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1973 Some Biological Implications of Polymorphism in the Bean Leaf Beetle, Cerotoma Trifurcata (Forster). Donald Charles Herzog Louisiana State University and Agricultural & Mechanical College
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Recommended Citation Herzog, Donald Charles, "Some Biological Implications of Polymorphism in the Bean Leaf Beetle, Cerotoma Trifurcata (Forster)." (1973). LSU Historical Dissertations and Theses. 2398. https://digitalcommons.lsu.edu/gradschool_disstheses/2398
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Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 HERZOG s Donald Charles, 1944- SOME BIOLOGICAL IMPLICATIONS OF POLYMORPHISM IN THE BEAN LEAF BEETLE, CEROTOMA TRIFURCATA (FORSTER).
The Louisiana State University and Agricultural and Mechanical College, Ph.D., 1973 Entomology
University Microfilms, A XEROX Company, Ann Arbor, Michigan
THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. Some Biological Implications of Polymorphism in the Bean Leaf Beetle, Cerotoma trifurcata (F o rste r ).
A Dissertation
Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy
in
The Department of Entomology
by Donald Charles Herzog B.S., Texas Technological College, 1966 M.S., Louisiana State University, 1968 May, 1973 ACKNOWLEDGEMENTS
The author wishes to express his sincere gratitude to Dr. L. D.
Newsom, Head of the Department of Entomology and Major Professor, who directed the writer’s doctoral research. His contributions of time, advice and constructive criticisms were invaluable in the course of these investigations and in the preparation of this manuscript.
Sincere thanks are also due to the members of the writer's advi sory committee composed of Drs. H. B. Boudreaux, D. F. Clower, J. B.
Graves and S. D. Hensley of the Department of Entomology, Dr. K. L.
Koonce of the Department of Experimental Statistics, and Dr. W. L.
French of the Department of Zoology for their advice and criticism s.
Further gratitude is expressed to Drs. Koonce and Graves for their assistance in the statistical analysis of data.
Grateful acknowledgement is made to the Messrs. Lawerence,
Ernest, Francis and Antonio Richard of St. Landry Parish for their consideration in allowing collections of bean leaf beetles on their land.
The author is indebted to Dr. R. L. Jensen, Assistant Professor, and Mr. J. W. Thomas, Jr., Research Associate, of the Department of
Entomology for their able assistance in field collections.
Very special thanks are due the w rite r's wife, Sandra, for typing the first draft of this manuscript and without whose endless patience, constant support and encouragement this dissertation could never have been completed. Finally, the author would like to thank Miss Mary Weber for typing the final draft of this manuscript.
i i i TABLE OF CONTENTS
Page
ACKNOWLEDGEMENT ...... i i
TABLE OF CONTENTS...... iv
LIST OF T A B L E S ...... v ii
LIST OF F I G U R E S ...... xi
ABSTRACT ...... x i i i
INTRODUCTION...... 1
LITERATURE REVIEW . . . 2
Bean Leaf Beetle ...... 2
Background and History ...... 2
Distribution ...... 2
Economic Importance ...... 3
Rearing ...... 4
Parasitism ...... 5
Insecticide Resistance ...... 6
Polymorphism ...... 7
Mimicry ...... 10
Current Research ...... 10
Polymorphism ...... 1 1
Seasonal Variability ...... 12
Geographic Polymorphism ..... 13
Sexual Dimorphism ...... 14
Sexual Selection and Assortive Mating . . . 16
iv Page
Size and Structure ...... 16
Sex Determination ...... 17
METHODS AND MATERIALS ...... 18
Color and Marking Pattern Variations .... 18
Bean Leaf Beetle Collections ..... 18
Rearing Technique ...... 19
Weight Determination ...... 23
Insecticide Study ...... 24
Temperature Tolerance Studies ..... 25
Parasitism ...... 27
Statistical Analyses ...... 28
Symbols and Abbreviations ...... 29
RESULTS AND DISCUSSION ...... 31
Laboratory Rearing ...... 31
Egg Storage ...... 31
Larval Food Source ...... 31
Matxng ...... 22
Adult Life History ...... 38
Longevity ...... 50
Egg Production ...... 56
Weight Determination ...... 71
Susceptibility to Methyl Parathion .... 75
Parasitism ...... • .75
v Page
Temperature Tolerance . . * . 79
Biological Implications .... 107
Miscellaneous Observations on Factors Concerning Bean Leaf Beetle Abundance 108
CONCLUSIONS ...... 110
LITERATURE CITED ...... 112
APPENDIX ...... 119
VITA ...... 176
v i LIST OF TABLES
Page
1. Dates of installation and number of bean leaf beetles included in replicated temperature tolerance studies. Baton Rouge, Louisiana. 1972...... 26
2. Total number of females, number and per cent reproductive females obtained in a bean leaf beetle laboratory-rearing program. Baton Rouge, Louisiana. 1970-71...... 34
3. Proportions of pairings resulting in mating for female and male laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71...... 35
4. Proportions of bean leaf beetle pairings by color and by marking pattern resulting in mating. Baton Rouge, Louisiana. 1970-71...... 36
5. Mean number of males required before impregnation of laboratory-reared female bean leaf beetles was achieved. Baton Rouge, Louisiana. 1970-71. . . 39
6. Mean duration of preoviposit ion, oviposit ion, and postoviposition periods of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. . . 40
7. Proportion of laboratory-reared female bean leaf beetle life cycle spent in preoviposit ion, oviposition, and postoviposit ion periods. Baton Rouge, Louisiana. 1 9 7 0 - 7 1 ...... 44
8. Oviposition initiation, oviposition cessation, and mortality of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. .... 46
9. Mean longevity of reproductive and nonreproductive laboratory-reared female bean leaf beetles. Baton Rouge, Louisiana. 1970-71...... 51
10. Comparative mortality of reproductive and non- reproductive laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. . . . . 55
11. Mean number of egg-laying days/female and the proportion of the oviposition period in which eggs were laid by laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71...... 57
v i i Page
12. Mean egg production of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. . . 58
13. Mean numbers of eggs laid/laying day, eggs laid/day in the oviposition period, and eggs laid/day through life in laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71...... 60
14. Mean daily of laboratory-reared bean leaf beetles from day 11 to day 50 after emergence. Baton Rouge, Louisiana. 1970-71...... 62
15. Mean daily oviposition of laboratory-reared bean leaf beetles through four 10-day intervals beginning day 11 after emergence and day 1 of the oviposition period. Baton Rouge, Louisiana. 1970-71. .... 63
16. Mean daily oviposition of laboratory-reared bean leaf beetles from day 1 to day 40 after the initiation of oviposition. Baton Rouge, Louisiana. 1970-71. . 66
17. Cumulative egg production of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. . 70
18. Oviposition data for 10 unmated laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1 9 7 0 - 7 1 ...... 72
19. Mean weights in milligrams of field-collected bean leaf beetles differing in color and marking pattern. St. Landry and Catahoula Parishes, Louisiana. 1971...... 73
20. Response of bean leaf beetle color forms to topical applications of methyl parathion. Baton Rouge, Louisiana. 1971...... 76
21. Percentage parasitism of bean leaf beetles by Celatoria diabroticae (Shimer). St. Landry Parish, Louisiana. 1972...... 80
22. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 100° F. Baton Rouge, Louisiana. 1972...... 81
23. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 95° F, Baton Rouge, Louisiana. 1972...... 84
v i i i Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 90° F. Baton Rouge, Louisiana. 1972. .... 85
Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 80° F. Baton Rouge, Louisiana. 1972-73. . 87
Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 70° F. Baton Rouge, Louisiana. 1972-73. 89
Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 60° F. Baton Rouge, Louisiana. 1972-73, 91
Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 50 F, Baton Rouge, Louisiana. 1972-73. . . 93
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 100° F. Baton Rouge, Louisiana. 1972. . 96
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 95° F. Baton Rouge, Louisiana. 1972. 98
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 90° F. Baton Rouge, Louisiana. 1972. 99
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 80° F. BatonRouge, Louisiana. 1972-73. 100
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 70° F. Baton Rouge, Louisiana. 1972-73. . 101
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 60 F, Baton Rouge, Louisiana. 1972-73. 103
Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 50° F. Baton Rouge, Louisiana. 1972-73. 104
ix Page
36, Bean leaf beetle m ortality after 7 days exposure to constant temperatures ranging from 50-100° F. Baton Rouge, Louisiana, 1972...... 106
x LIST OF FIGURES
Log time-probit lines showing progression of laboratory-reared bean leaf beetle adult life history stages over time after adult emergence. 47
Log time-probit lines showing progression of oviposition cessation and mortality of laboratory- reared bean leaf beetles after oviposition initiation. 48
Log time-probit line showing mortality of laboratory- reared bean leaf beetles over time after oviposition cessation. o...... 49
Relationship between longevity of reproductive and nonreproductive female laboratory-reared bean leaf beetles...... 52
Log time-probit lines showing mortality of reproductive and nonreproductive laboratory-reared bean leaf beetles...... 54
Mean daily oviposition of laboratory-reared bean leaf beetles through four 10-day intervals beginning (A) day 11 after emergence and (B) day 1 of the oviposition period...... 64
Proportion of laboratory-reared bean leaf beetles in oviposition and proportion of total egg production over time after adult emergence. . . . . 68
Log time-probit lines showing cumulative egg production of laboratory-reared bean leaf beetles. 69
Relative frequency histograms showing change in weight distribution of field-collected females bean leaf beetle populations over time. St. Landry Parish, Louisiana...... 74
P arasitizat ion of female bean leaf beetles by Celatoria diabroticae (Shimer), in collections from St. Landry Parish, Louisiana...... 77
Log time-probit lines showing bean leaf beetle color form mortality in response to exposure to a constant temperature of 100° F...... 82
x i Page
12. Log time-probit lines showing bean leaf beetle color form mortality in response to exposure to a constant temperature of 90° F...... 86
13. Log time-probit lines showing bean leaf beetle color form mortality in response to exposure to a constant temperature of 80° F...... 88
14. Log time-probit lines showing bean leaf beetle color form mortality inQresponse to exposure to a constant temperature of 70 F...... 90
15. Log time-probit lines showing bean leaf beetle color form mortality in response to exposure to a constant temperature of 60° F...... 92
16. Log time-probit lines showing bean leaf beetle color form mortality in response to exposure to a constant temperature of 50° F...... 94
17. Log time-probit lines showing bean leaf beetle mortality by marking pattern in response to exposure to a constant temperature of 100° F. 97
18. Log time-probit lines showing bean leaf beetle mortality by marking pattern in response to exposure to a constant temperature of 70° F. . 102
19. Log time-probit lines showing bean leaf beetle mortality by marking pattern in response to exposure to a constant temperature of 50° F. .... 105
x i i ABSTRACT
Polymorphic forms of the bean leaf beetle, Cerotoma trifu rcata
(Forster), were studied in an attempt to identify variant physiolo gical and behavioral characters and ecological tolerances correlated with observed polymorphisms.
Color forms and marking pattern types of laboratory-reared bean leaf beetles showed no statistically significant differences in mating frequency; however, trends were observed which were consistent between males and females. Polymorphic forms showed l i t t l e v a ria b ility in the duration of adult life history stages or in fecundity. Reproductive females lived significantly longer than the nonreproductive females, but within color forms this difference in longevity could be demon strated only within beige forms. Within reproductive females, only beige could be demonstrated as living longer than crimson, while within the nonreproductive females only salmon could be demonstrated as living longer than beige.
Bean leaf beetle color forms and marking pattern types collected from the field differed significantly in weight. Those beetles exhibiting marking confluence were smaller than those of other pat terns. Within the color forms, only the heaviest, crimson, could be separated from the lightest, salmon and beige.
Differences in parasitism by Celatoria diabroticae (Shimer) were noted among the color forms, yellow apparently being the most suscep tible, and pink the least susceptible to parasite attack.
x i i i Mating frequency, longevity of reproductive forms, and weight all showed strong correlations with intensity of red pigment. Corre lations among these variables attained significance only in longevity x weight.
D ifferential color form and marking pattern type response to temperature was observed, but no temperature-related trends could be established. Yellow forms showed most rapid progressive mortality at all temperatures. In general, marking pattern variants showed a greater similarity of response than did the color forms.
Techniques for rearing the bean leaf beetle in the laboratory are described.
x iv INTRODUCTION
Polymorphic animal suedes have long been of interest to naturalists, taxonomists, geneticists and students of speciation.
The bean leaf beetle, Cerotoma trifurcata (Forster),* an economically important pest of the soybean, occurs in a particularly interesting and complex polymorphic series. It is especially noticeable because of this insect's abundance in soybean fields of south-central
Louisiana.
An e a r lie r study showed that the sexes d iffered in the r e la tiv e frequencies of the various forms and that the relative proportions of these forms showed a cyclic change through the year.
This study was initiated to attempt to correlate polymorphism, the phenotypic manifestation of a genetic base, with physiological and behavioral characters and ecological tolerances within this species.
*Coleoptera: Chrysomelidae REVIEW OF LITERATURE
Bean Leaf B eetle
Background and H istory
The bean leaf beetle, C. trifurcata, was first named and described by Forster (1771) as Chrysomela trifurcata. It was later described by Fabrlclus (1801) and named Crlocerls camlnea. A third name, Galeruca flb u la ta . was published by Germar (1824). The sp e cie s was transferred to the genus Cerotoma by Gemminger and Harold (1876).
The name Cerotoma was first presented by Chevrolat in 1837 and the genus was subsequently described by him in 1843. Although a non- valid synonym, the Fabriclan C, camlnea was designated as the type species by Chapius (1875). The genus Cerotoma has also appeared in the literature in error as Cerotana (Bowditch, 1913) and Ceratoma
(Guerin, 1953).
The common name, the bean leaf beetle, was suggested by Fopenoe
(1889) in recognition of the habit of this species feeding on bean leaves. £. trifurcata has also been called the "three-spotted bean beetle" (Essig, 1926).
Distribution
C. trifurcata has been reported by Chittenden (1899) and Zappe
(1919) as a native North American species. Indeed, like the
Diabrotica (Webster, 1895, 1901, 1902), the genus Cerotoma is probably o f New World o r ig in (Barber, 1945). The bean le a f b e e tle i s a w idely distributed species, inhabiting the entire eastern one-half of the
2 3
United States as far north as New York (Schmelter, 1879) and south
eastern Canada (Horn, 1893; Wickham, 1897), and w ith the western
limits of its distribution being Arizona (Arnett, 1963), Kansas
(Popenoe, 1877), Nebraska (C hittenden, 1897), South Dakota (Anonymous,
1962) and Minnesota (Webster, 1888). The first published report of
th is in sect from Louisiana was by Townsend in 1885, with subsequent
references by Webster (1888) and Chittenden (1897).
Economic Importance
The first reference to a food plant of the bean leaf beetle was
made by Popenoe in 1877, who stated simply that it "eats holes in
the leaves of bush beans." Louisiana's first host record was given
by Webster (1888), who reported it as feeding on garden beans and
cowpeas. The food o f the bean le a f b e e tle is commonly considered to
be restricted to plants of the family Leguminosae (Chittenden, 1892;
Webster, 1894; Wilcox, 1954; and Milliron, 1958). There is, however,
one doubtful report of this beetle feeding on corn (Metcalf and
Flint, 1962), and another on cotton (Folsom, 1936). These are both probable cases of misidentification. Many early papers stressed the resemblance of this species to the spotted cucumber beetle, Diabrotica undecimpunctata howardi (Barber), and cautioned against confusing the
two species (Webster, 1888; Popenoe, 1889; and Chittenden, 1897).
