR - TR stages. At the same time, egg num-
1 ¿
bers decreased for planting 6 after the 10th week of plant growth. The
mean number of eggs in planting 8 increased slightly from week 3 (TV2) ,
through 7 (TR., ) . These data indicate that there may be several factors,

Figure 7. Frequency of tomato pinworm eggs at different distances (cm) between eggs when mean
eggs were A) 2 eggs per leaflet and B) 5 eggs per leaflet.

Frequency
CD
U1
Distance Between eggs

Table 18. TPW oviposition on tomato at different plant stages. Homestead, Florida, 1981.
Planting Date of Sampling
Date
4/1
4/a
4/16
4/24
VI
5/8
5/14
October
.05*(17)b*‘ TR
.3(IB)a TR
0.20(19)b
TR
0.30(20)b
S,
1.3 (21)b
s
3
3
3
1
1
November
.10 (I3)b TR
1.3114)a TR
0.25(15)b
TR
2.45 Í16)a
TR
0.1 (17)b
TR
0.7
(18) b
TR
0.8
(19) b
TR
2
2
2
2
3
3
3
December
1.9 ( 9)a TR t
1.2(10)a TR
0.75(11)ab
TR
0.30(12)b
TR
0.5 (13)b
TR
1.0
(14) b
TR
0.6
(15) b
TR
2
2
3
2
2
2
2
January
1.6 ( 6) a TR
1.5 ( 7)a
TR
1.90( 8)a
TR
1.55{ 9)a
TR
1.24 (10)a
TR
0.4
(11) b
TR
.1
i
2
2
2
February
0.4 ( 2) a TV
1.0 ( 3)ab
IV
1.1 ( 4)a
'IV
2.85( 5)a
TV
1.4
( 6)a
TR
2.65( 7)a
TR,
1
2
2
2
i
1
*
Numbers
in parenthesis are
weeks after emergence of crop.
* *
Numbers within a column followed by the same letters are riot significantly different according to Duncan's Multiple Range Test
(P Stages of the tomato plant according to Chapter 1.

87
such as presence of inflorescence and water content associated with
plant age, which account for frequency of TPW oviposition. The effect
of plant water content in oviposition will be discussed in chapter 8.
General Discussion and Conclusions
Tomato pinworm egg distribution is one of the least studied aspects
of this pest. In this research useful data were gathered about the spatial
pattern of TPW eggs in the tomato plant, effect of plant age on oviposition,
and use of these data for TPW egg sampling allocation.
The tomato pinworm tends to select certain leaf sides, plant strata
and plant age for oviposition. TPW prefers ovipositing on the leaf under¬
side. Ninety percent of the total eggs were laid in the leaf underside.
Tomato pinworm oviposited mainly in the upper and middle plant canopy.
Fifty-one percent of the eggs were laid in the upper canopy. There were
exceptions to this rule. For instance, a crop planted during Oct. 30,
1980, in which sampling was done during January and February, had a higher
than usual percent (17%), of eggs laid in the internal canopy.
The information described before was used to develop a sampling plan.
First, 10-20 plants/acre can be used as sample size if a practical equi¬
librium between sampling cost per acre (33) dollars, and index of precision
(54-31%), is desired. It was found that proportional sampling can be
allocated for the upper external stratum (n^=6), followed by the middle
internal stratum (n^=5) , and 3, 1, 4 and 1 samples for lower external,
upper, middle and lower internal canopy, respectively. Experiments also
showed that under high density (22 eggs/plant) the insect prefers the
middle leaves of the plant. These results were expected because TPW egg

88
population is considered clumped in the plant (k=0.01-0.45) . The strata
that has more eggs had higher variance than the strata with less ovi-
oosition. Therefore, more samples were allocated (n =4) to the middle
h
stratum.
Also, the shortest range for a confidence interval of the mean
was in the upper external stratum (0.51-1.7). At this time it is not
known which is the economic threshold based on egg counts. The larval
economic threshold is considered to be 1 larvae per plant (Chapter V),
therefore, if no egg mortality is expected 1 egg per plant will be the
economic threshold. The strata in which confidence intervals are less
fluctuating through time are the upper and middle external canopies.
Then, these strata are the ones to be selected for egg sampling in the
field.
As expected, oviposition increased as plant age increased. More
eggs were found on plants 4 weeks old (TV ) than on plants 2 weeks old
(TV ). Egg numbers increased for plants 8-16 weeks old (TR^ - TR^),
then decreased for plants 17-21 weeks old (TR -S). More research is
nececessary to evaluate attraction for female oviposition based on
physical and chemical qualities of the plant.
2
Leaflet area and egg density were unrelated (r =0.026). Distance
2
between eggs was not influenced by egg density (r =0.19-0.23). The
average distance between eggs per leaflet ranged from 0.5 to 0.75 cm.
These results indicated TRW tendency to oviposit eggs at a common
distance despite egg density and leaflet area.

CHAPTER IV
SPATIAL PATTERNS OF DISPERSION OF TOMATO PINWORM
LARVAE IN TOMATOES
Introduction
Dispersion patterns of Keiferia lycopersicella (Wals.), tomato pin-
worm (TPW), larvae in tomatoes have not been studied in great detail.
Knowledge of these patterns is necessary to develop a better sampling
procedure. Several techniques for population estimates such as ab¬
solute, relative estimates and population indices are used to determine
insect distribution (Southwood 1978).
The TPW larval population intensity method has been used by Wellik
et al. (1979) to determine sampling accuracy. However, TPW population
indices have only been used to measure economic damage (Wolfenbarger et
al. 1975) and plant resistance (Schuster 1977a).
Because leaf mining lepidopterous larvae do not move from a given
plant to neighboring ones (Dethier 1959, Nishijima 1960, Schoonhoven
1972) population indices can also be used to detect distribution pat¬
terns (Gomez and Bernardo 1974, Henson and Stark 1959, Condrashoff
1964). I used TPW damage index to (1) estimate dispersion patterns of
TPW larvae during different plant stages, (2) determine an appropriate
sample size and sample unit for TPW larval injuries, and (3) discuss
sampling strategy.
89

90
Materials and Methods
Sampling Methods
To determine the relationship between damage by TPW on different
strata of the tomato plant, 8 plantings of nonstaked tomatoes cv.
'Flora-Dade' (Nov. 3, 1979; Dec. 5, 1979; Jan. 8, 1980; Oct. 30, 1980;
Nov. 25, 1980; Dec. 30, 1980; Jan. 30, 1981; and Feb. 28, 1981) were
evaluated at the University of Florida Agricultural Research Center,
Homestead, Florida. Each planting (ca. 450-947 plants) was direct-
seeded in raised beds (3-5) (ca. 45 m long) of Rockdale soil, and
mulched with light colored plastic. Seedbeds midlines were 182 cm
apart. Plants were spaced 38 cm apart.
Sample size was selected by a preliminary random sampling of 50
plants on 2 dates. The method described by Elliott (1979) was adopted
in this selection. The sample size was chosen at the point when TPW
mine density variance stabilized. The percentage error that can be
tolerated in the estimation of the population mean was expressed as the
standard error of the mean. Finally, the cost of sampling within the
plant (C ) (Southwood 1978) and the cost of moving from 1 plant to
another (C ) were considered to estimate the relative net precision
P
100 —
value (RNP = — -■--) , RV being the ratio SE/x and C = C + C .
RV x C ups
u
Selection of the sampling unit for each plant was made by randomly
collecting 1 leaf/plant, 2 leaves/plant and inspection of the whole
plant (upper and lower canopy). The survey was made on 11 dates on

91
2 crops planted during Oct., 1979, and Dec., 1979. The mean per sample
and standard error of the mean were estimated. The proportion of the
true population collected per sample was determined by dividing the
number of individuals per sample unit by the number on the whole plant
and by comparing the RNP per sample unit.
After this, 20 plants in each planting were sampled by use of the
simple random sample technique (Cochran 1977). Sampling was done on a
weekly basis from Feb. 7, 1980, through May 29, 1981. The whole leaves
of the plant were first examined to determine differences in larval
injuries on lower and upper leaf surfaces (plantings 1-3) and then to
detect differences in larval injuries in different plant strata (plantings
1-8). The plant was divided into 2 sections (upper and lower) for the
first 3 plantings (1979-1980) and divided serially into 6 sections
(upper, middle, and lower part of the external and internal canopy) for
the last 5 plantings. After values of bilateral asymmetry of frequency
distribution (kurtosis) have been obtained, data can be normalized
(Sokal and Rohlf 1969). Data were transformed by replacing the value
(x) by logarithm (x+1) (Elliott 1979). The counts were assessed by a
t-test for results from plantings 1-3, and by analysis of variance
(plantings 4-8). Means were separated by Duncan's Multiple Range Test
(P=0.05).
Results and Discussion
Sample Size
Two goals were determined for sample size in tomatoes. One of them
was an index of precision (1^=25%) and the other was the cost of tomato

92
sampling in southern Florida ($7/acre). To accomplish this, sample size
was selected based on 3 major criteria.
First, following the criteria outlined by Elliott (1979) , a suitable
sample size can be selected when the mean ceases to fluctuate (Table
19). It is observed that when average larval injuries run as low as
2.12 or as high as 11.05, injuries, variances and means tended to
stabilize at ca. 20 plants.
Second, the use of index of precision (It,=SE/x) demonstrated that
for low populations (0.2-2.12), SE/x ratio was between 91-24 when 20-50
plants were sampled (Table 19). It fluctuated between 20-29% for the
20-15 plants at populations above 10.93 injuries per plant. These
results demonstrate that for lower insect populations it is necessary to
increase the number of plants up to 50 and to reduce it to 15 when the
population is as high as 11 injuries per plant.
Third, the sample number does reconcile with the actual budget. Cost
of sampling larval injuries is 1.26 dollars less than the actual budget
per acre if the sample size selected is 20. The use of relative net pre¬
cision (RNP = ) can also be used for sample size selection. The
u
larger the RNP of a sampling method, the greater the precision for the
same cost. RNP values when population is low can be selected for an
acceptable I . Therefore, RNP values can be accepted for 30-50 plants
under such conditions. RNP values when the population is high can be
selected for 25-50 plants. Nevertheless, to reconcile precision with
cost, sampling 20 plants gave an acceptable RNP for high or low TPW
populations.

Table 19. Sample size and relative net precision (RNP) for sampling tomato pinworm injuries at low
and high population densities. Homestead, Dade County, Florida, 1980.
Larval
Density
No. Plants
Sampled
X
s2
S
SE
(SE/S)*100
Cs3
cPb
C total
RNPC
Low
5
0.2
0.2
0.44
0.2
100
0.83
0.6
1.43
0.69
10
1.2
3.95
1.98
0.63
52.5
1.67
1.2
2.87
0.44
15
1.26
3.78
1.94
0.501
39.76
2.50
1.80
4.3
0.58
20
1.9
12.41
3.52
0.787
41.42
3.34
2.4
5.74
0.42
25
2.12
11.52
3.39
0.67
31.6
4.17
3.00
7.17
0.44
30
1.83
10.07
3.17
0.58
31.6
5.01
3.6
8.61
0.36
50
1.58
7.3
2.7
0.38
24.05
8.35
6.0
14.35
0.28
High
5
17.2
212.6
14.6
6.5
37.84
0.83
0.6
1.43
1.84
10
13.2
187.95
13.7
4.33
32.80
1.67
1.2
2.87
1.06
15
10.93
153.5
12.4
3.2
29.2
2.5
1.8
4.3
0.79
20
11.05
179.52
13.39
2.99
27.05
3.34
2.4
5.74
0.64
25
12.24
173.77
13.18
2.63
21.48
4.17
3.0
7.17
0.64

Table 19--Continued.
30
11.86
148.91
12.2
2.22
18.71
5.01
3.6
8.61
0.62
50
11.14
120.51
10.97
1.55
13.01
8.35
6.00
14.35
0.53
Cost per man h sampling: $7.00; ta; time per plant 1.43 min.; cost/plant =$0.16 = Cs.
Cp 1.06 min. to move from 1 pi. to another; cost in $0.12 = Cp.
C RNP
100
(RV)x(CV)
hO

95
Sampling Unit
Data from different numbers of leaves per plant in the upper and
lower portions of the plant are shown in Tables 20-21. In Fig. 8 it is
shown that it is necessary to select 2 leaves in the lower canopy if the
number of injuries is as low as 0.03-0.55. This is 20-30% of the total
number of injuries per plant. When the insect population increased, 2
leaves from the upper and lower parts were also necessary to obtain 32-
34% of the total population. Selection of 1 leaf per plant from the
upper portion gave as low percentages of the total damage as 1-2%, but
increased during later sampling dates up to 9%. The number of injuries
in 1 leaf per plant selected from the lower mid portion of the plant
remained stable at about 10% with the highest being 16%.
Another aspect to consider for sample unit evaluation is the confi¬
dence interval of sample mean (CI=x+taS"). According to the results
expressed in Tables 20-21, at low larval injury densities it is more
appropriate to sample the whole plant. The confidence interval remained
stable (0.24, -.08) for the whole plant until larval density was higher
than 1. When density increased above 1, 2 leaves per plant gave a more
stable confidence interval through time.
It is necessary to evaluate sample unit based on cost of sampling.
Cost per sampling unit is shown in Table 22. Relative net precision
was considered lowest when the whole plant was inspected. Two leaves
from the lower canopy gave an acceptable RNP during 5 of the sampling

Table 20. Mean number of TPW foliar injuries and standard error on different sampling units
at specified date.
Crop planted in
Nov., 1979
. Homestead
, Florida
•
Sampl i rig
Unit
2/3
Lowe r
2/21
Upper
Lower
3/14
Upper
Lower
3/20
Upper
Lower
4/4
Upper
Lower
1 Le a f
Ü.0040.00
0.00+0.00
0.0040.00
0.00+0.00
0.08+0.05
0.1610.09
0.59+0.38
0.54+0.22
0.32+0.lb
1.44+0.27
(0.1S, -0.02)*
(0.35, -0.3)
(1.37, -.19)
(.99, .08)
(.69, -.05)
(1.99, .88)
2 LfdVüS
0.00+0.00
0.04+0.04
0.00+0.00
0.00+0.00
0.24+0.11
0.55+0.21
0.76+0.24
1.16+0.35
0.88+0.26
1.8840.41
(.12, -.04)
(.46, .01)
(0.98, .11)
(1.25, .26)
(1.88, .43)
(1.41, .35)
(2.72, 1.03)
Whole Plant
0.00+0.00
0.12+0.00
0.00+0.00
0.0840.00
1.66+0.37
4.77+1.38
1.324 0.26
4.32+1.01
3.6+0.52
4.84+0.75
(.20, .0-1)
(.24, -.00)
(2.42, .00)
(2.87, 1.92)
(1.85, .78)
(6.39, 2.24)
(4.67, 2.52)
(6.38, 3.29)
lJ5‘i confiJence limits.

Table 21
Mean number of TPW foliar injuries and standard error on different sampling units
at specified date. Crop planted in Jan., 1980. Homestead, Florida.
Sainpl i ny
2/8
2/] 2
2/25
3/14
3/20
Unit
Upper
Lower
Upper
Lower
Upper
Lowe r
Upper
Lower
Upper
Lower
1 I.eaf
0.03+0.13
0.21+0.09
0.09+0.05
0.10+0.07
0.25+0.1
0.32+0.12
0.50+0.20
0.70+0.22
0.24+0.10
0.36tO.95
(.34, -.16)*
(.39, .02)
(.15, -.05)
(.23, .03)
(.45, .04)
(.56, .07)
(.83, .12)
(1.13, .22)
(.45, .02)
(.55, .16)
2 Leaves
0.16+0.10
0.41+0.20
0.10+0.10
0.15+0.08
0.64+0.3
0.68+0.18
0.72+0.14
1.16+0.24
0.44+0.20
1.1210.14
(.37, -.05)
(.81, .01)
(.30, -.10)
(.31, -.01)
(1.25, .02)
(1.05, .30)
(1.0, .43)
(1.65, .66)
(.85, .02)
(1.7, .83)
Whole Plant
0.76-1-0.10
1.33+0.43
0.55tO.24
0.45+0.18
1.44+0.44
2.00+0.42
5.40+0.71
6.72+0.97
3.92+0.34
6.12+0.38
(1.08, -.06)
(2.21,. 44)
(1.04, .05)
(.82, .07)
(2.34, .53)
(2.86, 1.13)
(6.94, 4.0)
(8.71, 4.72)
(4.62, 3.21)
(5.33, 6.9)
95% confidence limits.

Figure 8. Percentage of tomato pinworm (TPW) larval injuries in 2 sampling units
from different plant portions, related to number of injuries in the whole plant:
1) 1st planting, Nov. 3, 1979; 2) 3rd planting, Jan. 8, 1980.