Webster (1894), and Chittenden (1899) have theorized that before
the cultivation of beans, the bean leaf beetle fed upon several wild
food plants, notably tick trefoil, Desmodturn spp., bush clover,
Lespedeza spp., and hog peanut, Apion spp. 4
So far as can be determined, the first published report of the
soybean as a host for the bean leaf beetle was by McConnell (1915).
Both Eddy and N e ttle s (1930) and I s le y (1930) sta ted that the
bean leaf beetle most often attracted attention as a pest of soybean,
cowpea, and snap bean. This was at a time when soybean culture in
America was in its infancy. Later, Isley (1942) stated that the
"increase in soybean acreage has changed the status of this species
as a pest . . . as many as 1800 beetles were collected in 100 sweeps
of a net." Motsinger and his coworkers (1967) point out that this
species "is considered to be a major pest of soybeans in Louisiana" and "the most important insect vector of BPMtf" (bean pod mottle virus).
The feeding of bean leaf beetle larvae and adults is said to have greatest injurious effect on the growth of host plants during
dry weather (Eddy and Nettles, 1930; Isley, 1930), but this type weather is unfavorable for the development of eggs, larvae and pupae.
Isley (1930) reported this insect as being quite tolerant to high
temperatures. Both McConnell (1915) and Isley (1930) have found the bean le a f b e e tle to be more numerous in soybean and cowpea grown on
light, poor soils, but the latter believed this was a direct result of wild host plant abundaneec However, according to Wuensche (1970) and Newsom (1973), soybean grown in heavy cla y s o ils recen tly cleared
from hardwood forests produce the largest bean leaf beetle populations
in Louisiana.
Rearing
I s le y (1930) and Eddy and N e ttle s (1930) almost concurrently published a procedure for rearing the bean leaf beetle. Adults were 5
caged on plants using either lantern globes or battery jars. Eggs
were collected by examining the base of the plant and the upper inch
of soil surrounding the plant. Eggs were incubated on moist paper
toweling in test tubes. Larvae were held in tin salve boxes and fed
either cowpea radicles or bean cotyledons. Pupation occurred in
loose soil in the bottom of the salve boxes. Isley reared 457
beetles through to the adult stage.
For many years there have been no reports of attempts to rear
the bean leaf beetle. Recently, however, Eastman (1973) conducted
experiments with aseptic rearing of the bean leaf beetle on soybean,
pinto bean, and cowpea callus tissues. She discussed possible appli
cations of this method of rearing insects. Cotyledonary tissue of
soybean, cowpea, and pinto bean were compared as larval food for the
bean leaf beetle. She concluded that purple-hull cowpea cotyledons
were superior to either soybean or pinto bean. Feeding larvae on soy
bean and cowpea callus tissue compared favorably with feeding of
cotyledonary tissue of either.
The larvae has been adequately described and figured by Boving
(1930) and Peterson (1960).
Parasitism
Only one parasite is mentioned in the literature as attacking
the bean leaf beetle. McConnell (1915) first reported Celatoria
diabroticae (Shimer) as parasitizing adult bean leaf beetles in
Mississippi, where it killed "only a small percentage of the beetles."
Isley (1930) reared small numbers from field-collected beetles in
Arkansas. Eddy and Nettles (1930) found this insect parasitizing
about 20 per cent of the population in South Carolina. 6
C. diabroticae was first described by Shimer (1871) who named
it Tachina (Melanosphora) diabroticae. It had been found parasitizing
Diabrotica vittata (Fabricius). Since that time this species has been
found parasitizing several additional species of Diabrotica. including
D. undecimpunctata howardi. D. virgifera LeConte, D. trivittata
Mannerheim, and D. b a ltea ta LeConte (Thompson, 1943). Although f ir st
discoveries of C. diabroticae parasitizing C. trifurcata occurred much
e a r lie r , Sweetman (1958) reported that th is p arasite confined i t s
attacks to the genera D iabrotica and Acalymma.
Insecticide Resistance
Jensen and coworkers (1971) have shown that it may be possible
for the bean leaf beetle to build up low levels of resistance to
chlorinated hydrocarbon, organophosphate and carbamate insecticides.
This, in fact, has occurred in several species of Diabrotica. notably
the western corn rootworm, D. longicornis (Say), and the northern
corn rootworta, D. virgifera (Hamilton, 1965; Blair and Davidson, 1966;
Patel and Apple, 1966; and Ball, 1968). Both species have developed
resistance to aldrin and diazinon and D. virgifera has developed
resistance to phorate.
If pesticide resistance can be developed within a species, is it
not possible for this resistance to be correlated with other poly
morphisms? This has been demonstrated in at least three species.
McEwen and Splittstoesser (1964) have found that both blue and green
larvae of the cabbage looper, Trlchoplusla nl (Hubner), occur in nature.
Color is genetically controlled with green resulting from dominance.
This genetic color factor was found to be associated with DDT 7
"sensitivity" in this species. Blue larvae were more susceptible than green, with larvae being intermediate in their susceptibility.
Blood color differences were found to result from differences in the co lo r o f the haemolymph.
Reichmuth (1967a, 1967b) found DDT susceptibility in the larvae
Leptinotarsa decemlineata (Say) to be correlated with the presence of carotinoid pigments.
Gast (1961), studying differential susceptibility of larvae of the corn earworm, Heliothis zea (Boddie), to DDT, found that lighter- colored larvae "appeared" to be more susceptible than the darker larvae. All larvae were treated at the same rate (not cited) and percent mortality calculated. The mortality of yellow larvae was 64 percent, black 33 percent, with green and red forms intermediate.
The data were not subjected to statistical analysis.
Polymorphism
All authors state that color in the bean leaf beetle is quite variable. However, one paper shows an inconsistency with the others.
Anonymous (I960) described the spots as red or brown when they in actuality are black.
Most authors agree that the c o lo rs range from yellow to red or yellow to brown. Two papers, however, make mention of doubtful color forms--green (Petty and Wainscott, 1961) and light green
(Norman, 1963).
In a previous study, Herzog (1968) classified the bean leaf beetle color variations into 5 arbitrary color categories as follows: beige (probably the "wild type"), pink, salmon, orange, and crimson. 8
Horn (1893) has given a description of the bean leaf beetle marking pattern and its variants.
"The usual coloration of the elytra consists of a triangular scutellar spot, which often sends a narrow stripe along the base to the humeri. From the umbone a moderately wide stripe extends nearly to apex; this stripe is often interrupted. Near the suture are four subquadrate spots arranged in a quadrangle; near the apex are two smaller spots."
"The variations from this observed are as follows: the larger spots forming the quadrangle may be divided longitudinally, producing two linear spots in the place of one. On the other hand, these spots may be lo n g itu d in a lly con flu en t, so that the two on each elytron form a short vitta. Specimens may occur with pale elytra with merely a small scutellar triangle and a small humeral black spot."
The complex series of marking pattern variants was reduced by
Herzog (1968) to 4 for simplicity: normal, broken vitta, marking con fluence, and marking suppression. The "normal" marking is that pre sumed to have been the ancestral marking pattern. It consisted of 3 pairs of median spots and a pair of lateral elytral vittae. Two p attern types resu lted from reduction and one from extension and con vergence of these markings.
Herzog (1968) conducted the only series of experiments which were designed specifically to study color and marking pattern vari ations in the bean leaf beetle, A discussion of the marking pattern types of C. trifurcata and their sim ilarities with those of other species was presented. Statistically significant differences were reported in the proportions of different color and marking pattern 9 types between males and females, between collecting localities, and over tim e.
Results showed that the beige and pink color forms were found in greater proportions in the males while orange and crimson were found almost exclusively in females. The salmon form was found with equal frequency in both sexes. The frequencies observed for males and females, respectively, were as follows: beige, 80.5 and 56.7 per cent; pink, 9.0 and 5.9 per cent; salmon, 7.2 and 7.3 per cent; orange, 3.0 and 10.7 per cent; and crimson, 0.1 and 17.1 per cent.
The normal and confluent marking patterns were found with greater frequency in males, while the broken vitta character occurred with greater frequency in females. Marking suppression occurred with approximately equal frequency in both sexes. The frequencies observed for males and females, respectively, were as follows: normal, 59.7 and 43.1 per cent; broken vitta, 6.8 and 32.3 per cent; marking confluence, 17.9 and 9.5 per cent; and marking suppression
14.9 and 15.5 per cent. He concluded that sex linkage was indicated in the Inheritance of color and marking pattern in the bean leaf b e e tle .
Only the crimson form appeared to be associated with locality, while all four marking pattern types showed differing frequencies at different collection sites.
Analyses o f color pattern over time showed that there was a significantly larger proportion of the beige color form in late summer populations than in overwintering and spring populations.
Fluctuations of other color classes were not found to be significant as changes in the beige form were distributed across the four other 10 color classes. Marking pattern variations showed the same general trends, however, all 4 marking patterns showed significant change in frequency. The decrease in the frequency of the beige form over the winter would seem to substantiate Frost's (1959) statement that
"overwintering adults are often darker in color than the summer generations o f the same sp ecies ..."
Weight determinations were made for the 5 color classes. Signi ficant differences were found, with weight increasing with increases in red pigment.
Mimicry
Webster (1895, 1896) has brought up the subject of intrasub family mimicry between the genera Cerotoma and Dlabrotica. He has postulated that both Cerotoma ruficornis sexpunctata Horn and C. trifurcata mimic D0 undecimpunctata howardi. Similarly, he suggests that another species of Cerotoma mimics Paranapiacaba trieincta (Say).
Finally, Cerotoma arcuata Olivier is supposed to be mimicked by
Lema crucifera Clark.
Current Research
Most current research concerning the bean leaf beetle is centered around the development of alternate methods of control such as resis tant varieties (Jensen, 1971; Clark and coworkers, 1972) and trap cropping (Jensen and Newsom, 1972).
There is increasing concern over insect transmission of soybean virus diseases. As the bean leaf beetle has been shown to be a vector of virus diseases, particularly the bean pod mottle virus in 11
soybean, research Is underway to study plant-virus-vector relation
ship s (W alters, 1970; Horn and coworkers, 1970).
The genus Cerotoma is presently under taxonomic revision by
Dr. R. F. Ruppel of Michigan State University.
An inclusive bibliography of the literature concerning C.
trifurcata and other species of Cerotoma is in preparation under
the direction of Dr. M. Kogan of the University of Illinois and the
Illinois Natural History Survey.
Polymorphism
Richards (1961), in a discussion of insect polymorphism, stated:
"A random collection of any species of insects is likely to show extreme variation which may be due to all of, or any combination of, four principal causes: the occurrence together of developmental stages; differences between the two sexes; the effects of a variable environment on the developing population; the effects of inter breeding amongst a stock which is subject to recurrent mutation and is not genetically uniform." By Ford’s (1940) definition, genetic polymorphism is "the occurrence together in the same h a b ita t o f two or more discontinuous forms of a species in such proportions that the rarest of them cannot be maintained merely by recurrent mutation."
Color pattern polymorphisms have long been recognized among the insects, particularly among the Lepidoptera. There is also a con siderable body of literature concerning polymorphism in Coleoptera, particularly concerning species of the families Chrysomelidae and
Coccinellidae. Herzog (1968) presented a discussion of polymorphism in insects. 12
Seasonal Variability
Tower (1906, 1910) in a comprehensive study of 42 "species" of
Leptinotarsa found that all species studied exhibited seasonal changes in the frequency of some character. L. decemlineata Say was
found to exhibit the most extreme seasonal polymorphisms.
Timofeeff-Ressovsky (1940) studied the relative frequencies of
2 color forms of Adalla bipunctata Linnaeus. He found a significant change in the frequencies of the 2 forms from early spring to late autumn and concluded that these forms have opposing selective values at different seasons of the year, enabling the polymorphism to be maintained within the population.
Tan (1949) studied several elytral color pattern types in
Harmonia axyr id is P a lla s, the in h eritan ce o f which was governed by a multiple allelic series. Although he found no significant change in the frequencies of the different forms in populations from winter to spring ". . . some indication of seasonal variation has . . . been noticed , . , from A pril to December ..." T'lis appears ". . . to
indicate the adaptive nature of these types to the climatic conditions."
Seasonal variations in the frequency of some character have been observed also in vertebrates. Gershenson (1945) found that in the hamster, Cricetus cricetus Linnaeus, black form frequency underwent significant changes corresponding to the seasons. Although trends in different geographic regions were in opposite directions, the trends remained essentially constant from year to year. He concluded:
"These changes in the frequencies of black hamsters during winter are caused by th e ir lower (or . . . higher) v ia b ilit y compared to that o f 13
normal hamsters."
Merrell and Rodell (1968) studied populations of the leopard
frog, Rana piplens Schreber. They found that there was a change in
the frequencies o f spotted and unspotted forms over tim e, and con
cluded that there was selection against unspotted forms in over wintering populations (selection coefficient of 0.23-0.38).
Geographic Polymorphism
Numerous examples of microgeographic and geographic polymorphism have been reported in the literature. Most of the species of
Leptinotarsa studied by Tower (1906) exhibited geographic variations
to a greater or lesser extent.
Crioceris asparagi (Linnaeus) showed increasing marking confluence
in populations from New York to Washington D.C. (Lutz, 1908). A
similar but opposite change in elytra! spotting occurs in Epilachna
varivestis Mulsant, with marking reduction proceeding south to Mexico,
and marking confluence in creasin g northward to Ohio (Landis and Mason,
1938) .
Zulueta (1932) found that the various color forms of Phytodecta
varlabills Olivier were ", . . found together but in different propor
tions in different localities." In fact, this represents a geographic variation in the sexual dimorphism of the species, as the striped form
is found almost exclusively in the females near Madrid, Spain, while
Granada populations show th is form with almost equal frequency in both
s e x e s.
Tower (1906) found that the extreme seasonal variation of L. decemlineata differed from locality to locality. 14
In studying 55 species of Coccinellidae, Johnson (1910) found a tremendous amount o f lo c a tio n a l and geographic v a ria tio n , but e s p e c ia lly in Hippodamia convergens Guerin, Tan (1949) showed that populations o f Harmonia a x y rid is from d iffe r e n t geographic areas varied in the frequency distribution of various alleles which gave rise to elytral marking patterns in this species.
Eichhorn and Graf (1971) found a complete absence of dark forms of Aphidecta obiiterata Linnaeus in some areas while they occurred commonly in other areas.
A froghopper, the meadow spittlebug, Philaenus spumarius
(Linnaeus), is widely distributed in Europe and North America. Con siderable work has been done on this highly polymorphic species, concerning both the nomenclature of the polymorphic forms and deter mination of their comparative frequencies. Original descriptions of
33 varieties were catalogued by Metcalf (1962). Beregovoi (1970) recognized 14 forms in central Europe, all but 3 of which were found in the central Urals. Farish (1972) recognized 14 forms, all but 2 of which were found in North America, His criterion for the occurrence of a morph in North America was its presence in excess of 0.25 per cent of the total population.
Sexual Dimorphism
Bateson (1895) and Zulueta (1932) both working with Phytodecta variabills found that the frequencies of color and marking oattern types varied in males and females. They reported an almost continuous gradation from "greenish-gray” to red, with the 2 extreme forms predominating. Intermediate forms occurred with only low frequency. 15
The spotted form predominated in males, while the striped form was
prevalent in the females. Of the spotted form, red males and
greenish-gray females were present with greatest frequency.
Aphidecta obliterata Linnaeus shows a pronounced sexual dimor
phism in color (Eichhorn and Graf, 1971), The color of this species
ranges from light yellow to black. Five arbitrary color groups were
constructed to categorize the color variation. All males fell in
groups I and II, with 99.7 per cent in group I. The frequency dis
tribution of the female color groups was as follows: group I, 0.9
per cent; group II, 78.3 per cent; group III, 3.8 per cent; group IV,
10.0 per cent; and group V, 7.0 per cent. Their findings indicate
this insect to be a dimorphic species with dimorphic males and poly
morphic females.
Although Parish (1972) made no mention of sexual variability in
morph frequency in Philaenus spuroarius. Beregovoi (1970) stated:
T,The variation of the females in this species is much greater than
that of the males." Color pattern variation in this species was con
centrated on the dorsal surface of the body, the frons, and the
pleurites. The color pattern of the dorsal surface varied indepen
dently of the frons and pleurites. In the males the frons was invari
ably white, while in females it ranged from white to white with a
longitudinal black stripe. Female pleurites were invariably white, i while those of males were either white or black. There was a high
positive correlation (r=0.93) between the frequencies of black
frons in females and white pleurites in males. This strongly indicates
a unit inheritance of the two characters. 16
Sexual Selection and Assortive Mating
Tower (1906), in studying Lentinotarsa oblongata Tower, L. undecemlineata Stal, and L. decemlineata. determined that assortlve mating was not observed between individuals of different colors or between individuals exhibiting different marking pattern types. How ever, assertive mating was observed with respect to size, i.e ., individuals of the same size mated with greater frequency than did extreme individuals.
Shepoard (1952) studied sexual selection in the moth, Panaxia dominula (Linnaeus). Both males and females exhibit several forms.
He found that females prefer to mate with males belonging to forms other than their own. Males, on the other hand, showed no such preference.
Burns (1966), in a study of populations of Papilio glaucus
Linnaeus, concluded that the monomorphic yellow males mated preferen tially with the yellow form of the dimorphic female rather than with the black mimetic form. However, Pliske (1972), in a subsequent study on the same species, could not duplicate Burns' findings and concluded that no sexual selection by color occurs in P. glaucus.
S ize and Structure
Tower (1906) noted considerable d iv e r sity in the s iz e o f
Lertinotarsa oblongata, L. undecemlineata, and L. decemlineata but this difference in size could not be correlated with color. However, in all species of Leptinotarsa studied there was a "strong" correla tion between all variations in coloration and structural characters.
Eichhorn and Graf (1971) could find no correlation between size and color in Aphidecta obliterata. 17
Sex Determination
Smith (1950) studied the chromosomes of many species of
Coleoptera. He found that the family Chrysomelidae was the most heterogeneous group in the order, w ith d ip lo id numbers ranging from
16 to 36. He found 3 types of sex determination systems in males of this family: XO, XY, and Xy^ (small, parachute-shaped Y con sidered by Smith to be p r im itiv e ), METHODS AND MATERIALS
Color and Marking Pattern V ariations
The arbitrary color classification of Herzog (1968) was modified to include a sixth category--yellow. This category was comprised mainly of individuals which would have been classified as beige in the former study. However, since some marginal yellow- orange forms occurred, it is possible that some yellow individuals would have been classified as salmon in the original grouping. The marking pattern categorization followed was identical to that used
in the former study.
Bean Leaf B eetle C o llectio n s
Bean leaf beetles were collected from mixtures of white clover
(Trifolturn repens L.) and Persian clover (Trifolium resupinatum L.) or from Lee, Dare or Bragg variety soybean (Glycine max (L.) Merr.) by means o f a 1 5 -in . diameter sweep n et. B eetles were placed in
3 x 6 x 15-in. polyethylene bags. Each bag was provided with a
12 x 12-in. sheet of paper toweling and several soybean leaflets. The leaflets provided food and prevented dessication of the beetles by supplying moisture. The paper toweling absorbed any excess water of condensation and provided additional surface area for the beetles to prevent overcrowding. Bags containing beetles were placed in a
2 x 2 x 4-ft. styrofoam ice chest containing ice for field storage and transportation. This was necessitated due to excessive heat in the field which would have resulted in high mortality. It was also
18 19
deemed necessary to Inhibit beetle activity due to extreme crowding
conditions within the polyethylene bags.
The beetles were taken to the laboratory and stored in a
refrigerator at 45° F. until they could be examined.
Rearing Technique
Bean le a f b e e tle s were c o lle c te d from the Richard Farm near
Krotz Springs in St. Landry Parish, Louisiana, on June 20, 22, 27,
and July 7, 1970. Since the females were gravid, they were separated
from the males using sexual differentiation criteria described by
Eddy and Nettles (1930), and Herzog (1968), The beetles were
examined with the aid of a binocular dissecting microscope. The
females were numbered for id e n tific a tio n and co lo r and marking
pattern tyoes were recorded for each. The females were placed singly
in 5-dr. (26 x 55 mm.) shell vials which were subsequently capped with polymerized plastic stoppers. The inserts from the plastic
caps had been removed and several h o les were burned in to the caps with a redhot probe to provide for air exchange. Soybean leaflets
of the Bragg variety were collected from the H ill Farm on the
Louisiana State University campus, Baton Rouge, and placed in vials as food.
Females were held in an environmental cabinet at a temperature of 80+2° F, and 14-hr. light and 10-hr. dark periods. Additionally, all life cycle stages in the rearing process were maintained under the same temperature-photoperiod environmental regimen. Relative humidity remained uncontrolled, but ranged from approximately 25
to 75 per cen t. 20
Every 2 days the beetles were transferred to clean vials and
provided with a fresh food supply. Vials needed to be changed because the buildup of excrement provided a medium for fungal and mold development. Leaflets also required changing because of dessication, beetle feeding activities, and the likelihood of mold development. The vials were washed in a commercial detergent and
oven-dried at 212° F.
If tachinid parasites emerged from the females, the puparia were kept until the adult fly emerged. When the females died without
parasite emergence, they were autopsied by dissection to determine if
parasite larvae were present within the abdomen. Species identifi cation of larvae, pupae, and adults was made when p o ssib le .
The actual rearing technique was a modification of the procedures
used by Isle y (1930) and Eddy and N e ttle s (1930).
Vials and leaflets were checked for the presence of eggs every
other day. In the event that eggs were found, the number of eggs was recorded individually for each female. The eggs were transferred
to 1 -o z. p la s tic rearing cups by means o f a s iz e "00" cam el's hair brush, or very carefully with a fine precision jewelers forceps. A maximum of 5 eggs was placed in each cup for maximum utilization of
space and minimum crowding. The bottom o f each rearing cup was
provided with 5 pieces of 1 x 1-in. paper toweling. The paper towel
ing was saturated with water to maintain a high level humidity within
the cups, thereby keeping the eggs from drying out. A tight-fitting,
plastic-coated cardboard cap was then placed on each cup. After 2
days the eggs were moistened with a 0.5 per cent aqueous cupric 21 s u lf a te (CUSO4) solution to inhibit fungal growth on the eggs which would have resulted in egg mortality.
Approximately 1 day before the eggs hatched, the cups were provided with a larval food source, consisting of cotyledons from fr e sh ly sprouted Crowder pea (Vigna sin e n sis (L.) E n d l.), purple- hull cowpea (V. sinensis), or pinto bean (Phaseolus vulgaris L.) cotyledons. Attempts to use soybean (G. max) and black-eyed pea
(V. sinensis) cotyledons as a larval food source were abandoned.
The food plant sp ecies used depended upon the a v a ila b ility of seed during the lG-month period of this experiment. Seeds were placed between 2 layers of Cellucotton in halves of 6-in. petri dishes.
The Cellucotton was saturated with water and the seed allowed to germinate until it sprouted and the radicle emerged. The seed coat, r a d ic le , and plumule were removed. The cotyledons were immersed in a 0.5 per cent solution of cupric sulfate and allowed to soak for
5 to 10 minutes. This served to surface-sterilize the cotyledons and inhibited fungal growth. Soaking in this solution for a longer period of time resulted in absorption of sufficient cupric sulfate to cause the death of at least a portion of the cotyledonary tissue. The cotyledons were then placed over the eggs with the flat or concave median surface down to insure that the larvae found their food source.
The rearing cups were examined periodically and moisture and food were replinished as needed.
Approximately 1 to 2 days before the prepupal stage was attained the larvae and cotyledon were transferred to rearing cups which had been filled to %-in. with coarse sand to provide a suitable site for pupation. The sand was moistened to provide a high relative humidity. 22
Care was taken not to supersaturate the sand due to the possibility
of the larvae drowning. The cups were examined periodically and the
sand moistened as needed. After the prepupal stage had been attained
the cotyledons were removed.
No data were kept concerning the duration of the incubation
period or the duration of the immature stages.
Pupal holding cups were examined daily for adult emergence.
Emerging adults were transferred singly to holding vials containing soybean le a f le t s and were assigned id e n tific a tio n numbers which
facilitated the maintenance of lineage records. Marking patterns were evident almost immediately upon adult emergence, but i t was found
that the final background color was attained and remained constant only after 5 to 7 days. It was for this reason that pairings for mating attempts were made from 5 to 10 days after emergence of the a d u lts.
At intervals of from 2 to 7 days, as time permitted, the beetles were examined under a low-power binocular dissecting micro scope. The sexes were separated by a method described by Eddy and
Nettles (1930): "The male form possesses a more blunt last abdominal segment than does the female. The male also possesses one more visible segment in the abdomen than the female does." Color and marking pattern were recorded for each individual and pairs were selected for mating attempts. The selected pairs were placed in 5-dr. vials pro vided with soybean leaflets for mating and oviposition. The vials and leaflets were examined daily for oviposition initiation. In the event that the male died before mating occurred, another male was substituted with the hope that mating would occur. This was repeated 23 until mating was achieved or the female died. If the female died before the male, he was recycled in to the pool o f unmated fem ales.
If mating occurred, the male remained with the female until he died or until the female died. In the latter case, he was recycled into the pool of unmated females.
By October the field-collected leaves were senescent and drying, of little value as food. It was decided to use seedling soybean plants as an adult food source. Greenhouse flats, which measured
35.5 x 50.8 x 8.5 cm., were filled to a depth of approximately 5 cm. with commercially available vermiculite (expanded mica). Soybean seed, without regard for variety, were planted and covered with a thin layer of vermiculite. The vermiculite was then saturated with water and subsequently watered daily. After approximately 5 days of when the seed lin g s had reached a h eigh t o f 1 to 3 in . they were pulled from the vermiculite and placed in vials as adult food. The soybean seedling food source was replaced every 2 days as were the soybean leaflets.
As the rearing program neared its termination, egg production reached a point where time limitations no longer permitted handling of the eggs Immediately. Therefore, eggs were placed in a refrigerator at approximately 45° F. while still in the oviposition vials. They were held until time permitted their handling.
Weight Determinations
Beetles were collected from clover near the Richard Farm on
A pril 29 and May 7, from seed lin g soybean on June 8 and from soybean on June 25 and July 16, 1971. Another collection was made from 24 numerous soybean fields in the Harrisonburg--Jonesville--St. Joseph,
Louisiana area on July 14, 1971.
The beetles were separated into the proper color and marking pattern categories with the aid of a binocular dissecting microscope.
They were then weighed on a Mettler type B balance and weights were recorded to the nearest 0.1 mg.
Insecticide Study
Bean leaf beetles were collected 5 times during 1971 from the
Richard Farm in St. Landry Parish. Collections made April 30 and
May 7 by the sweeping method were from mixtures o f w hite (T. repens) and Persian clover (T. resupinaturn). One collection was made by hand-picking from seed lin g soybean on June 8. The fin a l two c o l le c tio n s were swept from soybean on June 25 and July 16. The b e e tle s were brought to the laboratory, sexed, and categorized as to color as described above. Marking pattern variants were disregarded in this study.
The beetles were treated topically with graded dosages of methyl parathion (0, 0-dimethyl 0-p-nitrophenyl phosphorothioate) to attempt to determine differential response of the various color forms to this commonly-used pesticide.
Recrystallized methyl parathion was dissolved in acetone to ob tain a weight by volume stock solution of the toxicant. From this stock solution serial dilutions were prepared by means of volumetric pipetting. The concentrations of toxicant which were used in this study were 0.01, 0.015, 0.02, 0.03, 0.04, and 0.1 .ug./ul. 25
The insecticide-acetone solutions were placed in a micrometer- driven 1 ml. glass syringe fitted with a 1 in., 27-ga. hypodermic needle and calib rated to d e liv e r a volume o f 1 u l. The b eetles were treated in ascending toxicant concentration series, beginning with an acetone control treatment. The beetles were hand-held and the I ul. droplets were applied at the apex of the elytra so that the toxicant in acetone was drawn up under the wing covers.
A maximum of 25 beetles were confined on 2-in. soybean seedlings in 1-pt. plastic-coated cardboard cups fitted with a plastic top. The beetles were held at room temperature (ca. 75° F.). Mortality counts were made at 24 and 48 hours after treatment. The criterion for death at 48 hours was set as the inability to retain or regain the upright p o sitio n .
Temperature Tolerance Studies
Seven temperatures were selected for study of temperature
tolerance response in the bean leaf beetle: 50, 60, 70, 80, 90, 95,
and 100° F, C o llectio n s were made from the Richard Farm on 14 c o l
lecting dates during 1972 allowing for 2 replications per temperature
studied. Date of initiation and the temperature to which each lot of
beetles was exposed are presented in Table 1.
The beetles were separated into their respective color and mark
ing pattern categories with the aid of a binocular dissecting micro
scope. Females were placed in 5-dr. shell vials provided with a food
source of soybean leaflets and capped with a polymerized plastic
stopper. Each lot of beetles was placed in an environmental cabinet which was programmed for the desired temperature. All temperatures Table 1. Dates of installation and number of bean leaf beetles included in replicated temperature tolerance studies. Baton Rouge, Louisiana. 1972.
Temperature Date o f Number o f Study (°F.) Installation B eetles
100° I June 12 701
II August 20 945
95° I July 4 1061
II August 25 1062
90° I June 21 994
II September 16 991
80° I July 12 1209
II July 25 972
70° I July 21 899
II July 31 1091
60° I May 26 734
II August 21 866
50° I June 5 317
II September 21 971 27 within +2° F, with the exception of the 50° test, which fluctuated within +5° F. All tests were conducted under a 14-hr. light and 10- hr, dark photoperiod, again with the exception of the 50° test. Due to lack of environmental cabinet space, this test was conducted in a refrigerator in total darkness.
The food source was replaced and clean holding vials substituted every 2 days.
Longevity was chosen as the criterion for temperature tolerance,
i.e., length of time from initiation to death. The tests were checked daily or every two days for beetle mortality. Dead beetles were removed and recorded at that time.
Again in 1972, field-collected leaves were rapidly approaching senescence by September and were drying rather rapidly, especially at the higher temperatures. Therefore, soybean seedlings were sub stituted as the food source. They were replaced every 2 days in the
80 and 90° t e s t s , tw ice weekly in the 60 and 70° t e s t s , and weekly
in the 50° t e s t . By th is time the 95 and 100° te s ts had been concluded,
P arasitism
Since a significant number of field-collected beetles were to be observed in the course of the temperature tolerance study, a means was provided whereby percentage parasitism for the various color and marking forms was determined. All parasite pupae found within the holding vials were recorded. Further, after death occurred all bodies were autopsied to determine if parasite larvae or pupae were 28 present within the abdomen. Species identifications were made when p o ssib le .
Statistical Analyses
The combination of the 6 color and 4 marking pattern types com
prised a 6 x 4 factorial arrangement of treatments in a complete block design replicated over 4 generations. Due to widely variant c la ss and subclass s iz e s , a l l data re su ltin g from rearing stu d ie s,
including life cycle and egg production, were subjected to least-
squares (LSMLGP) analyses of variance, adjusting for covariance as necessary (Least-Squares and Maximum Likelihood General Purpose
Program o f the Louisiana S ta te U n iversity Computer Research Center)
(Harvey, 1960, 1969).