% TPW Injuries
Sampling Oates

Table 22. Sample size and relative net precision (RNP) for sampling TPW larval injuries on upper
and lower plant canopy. Homestead, Florida, 1980.
Sampling
Sampling Date
Unit
2/8 2/1?
2/25
3/14
3/20
4/4
4/18
Upper Lower Upper Lower
Upper Lower
Upper Lower
Upper Lower
Upper Lower
Upper Lower
1 Leaf*
0.75**
0.20
0. so
0.12
0.80
0.22
1.41
0.25
1.16
0.31
1.09
0.76
1 . 33
0.2 3
2 Leaves
0.07
0.11
0.04
0. 10
0.09
0.20
0.22
0.25
0.10
0.42
0.21
0.37
0.14
0.19
Whole Plant
0.02
0.03
0.02
0.02
0.03
0.05
0.07
0.67
0.11
0.15
0.07
0.00
0.08
0.10
*
Cs = 1 loaf upper = 0.02 min-man
1 leaf lower = 0.12 min-man
2 loaves upper = 0.23 min-man
2 loaves lower = 0.10 min-man
Whole Pi. upper = 1.43 min-man
Whole PI. lower = 1.43 ini n-man
100

101
sampling dates. The highest RNP was obtained from sampling 1 leaf. A
sound sampling program requires precision and depends on resources. A
balance must be struck between the two to keep variance minimal for
fixed costs. Emphasis must always be given to practical considerations
(Ramsany 1980). In general, 2 leaves from the lower canopy should be
used as a sampling unit when a stable RNP is desired.
Statistical Distribution of TPW Larval Injuries
Different procedures were used to detect data normality. The sta¬
tistics summarized in Table 23 suggest a larval aggregation of TPW in the
tomato plant foliage. In general, as the mean increases, variance also
increases. The variance to mean ratio, or index of dispersion (I) will
approximate unity if there is agreement with a Poisson series. The (I)
values obtained were far from unity. Values ranged from 2.45 to 6.05.
Values of skewness and kurtosis were all positive. This means that fre¬
quency distribution of larval counts tails off among the higher counts.
Values of the k from negative binomial distribution are considered more
clumped when k approaches zero. The lowest value found in this data was
k=0.21. The Poisson distribution, however, fitted (P=0.005) 34% of the
sampling dates (see Appendix). In general, TPW injuries were considered
clumped in the tomato plant.
Distribution of Injuries on Upper and Lower Portions of the Plant
Statistically significant differences (P=0.05) were detected for
the injuries located in the lower part of the plant for the 5 sampling
dates in the first planting (Table 24). Statistical differences were

Table 23. Sample statistics: Mean tomato pinworm larval injuries per plant in 8 tomato
plantings. Homestead, Florida, 1979-81.
Planting
No.
/Date
N
X
s2
S
Skewness
Kurtosis
k*
J * *
1 Nov.
3,
1979
525
3.76
22.77
4.76
2.59
9.73
0.743
6.05
2 Dec.
5,
1979
240
2.49
14.65
3.82
3.33
18.22
0.509
3.83
3 Jan.
8,
1980
596
2.40
12.07
3.47
2.81
14.35
0.59
5.02
4 Oct.
30
, 1980
882
0.40
1.13
1.06
3.9
19.84
0.21
2.82
5 Nov.
25
, 1980
894
0.41
1.13
1.06
3.92
19.42
0.23
2.76
6 Dec.
30
, 1980
1019
0.49
1.37
1.17
3.25
12.78
0.27
2.79
7 Jan.
30
, 1981
953
0.39
0.97
0.98
3.35
13.30
0.26
2.48
8 Feb.
28
, 1981
717
0.35
0.86
0.93
3.65
17.33
0.24
2.45
*
x
102

Table 24. Mean tomato pinworm (TPW) foliar larval injuries at 2 different plant levels for 3
different plantings. Homestead, Dade County, Florida, 1980.
Planting Strata
Mean TPW Leaf Injuries*
k per Stratum on Specifi
ud Date
2/8
2/12
2/21
2/29
3/7
3/14
3/20
3/27
4/3
4/11
4/24
5/2
5/9
5/20
6/2
1 Upper
0.87b**
0.55a
1.44a
2.04a
5.48a
3.92a
8.68b
7.32b
5.04a
11.8a
8.64a
5. ba
10.4a
I ,owe r
1.33a
0.45a
2.00a
2.24a
6.72 a
6.12a
16.8a
11.84a
7.32b
11.76a
8.30a
5.7a
5.6a
2 llppor
0.00
0. ou
0.00
0.00
0.7a
1.0a
3.0b
2.7a
4.3a
7.4a
6.5a
4.8a
4.2a
3.4a
6.3a
Lower
0.04
0. 33
0.00
0.16
1.36
5.3a
4.7a
3.5a
8.1a
7.0a
5.2a
3.3a
3. la
1.9a
4.7a
2 Ui per
0.00
0.00
0.00
0.01a
0.08a
0.00
0.8b
2.3b
3.5b
13.2a
7.8a
7.6a
1 . 5a
1 . 3a
Lower
0.00
0. 00
0.00
0.12a
0.16a
0.08
1.7a
8.3a
6.6a
7.4b
4.3b
4.4b
0.36a
0.8a
A
Data transformed
back to the original unit
s after
statist.
leal analyses have been
carried
1 out from the transformed data.
Moans followed 1
>y the same letter
in the
same planting and date are not
significantly different
at P=0.
.05, according to
Students
t-tost..
103

104
only detected once for the second planting. In general, the lower part
of the plant had 60% more injuries than the upper half during the first
4 sampling weeks for the first planting (Nov., 1979). Also, the upper
half had zero injuries during the first 4 sampling weeks. When plantings
1, 2, and 3 were 16 (TR^), 12 (TR ), and 8 (TR0) weeks old, respectively,
there were similar numbers of injuries in the lower (55%) and upper
(45%) parts of the plant in planting 1. In planting 2, 75% of the
injuries occurred in the lower half and planting 3 had 67% in the lower
half. Nevertheless, the number of injuries in each stratum was not
significantly different. There were more injuries in the upper part of
the plant when the plantings were 21 (S^), 17 (TR^), and 13 (TR^) weeks
old. This may indicate more active larval consumption in the lower half
of younger plants and fewer injuries in that level in older plants.
These results disagree with the Florida results of Wolfenbarger et al.
(1975) , but are in agreement with those of Wellik et al. (1979) in
Texas.
Distribution of TPW Foliar Injuries in 6 Plant Strata
Statistically significant differences were detected for 3 of the 16
sampling dates for planting 4 (Nov., 1980) (Table 25). Greater numbers
of larval injuries were found in the middle internal, lower external,
and lower internal canopies. When data were converted into percentages
and analyzed, there were significant differences between the lower
portion of the plant and other strata (Fig. 9-10) . The number of foliar
injuries for planting 5 (Oct., 1980) showed statistically significant
differences in 1 sampling date. Greater damage was recorded from the

Table 25. Mean tomato pinworm (TPW) larval injuries in 6 plant strata for 5 plantings. Homestead,
Dade County, Florida, 1981.
Planting Stratum Sampling Date
1/27
2/4
2/10
2/16
2/25
3/3
3/11
3/18
4/1
4/8
4/16
4/24
5/1
5/8
5/15
5/29
4
Upper Ext.
o. no
0.00
0.00
0.05
0.1*
0.05
0.0b**
0.2a
0.4b
0.25a
0.3a
0.4a
0.8ab
0.6a
0.5a
0.7a
Upper Int
0.00
0.00
0.00
0.00
0.00
0.00
0.00b
0.05 a
0.3b
0. Oa
0.0a
0. Oa
0.0a
0.0a
0. Oa
0. Oa
Middle Ext.
0.00
0.00
0.00
0.00
0. 1
0.2
0.1b
0.3a
0.3b
0.55a
0.4a
0.5a
0.7ab
1 .2a
1.5a
1.2a
Middle Int.
0.00
0.00
0. 1
0.0
0.05
0.6
0.3a
0.9a
0.6b
0.4a
0.1a
0.3a
0.95ab
0.0a
0.0a
0.0a
Lower Ext.
0.00
0.00
0.00
0.00
0.05
0.4
0.1b
0.05a
0.9a
0.05a
0.3a
0.6a
0.3b
0.5a
2.6a
1.8a
Lower Int.
0.00
0.00
0.05
0.0
0.2
0.3
0.2ab
0.2a
0.0
0.2a
0.5a
0.1a
0.55ab
0.0a
0. Oa
0. Oa
5
Upper Ext.
0.00
0.00
0.00
0.00
0.00
0.00
0.05a
0 . la
0.0a
0.4a
0.35b
0.1b
0.9a
0.4a
0.2a
0.3a
Upper Int.
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.1a
0.0a
0.0b
0. Ob
0.3a
0.0a
0.0a
0.0a
Middle Ext.
0.00
0.00
0.00
0.00
0.00
0.00
0.2a
0.3a
0.4a
0.6a
0.3b
0.1b
1.2a
0.8 a
2.6a
0.8a
Middle Int.
0.00
0.00
0.00
0.00
0.00
0.00
0.05a
0.4a
0.2a
0.6a
0.35b
0.1b
0. la
0. 2a
0
0
Lower Ext.
0.00
0.00
0.00
0.00
0.00
0.00
0.05a
0.3a
0.2a
0.6a
0.75a
0.7a
0.3a
O.fia
3.5a
0.8a
Lower Int.
0.00
0.00
0.00
0.00
0.00
0.00
0.05a
0.05a
0.1a
0.5a
0.0b
0.4b
0.4b
0.0a
0.1a
0.0a
6
Upper Ext.
0.0
0.0
0.0
0.0
0.0a
0.0a
0.0a
0.2ab
0.9b
0.75a
1 . la
1. a be
0.15b
Upper Int.
0.00
0.0
0.0
0.0
0.0
0.0
0.0
0.0»)
0.0b
0.0a
0.1a
0.1c
0. Ob
105

Table 25—continued
Middle Ext.
0.00
o
o
0.0
0.0
0.0
C. la
0.15a
0.45ab
1.4a
1.15a
0.5a
1.9ab
0.85a
Middle Int.
0.00
0.0
0.0
0.0
0.1a
0. la
0.15a
0.101)
0.85b
J .7a
0.7a
0.6bc
0.3b
Lower Ext.
0.00
0.00
0.00
0.0
0.0a
0.1a
0.15a
0.6a
0.9b
2.5a
l.Ra
3. 3a
0.7a
Lower lnt.
0.00
0.00
0.00
0.00
0.1a
0. la
0.0a
0.2ab
0.25b
1.1a
0.4a
0.2c
0.2b
Upper Ext.
c
o
0.0a
0. lab
0.2a
0.05a
0.25b
0. 3a
0.2bc
Upper Int.
0.0
0.0a
O.Oab
0.0
0.0
0.0a
0.0
0.1c
Middle Ext.
0.0
0.05a
0. lab
1 .1
0.55a
0.5b
1. la
0.8a
Middle Int.
0.00
0.00a
0.45ab
0.15
0.35a
1.0b
0.7a
0.1c
Lower Ext.
0.00
0.00a
0.55a
0.25
1.3a
2.1a
1.7a
0.8ab
Lower Int.
0.00
0.00a
0.2ab
0.G5
0.7a
0.0b
0.1a
0.0c
Upper Ext.
0.0
0.05a
o
o
0.0
0.1
0.15
0.7b
0.7ab
Upper Int.
0
0.0a
0
0.1a
0.0a
0.0a
0.0b
O.Oab
Middle Ext.
0
0.0a
0
0.15a
0.3a
1.2a
1.55ab
0.3ab
Middle Int.
0
0.0a
0
0.05a
0.5a
0.8a
0.8b
0.6ab
Lower Ext.
0
0.0a
0
0.15a
0.7a
1.1a
1.8ab
0.9a
Lower Int.
0
0.0 a
0
0.1a
0.3a
0.3a
0.3a
0.2ab
Data retransformed to the original units after statistical analysis.
* *
Numbers within a column by planting followed by the same letter are not significantly different according to Duncan's Multiple
Range Test.
106

Figure 9. Percentage of tomato pinworm (TPW) foliar injuries found at upper, medium, and
lower stratum in 4 tomato plantings: 1) Oct. 30, 1980; 2) Nov. 25, 1980; 3) Dec. 30, 1980;
and 4) Jan. 30, 1981. Bars followed by different letters were significantly different
according to Duncan's Multiple Range Test (P=0.05). Percentages were previously trans¬
formed to arc sine. Percentages are expressed as actual numbers before transformations.

Sampling Dates
3
Upper Stratum
Medium
Lower
4
108

TPW foliar injuries/stratum
75 _
50 _
25 _
0
CZD
4*
F2
a
d
BIM
H IS
|ü£l
â–  ... i
■ ®
a
B 1;.*.
1
1 i i I
ii
—
p p
il
B
Si
Kl
il Ii
ii n
s
4
Jlil
s
â– 
®MJ
!
tv,
M4
Upper Stratum
Medium
Lower
1
2
Sampling Dates
109

Figure 10. Percentage of larval injuries at the external and internal canopy
evaluated from 5 tomato plantings. Plantings 4,5, and 6 planted in Oct., Nov.
and Dec., 1980; Plantings 7 and 8 planted in Jan. and Feb., 1981. Homestead,
Florida, 1980-81.

Planting Number
% Foliar Injuries
ITT
Internal Canopy

112
lower external canopy. During the 14-16 weeks of plant development, the
lower, middle, and upper canopy contained 91, 88, and 88% of larval
injuries, respectively. Larvae tended to occupy the upper part of the
plant in older plantings (18-20 weeks). Statistical differences for the
crop planted on Dec., 1980 were observed in 4 of the sampling dates.
More extensive larval injury was found in the lower external and middle
external canopy. When plants were 10-15 weeks old, higher proportions
of injuries (99-100%) again were found in the middle and lower canopy.
Statistical differences for the crop planted in Jan., 1981 were observed
in 3 of the 7 sampling dates. Higher percentages of numbers of injuries
were found (98-100%) when the crop was 9-15 days old. Larger numbers of
injuries were recorded for the upper external part of the plant for
planting 8 (Feb., 1981) during 1 sampling date.
Sampling 6 strata demonstrated that TPW larval injuries are signifi¬
cantly higher in the middle and lower canopy. Sample allocation (n^) as
2
outlined by- Cochran (1977) is necessary to minimize variance (S ).
Sample cost was considered equal for each stratum. Sample allocation
was estimated on dates in which statistical differences were detected.
In general, more samples should be allocated to the middle and
lower strata (Table 26). The average numbers (n=20) for all plantings
were 2, 3, and 5 samples for upper, middle and external canopy, and
0.09, 5, and 3 from upper, middle, and internal canopy. Allocation
ranged from 0-20 samples for the lower external canopy, and ranged from
0-9 samples for the lower internal canopy. I considered this sample
allocation to be the best, because standard error (SE) of the sample
mean was mere constant through time (range :0.20-0.66) .

113
Table 26. TPW larval injury sample allocation for 6 plant strata at 3
different plant stages: second reproductive (TR ), third re¬
productive (TR^) and senescent (S^) .
External
Stratum
Internal
Stage
Upper
Middle
Lower
Upper
Middle
Lower
S1
4*
3
6
0
4
0
S1
1
3
1
0
5
4
S1
0
5
9
0
6
0
tr2
0
0
20
0
0
0
TR
2
2
2
5
0
5
2
TR.,
0
0
0
0
12
4
TR
2
4
5
4
0
5
2
TR
3
2
1
2
0
2
9
TR,
3
2
4
5
1
4
2
TR,
1
2
5
0
5
4
tr3
5
5
2
0
2
6
Avg.
2
3
5
0.09
5
3
★
nh = (
N S
h h ) n;
n = 20, N
= 947.
SN’n Sh
n h

114
There were exceptions to these sample allocations. Allocation
increased for the upper canopy when plants were in the second repro¬
ductive stage (TR^) and during the senescent stage (S^). It is
necessary to correlate this sample allocation with fruit damaged in
different parts of the plant canopy. Knowledge of this relationship
will help in better prediction of yield losses.
Larval Injuries at Different Plant Growth Stages
The relationship between TPW foliar larval injuries and stage of
plant development was determined during the study of plantings 4-8
(Table 27). For the earlier planting (Oct., 1980), the mean number of
TPW injuries appeared during the second reproductive stage (TR^) and
peaked during the senescent (S^) stage. On later plantings (Nov.-Dec.,
1980) TPW larval infestation was not observed until the second repro¬
ductive stage (TR^) / when plants were 14 and 11 weeks old, respectively.
Higher foliar infestation was obtained when plants were in the late
reproductive stage (TR ) and also in the senescent stage. For the
winter season plantings (Jan.-Feb., 1981) injuries were first observed
during the second reproductive stage. The reason for differences in
larval infestation in different crops can be related to several factors.
First, there was a low oviposition rate in the earlier planting (Oct.-
Dec.). Secondly, the earlier fall crop (Oct., 1980) had a dense leaf
canopy and approached the senescent stage later than the other plantings
(Table 2). Thus, infestations in this planting were probably influenced
by a complex of factors, including increased food consumption by the
pest during the senescent stage due to reduction of food quality.

Table 27. Mean number and standard error of tomato pinworm (TPW) injuries in 5 different plantings at
specified date and plant growth stage.
P1 anti ny Sampling Dates*
1/27
2/4
2/10
2/16
2/25
3/3
3/11
3/18
Oct.
-80
0**TK (13)
0.0610.17 TH2(14)
0.15+0.08 TR^(15)
0.05+0.05
TR3(16)
0.55+0.3 TR ^(17)
1 . 4+0.4 5
TR 3(18)
0.6+0.23
s, (19)
1.7+0.72 G3(20)
Nov.
-80
0 TR? (B)
00+0.00 TR^ (9)
o.oo tr2(10)
0.00
tr2(H)
0 TR?(12)
0
TR2(13)
0.4+0.13
TR2(14)
1.05+0.69 TR (15)
Dec.
-80
0.00
TR1(7)
0.00 tr2(8)
0
TR2(9)
0
TR , (10)
0.20+0.15 TR2(1 l)
.Ian.
-81
0
TV?(5)
0 TR3(6)
Feb.
-81
0
TVj (1)
0 TV!(2)
115

Table 27--Continued
4/1
4/0
4/36
4/24
5/1
5/8
5/15
5/29
Oct.
.-80
2.1 + 1.11
Sj(22)
] .3+0.30 S (23)
) .6+0.53
(24)
1.7+0.39 Sj (25)
5.9+0.93 (2b)
2.3rn.63 (27)
4.6+1.21 S (28)
3.7+0.36 S
1 (2‘J,
Nov.
, -80
0.8+0.31
TR^(17)
3.55U.45
TR3(1B)
1.75+0.33 (19)
1.15+0.29 S3(20)
G.1+1.06 S3 (21)
2.0+0.66 S3(22)
6.6+0.20 (23)
1 .9+0.70 S
j (24)
Pec.
, -60
0.3+ü.i1
TR2(13)
0.45^0.22
TR, (13)
1.8+0.32
TR., (14)
4.25+_0.76TR2 (15)
10.15+_l. 1ÃœTR ^ (16)
4.4+0.88 TR^(17)
6.6 + 0.8 Tl<3 (18)
2.2+0.44 Sj
(19)
J a n.
-HI
0
TR, (8)
0.05m. 05
ti<2 (y)
1.3HJ. 37
TR,, (10)
2.25m. 5 TR, (11)
2.9 + 0.62 TR., (12)
3.85+0.33 TR (13)
3.8+0.7 TR2(14)
1.9+0.3 TR
j (15)
Feb.
-61
0
TV., (4)
0.05+0.05
TV, (5)
0
TR1(6)
0.55+0.2 TRj(7)
1.6m. 46 TR, (0)
3.85+0.45 TR2(9)
5.05+0.93TR (lo)
2.25+0.47
TR, (1 1)
A
Planti ncjs wore sampled during 1‘JBl .
A A
Mean number ot TPW foliar injuries, standard error of the mean plant growth stage, plant aye in weeks. Plant stages: TR - Reproductive,
= Senescent, TV - vegetative.
116

117
General Discussion and Conclusions
In this research useful data were gathered about the use of TPW
damage index to estimate TPW larval patterns during different plant
stages and about selecting an appropriate sample size and unit.
The population index used (TPW larval injuries/plant) indicated
that during plant stage TV2, TPW damage range (0.0-0.05) is lower than
that found in reproductive stages (TR^=0-0.55, TR2=0-5.05, TR^O.8-10.15) ,
and lower than the estimates from the senescent stage (S^=0.6-6.6).
Further information is needed to detect if nutritional changes in the
plant are the major factor for higher or lower number of injuries per
plant. The information is also necessary for detailed plant resistance
studies.
The information supplied by sample size can be used to develop a
sampling plan. For instance, 20 plants/acre can be used as sample size
when a practical equilibrium between index of precision (I ) and cost
ir
(RNP) is desired. The sampling unit to be selected is 2 leaves/plant
from the lower canopy when an acceptable (RNP=0.25 or less) is found.
This information can be used for commercial sampling for detection of
foliar injuries. One of the problems involving this selection is that
TPW foliar injury does not necessarily mean fruit injury. Therefore,
establishment of a relationship between fruit and leaf damage is needed.
Sample allocation resulted in a major proportion of samples allocated to
the middle and lower canopy. When total sample size equals 20, n, =8

118
samples should be taken from the middle as well as from the lower
canopy. An 21^=2 should be allocated for the upper canopy.
These results were expected because TPW injuries were considered
clumped in the tomato plant. The values obtained from the index of
s^
dispersion (1=—) ranged between 2.45-6.05. This departure from unity
x
means TPW larval aggregation in the plant. The data can be used to
transform larval counts in order to obtain normalization. If sequential
sampling is projected, it is necessary to find a common k value for TPW
injuries.
The population index should be incorporated into a more sophis¬
ticated damage and population evaluation method. For instance,
techniques such as degree of damage (Chapter 2) would be useful to¬
gether with the population index. The combination of these techniques
will help the scout in detection of larval populations approaching
economic thresholds.

CHAPTER V
TOMATO PINWORM ARTIFICIAL INFESTATION: EFFECT OF FOLIAR
AND FRUIT INJURY ON GROUND TOMATOES
Introduction
Economic losses resulting from insect injury to the foliage and
fruit are difficult to measure. One of the problems in determining an
economic threshold of a pest species is to distinguish between its
mere presence in a crop and the density that will cause an unacceptable
loss quality or quantity (Stern 1973) . Damage to the tomato fruit by the
tomato pinworm, Keiferia lycopersicella (Wals.), has been evaluated by
Poe and Everett (1974), Wolfenbarger et al. (1975) and Wellik et al.
(1979). The results were contrary in the three studies. Poe and Everett
(1974) found no correlation between leaves mined and presence of larvae
and fruit loss. Wolfenbarger et al. (1975) determined that TPW damage to
the 3 top leaves could be associated with fruit injury. Wellik et al.
(1978) indicated that damaged lower leaves and large fruit are the best
for estimating K. lycopersicella infestations. Waddill (1975) and
Schuster and Everett (1982) show estimates for losses from TPW natural
infestations.
In this study, two techniques for estimation of TPW fruit loss are
assessed: (1) use of TPW larval artificial infestation to measure yield
reduction and (2) use of the population index (TPW foliar injuries) and
119

120
its relation to yield losses. In addition, fruit damage related to time
of planting is examined, and the effect of plant pruning on TPW fruit
damage is evaluated.
Materials and Methods
First Experiment
Reductions in 'Flora-Dade' tomato yield caused by different popu¬
lation levels of TPW were measured during 1980 and 1981 at the Agri¬
cultural Research Center, Homestead, Florida. Crops were direct-seeded
on December 5, 1979; January 8, December 1980; December 30, 1980; and
January 30, 1981. Plots were thinned to one plant every 0.30 m. Seed¬
bed's midlines were 182 cm apart. Each crop was sprayed weekly with
fenvalerate 2.4EC at a rate of 0.064 kg ai/ha. Insecticide application
was discontinued 25 days before artificial infestation with TPW second
instar larvae. Larvae were reared on tomato plants in a greenhouse at
o
23+2 C and 75+4% RH. There were 2 replications per treatment in a
randomized complete block design. Artificial infestation was as
follows: First, plants were inspected to remove any TPW eggs
and larvae. Then, 10 plants 40-45 days old were infested once with
different numbers of larvae (1, 2, 4, 8, 12, 14 per plant). Plants
were inspected 1 day after infestation and larvae replaced if lost.
In a 2nd experiment, plants were infested twice with larval levels
mentioned above. An uninfested control was sprayed with fenvalerate
to keep it TPW free. A second control without insecticide or arti¬
ficial infestations was also used to compare with treatments. When
foliar and fruit injuries occurred in the unsprayed control, these
numbers were subtracted from the numbers found in the infested plots.

121
At harvest, counts of the numbers of leaves and fruits injured were
recorded together with the total number of fruits per plant from the
upper and lower canopy of the plant. Fruits were graded according to
USDA grading standards (Anonymous 1982) . In order to reflect market
standards, values of the fruit were modified for statistical analysis
by multiplying each fruit grade by a number. Numbers of extra large fruit
were multiplied by 5, large by 4, medium by 3, small by 2, and very
small by 1. Results were also analyzed before multiplication for fruit
size. Data were subjected to analysis of variance; treatment means
were separated by Duncan's Multiple Range Test at P=0.05.
Regression analysis was used to establish a possible relationship
between the number of larvae and the number of leaves and fruit damaged.
Data from leaf injuries in the upper plant canopy ( Y ), leaf injuries in
the lower canopy (Y7), number of fruits injured in the upper (F ), and
number of fruits injured in the lower (F ) canopy were regressed separately
on one independent variable: number of larvae per plant (x) .
number of fruits injured in the upper (F ) and lower (F,) canopy were also
regressed on Y and Y , respectively.
Finally, 2 regression models were used to find a statistical model
that related percent yield loss to the number of larvae per plant and
to number of foliar injuries. The main objective was to find a simple
model for infestation levels which could be used to establish economic
. . ' A A
injury levels. The regression model used was the form y =a+bx and y=
2 A
a+bx+cx . For the simple linear regression, y=estimated value of y,
a=the y intercept of the regression line, b the regression coefficient
and x the sample estimate. For the curvilinear regression, c=the second

122
regression coefficient preceding the second power of x. Results were
compared with the previous work by Poe and Everett (1974) , Wolfenbarger
et al. (1975) and Wellik et al. (1979).
Second Experiment
To determine the effect of planting date on damage to the tomato
fruit by TPW larvae, natural infestations with this insect were studied
in those plantings mentioned in Chapter 3. Crops were direct-seeded
during October, November and December, 1980 and January and February 1981.
Number of injuries per plant and number of tomato fruits damaged and not
damaged were recorded for each planting. A total of 20 randomly selected
plants was chosen and numbers of damaged tomatoes were compared. Secondly,
comparisons in damage estimates obtained from the customary sampling
method (6 contiguous plants per row) vs randomly selected plants per
row were made. Results were compared by use of relative net precision
(RNP=100/RVxCu), for both systems. Thirdly, to determine the effect of
pruning laterals on tomato fruit damaged by TPW, laterals from a total
of 20 randomly chosen plants (45 days old) were removed and yield was com¬
pared to that from 20 unpruned plants. Treatments were replicated 4
times. Analysis of variance and Duncan's Multiple Range Test were used
to compare treatments.
Results and Discussion
First Experiment
Single artificial infestation. After a one-time infestation of
plants with TPW larvae, the number of fruits damaged in the lower plant

123
canopy was significantly different from the sprayed control (P=0.05). In
the upper canopy average fruit damaged did not differ statistically among
infestation levels (Table 28). The number of fruit damaged in the
lower canopy when plants were infested with 1 larva was 10.5, 24.5 and
29.5% less than that found at 8, 12, and 14 larvae, respectively (Table
28). In general, damage from 4 larvae per plant and 14 larvae per plant
did not differ statistically. Results may indicate that 4 larvae per
plant represents the upper practical limit for fruit damage for 'Flora-Dade'
tomatoes. The numbers of injuries per fruit may increase with increased
larval infestation levels. Once populations reach ca. 4 TPW larvae per
plant, multiple injuries to the same fruit may become very common. There
is more fruit in the lower than in the upper part of the plant. Because
of larval positive geotaxis, larval activity seemed concentrated on the
lower plant parts. Consequently, fruit infestation in the lower canopy
is higher.
Marketable value of fruits on each canopy was significantly differ¬
ent for the infestation levels when different values were assigned to
fruit (Table 29). Values of fruit damaged by 1, 2 and 4 larvae per plant
differed from damage by 8, 12 and 14 larvae per plant. Eight, twelve
and fourteen larvae caused 1.63, 1.76 and 2.2 times more damage than 1
larvae.
Double artificial infestation. After a double infestation with TPW
larvae, the number of fruits damaged in the lower canopy at all levels
of infestation was again significantly different from the uninfested con¬
trol. Tomato fruit damaged in the upper canopy did not differ statis¬
tically among infestation levels (Table 30). There were no significant

Table 28. Tomato fruit damaged in the upper and lower plant canopy, after a single artificial
infestation with K. lycopersicella larvae on ground tomatoes.
Number Larvae Damaged Fruit Per Plant On*
Per Plant Upper Canopy Lower Canopy Total Damage % Fruit Loss Per Plant
1
0.70
+
0.92
2.30
+
1.4913
3.00
10.5
2
1.15
+
1.26
2.80
+
1.85b
3.95
10.5
4
1.05
+
1.23
3.10
+
2.61a
4.15
9.00
8
0.80
+
1.15
4.60
+
2.39a
5.40
19.00
12
1.15
+
0.93
3.10
+
2.73a
4.25
35.00
14
1.50
+
1.67
4.60
+
3.16a
6.10
40.00
* *
Control
0.35
+
0.488
0.90
+
1.31C
_
_
Values within a column followed by the same letter are not significantly different according
to Duncan's Multiple Range Test (P=0.05).
* *
Control sprayed weekly with fenvalerate; mean number from treatments was subtracted from
infestation that occurred in the untreated control without insecticides.
124

Table 29. Marketable value for tomato fruit damaged in the lower and upper plant canopy
after a single artificial larval infestation of K. lycopersicella on ground
tomatoes.
Number Larvae
Per Plant
Value of Damaged Fruit
Upper Canopy
*
Per Plant On
Lower Canopy
Total Damage
1
1.04°
7.06b
8.10c
ab
_ b
cd
2
1.80
5.64
7.44
ab
b
c
4
1.35
6.63
7.98
ab
a
b
8
1.65
11.57
13.23
a
a
ab
12
2.17
12.09
14.26
a
a
a
14
2.71
15.16
17.87
Control
0.55C
3.08°
4.63C
k
Average value of damaged fruit based on U.S.D.A. size-grade standards: extra large=
5, large=4, medium=3, small=2, very small=l, unit values used in the analysis.
â– k k
Values followed by the same letter are not significantly different according to
Duncan's Multiple Range Test (P=0.05).
125

Table 30. Tomato fruit damaged in the lower and upper plant canopy after a double
artificial infestation of K. lycopersice11a larvae on ground tomatoes.
Number Larvae Damaged Fruit Per Plant
Per Plant Upper Canopy Lower Canopy Total Damage % Fruit Loss
1
0.63
+
0.92b
2.13
+
C'k k
2.94C
2.76
22.45
2
0.50
+
b
0.82
2.73
+
be
2.40
3.23
27.86
4
0.70
+
0.95b
3.40
+
3.72abc
4.10
27.47
8
0.73
+
1.14b
3.23
+
. n . abc
4.14
3.96
41.28
12
1.10
+
1.12b
4.46
+
2.66a
5.56
44.45
14
1.80
+
1.47a
4.40
+
2.19ab
6.20
49.00
k
Control
0.35
+
0.48C
0.90
+
d
1.31
1.25
-
*
Damaged fruit from an absolute control with insecticide application; results
obtained from the treatments were subtracted from the results of the control
without insecticides.
•k k
Numbers within a column followed by the same letter are not significantly different
according to Duncan's Multiple Range Test (P=0.05).
126

127
differences (P=0.05) in the lower canopy observed between 1 larva in¬
festing the plant and the 2-14 larval infestation levels. Infesta¬
tion with 14 larvae per plant resulted in 2.24 times more damage than 1
larva. Marketable values of fruits on each canopy were significantly
different from the control (Table 31). Values were not significantly
different among the treatment levels.
In explanation of these results, successive generations of TPW
may increase damage per fruit. Since the damage index is the number
of fruit damaged per plant, the number of injuries produced per fruit
is not accounted for. Another explanation could be that competition
among larvae may displace the second larval infestation to the foliar
canopy, reducing injury to the fruit.
Relationship Between Leaf and Fruit Injury and Larval Infestation Levels
Single infestation. Factors that relate to the estimation of larval
presence were examined: the validity of using leaf injury and the fruit
injured respective to the number of larvae per plant, and the use of
foliar injury to detect fruit damage. A curvilinear and linear regres¬
sion equation indicating leaf and fruit injury expressed in function of
the number of larvae per plant as an independent variable is shown in
Figures 11-12. The relationship between leaf injuries found in the
lower canopy (Yn) and the number of larvae (x) per plant was best des¬
cribed by the following highly significant (P<0.009) linear regression
(y=2.3 + 0.41x, r2=0.511) (Fig. 11). The relationship between
fruit injured in the lower canopy (F^)_ and the number of larvae

Table 31. Marketable value for the tomato fruit damaged in the lower and upper plant
canopy after a double infestation of K. lycopersicella on ground tomatoes.
Number Larvae
Value of Damaged Fruit
★
Per Plant On
Per Plant
Upper Canopy
Lower Canopy
Total Damage
1
1.22ab**
5.60ab
6.83ab
2
1.34ab
7.78ab
9.13ab
4
1.33ab
8.39ab
9.73^
8
ab
1.17
8.93ab
10.11ab
12
2.32ab
11.IIa
13.43a
14
3.06a
10.23a
13.30a
Control
0.55b
3.08b
3.64b
*
Average damaged fruit as estimated from commercial value, extra large fruit=5,
large=4, medium=3, small=2, very small=l, market values used in this analysis.
* *
Values followed by the same letter are not significantly different according
to Duncan's Multiple Range Test (P=0.05).
128

Figure 11. Relationship between number of tomato pinworm larvae per
plant and number of injured fruits and leaves in the lower plant canopy
by a single artificial infestation with TRW harvae. Homestead, Florida,
1980-1981.

Number T PW Larvae
= 2.30+ 0.4 IX rl 0.511 RlO.46
Fruits Injured
Low e r Ca nop y
CD rsj cn *si
• • • •
CD l/l CD * n
130

Figure. 12. Relationship between number of tomato pinworm larvae per
plant and number of injured fruits and leaves in the upper plant canopy
by a single infestation of TPW larvae. Homestead, Florida, 1980-81.

Number TPW larvae
o
00-
IS5
Leaves injured upper canopy
NO on
cn cn
1 i i
O
X
ro
Fruits injured upper canopy
ro
In
cn
i
cn
j
j.
*0
N/
71
©
i—*
On
*T1
T1
â– i
c
N3
•1
Co
©
bo
Cn
r
©
©
to
i—»
N
©
X
CO
CO
£>
©
©
NO
X
N?
132

133
(x) was best expressed by the following significant (P<0.03) linear
2 2
regression. The r value (r =0.37) was less than intermediate. The
correlation coefficients for the lower canopy were intermediate, indi¬
cating that 51-37% of the variation in foliar and fruit injuries was
due to larval numbers infesting plants.
When data from the upper canopy were regressed against the number
of larvae per plant, curvilinear regression had a better fit than linear
regression. 'There was no significant relationship found between injuries
in the upper canopy (Y ) and number of larvae (x) infesting the
u
plant (Fig. 12). Again, no significant relationship was found between
injured fruits in the upper canopy (F ) and the number of larvae per
plant. The lack of significant relationship between larval numbers and
TPW injuries in the upper canopy indicate that upper canopy counts can
not explain number of larvae present in the plant at TR^ stage.
The relationship between fruits injured in the lower canopy (F^) and
foliar injuries in the lower canopy (Y ) was best expressed by a signifi-
1
2 2
cant (P=0.024) quadratic equation F^ =1.09 + 0.46Y^-0.02Y^ and r =0.56.
(Fig. 13). F^ began to decrease at a level of about 7.5 injuries per
plant. This may be an artifact, or it may be that beyond this level
of larval infestation of leaves, multiple injuries to fruit may be
more common than infestation of undamaged ones. Regression analysis of
numbers of fruit injured in the upper canopy and foliar injuries in the
same plant part did not indicate a significant relationship (Fig. 14).
Also, regression of fruit injured on lower canopy and the total plant
injuries was not significant. Therefore, TPW sampling by scouts is
probably best done in the lower plant canopy.