Due to the extremely large number of beetles involved in lon gevity studies (productive + nonproductive * 1717) and weight deter mination study (4945), the subclass (color-marking) means for each generation or replication, respectively, were used as the experimental units. This resulted in several missing generation x productivity x color x marking subclasses in the longevity study and 1 missing replication x color x marking subclass in the weight study. There
fore, these data were subjected to least-squares analyses of variance and corrected for unequal numbers o f observations per subclass by
fitting a partial linear regression for each set of data using the number of beetles that made up each experimental unit as the covariable.
Parasite data obtained in the course of the' temperature tolerance
study series were analyzed by means of LSMLGP, using overall per cent 29 parasitization for each color-marking subclass across all collection dates as the observational unit, and correcting for unequal subclass size as above.
Data concerning preovipodition, oviposition, and postoviposition periods, period of reproductive activity, reproductive maturity, longevity, and egg production were subjected to probit analysis
(Finney, 1947; Daum, 1970) to fit LTP (log time-probit) regression lines and derive ET (effective time) values. Data obtained from the temperature tolerance study were also subjected to probit analysis.
LTP regression lines and LT (lethal time) values were used to compare the response of the 6 color and 4 marking pattern types.
Mean difference separation was accomplished by means of Duncan's
M ultiple Range T est.
Correlation coefficients were calculated and analyses of corre lation were conducted by conventional methods.
Symbols and A bbreviations
Several symbols are used within the text without further explanation. The symbol ns indicates nonsignificant mean differences at the .05 level of probability; a single asterisk (*) indicates mean differences significant at the .05 level; and a double asterisk (**) indicates significance of mean differences at the .01 level of probability. For the sake of simplicity and the conservation of space, i t was deemed ad visab le to abbreviate color and marking pattern types in tables and figures. The abbreviations used are as follows: yellow (Ye), beige (Be), pink (Pi), salmon (Sa), orange (Or), and crimson (Cr) for the colors, and normal (No), broken vitta (Vi), 30 marking confluence (Co), and marking suppression (Su) for the marking p attern s. RESULTS AND DISCUSSION
Laboratory Rearing
The bean leaf beetle was reared through A generations in the
laboratory, a total of 5,407 Individuals attaining the adult stage.
A total of 111,776 eggs was collected from 820 reproductive females.
ERR Storage
At times during the rearing program egg production exceeded that which could be effectively handled. Eggs still contained in
the oviposition vials were placed in a refrigerator at approximately
45 to 50° F. until time permitted their handling. Although no quantification was made, observation showed that eggs could be refrigerated under these conditions for a maximum of 2 weeks before d e ssic a tio n became a lim itin g fa cto r. It is p o ssib le that eggs could
be held for longer periods of time in completely closed vials, or in an environment in which humidity was controlled at the optimum.
There seemed to be no apparent change in egg viability up to
two weeks of storage. Eggs hatched normally in 6 to 7 days after removal from cold storage and incubation at 80° F.
Larval Food Source
As mentioned in an earlier section, a variety of larval food sources was utilized in the rearing program. Larval rearing was
in itia te d using Crowder pea cotyledons as a food source. These proved satisfactory but were in short supply. There was some problem with fungal contamination. Purple-hull cowpea cotyledons
31 32 were substituted and proved very satisfactory. Developmental rates were comparable, but there was less trouble with fungal contamination.
When supplies of purple-hull cowpea ran short both, soybean and black eyed pea were used, but with little success. Neither proved satis factory. The majority of the larvae failed to initiate feeding on soybean cotyledons. Larvae fed on the black-eyed pea cotyledons, but the cotyledons deteriorated much too rapidly to support the larvae for an adeouate length of time. Pinto bean was the final food source used in this study. It proved to be a satisfactory larval food with both advantages and disadvantages over cowpeas. Cowpea cotyledons deteriorated a little more rapidly than did those of the pinto bean, the latter sometimes not needing to be replaced for a week or more.
There was almost no fungal contamination problem with pinto bean.
However, the developmental rate of the larvae was much more variable with pinto bean than with cowpea.
These observations support those of Isley (1930) who found that bean leaf beetle larvae develop more rapidly on cowpea cotyledons than on those of lima bean or soybean. In a recent study Eastman
(1973) Quantified this relationship, showing a larval development of 10.2 days on purple-hull cowpea, 13.6 days on soybean, and 14.8 days on pinto bean. She also showed variable larval mortality, 8.5 per cent on purple-hull cowpea, 37.5 per cent on soybean, and 44.8 per cent on uinto bean.
Mating
In this study a total of 1717 females reached the age of 5 to
7 days at which time pairings were made for mating. Of these, 820 33 females mated and produced eggs. Table 2 shows the number o f reproductive females obtained In the color-marking pattern categories.
A total of 2,546 pairings were made Including successive attempts with females for which previous pairings were unsuccessful.
Overall 36.2 per cent of the pairings were successful In mating. The proportions of mating success for male and female bean leaf beetles of the various color and marking pattern types are presented in Table
3. Although no statistical significance in mating response was obtained for color or marking patterns in either males or females, certain observations converning trends can be made. Crimson females showed the greatest mating frequency under conditions of forced mating (Table 3A). However, if crimson females are disregarded,* yellow forms, both male and female, apparently showed the greatest frequency of mating success. Both pink males and females showed the lowest mating frequency. Hales and females exhibiting the marking confluence pattern showed the greatest proportions of successful m atings.
Table 4 shows proportions of success for female color x male color and female marking x male marking pairings. Although sta tistical analysis showed no significance, there was a tendency for all females to mate more frequently with yellow males than with males of other colors. Males showed no such tendency.
Considering marking patterns, the male marking confluence x female marking confluence pairings showed the greatest positive mating response. With two exceptions, males and females tended to
*Only 1 crimson male was obtained in this study. It failed to mate, and was not Included in the statistical analysis. 34
Table 2. Total number of females, number of and per cent reproductive females obtained in a bean leaf beetle laboratory rearing program. Baton Rouge, Louisiana.. 1970-71.
Marking Pattern Color Color T otal No Vi Co Su
Ye Number Mated 55 46 12 32 145
Total Number 116 100 18 53 287
Per Cent Mated 47.4 46.0 66.7 60.4 50.5
Be Number Mated 119 102 10 29 260
Total Number 232 235 28 64 459
Per Cent Mated 51.3 43.4 35.7 45.3 46.5
Pi Number Hated 30 22 1 7 60
T otal Number 61 45 14 11 131
Per Cent Mated 49.2 43.9 7.1 63.6 45.8
Sa Number Mated 54 47 13 36 150
T otal Number 117 98 22 66 303
Per Cent Mated 46.2 48.0 59.1 54.5 49.5
Or Number Mated 68 69 13 13 163
T otal Number 137 147 23 27 334
Per Cent Mated 49.6 46.9 56.5 48.1 48.8
Cr Number Mated 13 26 2 1 42
T otal Number 39 57 2 5 103
Per Cent Mated 33.3 46.5 100.0 20.0 40.8
Marking Number Mated 339 312 51 118 820 Pattern T otal Total Number 702 682 107 226 1717
Per Cent Mated 48.3 45.7 47.7 52.2 47.8 35
Table 3. Proportions of pairings resulting in mating for female and male laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. '
(A) Females
Color Marking Marking Pattern Ye Be Pi Sa Or Cr Mean
No .321 .348 .379 .315 .373 .309 .341
Vi .360 .295 .340 .318 .369 .400 .347
Co .561 .200 .000 .371 .450 .767 .382
Su .451 .353 .377 .421 .396 .260 .376
Color Mean .423 .299 .260 .356 .397 .434 .362
(B) Males2)
Color Marking Marking Pattern Ye Be Pi Sa Or
No .403 .373 .277 .400 .304 .352
Vi .489 .410 .331 .398 .229 .373
Co .513 .363 .345 .323 .600 .429
Su .478 .354 .173 .314 .145 .293
Color Mean .473 .357 .282 .358 .320 .362
^Analysis of variance presented in Appendix Table 1.
Only 1 crimson male was obtained in this study; it failed to mate and was not included in the s t a t i s t i c a l a n a ly sis. 36
Table 4. Proportions of bean leaf beetle pairings by color and by marking pattern resulting in mating. Baton Rouge, Louisiana. 1970-71. '
(A) Color
Female Color Male Male Color Color Ye Be Pi Sa Or Cr Mean
Ye .509 .389 .311 .454 .530 .644 .473
Be .477 .342 .292 .383 .339 .418 .375
Pi .295 .214 .242 .286 .394 .259 .282
Sa .445 .334 .228 .241 .469 .435 .358
Or .390 .217 .226 .418 .252 .415 .320
Female Color Mean .423 .299 .260 .356 .397 .434 .362
(B) Marking Pattern
Male Female Marking Pattern Male Marking Marking Pattern Vi Co Su MeanNo
No .348 .313 .435 .311 .352
Vi .347 .307 .368 .471 .373
Co .432 .398 .508 .375 .429
Su .236 .370 .217 .348 .293
Female Marking Mean .341 .347 .382 .382 .362
^Analysis of variance presented in Appendix Table 1. 37 mate most readily with partners exhibiting marking confluence.
Reciprocal pairings of marking suppression x broken vitta attained greater success.
These apparent preferences cannot be construed to be color or marking pattern selection or assertive mating since the mating response was measured on a nonrandom mating b a sis. It i s p o ssib le that matings allowing free choice would show that there is actual color or marking pattern selection (assertive mating) in this species.
On the other hand, such a study may show that the apparent differences are merely artifacts of the experimental procedure.
Disregarding attempted matings between individuals of the same color, reciprocal color x color pairings showed a significant positive correlation (r=>,66l) in comparative mating success (Appendix Table 2).
This correlation indicates that males and females of the various colors react similarly with nonidentical individuals of the opposite sex. However, comparison of corresponding reciprocal marking x marking pairings yielded no significant correlation (r=.509) in their comparative mating success (Appendix Table 4), This indicates that there was some dissim ilarity in mating response of males and females with nonidentical individuals of the opposite sex.
There was a significant interaction in the mating response of female color-marking pattern combinations. In Table 3A it may be seen that there was a highly differential response of the marking confluence pattern within the 6 color forms. Table 2 shows that only
7 per cent of the pink females showing marking confluence mated, while both crimson females exhibiting this pattern were reproductively successful, 38
When the reproductive success of the 20 color-marking pattern
types occurring in males (Table 3B) was compared with that of the
corresponding female forms (Table 3A) (e.g., YeNo vs. YeNo, YeVi
vs. YeVi, etc.) the resultant low and nonsignificant correlation
coefficient (r<=.l86) (Appendix Table 6) indicated a dissim ilarity
in mating frequency of similarly-colored males and females.
Reproductive females required from 1 to 8 successive pairings with males before impregnation was achieved with a mean of 1.23
(Table 5); 83.2 r>er cent of the females required only 1 male. No
significant differences were detected between polymorphic forms.
The mean male age at mating in this study was 21.9 days, with a range of from 5 to 141 days. It was found that males were capable
of multiple matings. Of the 820 matings achieved, 67 were second matings for a male and 4 were third matings. It was not determined, however, if females were also capable of successive matings.
Adult Life History
No significant differences were detected among the various color pattern categories in the duration of the preoviposition, oviposition,
or postoviposition periods (Table 6). If indeed there were real differences, they were probably obscured by extreme within-class variability.
The mean preoviposition period was observed to be 20.9 days
(Table 6A), with a range of from 8 to 154 days. This was a con siderably longer time than those reported by Isley (1930) and Eddy and Nettles (1930) of 8.9 and 7.4 days, respectively. There are
several possible explanations for this wide discrepancy. First, the 39
Table 5. Mean number of males required before impregnation of laboratory-reared female bean leaf beetles was achieved. Baton Rouge, Louisiana. 1970-71.1)2)
Color Marking Marking Pattern Pattern Ye Be P i Sa Or Cr Mean
No 1.25 1.24 1.12 1.44 1.40 1.16 1.27
Vi 1.20 1.28 1.23 1.28 1.27 1.25 1.25
Co 1.16 1.26 1.11 1.29 1.21 1.16 1.20
Su 1.20 1.32 1.24 1.16 1.27 1.15 1.22
Color Mean 1.20 1.27 1.17 1.29 1.29 1.18 1.23
l^Range 1 - 8 males 2) Analysis of variance presented in Appendix Table 8. No statistical significance detected. 40
Table 6. Mean duration of preoviposition, oviposition, and post- oviposition periods of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.^
(A) Days in Preoviposit ion Period^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 20.5 24.1 19.0 22.3 18.1 12.9 19.5
Vi 18.3 28.7 35.4 22.2 21.0 20.9 24.4
Co 15.1 18.3 18.1 25.8 20.7 11.0 18.2
Su 24.0 19.4 28.1 17.4 19.5 20.6 21.5
Color Mean 19.5 22.6 25.2 21.9 19.8 16.3 20.9
(B) Days in Oviposition Period^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 14.2 12.4 13.0 12.2 11.9 9.6 12.2
Vi 10.9 12.3 12.1 14.5 16.2 10.5 12.7
Co 15.0 19.7 20.1 9.9 18.3 6.2 14.8
Su 13.4 11.6 12.2 14.7 11.4 13.8 12.8
Color Mean 13.4 14.0 14.3 12.8 14.4 10.0 13.2
■^Analyses of variance presented in Appendix Table 11. No statistical significance detected.
^Range 8 - 154 days 3) Range 1 - 112 days 41
Table 6 (continued).
(C) Days in Post oviposit ion Period^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 7.8 9.0 9.4 11.5 8.9 7.6 9.0
Vi 10.7 6 .1 8.2 6.7 8.7 19.9 10 .0
Co 3.4 25.5 7.9 10.4 6 .1 4.7 9.7
Su 5.4 8.9 8.7 8.4 5.8 4.5 7.0
Color Mean 6 .8 12.4 8.5 9.3 7.4 9.2 8.9
^Range 1 - 199 days 42 values presented by these authors are based on observations on 18 and
5 females, respectively, a rather small sample size. Secondly, all female beetles in their studies were fed on plant seedlings (cowpea by Isley and bean by Eddy and Nettles). This, according to Isley, greatly accelerates the initiation of oviposition. In the present study the first generation and a part of the second were fed on soy bean foliage. Finally, both Isley and Eddy and Nettles paired males and females for mating soon after emergence. In this study males and females were isolated for a period of from 5 to 10 days to allow for the development of the final background color before pairings were made for mating.
Isley (1930) states: "The duration of the preoviposition period was found to be dependent upon the availability of host seedlings as food for beetles when they first transformed to the adult stage." He did not secure mating with 11 pairs fed on cowpea foliage. Of 26 pairs fed on cowpea seedlings, 18 mated and the females laid eggs.
It has been demonstrated that bean leaf beetles fed on soybean leaves in the laboratory mated and laid eggs. This would be expected because adults emerging in the field in June and July cannot find seedling soybean for food, yet they mate and lay eggs.
Although bean leaf beetles w ill reproduce when fed a diet of soy bean leaves, frequency of mating decreases and mortality increases sharply with the onset of plant senescence. If soybean seedlings are substituted as food, mating frequency increases and the length of the preoviposit ion period decreases sharply.
Another observation from Isley's study indicated that when females were fed on cowpea foliage for a period of 3 weeks and then 43
transferred to cowpea seedlings little feeding occurred and there was
no oviposition. When the transition was made from soybean foliage to
seedling soybean in this study, a substantial number of females
ranging in age from 14 to 46 days had not yet mated. When they were
transferred to soybean seedlings many mated and oviposition was in iti
ated in from 4 to 10 days.
The proportion of the total adult life span spent in the preovi-
position period was .548 (Table 7A).
The mean duration of the oviposition period was found to be 13.2
days (Table 6B), ranging from 1 to 112 days. This corresponds rather well with Isley's and Eddy and Nettles' findings of 16.9 and 15.6
days, respectively. The proportion of the total adult life span *
spent in the oviposition period was .323 (Table 7B).
The mean postoviposition period was 8.9 days (Table 6C), with a
range of from 1 to 199 days. Isley 's value of 25.7 days is much
greater than this, while Eddy and Nettles reported 6.5 days. The
proportion of the total adult life span in the postoviposition period was .193 (Table 7C).
At the age of 8 days, 10 per cent of the females had begun ovi
position, while by 16 days, 50 per cent were laying eggs (Table 8 ,
Figure 1). At 2 days after the onset of oviposition, 10 per cent had
stopped laying eggs and 50 per cent after 8 days (Figure 2), or at age 13 and 26 days, respectively (Figure 1). One day after the cessation of oviposition 10 per cent of the females had died, and 50
per cent by 4 days (Figure 3). This was at 5 and 14 days, respec
tively, after oviposition was initiated (Figure 2), or at 15 and 33 days of age, respectively (Figure 1). u
Table 7. Proportion of laboratory-reared female bean leaf beetle life cycle spent in preoviposition, oviposition, and post oviposition periods. Baton Rouge, Louisiana. 1970-71.^
(A) Proportion of Life in Preoviposit ion Period2^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No .537 .563 .546 .568 .573 .528 .552
Vi .541 .569 .527 .564 .553 .610 .561
Co .541 .594 .452 .523 .516 .557 .530
Su .553 .602 .505 .573 .549 .517 .550
Color Mean .543 .582 .508 .557 .548 .553 .548
(B) Proportion of Life in Oviposition Period^)
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No .330 .313 .297 .309 .306 .326 .314
Vi .322 .306 .325 .325 .310 .336 .321
Co .353 .257 .399 .334 .324 .282 .325
Su ,317 .310 .361 .301 ,324 .382 .333
Color Mean .330 .297 .346 .317 .316 .332 .323
^Analyses of variance presented in Appendix Table 13. No statistical significance detected. 2>Range .07-.99 ■^Range .01-.96 45
Table 7 (continued).
(C) Proportion of Life in Postoviposition Period^)
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No .193 .194 .231 .196 .189 .221 .204
Vi .215 .193 .210 .166 .189 .152 .187
Co .154 .222 .172 .216 .200 .243 .201
Su .188 .172 .201 .186 .191 .134 .179
Color Mean .187 .195 .203 .191 .192 .188 .193
^Range .01-.96 Table 8 . Oviposition initiation, oviposition cessation, and mortality of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.1)
95% Confidence 95% Confidence " 1 0 " 5 0 Days Interval Days Interval
Oviposition Initiation^) 7.8 5.9- 9.4 16.1 14.2-17.8
Oviposition Cessation^) 13.0 12.4-13.6 26.5 25.9-27.0
Mortality 15.3 14.4-16.2 33.3 32.5-34.1
3) Oviposition Cessation 1 .8 1.4- 2.3 7.7 6 . 8 - 8.5
Mortality^) 4.6 4.2- 5.0 14.3 13.8-14.9
Mortality^) 0.9 0.7- 1.2 4.1 3.5- 4.7
^Computed from data shown in Appendix Tables 30 and 31.
Measured from adult emergence; see Figure 1.
Measured from oviposition initiation; see Figure 2. 4) Measured from oviposition cessation; see Figure 3. Per Cent In itiation 99 90 iue . o tm-rbt ie soig rgeso o laboratory-reared of progression showing lines time-probit Log 1. Figure dl emergence. adult en ef ete dl lf itr sae oe tm after time over stages history life adult beetle leaf bean as fe Tvmergence after Days 0 0 0 0 200 100 50 20 10 > I ' " B " B B " ■ "I ' I ' l> C B A - Oviposition - A - Mortality - C - Oviposition - B Cessation Initiation
--j -t> Per Cent In itia tio n 99 90 50 10 1 1 iue . o tm-rbt ie soig rgeso o oviposition of progression showing lines time-probit Log 2. Figure 2 ete atr vpsto i itiation in oviposition after beetles esto ad otlt o lbrtr-erd en leaf bean laboratory-reared of mortality and cessation as fe Oioiin Initiation Oviposition after Days 5 10 20 50 - Oviposition - A - Mortality - B 100 Cessation 200
500
Figure 3. Log time-probit line showing mortality of laboratory-reared bean leaf beetles over time after oviposition cessation.
m"" """ "i 1 i ' ii « " r 1" 1,1 n 1 i .1 .2 .5 1 2 5 10 20 50 Days after Oviposition Cessation 50
Although the mean preoviposition period was 20,9 days, examina
tion of the LTP oviposition initiation response curve (Figure 1)
shows that 50 per cent of the females had begun laying eggs in
approximately 16 days. Therefore, the rather long mean preoviposition
period was a probable reflection of several extreme values, e.g., 154
days.
Longevity
There was a significant difference in the longevity of reproduc
tive (those females which mated and laid eggs) and nonreproductive
females (those which failed to mate and laid no eggs). The mean lon gevity of reproductive females was 41.2 days and that of nonreproduc-
tive females 35.8 days. This difference is somewhat artificial in that all reproductive females were included in the analysis, but of the nonreproductive females only those were included which survived to achieve their final background color. Therefore, the nonreproduc-
tive longevity mean is probably biased upward, with the true value probably fa llin g somewhere below 35.8 days. These data are in direct contradiction to Isley 's (1930) report that reproductive female lon gevity was much shorter than that of nonreproductives.
A significant interaction occurred between productivity and color. This interaction is shown numerically in Table 9 and graph
ica lly in Figure 4. Yellow, salmon, orange, and crimson forms reacted similarly in longevity whether reproductive or nonreproductive. How ever, beige and pink reproductive forms lived much longer than the corresponding nonreproductive forms, 48,2 and 47.1 days as compared to
30.2 and 34.8 days, respectively. However, this difference was 51
Table 9, Mean longevity of reproductive and nonreproductive laboratory-reared female bean leaf beetles. Baton Rouge, Louisiana. 1970-71.1)2)
Longevity in Days^
Reproductive Nonreproductive Females Females
Color Ye 37.7 31.0
Be 48.2 ab 30.2 ac
Pi 47.1 34.8
Sa 42.6 45.6 c
Or 38.1 39.5
Cr 32.0 b 33.8
Marking Pattern No 38.7 36.6
Vi 46.1 35.1
Co 39.8 35.4
Su 39.2 36.1
Overall — 41.0 35.8
^A nalysis of variance presented in Appendix Table 15.
^Range for reproductive females 10 - 212 days. Range for nonreproductive females 8 - 198 days.
^Mean longevities connected by the same letter differ significantly at the 5% protection level according to Duncan's Multiple Range Test. F igure 4 Relationship between longevity of reproductive and nonreproductive female laboratory-reared bean leaf beetles. Reproductive 50 1 Females
0 - Nonreproductive Females
45 ■
CO » 40 ■ •He •U •H > 0) t>0 oC tJ 35 ■ < H >
30 ■ 52b Ye Be Pi Sa Or Cr Color 53 statistically significant only within the beige forms.
The series yellow, beige, pink, salmon, orange and crimson represents increasing red pigment. Although significant differences in longevity of reproductive forms were detected only between beige and crimson beetles, it is interesting to note that the longevity of these forms correlate well with the amount of red pigment present.
There was an apparent decrease in length of life with increases in red pigment. The yellow form was apparently discontinuous, as was hypothesized by Bateson (1895) for the yellow form of Phytodecta variabilis.
The correlation between longevity and the intensity of red pig ment was inconsistent in the nonreproductive forms. Significance in the nonreproductive forms was detected only between beige and salmon b eetles.
The relationship between the progressive mortality of reproduc tiv e and nonreproductive females is shown in Figure 5. Comparative m ortalities were significant from 0 through 60 per cent and from 90 through 100 per cent. Twenty per cent mortality occurred in non reproductive females at 13.7 days while 20.0 days were required in reproductive females; 50 per cent mortality occurred at 28.3 and 33.3 days, respectively; 90 per cent mortality occurred at 84.9 and 72.5 days, respectively. Thus, nonreproductives initially died more rapidly than reproductive forms, but a small percentage of non- reproductives survived for a considerably longer time than fertile females. This phenomenon was also observed by Tower (1906) in various species of Leptinotarsa. Per Cent M ortality 50 99 90 10 1 2 1 2 5 10 0 500 200 100 50 20 10 5 2 I 5 Cniec itras hw i Tbe 10. Table in shown intervals 95% Confidence ^ iue . o rm-rbt ie soig otlt o rpoutv and reproductive of mortality showing lines rime-probit Log 5. Figure ■ ■ "I " I « ""II ■" " ■■1 orpoutv lbrtr-erd en ef beetles. leaf bean laboratory-reared nonreproductive as fe Emergence after Days ...... I T I — — T T - Reproductive - A - Nonreproductive - B Females Females
55
Table 10. Comparative mortality of reproductive and non reproductive laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.^
95% l t 20 “ 50 95% Days Confidence Days Confidence Interval Interval
Reproductive Females 2 0 .0 19.1-20.9 33.3 32.5-34.1
Nonreproductive Females 13.7 12.8-14.6 28.3 27.4-29.2
^Computed from mortality data shown in Appendix Table 31. 56
There was a highly significant negative correlation (r=-.938) between the longevity of reproductively active females and the mating frequency of the various color forms (Appendix Table 17). Those color forms which exhibited a low frequency of mating (i.e., beige and pink) showed the longest duration of adult life. On the other hand, yellow and crimson forms showed the highest mating frequency and shortest longevity in reproductives.
Egg Production
Females did not produce eggs daily during the oviposition period. The mean number of egg-laying days/female was 10.6, which was 89.4 per cent of the oviposition period (Table 11). Egg-laying days ranged from 1 to 52 which was a range of from 1 to 100 per cent of the oviposition period observed. The longest consecutive period of time during the oviposition period in which eggs were not laid was 92 days, and the greatest number of non-laying days during the oviposition period was 97.
No statistically significant differences in fecundity as mea sured by to ta l egg production, eggs/laying day, eggs/day in the ovi position period, and eggs/day through l i f e were observed among the various forms. Again, i f true differences e x ist, they were obscured by excessive within-class variability. Bateman (1967) observed that high variability in egg production seems to be characteristic of laboratory populations of insects.
Mean egg production/female was 138.1 eggs (Table 12), with a range of from 1 to 896. This mean value is somewhat lower than those reported by Isley (1930) and Eddy and Nettles (1930) of 257.7 and 57
Table 11. Mean number of egg-laying days/female and the proportion of the oviposition period in which eggs were laid by laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.1)
(A) Egg-laying Days^)
Color Marking Marking Pattern Ye Be Pi Sa Or Cr Mean
No 1 0 .0 9.6 9.8 9.6 10.4 9.0 9.7
Vi 9.8 1 0.0 10.3 1 0 .0 10.7 10.4 10.2
Co 11.9 10 .1 14.9 10 .0 14.4 10.1 11.9
Su 8.9 11.1 10.5 8.9 11.3 11.9 10.4
Color Mean 10.2 10.2 11.4 9.6 11.7 10.4 10.6
31 (B) Proportiont of Oviposition Period in Egg-laying Days 1
Color Marking Marking m i ul, U vi il Pattern Ye Be Pi Sa Or Cr Mean
No .912 .894 .891 .867 .889 .857 .885
Vi .867 .909 .890 .867 .893 .879 .884
Co .899 .913 .902 .894 .925 .947 .913
Su .895 .925 .894 .874 .886 .882 .893
Color Mean .893 .911 .895 .875 .898 .891 .894
1)Analyses of variance presented in Appendix Table 19. No statistical significance detected. 2) Range 1-52 days.
^Range .01 - 1.0 0 58
Table 12. Mean egg production of laboratory-reared bean leaf b eetles. Baton Rouge, Louisiana. 1970-71.^ ^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 141.1 139.1 128.1 126.1 136.2 100.7 128.5
Vi 129.6 139.1 123.6 123.6 149.4 142.2 134.6
Co 181.5 127.4 214.3 98.5 237.4 153.4 168.7
Su 122.4 140.2 118.9 121.9 120.6 100.6 120.8
Color Mean 143.6 136.4 146.2 117.5 160.8 124.2 138.1
^Analysis of variance presented in Appendix Table 22. No statistical significance detected.
^Range 1 - 896 eggs 59
175.5, respectively. However, Eddy and Nettles state that their observed egg production was "somewhat too high for this insect."
The observed low mean egg production is the probable result of the fact that 13 females laid only 1 egg and over 10 per cent (86 ) laid less than 10 eggs.
The mean number of eggs/laying day was 13.1 (Table 13A), eggs/ day in the oviposition period, 11.5 (Table 13B), and eggs/day through life, 3.6 (Table 13C). These are almost the exact values obtained by
Eddy and N ettles (1930) of 13.9 eggs/laying day and 11.6 eggs/day in the oviposition period.
When the mean number of eggs laid/female/day was calculated for
4 intervals of 10 days beginning at day 11 after emergence*' a sig n i ficant difference was found both for color and for marking pattern
(Table 14). Yellow and orange females laid more eggs than did salmon and pink forms. Females exhibiting marking confluence were more productive than females of a ll other marking patterns. Females with marking suppression averaged fewer eggs/day than did a ll other females. Although there was an apparent decline in egg production over time (Table 15A, Figure 6), this was not found to be statis tically significant.
When the mean number of eggs laid/female/day was calculated in four 10-day intervals beginning at the onset of oviposition, analysis showed no significance either among forms (Table 16) or over time
(Table 15B). This result might be expected as this measurement is
*"0nly 31 eggs were laid through the fir st 10 days of lif e 60
Table 13. Mean numbers of eggs laid/laying day, eggs laid/day in the oviposition period, and eggs laid/day through lif e in laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. '
(A) Eggs Laid/Egg-laying Day2^
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 13.1 12.7 14.6 14.2 14.8 14.7 14.0
Vi 13.7 12.9 11.4 13.2 14.6 11.0 1 2 .8
Co 12.2 15.0 12.6 12.1 12.0 8 .8 12.1
Su 1 2 .0 14.0 13.9 13.3 13.5 13.6 13.4
Col or Mean 12.7 13.6 13.1 13.2 13.7 12.0 13.1
(B) Eggs Laid/Day in the Oviposition Period 3)
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 12.4 11.7 13.4 12.9 13.2 12.6 12.7
Vi 12.3 12.1 9.6 11.5 12.7 9.6 11.3
Co 11.3 13.9 10.2 9.9 1 1.0 8 .6 1 0 .8
Su 11.3 12.5 11.9 1 0 .8 10.3 9.3 1 1 .0
Color Mean 1 1 .8 12.6 11.3 11.3 1 1 .8 10.0 11.5
^Analyses of variance presented in Appendix Table 24. No statistical significance detected.
2^Range 1.0 - 87.0 eggs/day
Range 0.7 - 84.0 eggs/day 61
Table 13 (continued).