Figure 13. Relationship between number of leaves injured in upper and lower canopy and number
of fruits injured in upper and lower canopy by a single artificial infestation with TPW larvae.
Homestead, Florida, 1980-81.

Fruits injured Lower canopy
sei

Figure 14. Relationship between number of larvae per plant
of injured fruits and leaves (upper and lower canopy) after
artificial infestation with TPW larvae. Homestead, Florida
and number
a double
1980-81.

Number TPW larvae
Leaves injured lower canopy
N3
Cn
Cn
tn
Yl=2.06 + 0.91x-0.02 x
Fruits injured lower canopy
N3 CJl o
ui
137

138
Relationship Between Leaf, Fruit Injury and Larval Infestation Levels
Double infestation. The relationship between leaves injured in the
lower canopy (Y ) and number of TPW larvae (x) was best fitted to the
highly significant (P=0.0006) curvilinear regression, Y^=2.06+0.91x -
0.02x2, F=18.69, r2=0.805 (Fig. 14). The relationship between fruit
injured in the lower canopy (F^) and the number of larvae (x) was not sig¬
nificant (F=l.32, P=0.31). When data from upper canopy were regressed
against number of larvae (two-time infestation) per plant, there was no
significant regression found between injuries in the upper canopy (Y ) and
number of larvae (Fig. 14)_. No significant relationship was found between
injured fruits in the UDper canopy (F ) and number of larvae per
u
plant.
The relationship between fruits injured in the lower canopy and foliar
injuries in the lower canopy was not significant, F=1.21, P=0.341,
2
r =0.212 (Fig. 15). The regression analysis between fruits injured in
the upper canopy (F ) and leaves injured in the upper canopy (Y ) was
u u
not significant, F=0.98, P=0.33, r2=0.04 (Fig. 15). The reasons why
there was no relationship between fruits injured and foliar injuries
caused by a double TPW infestation may be due to an artifact, but it may
be that a double TPW larval infestation causes more injuries to the same
fruit, which will result in no increment of the total fruit damaged per
plant.
Yield Loss vs Density of TPW Larvae in Tomatoes
The second order model y=a+bx+cx , (y=percent of yield loss, x=
number of TPW larvae per plant), was fitted to the data on mean percent

Figure 15. Relationship between number of leaves injured in upper and lower canopy and number
of fruits injured in upper and lower canopy by a double artificial infestation with TPW larvae.
Homestead, Florida, 1980-81.

Fruits injured lower canopy
Ot'T

Number TPW Larvae
Leaves Injured Upper Canopy
Y: 2.47 0 3 9x
r2,0.29 R4.15
2.5 J
Fruits Injured Upper Canopy
p
(/>
141

142
yield losses for 3 tomato plantings attacked by TPW. The relationship
2
was highly significant, P=0.001 and had an intermediate r value of 0.64
(Fig. 16). The coefficient estimates b and c were significant. A
positive increment in yield losses was observed until 12 larvae were
infesting the plant; in contrast, beyond this infestation level, the
percentage of yield losses decreased.
Yield Losses vs Number of Plant Injuries
2
A significant curvilinear (y=8.76+4.21x - 0.14x ) regression was
found when mean percent yield losses combined for 3 plantings were
plotted vs TPW injuries per plant (P=0.001) (Fig. 17) for numbers of
foliar injuries up to 27 per plant. Regression analysis indicated a
positive relationship between yield losses and 10-15 injuries per plant.
The relationship turned negative when more than 15 injuries are found
2
per plant. Based on r value, the fitness of the model was intermediate
2
(r =0.608), but higher than that found with a simple linear regression
model (r^=0.322).
Actual yield losses may vary, depending on time of planting and
southern Florida environmental conditions. A more robust model relating
density and yield losses should be considered with lower larval infes¬
tation levels per plant. The models developed here provide
valuable information on the TPW-plant interactions. TPW larvae may bore
into the fruit for different reasons, yet high attraction to the fruit
was not found by Swank (1937). Thus, larval density, contact between
injured leaves and fruits, or positive geotaxis observed when the larva
suspends itself by the thread produced from the spineret, may account
for the fruit damage especially on the lower canopy.

Figure 16. Regression of percent of yield reduction against infestation
densities per plant of tomato pinworm larvae.

144
e
ro
â– a
oc
â– a
a<
>â– 
TPW Larval Density

Figure 17. Regression of percent yield reduction of tomato fruit against TPW number of foliar
injuries per plant.

146

147
Estimates of economic injury levels for TPW larvae may be made by
inspecting the equations describing the relationship between fruit damage
and larval density. The high market price of the tomato crop and cost of
controlling TPW larvae indicate again that infestation levels lower than
1 larva per plant should be used. The minimum economic injury level
damage determined for green cloverworm in soybeans by Stone and Pedigo
(1972) and modified by Hopkins et al. (1982) and Hall and Teetes (1982)
is defined as EIL= pest/ha, p=price or market value of the crop per ha, and b=the regression
value from the regression equation used. Gain threshold (Stone and Pedigo
1982) = (control cost/market value)x 100. For example, if the cost of con
trolling TPW is $50 per ha, and the market crop value of tomato is
$6629.5/ha, the percent gain threshold would be 0.0075x100=0.754. The
economic injury level for an infestation of TPW could be calculated by
using the regression coefficient (b=1.67) from the linear regression
equation y=15.08 +1.67 x; r2=0.45; F=17.72 P>F=0.004: EIL=^0.75/1.67 =
0.67 tomato pinworm larvae per plant. Similarly, the economic injury
level based on the population index (number of injuries per plant) can be
calculated using the regression coefficient (b=1.07) from the linear regres
sion equation y=16.82 + 1.07x; r2=0.32; EIL= Jo.75/0.07 = 0.83 tomato
pinworm larval injuries per plant.
There are constraints for these economic injury levels. They are
based on results from one phenological stage (TR^)- Therefore, it is
not known if they can be used for earlier stages (TV^ - TR^). Planting
time will also affect these results as demonstrated in the next
experiment.

148
Second Experiment
Effect of planting time on yield. Earliest plantings (October, 1980)
had the lowest TPW fruit damage (Table 32). Significant differences in
number of fruit damaged per plant were found between those crops planted
during October-December and crops planted during January-February. Largest
infestations occurred in the latest planting (February, 1981). The amount
of TPW damage in southern Florida depends on planting date with the
greatest threat during the winter-spring crops. Planting time might be
considered as part of an integrated control program for TPW. These re¬
sults call for further research on economic injury levels related to
planting time. For instance, TPW population peaks occur during March-
May (Chapter 7). At this time, those crops planted during the fall have
already been harvested or are in the second reproductive stage (TR^). The
use of an earlier planting date will be a common sense approach to pest
avoidance.
Effect of plant pruning and number of plants per row related to
fruit damage. Pruning plants had no effect on fruit damage (Table 33).
Trends in the data suggest that further studies should be done with more
levels of pruning in the experimental design. Differences in relative
net precision using 6 contiguous plants/row opposed to random plants per
row did not show any statistical difference (Table 34). Number of total
fruits per row in those plants selected at random was slightly higher
but did not lead to better RNP values. Therefore either method can be
used for selecting tomato plant for TPW assessment.

Table 32. Effect of planting time on fruit injured by K. lycopersicella larvae to
ground tomatoes, cv 'Flora-Dade1 during 1981.
Planting
Time
*
Total Fruit Damaged/Plant
Average Leaf Injuries/Plant
Oct.
30,
1980
0.667b**
0.6
Nov.
25,
1980
3.154b
2.52
Dec.
30,
1980
5.11b
8.35
Jan.
30,
1981
13.35a
10.80
Feb.
28,
1981
16.95a
12.57
*
Plantings without insecticide protection 20 days after emergence.
* *
Values followed by the same letter are not significantly different according to
Duncan's Multiple Range Test (P=0.05) .
149

Table 33. Differences in cost and relative net precision between sampling 6 plants per
row and 1 random plant per row.
Plants Per Row
N
Injuries Per Plant
★ *
Time
RNV
Total Fruit/Plant
6 plants
18
1.27 +
1.59
15
19.361
1 plant
18
1.33 + 0.40
2.38
12.34
25.08
*
Relative net
precision
= 100
RV x Cs
* *
Time expressed in minutes expended in detecting foliar injuries and walking from
1 plant to another.
^Numbers followed by the same letter are not significantly different according to
Student's t-test (P=0.05).
150

Table 34. Differences in mean fruit injured by K. lycopersicella in pruned and not pruned
tomato plants.
Treatment
Total Fruits/Plant
Injured Fruits
% Injured Fruits
Without pruning
26
a
6.90
26.95
Pruned
28
4.70
17.09
Mean values followed by the same letter are not significantly different according to
t-test (P=0.05).
a
151

152
Conclusions and General Discussion
The information generated in this study is useful to demonstrate
the complex effect of different TPW densities on tomato plants. First,
a single infestation of 1, 2, 4, 8, 12 and 14 larvae per plant resulted
in a 10.50 - 40% fruit loss. The result contrasted with 22.5 - 49% fruit
loss obtained when larval levels were doubled. This demonstrated that
only one generation of TPW is necessary to cause severe (40%) losses,
and also, data suggest that in spite of low larval levels (1 larva/plant),
yield losses are twice as much if two generations occur.
Second, regression analysis between number of fruit injured and
number of TPW larvae demonstrated the importance of plant stratum
selection for sampling TPW. Generally, the number of TPW larvae (x)
2
was better related (r =0.51) to the number of leaves injured in the
lower canopy than to the number of leaves injured in the upper canopy
2
(r =0.38). Also, number of leaves injured in the lower canopy (x)
2
and fruits injured in the lower canopy were better related (r =0.56)
than leaves and fruits in the upper canopy. These results were
expected because higher infestations occurred on middle and lower
canopies where most of the tomato fruit is located.
The relationship between number of TPW larvae per plant (x)
and percent yield loss was intermediate (r“=0.64). Data from a
simple linear regression (r =0.45) was used to determine EIL. EIL for
TPW was 0.67 larvae per plant. The relationship between number of
injuries per plant (x) and percent yield losses was fitted to a signi-
2
ficant curvilinear regression (y=8.76 + 4.21x - 0.14x ). The fitness
2
of the model was intermediate (r =0.608). Data from a simple linear

153
2
regression (r =0.322) was used to determine EIL based on the damage
index. The EIL was 0.83 TPW larval injuries per plant.
There are, however, some constraints for these economic injury-
levels. The EIL is based on results from one phenological stage
(TR2); therefore, it is not known if they apply to other stages (TV
TR^). Planting time will affect these results. Results concerning
effect of planting time suggest once again that the southern Florida
farmers should avoid planting during the later winter season because
of higher TFW population peaks during March-May.

CHAPTER VI
ADULT DISPERSION AND COLONIZATION OF TOMATO FIELDS
3Y THE TOMATO PINWORM
Introduction
Dispersion and field colonization should be among the first factors
considered with regard to insect distributional patterns (Price 1976).
Knowledge of spatial patterns is considered basic to design of appro¬
priate sampling plans and provides insight into the biology of the
species in question (Shepard and earner 1976).
Colonization patterns of members of the family Gelechiidae have
been studied mainly for the pink bollworm (Pectinophora gossypiella
Saunders). The Gelechiidae are considered weak fliers whose patterns of
dispersion are strongly influenced by wind, patchiness of the environ¬
ment, and also voltinism (Shelton and Wyman 1979a,b; Kaae et al. 1977;
Stern 1979).
Little has been done on the study of the factors affecting patterns
of dispersion of the TPW in commercial fields. It has been observed
that a common feature of TPW is its aggregation at the edges of fields
(V.H. Waddill, personal communication). The purpose of this study was
to determine the colonization pattern of TPW and to define the effect of
edges and hedgerows on the insect's distribution.
154

155
Materials and Methods
Dispersal of TPW Male Moths in the Field
This study was conducted during February through May, 1980, and
May-June, 1981, in order to determine short range pinworm dispersal
within the field. TPW was monitored by placing Pherecon lc® sticky
traps 60 cm above the ground and baiting them with sex attractant
(95.3%-(E)-4-tridecen-l-01 Acetate; 4.5%-(Z)-4-Tridecen-l-01 Acetate)
(personal communication, R. Heath, USDA, Gainesville, Fla.). In all
0.5-3 ha tomato fields of this study, three traps were placed in each of
the major compass directions (north, west, east, and south); the trap
lines and distances between them were 12, 60, and 300 m from the center
of the first field, and 12, 30, and 48 m for the remaining ones.
Trapped TPW male moths were recorded and removed daily from each trap
during February-March and April-May. This survey determined male moth
activity from the border to the center of the field.
Colonization Pattern of Tomato Fields by TPW
Tomato pinworm egg distribution can be considered an indicator of
female moth activity in a field. Because of low TPW larval motility
within the fields, immature stages were also taken as an index of popu¬
lation dispersion and colonization in the tomato fields. Four fields
ca. 1-3 ha located near Homestead, Florida were the study sites. The
fields had been under cultivation for several years in a corn- tomatoes-
bean- or squash rotation. No insecticides were applied during the

156
sampling time. Three sampling sites were located at rows 0, 30, and 120
m from the edge to the center of the field. Twenty randomly selected
plants from each row site were selected weekly, until the farming
practices (tillage, discing, or insecticide application) interfered
with sampling. The sampling unit used was larval injuries per plant.
Data from each field were analyzed by analysis of variance and means
from each site were separated by Duncan's Multiple Range Test (P=0.05).
Effect of Edges and Hedgerows of TPW Distribution
Three post-harvested tomato fields abandoned after the 2nd com¬
mercial harvest were sampled on May 4-22, 1981. Field sizes fluctuated
2
between 0.88-1.32 ha. These were divided into (ca. 1100 m ) quadrats.
Two middle leaves per plant were taken as the sampling unit from 20
randomly selected plants per quadrat, to detect the presence of TPW
larval injury. The first field was bordered to the northwest by mango
trees (Mangifera indica L.), to the north and northeast by tomatoes, and
to the south by a main road and grassland. The second field was bor¬
dered to the northwest by grassland, to the south by a hedgerow or
windfall of Casuarina equisetifolia Forst, and to the east and north by a
com field and a main road, respectively. The third field was separated
to the north and south by a windfall of C. equisetifolia, and bordered
to the east by a sweet potato (Ipomoea batata (L.)) field, and to the
west by a main road and grassland. For each field, mean numbers of TPW
foliar injuries were averaged per quadrat. Each quadrat was considered
a treatment. Differences among quadrats were defined by use of analysis
of variance and statistical differences were determined by use of
Duncan’s Multiple Range Test (P=0.05).

157
Results and Discussion
Dispersal of TPW Male Moths in the Field
The results presented in Fig. 18 reveal a pattern of K. lycopersi-
cella male distribution which indicate that males are mainly found in
the borders of the field. Data indicate male distribution only, al¬
though I hypothesize (without evidence) that female distribution follows
the same pattern. Fewer males (10-50) were captured during the early
part of the tomato growing season, February-March, than later, May-June
(75-395). The data probably reflect several successive generations
being produced in the field. During February-March, 1980, males were
more abundant (62%) in traps located farther from the center of the
field (300 m) than in those located closer (12-26%). The same negative
trend toward the center of the field (17-42%) was observed during May-
June, 1980-81. The predominant wind direction during February-March was
NE, and the proportion of moths captured corresponded to those traps
oriented N, W, E, and S, respectively. The behavior of TPW males in¬
dicated that there may be a dispersal tendency toward areas close to the
edges of the field. If the trapped moths were not part of the natal
population, data may indicate that TPW is a good colonizer which con¬
centrates mainly in the field edges, having a slower dispersion within
the field. Perhaps the migratory TPW population initiates a fast field
colonization at the borders, but later generations colonize the field
slower, since there is no need to cover long distances inside the tomato
field if food is available.

Figure 18. Abundance of tomato pinworm male moths at three different field sites.
A) Field 1, 1980; B) Field 2, 1980; C) Field 3, 1981. Mean number of moths at each
site corresponds to the average from 4 pheromone traps in north, south, west, and east
directions, Homestead, Florida, 1980-1981.

Moths captured
A
Days of capture

B
T3
V
i-
3
a
«
U
in
X
O
£
250
12 14 16 23
8
Ma y
Days of capture
• o
Field 2,1980
_ JO
~k
48
m from
field
Center
30
I»
H
I»
1»
12
1»
n
If
If
160

Moths Captured
c
Field 3,1981
8
May
10 . 17
June
20
12
30
48
m from field Center
Days of capture
161

162
Colonization of Tomato Fields by TPW
The results obtained in this experiment (Fig. 19) demonstrated
statistical differences (P=0.05) in number of injuries per plant during
12 of the 17 sampling dates among the border and rows located 30 and 120
m from the edges of the 4 fields sampled. The first field, sampled from
January-April, had a lower infestation (less than 5 injuries/plant) than
the others. The border of the field had the highest counts (15-19) and
did not increase sharply toward inner areas until April, 1980. The
second field, sampled in April, had higher counts (4-10 injuries/plant)
at the border, than at 30 m (1-4 injuries) or at 120 m (0.5-1 injuries)
from the border. The third field, sampled from February-April, showed
a larger increment toward the inner parts of the field during the first
2 weeks of sampling and remained as high as the border for the subse¬
quent sampling dates. However, the infestation obtained at the border
(1.5-3 injuries/plant) was significantly higher than that obtained in
the inner areas (0-2.5). There are several factors that may influence
these patterns, such as plant growth and temperature. Also, these data
may reflect different behavioral characteristics of different TPW genera¬
tions. I hypothesize (again without evidence) that moths which colonize
the field settle on field edges for several reasons (wind drift, etc.).
The next generations produced on each field move inside the field
depending on depletion of resources by previous larvae. The faster or

Figure 19. Mean number of tomato pinworm larval .injuries occurring in 4 com¬
mercial fields at 0, 30, and 120 m from the field border. Bars with the letter
"a" denote statistical differences at 0.05% difference level for a particular
date.