(C) Eggs Laid/Day through Life
Color Marking Marking Pattern Pattern Ye Be Pi Sa Or Cr Mean
No 3.9 3.7 3.3 3.6 3.5 3.6 3.6
Vi 3.5 3.7 3.8 3.6 3.4 3.8 3.7
Co 4.2 3.0 4.1 3.4 3.2 3.5 3.6
Su 3.8 3.6 3.9 3.4 3.5 3.6 3.6
Color Mean 3.8 3.5 3.8 3.5 3.4 3.6 3.6
^Range 0.01 - 17.0 eggs/day 62
Table 14. Mean dally oviposition of laboratory-reared bean leaf beetles from day 11 to day 50 after emergence. Baton Rouge, Louisiana. 1970-71. '
Marking Color Marking Pattern Pattern Ye Or Be Cr Sa Pi Mean^)
Co 14.0 13.3 8.9 12.5 8 .8 9.3 1 1.1 a
Vi 9.1 11.1 11.1 9.4 9.4 8.4 9.7 b
No 12.3 1 0 .8 10.7 7.4 8 .2 8 .0 9.6 b
Su 8 .6 7.6 9.3 5.8 8 .1 6 .6 7.7 c
Color Mean L} 1 1 .0 10.7 10.0 8 .8 8 .6 8 .1 9.53) a a ab be c c
1)Analysis of variance presented in Appendix Table 26.
^Mean number of eggs not connected by the same letter differ significantly at the 5% protection level accbrding to Duncan's Multiple Range Test.
^Range 0 .0 - 28.8 eggs /day. Table 15. Mean daily oviposition of laboratory-reared bean lea f beetles through four 10-day inter vals beginning day 11 after emergence and day 1 of the oviposition period. Baton Rouge, Louisiana. 1970-71.^
(A)
10-day Interval Overall 11-20 21-30 31-40 41-50
10.3 10 .0 9.8 8 .0 9.5
(B)
10-day Interval Overall 1-10 11-20 21-30 31-40
9.6 10 .0 9.1 10.5 9.8
Analyses of variance presented in Appendix Tables 26 and 28. 64
Figure 6 . Mean daily oviposition of laboratory-reared bean leaf beetles through four 10-day intervals beginning (A) day 11 after^emergence and (B) day 1 of the oviposition period. Mean Daily Oviposition 10 11 ' 9 8 ' Nmes n aetee idct aeae ubr f females number of average indicate parentheses in ^Numbers ovipositing in a given given a in ovipositing b " / \ / X a" - a 1 t dy 0 after 50 day to 11 Day - A - a 1 o a 4 after 40 day to 1 Day - B 67 X / * / X (607) vpsto i tain ( itiation in oviposition emergence 87)\ / '^ (203) ' ^ " / \ ) 7 8 0 0 0 0 50 40 30 20 10 A (290) (290) 10 dy interval. -day \ V 15 \ (135)
30 / (370) Days (52) P \ ♦ \ * \ o 95
) U 1 66
Table 16. Mean daily oviposition of laboratory-reared bean leaf beetles from day 1 to day 40 after the initiation of oviposition. Baton Rouge, Louisiana. 1970-71. '
Color Marking Marking Pattern Pattern Be Pi Sa Or Cr MeanYe
No 11.6 9.6 9.0 8.5 10.4 9.1 9.7
Vi 8.7 10.2 8.3 8 .8 10.4 12.1 9.8
Co 13.1 8.7 10.0 11.2 13.9 11.9 11.5
Su 10.1 1 0 .0 5.7 8 .0 6.9 8.5 8.2
Color Mean 10.9 9.6 8.3 9.1 10.4 10.5 9.82)
Analysis of variance presented in Appendix Table 28. No statistical significance detected. 2) Range 0.2 - 34.0 eggs/day 67 rather artificial in that individuals of identical physiological or reproductive age may have widely differing chronological age.
Figure 7 gives a graphic illustration of the percentage of females in oviposition over time as compared to egg production over time after adult emergence. Oviposition was not initiated until day
9 after emergence. The number of females in oviposition rose rapidly to a peak at day 20. Maximum egg production occurred at this time.
Egg production and the number of females in oviposition then declined rapidly to day 40 and gradually diminished to zero 90 days later.
Figure 8 shows the relative cumulative egg production of all females through their life and beginning at the initiation of ovi position. Ten per cent of total egg production was completed by age
14.3 days, 20 per cent by 17.7 days, 50 per cent by 26.7 days, and
95 per cent by 2 months (59.7 days). Twenty per cent of all eggs had been laid by 3.5 days after the initiation of oviposition, 50 per cent by 7.9 days, and 95 per cent by 39.0 days.
Occasionally females began laying eggs before pairings for mating were made. It is not uncommon for virgin female insects to lay infertile eggs. That was the case with these beetles. Of the
897 nonreproductive females, ten females initiated oviposition before being exposed to males. All eggs which were produced by unmated females failed to hatch after incubation at 80° F. These females were provided with males in an attempt to secure mating. Although dissections were not made to determine the presence or absence of sperm in the spermatheca, it is assumed that none of the females mated as in no case did those which initially laid infertile eggs subsequently lay fertile eggs after exposure to males. Figure 7. Proportion of laboratory-reared bean leaf beetles in oviposition and proportion of total egg production over time after adult emergence, .60 .30
£ .50 oa Eg Production Egg Total •HO of Proportion (B) •H4 -1 U-i (0 .40 O O ‘ .20 A, !\ •Ho O > 4-) >-1 c .30 O t 4 ODu » n
00 Figure 8 . Log time-probit lines showing cumulative egg production of laboratory-reared bean leaf beetles.^
A - Measured from adult emergence
B - Measured from oviposition initiation
1 2 5 10 20 50 100 200 500 Days
^95% confidence intervals shown in Table 17. Table 17. Cumulative egg production of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. '
ET 2) ET 3) "10 95% 95% Days Confidence Daysn 50 Confidence In te rv a l In terv al
Measured from adult emergence 14.3 13.5-15.0 26.7 26.1-27.3
Measured from oviposition initiation 2.3 1.9- 2.7 7.9 7.3- 8.5
^Computed from data shown in Appendix Table 32.
^Time required for 10% of total egg production to occur.
■^Time required for 50% of total egg production to occur. 71
Oviposition data for these unmated females is shown in Table 18.
Egg production ranged from 1 to 600, with a mean of 145.5. The ovi position period averaged 10.9 days duration, yielding a mean daily oviposition of 11.5 eggs/day.
Weight Determination
Significant weight differences were found between the various color and marking pattern types (Table 19). Beetles exhibiting the marking confluence pattern weighed significantly less than did bee tles of the 3 other pattern types. Differences in weight between the color categories closely corresponded with intensity of red pig ment, weight increasing as red pigment increased. Only the heaviest, crimson, could be separated from the lightest, salmon and beige.
As there was a similar although opposite correlation between longevity and red pigment intensity, a correlation coefficient was calculated comparing the mean longevity of the 6 color forms in the laboratory-reared population with corresponding mean weights of field-collected populations. The resultant correlation coefficient, although quite high (r=-.720), was not statistically significant
(Appendix Table 35). The lack of significance in the correlation coefficient may be due to the fact that only 6 pairs of numbers were available for comparison, leaving 4 degrees of freedom for error.
Further study may show that this correlation is, indeed, significant.
Statistical analysis showed a significant difference in mean weight between sampling dates. There was an apparent positive shift in the weight distribution of samples collected from St. Landry
Parish, as evidenced in Figure 9. 72
Table 18. Oviposition data for 10 unmated laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Female Egg Days in Mean Daily No. Production Oviposition Oviposition
1 110 4 27.5
2 6 2 3.0
3 35 16 2.2
4 6 2 3.0
5 208 10 28.0
6 56 4 14.0
7 83 6 13.8
8 1 1 1.0
9 350 33 10.6
10 600 31 19.4
Mean 145.5 10.9 11.5 73
Table 19. Mean weights in milligrams of bean leaf beetles differing in color and marking pattern- St. Landry and Catahoula Parishes, Louisiana. 1971.
Color Marking Marking P a tte rn P a tte rn Cr Or Pi Ye Sa Be Mean^)
Vi 15.1 14.7 14.1 15.1 14.3 14.7 14.7 a
Su 14.8 14.5 14.9 14.6 14.2 14.1 14.5 a
No 14.8 14.3 14.3 14.7 14.3 14.4 14.5 a
Co 14.8 14.6 14.6 12.9 13.2 12.8 13.8 b
Color Mean ' 14.9 14.5 14.5 14.3 14.0 14.0 14.43) a ab ab ab b b
^Analysis of variance shown in Appendix Table 33. 2 ) 'Mean weights not connected by the same letter differ significantly at the 5% protection level according to Duncan's Multiple Range T e s t. 3) Range 4.4 - 25.2 mg. Relative Frequency .10 “ .15 2 " .20 0 - .05 - iue 9, Figure 6 7 8
9 el ppltos vr i . . ady rs Louisiana. , arish P f Landry a le t. S bean female e. d tim te c weight over lle o in -c ld change fie populations showing of s n eetle b tio u histogram trib is frequency d elative R 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 i egt n illigrams M in Weight * - Jn 25 Jun —-A A* A Jun Jun p. 9 My 7 May + 29 Apr. 8 75
Susceptibility to Methyl Parathion
Topical applications of graded dosages of methyl parathion to
female bean leaf beetles resulted in values comparable to those
obtained by Jensen and coworkers (1970), of the order of 0.022-0.03 u g /b e e tle .
Although the color forms exhibited slight differences in 1^)50 and LD^q response (Table 20), these differences were insignificant as all 95 per cent confidence intervals overlapped. The yellow form
yielded the highest value, 0.03 ug/beetle, but also exhibited
the widest confidence interval, 0.025 ug. The salmon color form responded with the lowest I^D^q value, 0.022 ug/beetle, and had the narrowest confidence interval, 0.003 ug.
It is probable that a more intensive study of bean leaf beetle response to methyl parathion would show these differences to be sig nificant. It is suggested that this receive further study and that
additional studies be initiated to investigate relative susceptibility
of bean leaf beetle color forms to the chlorinated hydrocarbon and
carbamate classes of insecticides.
P arasitism
P arasitism of the bean le a f b e e tle by £ . d ia b ro tic a e showed
significant seasonal fluctuations through 1972 (Figure 10). Per centage parasitization was low early in the season (late May to early
June), increasing rapidly to a peak in mid- to late June. Maximum
parasitism at this time reached 22 per cent. There was an immediate rapid decline followed by a gradual decrease to 0 in late season. Table 20. Response of bean leaf beetle color forms to topical applications of methyl parathion. Baton Rouge, Louisiana. 1972.I ) 2)
957. Confidence LE^O 90 Color u g /b e e tle In te rv a l u g /b eetle Slope
Ye .030 .019-.044 .058 4.37+1.11
Be .023 .014-.033 .040 5.43+1.46
Pi .025 .023-.028 .048 4.44+0.60
Sa .022 .020-.023 .040 4.78+0.47
Or .023 .021-.025 .045 4.52+0.48
Cr .025 .022-.027 .042 5.63+0.83
Computed from 48-hour m o rtality data shown in Appendix Table 37.
^No statistical significance detected. 77
Figure 10. Parasitization of female bean leaf beetles by Celatoria diabroticae (Shimer) in collections from St. Landry Parish, Louisiana. Per Cent P arasitism 25 20 15 10 5 20 May June July August 28 September 79
Overall parasitism varied significantly between the various color forms (Table 21). The highest per cent parasitism occurred in the yellow color form and the pink form showed the lowest percentage parasitism. Mean separation between the intermediate forms was not complete.
Temperature Tolerance
Temperature tolerance studies were initiated in an attempt to determine if observed cyclic fluctuations in color and/or marking frequency over time (Herzog, 1968) were temperature-related.
Results of probit analyses showed measurable and significant differences in mortality occurred at all temperatures except 95°.
However, at higher temperatures, mortality proceeded much too rapidly for meaningful differences to be detected. With decreasing tempera ture differential time-mortality response became evident.
The most obvious overlying fact was that, although not always s ig n ific a n tly so, the yellow form with few exceptions showed more rapid progressive mortality at all temperatures.
At a constant temperature of 100° F. mortality proceeded very rapidly, as evidenced by the range of regression coefficients
(b=5.01-8.30). Only 2 beetles survived for more than 5 days. Of the colors, only yellow and salmon could be separated at the median lethal time (Table 22, Figure 11), their confidence intervals sepa rated by mere hundredths of a day.
At a constant temperature of 95° F. mortality again advanced rapidly, 100 per cent mortality occurring in 15 days. No significant differences in mortality could be detected at LT^q, LT^q, or LT^q Table 21. Percentage parasitism of bean leaf beetles by Celatoria diabroticae (Shimer). St. Landry Parish, Louisiana. 1972.
Color
Ye Or Sa Cr Be Pi
Per Cent Parasitism ' 6.7 4.8 4.8 3.8 3.8 2.5. a ab ab be be c
Marking Pattern
No Vi Co Su
Per Cent Parasitism ' 4.0 4.7 4 .8 4.1
■^Analysis of variance presented in Appendix Table 38. 2 } Mean percentages not connected by the same letter differ significant ly at the 5% protection level according to Duncan's Multiple Range T est.
^No statistical significance detected. Table 22. Time-mortality resgonse of bean leaf beetle color forms subjected to a constant temperature of 100 F. Baton Rouge, Louisiana. 1972. ^ ^ ) 3 j
Color LTlO 95% Confidence LT50 95% Confidence LT90 957. Confidence Days In te rv a l Days In te rv a l Days In te rv a l
Ye 0.9 a 0. 6- 1.1 1.5 a 1. 2- 1.7 2.4 a 2 .0 - 3.1
Be 1.1 a 0. 8 - 1.3 1.8 ab 1.5-2.0 2.9 a 2.5- 3.5
Pi 1.1 a 0.7-1.3 1.8 ab 1.5-2.1 3.0 a 2.6- 3.9
Sa 1.0 a 0.9-1.1 1.7 b 1.7-1 .8 3.0 a 2.8- 3.2
Or 0.9 a 0. 6- 1.1 1.6 ab 1.3-1.9 2.9 a 2 .4 - 3.8
Cr 1.2 a 0. 1- 1.6 1.7 ab 0 .4 -2 .6 2.4 a 1.7-38.1
^Computed from mortality data shown in Appendix Table 40.
^LT values not connected by the same letter differ significantly by 95% confidence interval separation.
"^Slope of log time-probit lines shown in Appendix Table 47. Per Cent Mortality Figure 11. Log tim e-probit lin e s showing bean le a f b eetle color color eetle b f a le bean showing s e lin e-probit tim Log 11. Figure form m o rtality in response to exposure to a constant constant a to exposure to response in rtality o m form e eaue f 0° F. 100° of perature tem as er I taton tio itia In r te f a Days 83
(Table 23).
After treatment of the beetles at 90° F., yellow could be sepa rated from all other colors excepting orange by time LT^q, and from all other colors by time LT^q (Table 24, Figure 12). By the time
90 per cent mortality occurred, only orange and crimson could be se p a ra te d .
A mortality level of 10 per cent occurred more rapidly in yellow than in beige forms at 80° F,, but neither could be separated from the remaining forms (Table 25, Figure 13). Mortality proceeded most rapidly to the median lethal time level in yellow and pink beetles, while the death of 50 per cent of the crimson beetles required the longest time. By the time that 90 per cent mortality had occurred c o n siste n t d ifferen c es had become obscured.
No significant differences could be detected in the time required for 10 per cent mortality at 70° F. (Table 26, Figure 14). and
LTgQ values were significantly lower in yellow forms as compared with all other colors.
Again at 60° F. no significant LTjq values could be detected
(Table 27, Figure 15). Yellow, pink, and salmon forms attained 50 per cent mortality more rapidly than did those of beige or crimson.
The orange form could not be separated from any of the other colors, but a t LT^q it, with yellow and pink, showed significantly lower values than beige and crimson.
The most complex series of interrelationships occurred under a
50° F. constant temperature regimen (Table 28, Figure 16). Although it had been hypothesized that lighter-colored forms would be least tolerant of low temperatures, results showed that the beige form, Table 23. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 95° F. Baton Rouge, Louisiana. 1972.1)2)3)
95% Confidence LT 95% Confidence LT 95% Confidence Color Day9 In terv a l Da£§ In terv a l Da $9 In terv a l
Ye 2.1 1.4-2.7 4.3 3.5-5.1 9.0 7.6-11.4
Be 2.5 1.5-3.4 4.9 3.8-5.9 9.4 7.7-12.9
P i 2.8 1 .3 -3 .8 5.4 3.9-6.7 10.6 8.3-16.7
Sa 2.6 1.8-3.2 5.3 4.5-6.1 10.9 9.3-13.9
Or 2.3 1. 6-2.8 4.8 4.1-5.5 10.1 8.7-12.5
Cr 2.2 1.3-3.0 4.9 3.8-5.5 10.6 8.5-14.9
^Computed from mortality data shown in Appendix Table 41.
^^No statistical significance detected.
Slope of log time-probit lines shown in Appendix Table 47.
-p*00 Table 24. Time-mortality response of bean, leaf beetle color forms subjected to a constant temperature of 90° F. Baton Houge, Louisiana. 1972.1)^)3)
957, Confidence LT 95% Confidence LT „ 95% Confidence Color Days> In terv al * a » In te rv a l Da$8 In terv al
Ye 1.7 a 1.0- 2.4 5.7 a 4 .4 - 6.9 19.0 ab 15.9-23.6
Be 3.7 b 2 .6 -4 .8 9.9 b 8.4-11.3 26.3 ab 22.8-31.6
Pi 3.9 b 3,0-4.7 9.7 b 8 . 6- 10.8 24.6 ab 21.8-28.4
Sa 4.0 b 3.0-4.9 9.8 b 8 . 6- 11.1 24.5 ab 21.6-28.7
Or 3.1 ab 2.4-3 .8 9.1 b 8 , 0- 10.1 26.5 b 23.6-30.6
Cr 3.8 b 3.1-4.5 8.8 b 7 .9 - 9.6 20.2 a 18.2-22.9
1)Computed from m o rtality data shown in Appendix Table 42.
2)lT values not connected by the same letter differ significantly by 95% confidence interval separation.
"^Slope of log tim e-probit lines shown in Appendix Table 47. Per Cent M o rtality 9 . 99 0 - 50 - 1 iue 12 Figure T m o rtality in response to exposure to a constant tem perature perature tem constant a to exposure to response in rtality o m Log tim e-probit lin e s showing bean le a f b eetle color form form color eetle b f a le bean F. showing 90° s of e lin e-probit tim Log 5 2 1 2 50 20 10 5 2 1 .5 IB B B r B ■ B B I ■ as t I taton tio itia In r fte a Days Pi+Sa Or Cr - Be Ye oo Table 25. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 80° F. Baton Rouge, Louisiana. 1972-73,^^)3)
95% Confidence 957, Confidence LT 95% Confidence Color Days In te rv a l Days In te rv a l Days In te rv a l
Ye 3.1 a 2.7-3.5 16.5 a 15.6-17.5 88.8 a 79.8-100.2
Be 4.2 b 3.9-4.5 21.7 b 21.1-22.4 113.3 be 105.8-121.9
Pi 3.7 ab 3.1-4.2 18.6 a 17.4-19.8 93.6 a c 82.5-108.2
Sa 3.6 ab 3.1-4.0 21.6 b 20.5-22.8 130.3 b d 115.4-149.4
Or 3.8 ab 3,2-4.5 21.6 b 20.0-23.4 121,7 bed 103.1-148.1
Cr 4,1 ab 3.4-4.7 26.3 c 24.6-28.3 168.8 d 143.3-207.2
^Computed from mortality data shown in Appendix Table 43.
2)LT values not connected by the same l e t t e r d if f e r s ig n ific a n tly by 95% confidence in te rv a l separation.
Slope of log time-probit lines shown in Appendix Table 47. Per Cent Mortality 10 - 1 iue 3 Lg i -rbt ie soig en ef el clr form color eetle b leaf bean showing lines e-probit tim Log 13. Figure 2 m ortality in response to exposure to a constant temperature temperature constant a to exposure to response in ortality m f 0 F. 80° of 5 as t I taton tio itia In r fte a Days 10 20 50 200100 Sa Ye Be Pi 500 00 00 Table 26. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 70° F, Baton Rouge, Louisiana. 1972-73.'^'^
95% Confidence T ^ 95% Confidence LT 95% Confidence Color f i o , 5° Days In te rv a l Days In te rv a l Daysn 90 In terv al
Ye 10.4 a 8.5-12.3 38.1 a 35.0-41.1 139.2 a 125.6-157.0
Be 15.5 a 11.4-19.4 55.5 b 49.6-61.6 199.0 b 165.2-255.6
Pi 15.6 a 11.5-19.5 55.0 b 49,1-61,0 193.8 b 161.4-247.6
Sa 14.3 a 10.4-18.0 53.8 b 48.1-59.7 203.0 b 168.3-260.5
Or 13.9 a 10.7-17.1 56.0 b 5C.9-61.2 224.5 b 188.6-280.0
Cr 10.5 a 7.5-13.6 52.1 b 46.5-57.8 258.1 b 208.7-341.7
1 )Computed from m o rtality data shovm in Appendix Table 44.
2) LT values not connected by the same letter differ significantly by 95% confidence interval separation.
3) Slope of log time-probit lines shown in Appendix Table 47. Per Cent M ortality 99 90 50- 0 1 - ______iue 4 Lg i -rbt ie soig en ef ete oo form color beetle leaf bean showing lines e-probit tim Log 14. Figure V 2 y raiy n epne o xoue o cntn tmperature tem constant a to exposure to response in ortality m 9L n 1 2 5 10 0 500 200 100 50 20 10 5 I ■ . . ■ . II Ei i lE D's er Inii i n tio itia n I r te f a -Da'^s ?y ______.. ■— ______Be+Pi 1 ...... Ye Cr Or Sa
4 ______... o Table 27. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 60° F. Baton Rouge, Louisiana. 1972-73.^^^
95% Confidence 95% Confidence LT 95% Confidence Color Days> In te rv a l Days In terv al Da?s In te rv a l
Ye 15.9 a 13,1-18.6 40.8 a 37.3-44.3 105.0 a 95.4-117.6
Be 18.2 a 15.2-21.0 51.7 b 48.0-55.4 147.2 b 132.6-166.9
Pi 16.9 a 14.3-19.4 42.2 a 39.0-45.3 105.3 a 96.6-116.4
Sa 16.4 a 14.5-18.2 45.1 a 42.7-47.5 124.0 ab 115.7-134.0
Or 19.9 a 15.7-23.8 47.6 ab 42.7-52.5 113.4 a 100.7-131.8
Cr 18.3 a 16.0-20.5 50.7 b 47.9-53.6 140.6 b 129.6-154.2
^Computed from mortality data shown in Appendix Table 45.
2)l T values not connected by the same letter differ significantly by 95% confidence interval separation.
Slope of log time-probit lines shown in Appendix Table 47. Per Cent M ortality 90 10 50 99 1 2 1 2 5 10 0 500 200 100 50 20 10 5 2 1 iue 5 Lg i -rbt ie soig en ef ete oo form color beetle leaf bean showing lines e-probit tim Log 15. Figure raiy n epne o epsr t a osat temperature constant a to exposure to response in ortality m t " I I H II 1 " 'I tt I as t I taton tio itia In r fte a Days ...... ' 1 ' 11 ' 1 ' r / Cr Ye Or Sa Be i \D !s» Table 28. Time-mortality response of bean leaf beetle color forms subjected to a constant temperature of 50° F. Baton Rouge, Louisiana. 1972-73.1)2)3)
95% Confidence 95% Confidence 95% Confidence Color “ w Days In te rv a l Days In te rv a l Dajrsi f 90 In te rv a l
Ye 5.4 a 3.2- 7.9 41.8 a 35.8-47.7 322.1 ab 244.1-472.0
Be 13.3 b 10.5-16.0 82.0 b 75.7-89.4 507.4 a 403.6-677.3
Pi 8.3 a c 6.7- 9.9 55.4 cd 51.9-59.0 369.2 ab 315.3-444.9
Sa 9.6 abc 7.1-12.3 49.9 a c 45.0-55.0 258.6 b 213.0-331.5
Or 10.0 be 8.4-11.7 60.2 d 56.7-63.8 361.3 ab 311.9-429.3
Cr 14.9 b 11.0-18.7 85.4 b 77.0-95.9 489.5 a 367.3-723.1
^Computed from mortality data shown in Appendix Table 46.
2) LT values not connected by the same letter differ significantly by 95% confidence interval separation. 3) Slope of log tim e-probit lines shown in Appendix Table 47. Per Cent M ortality 50 10 0 9 99- - 5 1 iue 6 Lg i -rbt ie soig en ef ete oo form color beetle leaf bean showing lines e-probit tim Log 16. Figure 2 m ortality in response to exposure to a constant tem perature perature tem constant a to exposure to response in ortality m of 50° F. F. 50° of 10 ______as t I taton tio itia In r fte a Days 20 50 0 500 100 200 Ye Cr Or Pi Be Sa 95
together with crimson, showed the greatest tolerance as evidenced by
values of 82 and 85.4 days, respectively. Yellow and salmon
forms appeared least tolerant, exhibiting values one-half the
magnitude of the above.
No definite trends could be discerned concerning time mortality
response of the various color forms. The overlying fact remains that
yellow forms consistently showed more rapid progressive mortality at
all temperatures.
In only 1 case was differential response detected among marking
pattern types in the time required for attainment of 10 per cent mor
tality. This occurred at 100° (Table 29, Figure 17), with 0.01 day
separating the 95 per cent confidence intervals between marking con
fluence and marking suppression. No significant differences in mor
tality were detected at 95° (Table 30), 90° (Table 31), or 80°
(Table 32). At 70° the marking confluence pattern attained 50 per
cent mortality more rapidly than did the other forms (Table 33, Figure
18). No meaningful differences occurred at 60° (Table 34). The
marking confluence and broken vitta patterns were more tolerant to a
constant temperature of 50° than were the normal and marking suppres
sion pattern types (Table 35, Figure 19).
In general, beetles exhibiting the various marking patterns
showed greater similarity of response than did the 6 color forms.
Examination of mortality data after 7 days exposure to each tem
perature studied showed that the median lethal temperatures for all
color and marking pattern variants were quite similar (Table 36).
Marking p a tte rn v a ria n ts showed g re a ter homogeneity of both LT^q and
LTgg response than did the 6 color forms. The median lethal Table 29. Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 100° F. Baton Rouge, Louisiana. 1972.1)2)3)
Marking “ lO 957c Confidence LTCrt 957, Confidence LT 95% Confidence P attern Days In te rv a l Days In terv al Da$§ In terv al
No 1 .0 ab 0.6-1.3 1.6 ab 1.3-2.0 2.7 a 2.2-3.9
Vi 1.0 ab 0.7-1.2 1.7 ab 1.4-1.9 2.8 a 2.4-3.5
Co 1.2 a 1.1-1.4 1.9 a 1.8-2.0 2.9 a 2.7-3.2
Su 1.0 b 0.9-1,1 1.6 b 1.6-1.7 2.7 a 2.6-2.9
Computed from m o rtality data shown in Appendix Table 40. 2 ) LT values not connected by the same letter differ significantly by 95% confidence interval separation.
^)Slope of log time-probit lines shown in Appendix Table 47. Per Cent M ortality 10 9 - 99 0 " 50 ■ 90 " 1 " iue 17 Figure y akn patr n epne o xoue o constant a to ortality m exposure eetle to b leaf response bean in showing attern p lines marking by e-probit tim Log 5 2 1 20 10 5 2 1 .5 as t Inii i n tio itia n I r fte a Days No+Vi+Su Co- T 50 J - ' Table 30. Time-mortality response of bean leaf beetle marking pattern types, subjected to a constant temperature of 95° F. Baton Rouge, Louisiana. 1972. ' ' '
Marking 95% Confidence 95% Confidence LT90 95% Confidence P attern DaysIf10 In te rv a l Sjs In terv al Days In terv al
No 2.7 1.8-3.3 5.2 4 .3 -6 .0 10.0 8.5-12.6
Vi 2.3 1.4-3.0 4.7 3 .7 -5 .6 9.8 8.0-13.2
Co 1.8 1.1-2.4 4.3 3.4-5.0 10.2 8.2-14.4
Su 2.5 1.6-3.2 4.9 4 .0 -5 .8 9.8 8.2-12.9
^Computed from mortality data shown in Appendix Table 41.
No statistical significance detected.
^Slope of log time-probit lines shown in Appendix Table 47. Table 31. Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 90° F. Baton Rouge, Louisiana, 1972. '
Marking 957. Confidence 957, Confidence LT 957o Confidence “ lO P attern Days In te rv a l Days> In terv al Days. 9 0 In terv al
No 3.1 2.2-3.9 8.4 7.2- 9,6 23.0 20.1-27.1
Vi 3.1 2.2-3.9 8.8 7.5-10.0 25.3 22.1-29.7
Co 3.1 2,1-4.1 8.2 6. 8 - 9.5 21.5 18.4-26.6
Su 3.5 2 .5-4.5 9.8 8.4-11.1 27.2 23.6-32.5
^Computed from mortality data shown in Appendix Table 42. 2) No statistical significance detected. 3j Slope of log time-probit lines shown in Appendix Table 47. Table 32. Tima-mortality response of bean leaf beetle narking pattern types subjected to a constant temperature of 80° F. Baton Rouge, Louisiana. 1972-73. ' ' '
Marking 95% Confidence • 957. Confidence LT10 rLT50 LT90 95% Confidence P attern Days In te rv a l Bays In terv al Days In te rv a l
No 3.6 3.2-4.1 20.7 19.7-21.8 118.5 106.4-133.8
Vi 3.7 3 .3-4.1 19.4 18,5-20.3 101.9 92.5-113.3
Co 3.8 3 .4-4.2 20.5 19.0-22.1 110.8 94.6-133.6
Su 3.8 3 .4-4.2 20.9 20.0-21.9 114.5 103.6-127.9
Computed from m o rtality data shown in Appendix Table 43.
2 ") 'No statistical significance detected.
3}Slope of log tim e-probit lines shown in Appendix Table 47. Table 33. Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 70° F. Baton Rouge, Louisiana. 1972-73. ' ' '
Marking 95% Confidence 95% Confidence 95% Confidence LT10 LT50 nT90 P attern Days In te rv a l Days In te rv a l Days In te rv a l
No 13.7 a 9.7-17.5 50.3 a 44.4-56.2 185.5 ab 153.7-239.0
Vi 13.1 a 10.2-15.9 50.6 a 46.2-55.1 195.7 ab 168.4-236.0
Co 10.3 a 8 . 6- 12.0 39.9 b 37.1-42.6 154.6 a 140.1-173.3
Su 14.9 a 11.8-17.9 56.9 a 52.2-61.8 218.0 b 186.1-266.0
^Computed from mortality data shown in Appendix Table 44. 2) LT values not connected by the same letter differ significantly by 95% confidence interval separation. 3) Slope of log time-probit lines shown in Appendix Table 47. Per Cent M ortality 0 ■ 90 9 . 99 0 " 50 10 " 2 1 2 5 10 0 500. 200 100 50 20 10 5 2 1 iue IS Figure „"T 1 I ■ I " l I"" 'I '■ I I 1 „""T— I y akn pten n epne o xoue o constant a to ortality m exposure F. to beetle 70° of response leaf in bear temperature showing pattern marking lines by time-probic Log as t Inii i n tio itia n I r fte a Days Vi No <-o u S ------Table 34. Time-mortality response ofQbean leaf beetle marking pattern types subjected to a constant temperature of 60 F. Baton Rouge, Louisiana. 1972-73.