Mean Injuries /Plant
Site 3
-P-
Feb. 8
f | 0 m from border
mini 30 -
Murió Apr.9
Sampling Dales
164

165
slower colonization of tomato fields will reflect the density of the 1st
TPW generation.
Experiment 2
The combination of data from the three fields sampled during 1981
enabled a fair analysis to be made of the importance of edgerows and
hedges in the process of any TPW infestation (Table 35, Fig. 20).
There were three broad edge categories: 1) hedgerows or windfalls, 2)
road or pastures adjacent, and 3) edge surrounded by the same crop or
other crops (com, sweet potato) . In general, TPW larvae showed maximum
density near windfalls. Also, density increased in different parts of
the fields surrounded by grassland and separated by a road. Two fields
had the highest population at the eastern edge of the field. Hedgerows
had the highest infestation (3.2-6.25 injuries/plant) compared to road
edges (1.20-4.10) and other crops or tomatoes as edge (0.33-4.15).
Edges surrounded by pastures had a similar infestation range (4.47-5.82)
as those surrounded by hedgerows.
The presence of hedges around fields probably increases the chances
of cumulative infestation by TPW. Because of lack of sheltering vegeta¬
tion, edges surrounded by pastures and roads are probably the entrance
for insects from neighboring fields. Lewis (1969) suggested, also, that
the pattern followed by flying insects depends on a process of uniform
delivery followed by differential removal from the air above different
parts of the field. Since TPW larvae are slow movers within the
field, the measure of immature insect stages could be taken as a result
of adult dispersal. The results in this study again support the idea of

166
Table 35. Effect of hedges and edgerows on tomato pinworm field in¬
festation at three fields in Homestead, Florida, 1981.
Edge Category
Field 1*
Field 2
Field 3
Hedgerow
6.25a**
4.33ab
4.90a
5.00a
4.00b
3.85a
4.65a
—
3.20bc
Road
2.35b
4.58ab
4.10a
0.60bc
0.20c
3.40ab
Pastures
—
5.82a
—
—
4.47ab
—
Other crop or tomatoes
1.33b
0.46c
4.15a
0.95bc
0.33c
1.70bc
0.35c
—
1.45c
â– k
Mean values of tomato pinworm injuries per plant. Values on respective
quadrat adjacent to the edge specified.
* * ....
Values with the same letter within a column are not significantly dif-
ferent at the 0.05% level.

Figure 20. Field plan: Fields 1, 2, and 3, respectively, each field was
divided into 8-9 quadrats. The shaded areas represent higher insect
populations. Homestead, Florida, 1981.

168

169
influence of edges on dispersion of TPW adults and that aggregation of
TPW is also a function of density. TPW gregariousness was evident in
resource exploitation.
Conclusions and General Discussion
In this research useful data were gathered about the pattern of
colonization and dispersion of TPW in tomato fields, as well as data
about the effects of edges and hedgerows on TPW population dispersion.
TPW adult male moths were generally (62%) more abundant on borders than
in inner regions of tomato fields (12.4%). TPW foliar injuries were
generally significantly higher at the field borders than at 30 and 120 m
from the border of the field. The infestation rate progressed toward
inner areas depending on initial infestation of the border and sampling
date. For instance, fields infested during January had lower infesta¬
tion than those sampled during March to May.
Three fields showed significantly higher populations on edges
surrounded by windfalls, roads, or pastures, and lower populations on
edges surrounded by the same crop or other crops.
The accumulation of TPW moths, eggs, and larval injuries on field
edges indicate the importance of intensifying sampling in those areas in
order to apply control measures to stop infestation progress. There are
still many questions unsolved about TPW dispersal patterns. One of them
is: Do these moths have a similar dispersal pattern to the one found
for P_. gossypieila by Flick and Noble (1961)? It is possible that
TPW generations are divided into colonizers, those that migrate at

170
higher altitude locating new food sources, and local individuals whose
movement inside fields is characterized by a low altitude and erratic
flight.

CHAPTER VII
EGG AND LARVAL PARASITISM OF TOMATO PINWORM IN SOUTHERN FLORIDA
Introduction
The tomato pinworm (TPW), Keiferia lycopersicella (Walsingham) has
been a serious pest of tomatoes in southern California (Oatman 1970) ,
Texas (Wellik et al. 1979) and in Florida (Poe 1974). In southern
Florida, TPW is a more serious pest during spring when populations build
up as the crop matures. Tomato pinworm larval parasites have been
studied by Cardona and Oatman (1971) , and seasonal occurrence of para¬
sites has been investigated in southern California by Oatman et al.
(1979). Hymenopterous larval parasites attacking TPW in Florida are
Apanteles spp. (Poe 1974b) , Temelucha spp., Sympiesis stigmatipennis
Girault, Zagrammosoma multilineatum (Ashmead), and Parahornius palli-
dipes Ashmead (Krombreii et al. 1979). The only egg parasites reported
have been Trichogramma spp. in southern California (Oatman et al. 1979)
and in Colombia attacking the South American pinworm Scrobipalpula
absoluta (Meyr.) (Garcia et al. 1974).
Reported here are investigations of 1) parasitism of TPW larvae in
tomato fields during 1980-81; 2) the parasitism of TPW eggs in the
laboratory by the naturally-occurring T. pretiosum (Homestead strain)
and by a laboratory reared strain of T. pretiosum (Texas strain); 3) the
effect of host distribution and host density on the parasitization by
171

172
Trichogramma spp.; and 4) the seasonal occurrence of the Trichogramma
spp. in several tomato fields.
Materials and Methods
Larval Parasitization
Twelve to fifteen commercial fields were surveyed for larval
parasitism from January 14 through September 5, 1980 and from January 23
through July, 1981. Mined leaves were examined in the fields and taken
to the laboratory. Occupied TPW mines were held for adult parasite or
moth emergence in an ice cream carton which had about 5 g of white sand
in the bottom. No additional food was provided. Therefore, most early
TPW instars probably died of starvation. Hosts and parasites were
recorded and identified. Percent parasitism was calculated from the
numbers of adult moths and parasites which emerged.
Egg Parasitism
Laboratory studies. The parasitism of tomato pinworm eggs by 2
strains of T. pretiosum was studied in a laboratory at ca. 25°C, 75 +_
2 RH, and scotophase of 11 h. 'Flora-Dade' tomato plants were exposed
to oviposition by TPW adults in an insectary. Eight plants with a total
of 405 TPW eggs were exposed to ca. 300 females of T. pretiosum (Texas
strain) in a cage (24 x 24 x 24 cm) for 24 h. At the same time, excised
leaves with a total of 51 eggs were placed in 6 petri dishes (100 x 25 mm)
which contained moist filter paper and ca. 3 field-collected females of
T. pretiosum (Homestead strain) per dish. Eggs were removed after 24 h.
exposure and placed (1 egg per dish) in smaller petri dishes (50 x 9 mm).

173
The eggs were examined initially to determine whether they were para¬
sitized and then observed daily for parasite adult emergence.
Field studies. Three tests were conducted from May to July, 1981
in abandoned tomato fields near Homestead, Florida, to determine ef¬
fectiveness of naturally occurring Trichogramma spp. In the first test,
I studied the spatial distribution on the plant of both parasitized and
nonparasitized TPW eggs. Only 2 leaves from the upper 1/3 and middle 1/3
portions of the plant were sampled since preliminary sampling indicated
few eggs were present on the lower leaves. Fifteen plants were collected
randomly from a field which had an avg. of 35.2 eggs/plant. Possible
differences in egg density and percentage of parasitism at each level
were determined by a student's t-test.
The second test evaluated the effect of host distribution on the
level of parasitism, and the relationship between host density and
percent parasitism. Two fields were selected as sites for this study:
(Field 1) had an avg. of 3.10 eggs per 2 leaves per plant and (Field 2)
had an avg. of 0.70 eggs per 2 leaves per plant. The fields were di¬
vided into 3 (2 border and 1 middle) sections lengthwise of 0.44 ha
each. Two leaves were collected from the middle portions of 80 plants
randomly selected per section. Egg parasitism was determined in the
laboratory by examining the eggs under a dissecting microscope. Fre¬
quency distributions of parasitized and nonparasitized TPW eggs were
compared with 3 types of distribution (Poisson, Negative Binomial, and
2
Positive Binomial) and tested by x for goodness of fit. Data were
transformed to log (x+1) prior analysis. Differences in egg density
among sections of each field were determined by the use of Duncan's

174
Multiple Range Test. Correlation between egg density and parasitism was
analyzed following arc sine transformations.
The third test was conducted to determine the extent of TPW egg
paraby T. pretiosum in Dade County. Fourteen tomato fields were sampled
weekly by collecting 2 middle leaves from 10 randomly selected plants
per field. Six of the fields were located in the northern part of the
farming area and the rest in the southern portion. Samples were taken
to the laboratory to determine degree of parasitism. The survey was
suspended as the plantings were destroyed by discing or mowing.
Results and Discussion
Larval Parasitism
Parasitization of tomato pinworm larvae ranged from an avg. of
39.34% in 1980 to 46.26% in 1981. During the study, parasitization
averaged ca. 2.5, 22, 51, 40, and 48.27% in January, February, March,
April, and May, 1980, respectively. Parasitization averaged ca. 49, 46,
53, and 46% in April, May, June, and July, 1981, respectively. Although
parasitism ranging from 40-60% was common during 1980-81, there was not
a consistent corresponding increase in parasitism with an increase in
host density (Table 36).
Of 3 primary parasites reared from tomato pinworm during 1980,
Apanteles spp. was the most important, parasitizing 10-66.66% of the
larvae. Temelucha spp. and Sympiesis stigmatipennis were the next most
abundant parasitoids. During 1981, (Table 37) 4 parasites were reared.
Apanteles spp. was again the most important, parasitizing 46% of the
larvae, followed by Parahormius pallidipes, Sympiesis stigmatipennis and
Chelonus phthorimae in order of importance.

Table 36.
Parasitism of the tomato
County, 1980.
pinworm larvae in
tomato fields in southern
Florida, Dade
Survey
Total
% Total
%Parasitized by
Week
Collected
Parasitized
Apanteles spp.
Sympiesis stigmatipennis
Temelucha spp.
Jan. 14
25
0
22
25
0
31
25
10
10
Feb. 6
46
3.57
0
0
3.57
13
20
33.33
33.33
0
0
19
30
33.00
33.00
0
0
25
25
17.39
17.39
0
0
Mar. 4
35
10.52
10.52
0
0
12
44
62.50
62.50
0
0
19
45
66.66
66.66
0
0
26
25
62.53
58.33
4.20
0
Apr. 2
50
48.26
41.37
6.89
0
7
30
49.07
45.45
1.81
1.81
16
78
38.77
29.52
5.55
3.70
175

Table 36--continued.
Apr. 3
10
60
60
9
5
40
40
22
27
47
47
May 1
15
35
35
7
25
40
40
14
31
45
22.43
26
155
65.16
63.87
Jun. 2
28
46.42
42.85
16
109
56.88
55.96
25
65
56.92
55.38
July 1
23
56.00
50.00
8
100
73
72.00
17
34
10
9.00
Total
627
Avg.
42.26
46.03
0
0
0
0 0
0
0 0
0
0 0
0
0 0
0
0 0
3.22
19.35 0
0.64
0 0.64
f—1
0
3.56 0
0.91
0 0
1.53
0 0
6.00
0 0
1.00
0 0
1.00
0 0
0.62
0.99 0.027
0.62
0.027
176

Table 37. Parasitism of the tomato pinworm larvae in tomato fields in southern Florida, Dade
County, 1981.
Survey Total % Total %Parasitized by
Week Collected Parasitized Apanteles spp. S^. stigmatipennis Temelucha spp.
Apr.
23
67
32.38
22.53
2.81
7.04
30
60
34.7
24.39
0.00
7.31
May
7
130
49.31
42.74
0.00
6.87
13
91
51.3
44.73
0.00
6.57
20
130
30.41
29.41
0.00
1.00
Jun.
2
72
62.06
58.62
3.44
0.00
July
4
47
70.00
70.00
0.00
0.00
Aug.
9
40
72.50
72.50
0.00
0.00
Sep.
5
3
66.66
66.66
0.00
0.00
Total
847
Avg.
39.34
37.50
1.07
1.64
177

178
Egg Parasitism
Laboratory studies. The results of the laboratory experiment
(Table 38) demonstrated that both strains of T. pretiosum can oviposit
and develop in TPW eggs. Although the percent parasitism and parasite
emergence from eggs parasitized by the Texas strain were 1.2 times
greater and 71% less, respectively, than from eggs parasitized by the
Homestead strain, these differences may have been due to differences in
egg density and searching area rather than differences between strains.
It is known that kairomones play an important role in successful para¬
sitism by Trichogramma (Seabrok 1977), and this could also account for
the observed differences since the Texas strain was reared on Angoumois
grain moth (Sitotroga cerealella) eggs. The sex ratio was 50:50 (males
and females) for the Texas strain and ranged between 50:50-60:40 (males
and females) for the Homestead strain.
Field studies. The mean number of TPW eggs and percent parasitism
were significantly higher on the middle leaves than upper leaves in test
1 (Table 39). When the egg density per plant was regressed on percentage
2
parasitism from both levels, the r“ obtained was 0.0181 which indicates
lack of correlation. This may indicate a complex of hosts other than K.
lycopersicella (including Heliothis zea [Boddie]). It may also in¬
dicate that searching capacity of T. pretiosum is a limiting factor in
parasitism.
The results obtained from the interaction between host distribution
and level of parasitism by T. pretiosum is reported in Table 40. The

179
Table 38. Keiferia lycopersicella eggs parasitism by 2 strains of
Trichogramma pretiosum in the laboratory. T 25+l°C;
75+2% RH.
Parasite No. Eggs
Strain Exposed
Parasitism
X + SE
%
Emergence
X + SE
Days to
Emergence
X + SE
Texas
405
68.93+2.91
27.47+1.58 8.33+0.862
Homestead
51
57.96+5.92
93.10+1.36
8.50+2.67

180
Table 39. Number of Keiferia lycopersicella eggs collected from two
strata and percent of parasitism by Trichogramma pretiosum.
Leaf
Location
Total
Eggs
Avg. per
21 Leaves
Range
% of Parasitization
x/21 Leaves Range
Upper
104
6.93 a*
0-19
39.73 a
0-100
Middle
281
18,66 b
0-50
57.80 b
26-100
*
Values followed by different letters are significantly different
(P<0.05) by Students t-test.

181
Table 40. Distribution of normal and parasitized tomato pinworm
eggs in 2 tomato fields.
No. Plants
X
2
s“
k
2
X
Field 1
Total TPW
eggs
80
3.09
12.76
1.43
19.39a
Parasitized
TPW eggs
2.16
5.45
Field 2
1.65
11.97b
Total TPW
eggs
80
0.659
1.35
0.526
3.12b
Parasitized
TPW eggs
0.42
0.74
0.45
5.46b
ax value of 19.39 is not below the 5% point of 19.67 (V=9, a=0.05).
Therefore, the model is not a good fit to the original counts and
agreement with a negative binomial distribution is not accepted at
the 95% probability level,
b 2
X values of 11.97, 3.12 and 5.46 were well below the 5% point of
19.67 (V=9, a=0.05). Agreement with a negative binomial distribu¬
tion accepted at the 95% probability level.

182
distribution of total TPW eggs only fit the negative binomial for the
second field. Parasitized eggs in both firled were fit to the negative
binomial distribution. Thus, there is an aggregative distribution of
TPW eggs and a concomitant distribution of T. pretiosum only when low
egg densities per plant are found. These results agree with Morrison
and Strong (1980). There were significant differences in host density
among the 3 areas of Field 2 but not in Field 1, despite a higher number
of host eggs (x=4.04) (Table 41). The major difference between the
fields was that Field 1 was apparently more completely colonized by TPW
than Field 2. Egg parasitism ranged from 0-55% for the first field and
0-70% for the second field, despite the lack of correlation (Fig. 21 A
2 2
and B) (r =0.026 and r =0.018, respectively) between the percentage of
parasitism and the mean egg density for both fields. There was a
negative trend in parasitism for egg density higher than 3 eggs/2 leaves
(Field 1), and a positive trend when TPW egg density was less than 3
(Field 2). Spatial variations in host density (Morrison and Strong
1980) may result in spatial variations in parasite activity. These
results indicate that the distribution of T. pretiosum followed the same
distribution as the TPW eggs; however, there was not a consistent
increase in percent parasitism with increased host density. Since TPW
eggs are not the only hosts for T. pretiosum, the response of the para¬
site might be the result of spatial differences in the densities of
alternate hosts. The limitation of searching capacity, if a reality,
or the response to kairomones, if any, may also play important roles in
the degree of successful parasitism.
The field survey indicated higher parasitism in the fields located
in the northern part of Dade County (Fig. 22A) compared to the fields

183
Table 41. Parasitism of tomato pinworm eggs by Trichogramma pretiosum
in 2 fields with different host densities.
Area
No. eggs in
sample x of
eggs/2 leaves
% Parasitism
Field Ia
Border
1
363
3.5 a
55
Middle
424
5.0 a
39
Border
2
367
3.5 a
30
Field 2
Border
1
51
0.6 a
58
Middle
22
0.3 a
57
Border
2
91
1.00 b
70
Values followed by different letters are significantly different at
P^0.05, Duncan's Multiple Range Test.

Figure 21 A-B. Relationship of Keiferia lycopersicella egg density to
percent parasitism by Trichogramma pretiosum in 2 fields.

Percentage Pa rasitisrn(arcs¡n)
Field 1
80.
60.
Y=4 9.46-1.23x
4 0
' II# II
2 4 6
Number TPW Eggs
I
8
185

Pe re en ta ge Pa rasi ti s m(arc si n)
►o a» od o
o o o o o
981

Figure 22 A-C. Seasonal occurrence of Keiferia lycopersice11a
parasitization by T. pretiosum in tomato fields located at the
(b) middle, and (c) southern areas of Dade County, Florida.
eggs and
(a) northern,

mean tpw eggs
S AMPLING D AT ES
188

189
located in the southern part (Fig. 22C) . Parasitism of TPW eggs ranged
from 30% in May to 65% in July for the northern fields. In the southern
fields parasitism ranged from 0% in May to 15% in July. The average
parasitism for both areas for the study period was 26.9%. Although
parasitism ranging from 33-73% was not uncommon when density was higher
than 5.2 eggs/2 leaves, there was not a consistent corresponding in¬
crease in parasitism with an increase in host density except for 1 field
surveyed (Fig. 22B) . It is known that the frequency with which a para¬
site finds hosts is a product of several components. Among them, Burnet
(1958) and Hassell (1966) found that the nature of host distribution has
considerable influence on the response of entomophagous insects.
Studies of the incidence of T. pretiosum on other hosts (Heliothis spp.)
should be conducted in tomato fields. This information is needed to
understand better how T. pretiosum can be manipulated for maximum benefit
of the tomato producer.
General Discussion and Conclusions
Data obtained from research on parasitism of TPW eggs and larvae in
southern Florida indicate that Trichogramma pretiosum, as an egg para¬
site, and Apanteles spp., as a larval parasite, are the most abundant
natural enemies. The parasitoid population increased during May-June,
when most southern Florida tomato fields are at post-harvest. This may
also indicate insecticide resistance of the parasite species from
continuous insecticide applications. Further research is suggested on
releases of these parasites on post-harvested fields to reduce TPW
resurgence during the following season.

CHAPTER VIII
EFFECTS OF RAINFALL AND RELATIVE HUMIDITY ON IMMATURE STAGES OF
THE TOMATO PINWORM UNDER GREENHOUSE AND FIELD CONDITIONS
Introduction
Some information concerning the effect of temperature and various
environmental factors on Keiferia lycopersicella (Walsingham), the
tomato pinworm (TPW), is available in the literature (Weinberg and Lange
1980, Poe 1974b, McLaughlin et al. 1979). Little has been published,
however, concerning the study of rainfall and humidity influencing the
rate of population increase of this insect. Effects of rainfall and
humidity on pests of the same family have been studied by Simmons and
Ellington (1933) , Hofmaster (1949) , Warren (1956) , and Clayton and
Henneberry (1982). In general, they found that rainfall and humidity
may reduce or increase abundance of different stages of the Gelechidae
species studied. This paper describes a study of the relationship
between rainfall and survival of the egg, larval, and pupal stages of
TPW the laboratory and in the field.
Materials and Methods
Seasonal Occurrence of TPW in Experimental Plots
To find a possible relationship between temperature and rain¬
fall and egg deposition by the TPW, oviposition was studied on 8
non-staked tomato plantings cv. Flora-Dade (Nov. 3, 1979; Dec. 5, 1979;
Jan. 8, 1980; Oct. 30, 1980; Nov. 25, 1980; Dec. 30, 1980; Jan. 30, 1981;
190

191
Feb. 28, 1981). Tomatoes were seed-planted at the Agricultural Research
and Education Center, University of Florida, Homestead, Florida. Each
planting (ca. 450-947 plants) was set in raised beds (3-5) (ca. 45 m
long) of Rockdale soil and mulched with light colored plastic. Plants
were spaced 38 cm apart. Oviposition per plant was averaged weekly for
3 plantings during 1980 and for 5 in 1981. Each weekly sample consisted
of 10-20 plants selected at random. Temperature and rainfall regimes
during February through May, 1980 and January through May, 1981 were
determined by the Climatological Weather Station at the Agricultural
Research and Education Center, Homestead, Florida.
Seasonal Occurrence of TPW in Southern Florida
Studies were conducted in 12-15 commercial fresh market tomato
plantings in Homestead, Dade County, Florida during 1980-81. The fields
were part of a survey program. Every crop differed in size and prac¬
tices such as insecticide application, but horticultural practices were
similar. Each planting during 1980 was systematically sampled for
larval injuries by inspecting the whole plant. During 1981, I limited
the sampling to inspection of 2 of the middle leaves on each plant. A
total of 20 plants per field was sampled. Each planting was sampled
before and after harvest, and sampling was suspended when the grower
disced or mowed the crop. Surveys continued when new plants emerged in
those fields. The survey during 1980 was concluded during October, when
the fall tomato crop was planted. The survey during 1931 was concluded
in July, 1981. During 1981, pheromone sticky traps (Pherocon lc®)
baited with pheromone (95.3%-(E)-3-Tridecen-l-01 Acetate; 4.5%-(Z)-4-
Tridecen-1-01 Acetate) were placed in 6 fields. Male adults trapped

192
were recorded weekly, and the mean number compared with immature stage
infestation levels in the field. Again, a possible relationship between
population increase and environmental factors is expressed.
Effect of Plant Water on Oviposition
Ovipositional preference related to water content was studied by
collection of 2 tomato plants of 5 different ages. Plants were natu¬
rally infested with TPW eggs in the experimental field mentioned above.
The plantings were 6, 5, 4, 3, and 2 months of age. Every week, plants
were pulled and taken to the laboratory where the numbers of eggs were
recorded. After this, the fresh weight of plant leaves was measured
(g), and 2 days later, the dry weight measured. Water content per plant
was estimated as the difference between fresh weight and dry weight.
The experiment lasted 6 weeks until plants approached 7.5, 6.5, 5.5,
4.5, and 3.5 months old. Plant water content and oviposition with
respect to planting time were analyzed and compared by Duncan's Multiple
Range Test at the 0.05 level.
Effect of Simulated Rainfall on Larvae
An average of 5 second instar larvae per plant, reared on 30 day-
old potted tomato plants, was used to determine the effect of rainfall
on larval mining. Each plant had 3+^1 leaves. The first treatment
consisted of a simulated continuous drizzling (mist) for 24 h. Total
water sprayed per plant was 100 cc per day. Drizzling was simulated by
use of an automatic mister that sprayed fine droplets for 1 minute every
5 minutes over the foliage. The second treatment consisted of spraying

193
the foliage with 50 ml of water 2 times a day. Rainfall was simulated
by use of a manual sprinkler. The third treatment was soil irrigation
with 100 cc of water per day. The number of injuries per leaf was
counted 2 days after the experiment was set, and counts countinued every
other day for during 9 days. Treatments were replicated 3 times. The
mean numbers of injuries per plant were obtained and compared by analysis
of variance (ANOVA). Fresh weight of leaves consumed was recorded (5
days after the experiment started) from randomly selected infested
leaves, following the procedure explained in Chapter I, in order to
determine leaf consumption under different water regimes. The larval
head capsule width was measured to determine larval instar.
Effect of Simulated Rainfall on Pupae
Newly formed pupae of TPW were subjected to the following condi¬
tions: a layer of coarse white sand was placed in a series of boxes
(21 x 30 x 23 cm), constructed of dry lumber, each fitted with a solid
lid and a screen (1 mm diam) in the bottom to help water drainage. Boxes
were surrounded with Tanglefoot® to avoid predation by ants. The boxes
were held in the greenhouse at 24 _+ 3°C. The pupae used in this study
were taken from a culture of 1st generation insects. Insects were
reared in a 12:12 light-dark cycle at a 24 + 3°C and 75 +_ 2% RH. A
total of 20 pupae was placed in each box. Two hundred ml (97 + 2% RH),
100 ml (80 + 1% RH) , 50 ml (60 + 10% RH) , and 0 ml (27 +_ 3% RH) of water
were applied every other day with a manual sprinkler. Treatments were
replicated 3 times. Percentages of emergence were taken of the number of
adults emerging per box. Data were transformed by arc sine procedures
before the analysis. Data are presented as actual percent emergence.

194
Results and Discussion
Seasonal Occurrence of TPW Eggs in Experimental Plots
TPW oviposition peaked early during 1980 (Fig. 23). Numbers during
this year were higher than during 1981. The number of eggs increased
sharply during the first 3 months of 1980, and this increase was main¬
tained until mid-May, 1980, when the tomato leaf area decreased sharply.
Average daily temperature in the field increased after the middle of
March, 1980, but remained stable around 20-25° C during March through
April, 1980. The amount of rainfall was negatively correlated with
oviposition. Oviposition increased when rainfall was zero during March
and decreased at the beginning of April when rainfall increased. The
increase in rainfall seemed to be related to low ovipositional activity
during April, and its delayed effects can be seen in Fig. 23.
During 1981, increased oviposition was observed as temperature in¬
creased and rainfall decreased. The temperature increased during
January through April, stablizing around 21-24° C during the later
months. When rainfall was low the pattern of oviposition increased
abruptly. This may suggest a pattern in which moth activity can be
related to oviposition patterns. This result agrees with the results
found by Simmons and Ellington (1933) who stated that high humidity
reduced 13 times the average number of eggs laid by Sitotroga cerealella
(Gelechiidae).

Figure 23. Seasonal abundance of TPW eggs in experimental fields,
related to temperature and rainfall regimes during A) 1980 and B)
1981, in Homestead, Florida.

Rainfall (mm) T °C 1* pw eggs/plant
196
Sampling Week

Sampling Week
TPW eggs/plant
Rainfall (mm) T °C
Z.6T

198
Seasonal Occurrence of TPW Larvae in Southern Florida
TPW larval density decreased from January 3 through the 2nd week of
February, then peaked at the beginning of March, 1980 (Fig. 24). The
population declined through March and the 1st weeks of April, but
steadily increased during the 3rd week of this month, to reach the
highest density in May. Most growers disced and mowed the tomato fields
during the 4th week of May, causing populations to decrease through
June-October. Despite this, larval injuries (Fig. 24) were found on
volunteer plants during these months. I consider that field infestation
is dependent on temperature and rainfall regimes and crop management.
It is observed in Fig. 30 that despite an increase in temperature
during June-August, rainfall also increased during these months, in¬
dicating a possible negative effect on pest abundance.
During 1981, TPW larval density in commercial tomato fields was
almost negligible. Population densities were low as late as February 25,
and population growth was not evident until early April. Early, low
temperatures seemed to be the factor suppressing the pest this year;
however, the high temperatures and low rainfall coincided with the pest
peaks.
Effect of Plant Water on Oviposition
Effect of oviposition response to leaf water content of the whole
plant has been demonstrated for Pieris rapae (L.) by Wolfson (1980).
Highest water content of each plant was found in those plants planted

Figure 24. Seasonal abundance of TPW larvae in tomato fields, related
to temperature and rainfall regimes during 1980-81, in Homestead,
Florida.

Mean injuries /Plant
200
Sampling Week

Rainfall (mm) T°C TPW moths/trap Mean injur ies/plant
201

202
latest (4 weeks old). Greater mean numbers of eggs obtained (Table 42)
corresponded again to the youngest crop followed by those crops 3, 4, 5,
and 6 months old. The numbers of eggs oviposited among the oldest crops
were not significantly different. When plantings were 3, 4, 5, and 6
months old, the water content of plants was 1.46, 2.69, 9.79, and 8.72
times less, respectively, than that found on 2-month old plants. These
results may indicate the possible relationship between water content and
oviposition; however, when these parameters were regressed, the co¬
efficient of determination was as low as 0.27, despite an F value of
7.06. Perhaps the high (71.37%) CV obtained tended to minimize the
effect of this relationship. Oviposition preference is not only related
to water content, but possibly to the amounts of other substances in the
plant leaf.
Effect of Simulated Rainfall on Larvae
The data in Table 43 show the effects of rainfall on the number of
injuries on each plant. The numbers of larval injuries per plant were
always significantly lower when water was sprayed on the foliage. Thus,
compared to the soil irrigated treatment, the numbers of injuries were
reduced 53% and 48% when water was applied to the foliage. The dif¬
ferences during 10 days after treatment are shown in Fig. 25. It was
found that when water was applied continuously to the foliage, the
larvae stopped mining the leaves and started feeding externally on the
leaf, or constructed a silk tent to protect against the excessive amount
of water. Leaf consumption was also reduced 1.46 and 3.57 times when
the double and continuous water spray was used. Since there was no

20 3
Table 42.
Plant water content in five tomato plantings related to
oviposition by the tomato pinworm.
Planting No.
Leaf Water Content (Fresh
Weight - Dry Weight) (g)
Mean Eggs/Plant
1
11.05 d*
0.56 d
2
9.05 e
0.46 e
3
35.83 c
0.63 c
4
65.99 b
1.51 b
5
96.43 a
1.84 a
* Numbers followed by different letters were significantly dif¬
ferent by Duncan Multiple Range Test (P=0.05).

204
Table 43. Effect of simulated rainfall on foliar larval injuries caused
by the tomato pinworm Keiferia lycopersicelia on plants grown
under greenhouse conditions.
Treatment
Average Number of
injuries/plant
Leaf Consumption
(mg) *
100 cc water applied
daily as a mist
45.78 b**
353.2 b
200 cc water applied
spray twice a day
as
42.071 b
853.2 ab
200 cc water applied
the soil every other
to
day
86.667 a
1261.0 a
* Leaf consumption measured days after treatment on larvae.
**Values followed by different letters were significantly different
by Duncan Multiple Range Test (P=0.05).

Figure 25. Mean number of TPW injuries per plant during 9 days of
simulated rainfall under greenhouse conditions, avg. daily temperature
25 + 2° C.

Mean TPW injuries/Plant
100
80
60
40
20
0
i I I I|
1 3 5 7 9
Days after Treatment
206

207
water stress in the treated plants, we may consider the physical action
of the simulated rainfall as a negative factor that may cause a larval
reaction to stop or delay feeding.
Effect of Simulated Rainfall on Pupae
An intermediate level of water in the soil increased TPW emergence
(Table 44) (Fig. 26). Analysis of the data also indicated that when
moisture levels increased (100-200 ml water) , or decreased (0 ml water) ,
emergence was 93%, 66%, and 48% less, respectively, than when 50 ml of
water were applied. Emergence from the control (no water) was 6.92 and
3.54 times greater than when 200 and 100 ml were applied. These results
are similar to those found for P_. gossypiella by Clayton and Henneberry
(1982). Other factors, such as soil type or depth of burial, may be
expected to have an effect on emergence. The ability of TPW larvae to
locate places to pupate may indicate a characteristic common among some
members of the Gelechiidae. It was observed that the South American
pinworm (Scrobipalpula absoluta) does not pupate in heavy clay-wet soil,
but prefers dry leaves in the canopy as a substrate (Garcia et al.
1974). The combined effect of temperature and high relative humidity
upon mortality factors of the TPW in southern Florida should be inves¬
tigated. It has been demonstrated by Simmons and Ellington (1933) that
egg laying of the angoumois moth is reduced when humidity increases, but
adult longevity is extended. Furthermore, it was determined by Hof-
master (1949) that outbreaks of the potato tuberworm occurred regularly
after unusually hot and dry seasons, and such conditions are considered
as the most favorable for potato tuberworm development.

208
Table 44. Mean percentage of tomato pinworm adults emerged by day after
pupal treatment with different simulated rainfall regimes.
Treatment
ml Water
Mean Percent of Emerged
Adults/Day
200
1.250 b*
100
2.94 b
50
17.76 a
0
8.67 b
* Mean values followed by different letters were significantly different
by Duncan Multiple Range Test (P=0.05).

Figure 26. Percentage of TPW adult emergence under greenhouse conditions
after treatment of pupae with 3 regimes of artificial rainfall (200, 100,
50, and 0 ml water), temperature 24 + 3° C.

% TPW emergence
Days after Treatment
210

211
Conclusions and General Discussion
In conclusion, it is clear that we are just beginning to understand
the influence of the environment on TPW populations. Useful data were
gathered in this research about effects of rainfall on TPW immature
stages.
For instance, during 1980, the highest (7-14) number of eggs per
plant coincided with lower rainfall (2.5-11 mm). This result was rein¬
forced by the increase in oviposition (2-7 eggs/plant) during 1981
compared to the low amount (0-8 mm) of rainfall obtained. This indi¬
cates that oviposition is reduced when rainfall is higher than 10 mm.
Research on female ovipositional behavior related to rainfall patterns
needs to be studied more closely. Knowledge of this relationship will
permit better egg forecasting.
The study of plant water effect on oviposition demonstrated that
a larger amount (96.43 g) of plant water coincided with a greater (1.84)
number of eggs per plant. At this point, it is not clear if this result
is related to the presence of a soluble plant chemical or physical plant
characteristics (e.g. leaf turgidity). Presence of chemicals at dif¬
ferent plant stages should be investigated to relate them to TPW
oviposition. For instance, higher water content occurred when the plant
was at the first reproductive stage (TR^). Plant water decreased during
the second reproductive stage (TR?) through senescence (S^). This
knowledge will be useful for TPW plant resistance studies as well as
better monitoring of TPW eggs on tomato plants.

212
The lowest amount of rainfall (0 mm) during 1980 coincided with the
greatest number of injuries per plant (10-15). During 1981, more (1.25-
2.5) TPW injuries occurred per plant where there was little rainfall
(0-0.5 mm). Data from the effect of simulated rainfall on TPW injuries
reinforced the results obtained in the field. Simulated rainfall re¬
duced by 50% the number of injuries per plant. The possible use of this
practice as a TPW reduction factor is not advised because of the ten¬
dency to increase plant pathogen virulence. The results, however,
partially explain why during a prolonged rainy season, the number of TPW
injuries per plant are reduced.
Simulated rainfall affected adult emergence. The highest (17.76)
mean number of TPW adults emerged when an intermediate amount of rain¬
fall (50 ml) was applied to TPW pupae. Only 8.67, 1.25, and 2.94 TPVJ
adults emerged per day when 0, 200, and 100 ml of water were applied.
Therefore, lower adult emergence is expected during the rainy season in
southern Florida.

CHAPTER IX
INFLUENCE OF POST-HARVEST TOMATO FIELDS ON
THE POPULATION DYNAMICS OF TOMATO PINWORM
Introduction
Elmore and Howland (1943) and Poe et al. (1975) mentioned that
cultural practices such as burning crop residues and discing the tomato
fields helped reduce Keiferia lycopersicella (TPW) populations. The
effect of post-harvest practices on survival of this insect during
noncropping periods is important in a pest management program for TPW in
southern Florida, since larger insect populations appear to coincide
with the post-harvest field season. Reported here are investigations on
the effect of cultural practices on oversummering populations of TPW in
southern Florida.
Materials and Methods
Post-harvest survival of TPW related to cultural practices in the
field was investigated in two separate experiments in Homestead, Dade
County, Florida. The first experiment was conducted during 1980 in 17
fall-winter planted commercial fields. The investigation is summarized
here only for 2 typical fields.
A tomato field in southern Florida can be classified after the main
harvest as: 1) "U-pick" field, 2) abandoned field, and 3) disced and
mowed. The "U-pick" fields are those plantings which are often rented
to a low income person who allows the public to harvest tomatoes for a
213

214
reasonable fee. Generally, the field is not sprayed with insecticides
after commercial harvest. The second category corresponds to those
fields which are left without any supervision for at least 1-2 months.
the third category is fields which are mowed, burned and finally disced
after the main harvest. Each category can become a subcategory (Fig.
27). The number of tomato plants and number of TPW injuries were
2
monitored in those fields by randomly selecting 10 m in each field.
2
The presence of plants and TPW injuries per m measured and averaged for
2
the 10 m per field. This survey was carried out over 4 months, de¬
pending on the cultural activities practiced.
The objective of the second experiment (1981) was to determine
which cultural practices increased the TPW population. The experiment
was done at the Agricultural Research and Education Center, in Home¬
stead, Florida. Experimental fields were planted during October 30,
November 25, and December 30, 1980. Each planting was set in raised
beds and mulched with light colored plastic. Plants were spaced 38 cm
2 2
apart. Each planting was 324 m and split into 3 treatments (108 m
each). Three weeks after treatments were set, the numbers of tomato
2
plants and TPW injuries on 5 m randomly selected were monitored on each
subplot. The survey was conducted biweekly for 1.5 months. Means were
analyzed by a nested analysis of variance (Sokal and Rohlf 1969). Mean
2
number of plants and number of injuries per m were separated by use of
Duncan Multiple Range Test.

Figure. 27. Tomato field status following the main harvest under S. Florida
conditions. Homestead, Florida, 1980.

216

217
Results and Discussion
Survey—1980
The survey demonstrated that 7 (41%) of the fields inspected
during 1980 were disced and mowed (Category 1) immediately after harvest.
Of the 10 remaining fields, 7 became "U-pick" fields (Category 2) and
2 (17.64%) were considered abandoned (Category 3). Two fields from
Category 1 were planted again with a summer crop (bean or squash), the
rest of the fields (5) from this category had tomato plant emergence and
regrowth of plants during a 2-month period. In Category 2, 4 of the 7
"U-pick" fields were disced and mowed in a 2-month period; the remaining
ones (2) were abandoned. During the new fall tomato growing season, 47%
of the 17 inspected fields in the previous season were planted to tomato
again.
In Fig. 28 are shown the mean number of plants and foliar injury
2
per m in 2 fields. Field 1 had an increase in the number of injuries
per plant until the field was disced. When the field was planted with
beans (Phaseolus vulgaris), the number of tomato plants germinating and
2
the number of TPW injuries per m increased up to 13 and 14, respec¬
tively. The second field ("U-pick") showed a slight increase in number
2
of plants per m until discing. After this practice, the number of TPW
injuries increased to 8 times greater than before discing. It was
observed that when beans and squash followed tomato planting, volunteer
tomato plants were more numerous than when the field was disced and
abandoned. This effect is possibly related to the irrigation and type

2
Figure 28. Number of tomato plants and TPW injuries per m in 2 post-
harvested tomato fields. Homestead, Florida, 1980.

20
10
5
0
10
5
0
F
1
2
M
2 3 A 1
A
Aug1
S
2

220
Table 45. Effect of crop age of post-harvested tomato plants on volun¬
teer plants and number of tomato pinworm larval injuries.
Crop
Planted
Months After
Harvest
Mean Plants/
m
Mean TPW Injuries/
m2
Oct.
30,
1980
3
0.1222 b*
0.1389 b
Nov.
25,
1980
2
0.41 b
0.641 b
Dec.
30,
1980
1
2.53 a
4.006 a
* Values followed by different letters were significantly different
according Duncan Multiple Range Test (P=0.05).

221
of herbicide used in the new crop. Field 2 had a slower increase of
plant and larval levels after entering a period of abandonment.
Experiment—1981
Results of the second experiment are expressed in Table 45. Tomato
crops planted earlier (Oct.-Nov., 1980) has a significantly lower mean
number of volunteer plants than the younger one (Dec., 1980). The mean
number of TPW injuries was significantly larger in the younger planting
than in the others. Abandoned fields had generally lower numbers of
plants than mowed and disced fields. The numbers of injuries were
similar for any treatment. Effects of planting time on the treatments
were obvious (Tables 46-47). The older planting had a greater number of
injuries per plant when abandoned than when the planting was mowed or
2
disced. The second planting (Nov., 1980) again had more injuries per m
if abandoned compared to the other treatments. In contrast, the younger
planting had a higher number of injuries when disced and mowed than when
abandoned. The effects are explicable. Older fields have fewer viable
seeds that will germinate than those from younger fields in which seeds
are immediately incorporated into the soil.
Effects from secondary host plants as oversummering sites have been
suggested for the TPW. The availability of off-season tomatoes helps
to maintain the TPW population but also helps in build up of natural
enemies. Use of abandoned or "U-pick" fields for pest management of
TPW by constant release of parasitoids would be a practice to reduce
pinworm populations for the next season, without interfering with the
farmer interests and environmental concerns.

Table 46. General effect of cultural practices on volunteer tomatoes
and infestation by tomato pinworm.
Treatment
. 2
Mean Plants/m
2
Mean TPVJ Injuries/m
Disced
1.377
a*
1.46
Mowed
1.15
a
1.70
Abandoned
0.54
b
1.67
* Values followed by the same letter are not significantly different
according to Duncan's Multiple Range Test.

223
Table 47. Effect of planting age and cultural practices on volunteer
tomato plants and number of TPW injuries.
Planting
Treatment
2
Number plants/m
... ,2
Number injuries/m
1) Oct. 30,
1980
disced
0.00
0.00
mowed
0.075
b*
0.13
abandoned
0.316
a
0.28
2) Nov. 25,
1980
disced
0.017
c
0.16
b
mowed
0.283
b
0.16
b
abandoned
1.066
a
1.51
3) Dec. 30,
1980
disced
2.333
b
4.233
mowed
2.166
a
4.55
abandoned
1.333
a
3.23
* Values followed by the same letter are not significantly different
according to Duncan's Multiple Range Test (P=0.05).

224
Consequently, effects of cultural practices to help control the TPW
in tomatoes can be considered in two ways. First, these practices may
be considered in relation to the pest density or, second, in relation to
the impact on natural control agents. For the most part, cultural
measures usually modify the environment to the disadvantage of the pest
(Anonymous 1979). However, cultural practices can destroy the host and
the parasites or natural enemies of the host. In southern Florida, the
widespread cultural practices could account for part of the insect
reduction during the past 10 years. The impact of some of the post¬
harvest practices related to pest density has been demonstrated in this
study.

GENERAL DISCUSSION AND CONCLUSIONS
In this research useful data were gathered about tomato plant
phenology, spatial distribution of eggs and larval populations of
Keiferia lycopersicella (Walsingham), and effects of tomato pinworm
(TPW) larvae on tomato yield. Studies of the effects of natural
enemies, rainfall, edgerows and cultural practices on the population
dynamics of this insect provided important data for better management of
TPW.
A different IPM approach is suggested based on data about different
stages of development of tomatoes. 'Flora-Dade' tomato phenology was
described based on 3 major stages: vegetative, reproductive and senescent.
Vegetative stages (TV^-TV ) were determined by presence of primary-
leaves and secondary vegetative growth on plants 1-35 days old. Repro¬
ductive stages TR^-TR^ were based on numbers of flowers and fruits on
plants 40-110 days old. The results indicate that reproductive stages
(TR^-TR^) should be subdivided so more precise pest management decisions
can be made. This classification can be refined by dividing the TR?
stage into 2 substages. One stage is during fruit formation and the
other stage is during harvest maturity. The rationale behind this is to
improve economic thresholds during fruit formation and just before
harvest.
Research on sampling TPW eggs indicated that high cost ($8.4-84) of
egg sampling combined with high SE/x ratio (20-100) reduce the practicality
225

226
of sampling this immature stage in the field. Data on TPW distribution
indicated that eggs were found mainly (44-68%) in the middle-upper
canopy of the plant. Therefore, proportional sampling can be allocated
for the upper external stratum (nh=6) followed by the middle internal
stratum (nh=5), and 3,4,4 and 1 samples for lower external, upper,
middle and lower internal canopies, respectively. Apparently, the
research of Burton and Schuster (1981) provided similar data that allow
the hypothesis of female attraction for oviposition in the upper plant-
part. Future research must be carried out on the relationship between
female trapping and egg presence in the field. The research reported
here indicates the need for more evidence on TPW oviposition on dif¬
ferent tomato plant stages.
Research on sampling TPW injuries per plant indicated that the
percent of the SE corresponding to the mean was 11-31% for 20-25 plants
sampled when the population was low (0.2-2.12 injuries per plant) and
was 21-29% for 15-20 plants when the population was high (11.05-17.2).
By sampling two leaves from the upper and lower canopies I account for
32-34% of the total larval injury per plant. Results indicated a
measure of efficiency (RNP) fluctuating between 0.49-1.20 for 50 and 5
plants inspected when the population was small and 0.49-3.18 for 50 and 5
plants when the population was large. Results generally agree with
those of Wellik et al. (1979) indicating that larvae are mainly (50-75%)
located in the middle-lower plant canopy. Therefore, more samples
should be allocated to the middle and lower strata. The average number
(n=20) was 2, 3, and 5 samples for upper, middle and lower external
canopies, and 0, 5, and 3 from upper, middle and lower internal canopies,

227
respectively. I recommend more research on the relationship between
oviposition and the population index (number of injuries per plant).
These data are necessary to establish the prediction of economic injury
levels.
Economic injury level studies provided useful data to determine
yield losses from high levels of larvae (1-14) per plant. The largest
yield reduction was from 12-14 TPW larvae per plant. Since the TPW
larva attacks both leaves and fruits, the results suggest that sampling
from the lower canopy will be more useful than sampling from the upper
canopy. These results disagree with those obtained by Wolfenbarger et
al. (1975). Conflicting views about sampling to detect an economic
injury level will be fewer if EIL is determined for every stage of
tomatoes. I suggest further research to develop economic injury levels
for the 4 main stages of development. In this case, it would help to
avoid relying on the stages of the plant close to harvest (TR,,) > when it
is too late to apply control measures.
The role of several factors (parasitoids, field edgerows, rainfall,
and horticultural practices) influenced TPW population dynamics in
southern Florida. For instance, the effect of natural enemies was an
important TPW mortality factor. The role of larval and egg parasitoids
increased after the main crop harvest. Levels of TPW larval para-
sitization fluctuated between 39.3-42.3% during 1980-81. I consider the
larval parasitoid Apanteles spp. and the egg parasitoid Trichogramma
pretiosum Riley the most promising biological control agents for TPW in
southern Florida. Apanteles spp. appeared earlier during the winter
season, increasing in density during the months of April-June. T.

228
pretiosum was found to be a TPW egg parasitoid with an intermediate
level (33-73%) of parasitism. A conclusion from these data is that
studies should be encouraged which focus on the effect of parasitization
of larvae and eggs following releases of these beneficiáis in post-
harvested fields. I consider post-harvest agroecosystem management to
be a good strategy to assure a low pest density. I recommend further
study of TPW parasitoids to find pesticide resistant strains.
Results describing the patterns of colonization of TPW in the field
provide answers to the accumulation of TPW in several areas of the
field. Data suggested that this microplepidopteran tends to aggregate
near field borders, especially near windfalls. These results may en¬
courage further research in dispersal of gelechiids. To understand the
dispersion of the different TPW generations, it is important to estab¬
lish which generation migrates over long distances and which disperses
over nearby fields. From a practical standpoint, such knowledge of TPW
aggregation can be used for control and monitoring of TPW populations.
Research on effects of abiotic factors such as rainfall indicated
reasons for TPW population reduction during 1980-81. The use of arti¬
ficial rainfall on TPW larvae and pupae demonstrated that when plant
foliage was irrigated there was a behavioral change in larval foliar
consumption which resulted in 50% reduction of injuries by larvae
compared to injuries on soil-irrigated plants. Adult emergence was
reduced 93% when high levels of water were applied to the soil. Never¬
theless, these experiments require a more elaborate microclimatic study
of the pest. I consider it useful to link the similarities of popula¬
tion dynamics of this pest and the potato tuberworm because of their
parallel activities related to temperature and rainfall regimes.

229
The results from evaluation of cultural practices on populations of
TPW indicate that post-harvested tomato crops planted earlier (Oct.-
Nov., 1980) had a significantly lower mean number of volunteer plants
than did the crop planted later (Dec., 1980). Mean numbers of injuries
2
per m were higher in crops planted later (Dec., 1980) than in crops
planted earlier (Oct.-Nov., 1980). Despite the complexity and dif¬
ficulty of proving which cultural practices are most adequate for a
sound TPW management program, two different approaches could be taken.
First, practices such as mowing, burning, and discing may eliminate
tomatoes as a source of TPW infestation. This implies more supervision
from farmers and agricultural agents of post-harvested fields.
Secondly, if habitat management of tomatoes is desired the use of
"U-pick," abandoned fields or fields where tomatoes grow voluntarily,
should be used as a source for gathering parasites and predators.
Natural enemies augmentation could be used to reduce TPW infestations
for the next tomato growing season. This may be a large step toward
improved management of the TPW.
Several additional studies are needed so that TPW population
assessment can be conducted most efficiently. At present, entomologists
must continue research on TPW monitoring as well as studying the pos¬
sible relationship between egg numbers and adults caught by pheromone
trapping. In general, a better IPM program in tomatoes will develop
when sampling techniques as well as multiple economic injury levels are
determined for TPW and other direct pests of tomatoes.

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APPENDIX
EXPLANATORY TABLES FOR CHAPTERS II AND III

Table 48. Tomato pinworm egg frequency distributions determined on tomato plants during 1981.
Planting3 Date
N
X
2
s
k
P
9
chi
2
p X
df
Distribution
4 March
18
18
0.277
0.564
—
—
1.288
0.2564
1
Poisson
0.234
1.19
1.235
0.2664
1
Negative
binomial
April
1
20
0.050
0.050
—
—
0.0006
1.0000
0
Poisson
8
20
0.300
0.450
—
—
1.7960
0.1800
1
Poisson
0.469
0.639
1.1650
0.2800
1
Negative
binomial
16
10
0.200
0.40
—
—
5.6000
0.0179
1
Poisson
0.097
2.011
1.2900
1.0000
1
Negative
binomial
23
10
0.300
0.233
—
—
0.0860
1.0000
0
Poisson
1.350
0.222
0.0070
1.0000
1
Positive
binomial
May
1
10
1.300
0.900
—
—
16.2800
0.0002
2
Poisson
0.084
15.36
1.5500
0.2100
1
Negative
binomial
15
10
1.000
1.470
—
—
1.2800
0.2500
1
Poisson
1.240
0.30
0.54
0.4600
1
Negative
binomial
244

Table 48--Continued.
March
18
20
0.10
0.097
1.90
April
1
20
0.10
0.094
1.9
8
20
1.30
7.789
0.469
16
10
0.20
0.178
1.800
23
10
0.40
1.60
0.049
May
1
10
0.10
0.10
—
8
10
0.70
2.45
0.270
15
10
0.90
2.32
0.860
—
0.005
1
0
Poisson
0.052
0.001
1
-1
Positive binomial
—
0.005
1
0
Poisson
0.052
0.0012
1
-1
Positive binomial
—
1.7960
0.18
1
Poisson
0.639
1.1650
0.28
1
Negative binomial
—
0.0200
1.00
0
Poisson
0.111
0.0050
1.00
1
Positive binomial
—
3.7080
0.054
1
Poisson
8.147
1.2310
0.267
1
Negative binomial
—
0.0020
1.00
0
Poisson
—
1.6600
0.197
1
Poisson
2.530
0.9000
0.340
1
Negative binomial
—
0.9600
0.320
1
Poisson
1.040
2.2700
0.130
1
Negative binomial
245

Table 48—Continued.
6 April
May
1 10 1.9
8 10 1.2
16 20 0.789
23 10 0.300
1 20 0.500
8 10 1.000
15 10 0.600
0.54
2.4
2.509
0.450
1.105
2.444
0.933
0.38
1.536
0.29
0.469
0.306
0.850
1.408
—
11.69
0.008
4.99
2.65
0.017
—
0.47
0.780
0.78
0.358
0.540
—
4.430
0.035
2.69
3.590
0.162
—
1.796
0.180
0.64
0.375
1.000
—
4.190
0.040
1.633
0.905
0.340
—
0.750
0.380
1.170
0.250
0.610
—
0.110
0.730
0.426
1.620
0.203
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
3
4
2
1
1
2
1
0
1
1
1
1
1
1
246

Table 48--Continued.
7 April 8 10 1.60
16 20 0.80
23 20 2.05
May 1 20 1.50
8 20 2.05
4.48
1.221
4.680
1.421
7.52
0.584
1.262
1.032
28.500
0.45
15
10 0.80
0.178
—
7.230
0.026
2
Negative
binomial
2.72
1.218
0.740
3
Poisson
—
1.120
0.280
1
Poisson
0.639
1.850
0.170
1
Negative
binomial
—
19.43
0.0006
4
Poisson
1.987
12.05
0.0300
5
Negative
binomial
—
0.8546
0.8360
3
Poisson
0.052
0.8360
0.6500
2
Negative
binomial
—
42.7900
0.0000
4
Poisson
4.510
12.550
0.0800
7
Negative
binomial
2.510
1.0000
0
Poisson
247

Table 48--Continued.
8 April
8
10
0.111
0.111
—
—
0.064
0.790
1
Poisson
16
20
1.000
1.895
—
—
8.447
0.014
2
Poisson
0.981
1.019
3.580
0.163
2
Negative
binomial
23
20
0.900
1.982
—
—
7.310
0.025
2
Poisson
0.559
1.668
1.280
0.520
2
Negative
binomial
May
1
20
1.400
3.200
—
—
7.27
0.063
3
Poisson
1.360
1.320
3.460
0.320
3
Negative
binomial
8
20
2.737
8.427
—
—
17.99
0.0002
5
Poisson
1.503
1.321
15.440
0.0100
6
Negative
binomial
15
20
2.260
14.760
—
—
12.410
0.0100
4
Poisson
0.650
3.430
4.360
0.6200
6
Negative
binomial
aTomato plantings correspond to tomato crops planted: 4) Oct., 1980; 5) Nov., 1980; 6) Dec., 1980;
7) Jan., 1981; and 8) Feb., 1981.
248

Table 49. Tomato pinworm foliar injury frequency distributions determined on tomato plants during
1980.
Planting3 Date
N
X
2
s
k
P
chi2
v,-2
p chi
df
Distribution
1 Feb. 8
19
1.211
3.064
—
—
3.77
0.149
2
Poisson
0.78
1.53
1.79
0.61
3
Negative
binomial
12
18
0.888
2.693
—
—
10.6
0.004
2
Poisson
0.31
2.833
2.41
0.295
2
Negative
binomial
21
18
2.056
4.291
—
—
9.771
0.04
4
Poisson
2.176
0.9448
8.481
0.075
4
Negative
binomial
Mar. 14
17
9.882
107.2
—
—
64.69
0
9
Poisson
0.81
12.11
29.93
0.41
29
Negative
binomial
20
20
7.52
23.76
—
—
18.42
0.010
7
Poisson
3.39
2.22
13.26
0.42
13
Negative
binomial
Apr. 4
22
23.45
75.59
—
—
54.56
0
15
Poisson
11.7
2.034
132.8
0
22
Negative
binomial
24
19
13.63
74.36
—
—
24.44
0.0109
11
Poisson
3.61
3.75
16.52
0.86
24
Negative
binomial
249

Table 49--Continued.
1 May 2 21 21.43 103.8
2.92 7.31
10 19 17.26 109.8
2.66 6.46
20 15 13.00 84.43
1.71 7.56
June 1 20 5.2 22.69
1.42 3.65
2 Feb. 12 20 0.15 0.239
0.16 0.89
Mar. 14 20 8.2 85.12
0.56 14.59
1.06 6.37
20
19
6.78 64.95
binomial
71.92
0
14
Poisson
43.24
0.032
28
Negative
46.66
0
12
Poisson
23.1
0.811
30
Negative
19.52
0.034
10
Poisson
25.13
0.56
27
Negative
12.73
0.047
6
Poisson
16.04
0.247
13
Negative
4.118
0.042
1
Poisson
0.37
1
0
Negative
46.27
0
8
Poisson
30.35
0.396
29
Negative
30.64
6.34xl0~5
7
Poisson
17.03
0.58
19
Negative
binomial
binomial
binomial
binomial
binomial
binomial
250

Table 49--Continued.
2 Apr.
3 Mar.
Apr.
4 20
18 17
11 20
14 20
20 20
11 16
18 25
11.5
10.24
14.57
0.30
0.1
11.0
14.64
44.16
37.57
46.96
0.274
0.094
30.13
42.91
5.38
2.79
7.46
0.05
1.9
6.793
8.103
1.37
3.65
1.95
5.14
0.05
1.61
1.307
11.19
0.34
14.02
0.66
28.25
0.0004
18.53
0.48
14.7
0.19
20.66
0.41
3.446
0.063
0.496
0.48
0.114
0.735
0.05
1.00
11.52
0.242
15.84
0.323
25.01
0.022
24.83
0.208
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Negative binomial
Poisson
Positive binomial
Poisson
Negative binomial
Poisson
Negative binomial
10
17
8
19
11
20
1
1
1
0
9
14
13
20
251

Table 49--Continued.
Apr.
25
23
11.17
56.33
—
—
25.45
0.004
10
Poisson
3.00
3.253
20.22
0.507
21
Negative
binomial
May
10
25
2.04
8.54
—
—
2.04
55.61
4
Poisson
0.366
5.57
6.41
0.49
7
Negative
binomial
2
18
11.61
22.13
—
—
12.5
0.252
10
Poisson
13.12
0.38
21.85
0.05
13
Negative
binomial
Planting numbers correspond to those tomato crops planted during Nov., Dec., 1979 and Jan., 1980.
252

Table 50. Tomato pinworm foliar injury frequency distributions determined on tomato plants during
1981.
Planting3 Date
N
X
2
s
k
P
chi2
. .2
p chi
df
Distribution
4 Mar.
11
20
0.6
1.095
—
—
5.027
0.02
1
Poisson
0.44
1.35
1.136
0.286
1
Negative binomial
18
20
1.5
0.684
—
—
26.47
0
3
Poisson
0.204
7.32
15.13
0.019
6
Negative binomial
Apr.
1
8
2.75
1.786
—
—
0.996
0.802
3
Poisson
3.09
0.38
2.64.
0.44
3
Negative binomial
8
20
1.25
2.09
—
—
3.171
0.20
2
Poisson
1.64
0.75
1.931
0.37
2
Negative binomial
16
20
1.77
4.69
—
—
6.100
0.046
2
Poisson
0.76
2.33
1.083
0.780
3
Negative binomial
24
20
1.7
0.063
—
—
7.61
0.050
3
Poisson
1.60
1.60
0.91
0.82
3
Negative binomial
May
1
19
5.84
17.36
3.14
1.35
28.64
0.009
10
Negative binomial
28.64
0.0001
7
Poisson
253

Table 50—Continued.
4 May
5 Mar.
Apr.
8 9
15 10
11 20
18 20
8 15
16 20
24 20
2.22 4.44
4.6 18.7
0.4 0.35
1.05 2.15
3.6 32.67
1.65 2.55
1.3 1.8
2.87
2.39
3.8
0.73
0.71
4.46
3.67
—
0.44
0.80
2
Poisson
0.77
2.63
0.45
3
Negative binomial
—
11.46
0.02
4
Poisson
1.98
8.72
0.27
7
Negative binomial
—
0.131
0.716
1
Poisson
0.105
0.075
0.780
1
Negative binomial
—
6.54
0.03
2
Poisson
1.42
5.96
0.049
2
Negative binomial
—
20.4
0.001
5
Poisson
5.11
21.51
0.017
10
Negative binomial
—
3.24
0.35
3
Poisson
0.36
4.47
0.104
2
Negative binomial
—
1.09
0.57
2
Poisson
0.35
0.47
0.68
2
Negative binomial
254

Table 50--Continued.
5 May
6 Mar.
Apr.
1 19
8 10
15 9
18 20
1 10
8 20
16 15
5.63
2.7
6.77
0.20
0.30
0.45
1.93
10.13
9.56
33.19
0.48
0.45
0.99
1.35
6.77
0.85
1.41
0.09
0.46
0.345
6.43
—
14.82
0.038
7
Poisson
0.33
8.41
0.39
8
Negative binomial
—
8.25
0.04
3
Poisson
3.14
4.09
0.53
5
Negative binomial
—
17.34
0.0038
5
Poisson
4.778
10.62
0.476
11
Negative binomial
—
2.94
0.290
1
Poisson
2.165
1.08
0.298
1
Negative binomial
—
1.79
0.180
1
Poisson
0.63
0.37
1.00
0
Negative binomial
—
1.86
0.17
1
Poisson
1.301
0.015
0.902
1
Negative binomial
—
1.78
0.610
3
Poisson
0.30
0.87
0.64
2
Negative binomial
255

Table 50--Continued.
6
Apr.
24
19
3.78
9.731
—
2.98
May
1
16
7.81
28.16
—
3.06
8
24
5.25
16.28
—
1.67
15
16
7.56
8.66
—
102.4
7
Apr.
8
20
0.05
0.05
—
16
17
1.23
2.69
—
0.87
24
20
2.25
4.93
—
2.03
May
1
17
3.00
8.075
—
1.08
8
15
4.2
10.46
4.83
—
9.35
0.09
5
Poisson
1.27
7.37
0.39
7
Negative binomial
—
10.56
0.15
7
Poisson
2.54
12.29
0.58
14
Negative binomial
—
36.89
0
7
Poisson
3.12
19.9
0.09
13
Negative binomial
—
2.53
0.92
7
Poisson
0.073
4.76
0.68
7
Negative binomial
—
0.026
0.87
1
Poisson
—
3.75
0.15
2
Poisson
1.40
2.37
0.30
2
Negative binomial
—
7.94
0.09
4
Poisson
1.104
7.22
0.12
4
Negative binomial
—
8.24
0.08
4
Poisson
2.76
7.67
0.36
7
Negative binomial
—
14.09
0.014
5
Poisson
0.33
9.62
0.21
7
Negative binomial
256

Table 50--Continued.
May
15
15
4.2
10.46
—
—
14.09
0.0149
5
Poisson
1.76
2.37
13.17
0.10
8
Negative binomial
Apr.
24
20
0.55
1.52
—
—
6.59
0.01
1
Poisson
0.152
3.60
3.06
0.07
1
Negative binomial
May
1
20
1.7
1.74
—
—
16.13
0.001
3
Poisson
0.71
2.39
5.92
0.31
5
Negative binomial
8
20
3.18
5.76
—
—
12.31
0.015
4
Poisson
4.58
0.69
10.05
0.07
5
Negative binomial
15
16
3.37
5.98
—
—
3.74
0.44
4
Poisson
4.76
0.70
8.39
0.135
5
Negative binomial
a
Planting numbers correspond to those tomato crops planted during: 4) Oct., 1980; 5) Nov., 1980;
6) Dec., 1980; 7) Jan., 1981; and 8) Feb., 1981.
257

Table 51.
TPW egg allocation
development TV^.
sample for 6
plant strata.
Planting 8, 1981.
Age: 38 days.
Stage of
External
Stratum
Internal
Parameter
Upper
Middle
Lower
Upper
Middle
Lower
X
0.6
0.1
0
0.2
0.4
0
s2
1.6
0.1
0
0.17
1.6
0
SE
0.4
0.1
0
1.13
0.4
0
SE/x
0.6
1.0
0
0.65
1.0
0
Sh
1.26
0.31
0
0.41
1.26
0
nhS
8.59
2.10
0
2.6
8.59
0
cib
0.6+1.35
0.1+205
—
0.2+1.33
0.4+2.05
—
a
nh
NhSH
ZNhSh )n'
N=947,
n=20
Cl = x+(SE)ta
b
258

Table 52. TPW egg allocation sample for 6 plant strata. Planting 8, 1981. Age: 46 days. Stage
of development TR^.
Parameter
External
Stratum
Internal
Upper
Middle
Lower
Upper
Middle
Lower
X
0.4
0.2
0.4
0
0
0
s2
0.46
0.48
0.98
0
0
0
SE
0.15
0.15
0.22
0
0
0
SE/x
0.37
0.75
0.55
0
0
0
Sh
0.67
0.69
0.99
0
0
0
nha
5.70
5.75
8.24
0
0
0
nh = (ENhSh ^n' N=947' n=20
259

Table 53.
TPW egg allocation
of development TR^.
sample for
6 plant strata.
Planting 7, 1981.
Age: 68 days.
Stage
Stratum
External
Internal
Parame ter
Upper
Middle
Lower
Upper
Middle
Lower
X
0.7
0
0.4
0
0.4
0.1
s2
â–  3.56
0
0.71
0
1.6
0.1
SE
0.59
0
0.26
0
0.4
0.1
SE/x
0.84
0
0.65
0
1.0
1.00
Sh
1.88
0
0.84
0
1.26
0.31
nha
8.76
0
3.81
0
5.72
1.40
cib
0.7+1.72
0
0.4+1.33
0
0.4+2.05
0.1+2.05
a
nh
NhSh
iNhSh ,n'
N=947, n=20
b
Cl = x+(SE)ta
260

Table 54.
TPW egg allocation sample for 6
Stage of development TR^
plant strata.
Planting 4,
30 Oct. 1980. Age:
77 days.
External
Stratum
Internal
Parameter
Upper
Middle
Lower
Upper
Middle
Lower
X
0
0.2
0.15
0
0.05
0.4
s2
0
0.8
0.23
0
0.05
1.41
SE
0
0.2
0.11
0
0.015
0.26
SE/x
0
1
0.73
0
0.3
0.65
Sh
0
0.90
0.48
0
0.22
1.18
nh3
0
6.47
3.44
0
1.58
8.49
cib
0
0.2+0.41
0.15+0.22
0
0.05+0.03
0.4+0.53
a
nh
NhSh
ENliSh
N=94 7,
n=20
b
Cl = x+(SE)tct
261

Table 55.
TPW larval injury sample allocation
Stage of development TR^.
for 6
plant strata. Planting
6, 1981.
Age: 78 days.
External
Stratum
Internal
Parameter
Upper
Middle
Lower
Upper
Middle
Lower
X
0
0
0
0
0.1
0.1
-2
S
0
0
0
0
0.20
0.04
SE
0
0
0
0
0.10
0.04
SE/x
0
0
0
0
1.00
0.44
Sh
0
0
0
0
0.44
0.20
a
nh
0
0
0
0
12.00
4.00
b
Cl
0
0
0
0
nh *ENhSh ^n' Nh 947' n 20
b
Cl = x+ta(S~)
— x
262

Table 56.
TPW larval injury sample allocation
Stage of development, TR^ •
for 6 plant strata. Planting
5, 1981.
Age: 108 days
Parameter
External
Stratum
Internal
Upper
Middle
Lower
Upper
Middle
Lower
X
0.1
0.3
0.3
0
0.4
0.05
s2
0.2
0.35
0.3
0
0.35
0.05
SE
0.1
0.13
0.12
0
0.13
0.05
SE/x
1.00
0.44
0.40
0
0.33
1.00
Sh
0.44
0.59
0.54
0
0.59
0.22
nha
4
5
4
0
5
2
a
nh
NhSh
Nh=947 ,
n=20
iNhSh
263

Table 57. TPW larval injury sample allocation for 6 plant strata. Planting 4, 1981. Age: 120 days.
Stage of development, TR^.
Parameter
External
Stratum
Internal
Upper
Middle
Lower
Upper
Middle
Lower
X
0.05
0.20
0.4
0
0.6
0.3
s2
0.05
0.134
0.87
0
0.78
0.43
SE
0.01
0.08
0.208
0
0.20
0.14
SE/x
0.22
0.40
0.52
0
0.33
0.48
Sh
0.22
0.36
0.93
0
0.88
0.65
Nha
1.25
2.04
5.28
0
5
4
= 264

BIOGRAPHICAL SKETCH
Jorge E. Pena was born on April 8, 1948, in Cali, Colombia. He
received his high school certificate in 1967 from Colegio Benjamin
Herrera in Cali, Colombia. He began his undergraduate studies in 1968
at the Universidad Nacional de Colombia, Facultad de Agronomía, and re¬
ceived the Bachelor of Science degree with a major in agronomy in
January 1973.
In March 1973, he started working for CIAT (Centro Internacional de
Agricultura Tropical) as a research assistant in a cassava entomology
program. In 1977 he was awarded a scholarship to pursue a Master of
Science degree in entomology at the University of Florida. He graduated
in 1979. He is currently a candidate for the degree of Doctor of Philos¬
ophy in the Department of Entomology and Nematology at the University of
Florida.
He is a member of Entomological Society of America, Florida Ento¬
mological Society, Root Crops Society and Sociedad Colombiana de
Entomología.
265

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree Doctor
of Philosophy.
V.H. Waddill, Chairman -
Associate Professor of
Entomology and Nematology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree Doctor
of Philosophy.
J.L. Stimac
Associate Professor of
Entomology and Nematology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality, as a dissertation for the degree Doctor
of Philosophy.
D.J. Schuster
Associate Professor of
Entomology and Nematology

I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
of Doctor of Philosophy.
S.H. Kerr
Professor of Entomology and
Nematology
I certify that I have read this study and that in my opinion
it conforms to acceptable standards of scholarly presentation and is
fully adequate, in scope and quality, as a dissertation for the degree
of Doctor of Philosophy.
K.L. Pohronezny
Assistant Professor of Plant
Pathology

This dissertation was submitted to the Graduate Faculty of
the College of Agriculture and to the Graduate Council, and was
accepted as partial fulfillment of the requirements for the
degree of Doctor of Philosophy.
April 1983
cT-
Dean,/College of Agf/culture
Dean for Graduate Studies and
Research

UNIVERSITY OF FLORIDA
3 1262 08553 4237




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