Marking 95% Confidence 957. Confidence 95% Confidence P a tte rn Days In te rv a l Days In te rv a l DaysnT’ ° In te rv a l
No 17.5 ab 14.6-20.3 47.7 a 44.1-51.3 130.4 ab 118.0-147.1
Vi 17.9 ab 14.6-20.9 43.4 a 39.4-47.1 105.2 a 95.2-118.9
Co 13.2 a 11.2-15.1 42.8 a 39.8-45.6 138.7 b 126.6-154.2
Su 16.5 b 15.2-17.8 45.9 a 44.2-47.6 127.7 b 121.5-134.8
^Computed from mortality data shown in Appendix Table 45. 21 ■'LT values not connected by the same letter differ significantly by 95% confidence interval separation.
Slope of log time-probit lines shown in Appendix Table 47. Table 35. Time-mortality response of bean leaf beetle marking pattern types subjected to a constant temperature of 50° F. Baton Rouge, Louisiana. 1972-73. ' ' '
Marking 95% Confidence 95% Confidence 95% Confidence > LT50 P attern Days In te rv a l Days In te rv a l Daysi f 90 In te rv a l
No 8.4 a 6. 0- 10.8 53.8 a 48.6-59.3 346.3 ab 276.1-463.8
Vi 9.9 a 7.5-12.4 70.2 b 64.3-76.4 493,8 b 388.8-668.6
Co 9.2 a 6.9-11.6 72.3 b 66.3-79.2 568.4 b 439.9-787.6
Su 11.0 a 9.3-12.7 51.7 a 48.7-54.8 243.0 a 216.0-278.6
^Computed from mortality data shown in Appendix Table 46.
LT values not connected by the same letter differ significantly by 95% confidence interval separation.
^Slope of log time-probit lines shown in Appendix Table 47. Per Cent M ortality 99- 500 1 iue 9 Lg i -rbt ie soig en ef ete ortality m beetle leaf bean showing lines e-probit tim Log 19. Figure 2 y akn pten n epne o xoue o constant a to exposure to response in pattern marking by eprtr o 5 F. 50 of temperature 10 as t I taton tio itia In r fte a Days 20 50 100 2005 Vi No Su 106
Table 36. Bean leaf beetle mortality after 7 days exposure to constant temperatures ranging from 50-100° F. Baton Rouge, Louisiana. 1972.^-'*'
l t 50 95% Confidence Slope °F. In te rv a l oF. l 9 0
Color
Ye 85.6 77.0- 94.1 103.0 15.9+3.6
Be 89.9 66.7-898.4 108.4 15.7+5.6
Pi 90.7 77.8-129.6 111.7 14.2+4.6
Sa 90.4 69.4-619.4 111.3 14.2+5.0
Or 89.0 73.6-111.3 107.4 15.6+4.9
Cr 89.6 80.1-108.2 111.8 13.3+3.7 liar king P attern
No 87.0 43.8-121.5 103.9 16.6+5.8
Vi 88.6 77.4-106.2 107.3 15.4+4.5
Co 87.5 72.1-116.0 109.3 13.3+4.2
Su 89.1 71.3-113.6 105.6 17.4+5.7
Computed from m o rta lity data shown in Appendix Table 48.
^No statistical significance detected. 107 temperature ranged from 87.0 to 89.1° F. for the normal marking pattern and marking suppression form, respectively, while color forms exhibited a range of ca. 5°, ranging from 85.6° F. for yellow beetles to 90.7° F. for the pink.
Biological Implications
"A difference between two . . . populations can always be established, provided the observer is willing to go to enough trouble to refine and increase the number of his measurements . , . What the zoologist must be first concerned with is the biological significance of a demonstrated difference, and for this kind of significance there is no test but that of intuition, experience, and intelligence"
(Simpson, ait al., 1960).
Color-marking pattern polymorphisms in the bean leaf beetle probably have little significance per se, other than the possible existence of assertive mating. Rather, the importance lies with other characters which are correlated with these polymorphisms. Color forms and marking patterns are, in effect, genetic markers for vari able physiological characters and ecological tolerance levels which are much more difficult to discern. In reality, color pattern poly morphism is the probable result of physiological variability rather than the converse being true. The correlation of a particular poly morphic form with a particular physiological character variant or ecological tolerance level is probably a chance occurrence. 108
Miscellaneous Observations on Factors
Affecting Bean Leaf Beetle Abundance
In the course of field surveys and laboratory and field research on this study and an earlier one (Herzog, 1968), several observations concerning factors affecting bean leaf beetle abundance have been made.
Two different species of tachinid parasites have been found attacking bean leaf beetles. The first, C. diabroticae. is the most common and is the only bean leaf beetle parasite reported in the literature (McConnell, 1915; Isley, 1930; Eddy and Nettles, 1930).
Only two adults of the second have been reared from field-collected bean leaf beetles. It has been tentatively identified as Hyalomyodes triangulifer (Leow).*
The larvae and pupae of both species are typically dipteran. The larva and puparium of C. diabrotlcae are heavily setose, while the larva of the latter is more lightly setose and the puparium smooth and glabrous. The larva of C. diabroticae completely consumes the contents of the abdominal cavity of the beetles then separates the abdomen from the r e s t of the body when i t emerges to pupate. The larva of the Hyalomyodes does not consume all of the body cavity contents of i t s host and e x its the abdomen leaving i t in ta c t w ith the thorax. The mean duration of the pupal period of 24 individuals of
The author expresses his gratitude to Dr. C. W. Sabrosky of the Systematic Entomology Laboratory, USDA, Washington, D.C. for identification of this species, and to Dr. J. B. Chapin for assis tance in obtaining the identification. 109
£• diabroticae was 8.1 days (ranging 5-10). Two adults of the
Hyalomyodes emerged in 9 and 10 days.
Typically, very few eggs are laid by parasitized beetles; a maximum of 5 observed from a field-collected parasitized specimen.
Most lay no eggs.
Dissections of numerous field-collected beetles have shown:
1) 1 case of a parasite larva within an overwintering, diapausing b e e tle ; 2) two cases of 2 parasite larvae found within a single host;
3) a small percentage (<1) pupate within the host; and 4) some para sites die within the host before they have fully matured.
One or possibly 2 unidentified species of nematodes have been found emerging from living beetles.
Several unidentified fungi have been found infesting dead adults.
It is not known if they were a primary mortality factor or merely secondary invasions.
In the course of the rearing program it was observed that some eggs and larvae suffered from a bacterial disease, probably Serratia marcescens Bizio (Newsom, 1973). Eggs turned red, and did not hatch.
If larvae, prepupae, pupae or even adults turned red, they died almost
immediately. Some, especially pupae and adults, died and then turned r e d . 110
CONCLUSIONS
Of 2,546 male-female pairings in the laboratory, 820 or 36.2 per cent were successful. If crimson females are disregarded because they had no male counterpart for comparison, yellow forms, both male and female showed the greatest mating frequency. Both sexes exhibit ing marking confluence mated most frequently. All females tended to mate more frequently with yellow males, and most forms of both males and females attained greatest mating success with partners exhibiting marking confluence.
No differences were detected in the duration of adult life his tory stages, regardless of color or marking pattern. However, reproductive females lived significantly longer than nonreproductives.
Beige and pink reproductive forms lived much longer than correspond ing nonreproductives, while other forms reacted similarly whether fecund or not.
Polymorphic forms showed no significant differences in fecundity as measured by total egg production, eggs/laying day, eggs/day in the oviposition period, or eggs/day through life. However, mean daily oviposition through the first 50 days of adult life showed yellow and orange females laid more eggs than did the salmon and pink, while marking confluence females laid more eggs than all others.
Significant differences in weight were detected among both colors and marking patterns.
Differences in parasitism by C_. diabroticae were noted among color forms, yellow apparently being the most susceptible to parasite attack, and pink the least susceptible. The frequency of the yellow I l l color form In a population may reflect the incidence of parasitism by this species.
Hating frequency, longevity, and weight all showed strong corre lations with intensity of red pigment present. Correlation among these variables were high only for mating x longevity and longevity x weight, only the former attaining significance.
Results concerning identification of differential susceptibility of color forms to methyl parathion were inconclusive.
Differential response of color forms and marking pattern types to temperature was observed, but no temperature-related trends could be established. Yellow forms consistently showed the most rapid progressive mortality at all temperatures studied. Marking pattern variants showed a greater similarity of response than did the 6 color forms. These are indications of subtly differing selective values for the many forms under a variety of ecological conditions.
Finally, from the data presented it can be concluded that there are subtle variations in physiological and behavioral characters and ecological tolerances in the bean leaf beetle and that these vari ations can be correlated with color and marking pattern forms within this polymorphic species. LITERATURE CITED
Anonymous. 1960. These are some pests th a t damage soybeans. Soybean Dig. 20(8):16-19.
Anonymous. 1962. Soybeans. USDA ARS Coop. Econ. Ins. Rept. 12:547.
Arnett, R. H. 1963. Beetles of the United States. The Catholic University of America Press: Washington. 1112pp.
Ball, H. J. 1968. A five-year study of potential western corn rootworm resistance to diazinon and phorate in Nebraska. J. Econ. Entomol. 61:496-498,
Barber, H. S. 1945. Note on Cerotoma and Andrector (Coleoptera, Chrysomelidae). Bull. Brooklyn Entomol. Soc. 40:121-122.
Bateman, M. A. 1967. Adaptations to temperature in geographic races of the Queensland fruit fly, Dacus (Strumeta) tryoni. Aust. J. Zool. 15:1141-1161.
Bateson, W. 1895. On the color variation of a beetle of the family Chrysomelidae, statistically examined. Proc. Zool. Soc. London 65:850-860.
Beregovoi, V. E. 1970. Differences in color variation in males and females of the common froghopper, Philaenus spumarius (L.). Doklady Biol. Sci. Proc. Acad. Sci. USSR. Biol. Sci. Sect. 191:201-203. (English translation).
B la ir, B. D. and R. H. Davidson. 1966. S u sc e p tib ility of northern corn rootworm adults to aldrin in Ohio. J. Econ. Entomol. 59:608-609.
Boving, A. G. 1930. Description of the larva of Cerotoma trifurcata Forster (Coleoptera: Chrysomelidae). Proc. Entomol. Soc. Washington 31:51-59.
Bowditch, F. C. 1913. The Phytophaga (except Cassidae and Hispidae) of the Stanford expedition to Brazil. Psyche 20:125-131.
Burns, J. M. 1966. Preferential mating versus mimicry: disruptive selection and sex-limited dimorphism in Papilio glaucus. Science 153:551-553.
Chapius, F. 1875. Histoire naturelle des insectes. Genera des coleopteres. 11:1-420.
Chevrolat, L. A. A. 1837. In Dejean, Catalogue des coleopteres de la collection de M. le compte Dejean 5:385-503.
112 Chevrolat, L. A. A. 1843, In d'Orbigny, Dictionnaire universal d'histoire naturelle . . . 3:1-744.
Chittenden, F. H. 1892. Notes on the food habits of some species of Chrysomelidae. Proc. Entomol. Soc. Washington 2:261-267.
Chittenden, F. H, 1897. The bean leaf beetle. USDA Bur, Entomol. Bull. 9:64-71.
Chittenden, F. H, 1899. Insects injurious to beans and peas. USDA Yearbook 1898:233-266.
Clark, W. J., F. A. Harris, F. G. Maxwell, and E. E. Hartwig. 1972. Resistance of certain soybean cultivars to bean leaf beetle, striped blister beetle, and boll worm. J. Econ. Entomol. 65:1669-1672.
Daum, R. J. 1970, Revision of two computer programs for probit analysis. Bull. Entomol. Soc. Amer. 16:10-15.
Eastman, C„ E. 1973. Aseptic rearing of the bean leaf beetle, Cerotoma trifurcata (Forster), on soybean callus tissue. , Unpublished master's thesis, Louisiana State University, Baton Rouge, 93pp.
Eddy, C. C,, and W. C, Nettles. 1930. The bean leaf beetle, South Carolina Agr. Exp. Sta. Bull. 265:1-25.
Eichhorn, 0., and P. Graf. 1971. Sex-linked colour polymorphism in Aphidecta obliterata L. (Coleoptera: Coccinellidae). Z. angew, Entomol. 65:225-231.
Essig, E. 0. 1926. Insects of western North America. Macmillan Co.: New York. 1035pp.
Fabricius, J. C. 1801. Systema eleutheratorum 1:1-506.
Farish, D. J. 1972. Balanced polymorphism in North American popu lations of the meadow spittlebug, Philaenus spumarius (Homoptera Cercopidae). 1. North American morphs. Ann. Entomol. Soc. Amer 65:710-719.
Finney, D. J. 1947. Probit analysis, a statistical treatment of the sigmoid response curve. Cambridge Univ. Press: Cambridge, England. 318pp.
Folsom, J. W. 1936. Additional notes on little-known cotton insects J, Econ. Entomol. 29:1066-1068.
Ford, E. B. 1940. Polymorphism and taxonomy, pp 493-513. In: Huxley, J. E., Ed. (The new systematics.) Oxford University Press: New York. 583pp.) 114
Forster, J. R. 1771. Novas species insectorum 1:1-100.
Frost, S. W. 1959. Insect life and natural history. Dover Publi cations, Inc.: New York. 526pp.
Gast, R. T. 1961. Factors involved in differential susceptibility of corn earworm larvae to DDT. J. Econ. Entomol. 54:1203-1206.
Gemminger, M., and E. von Harold. 1876. Catalogus coleopterorum hucusque descriptorum synonumicus at tomaticus 12:3479-3822.
Germar, E. F, 1824. Insectorum species novae aut minus cognitae, descriptionibus, illustratae 1:1-624.
Gershenson, S. 1945. Evolutionary studies on the distribution and dynamics of melanism in the hamster (Cricetus cricetus L.). II. Seasonal and annual changes in the frequency of black hamsters. Genetics 30:233-251.
Guerin, J. 1953. Coleopteros do Brazil. Bol. Fac. Files. Cienc, Let., Univ. Sac Paulo 1953:1-356.
Hamilton, E. W. 1965. Aldrin resistance in corn rootworm beetles. J. Econ. Entomol. 58:296-300.
Harvey, W. R. 1960. Least-squares analysis of data with unequal subclass frequencies. USDA ARS 20-8:1-130.
Harvey, W. R. 1969. Instructions for use of LSMLGP (least-squares and maximum likelihood general purpose program). Louisiana State University Computer Research Center: Baton Rouge. 22pp. mJmeo.
Herzog, D, C. 1968, Seasonal locational and sexual variation in the coloration and marking pattern of the bean leaf beetle, Cerotoma trifurcata (Forst.), in Louisiana. Unpublished master's thesis, Louisiana State University, Eaton Rouge. 86pp.
Horn, G. R. 1893. The Galerucini of boreal America. Trans. Amer. Entomol. Soc. 20:57-144.
Horn, N. L ., L. D. Newsom, R. G. Carver, s/nd R. L. Jensen. 1970. •■S-f^cts of virus diseases on soybeans in Louisiana. Louisiana Agr. 13(4):12-13, 15.
Isley, D. 1930. The biology of the bean leaf beetle. Arkansas Agr. Exp. Sta. Bull. 248:1-20.
Isley, D. 1942. Insect problems resulting from changes in agricul ture in Arkansas. J. Econ. Entomol. 35:473-477. 115
Jensen, R. L. 1971. Evaluation of soybean germ plasm for resistance to the bean leaf beetle, Cerotoma trifurcata (Forster), the southern green stink bug, Nezara viridula (Linnaeus), and the soybean looper, Pseudoplusia includens (Walker). Unpublished doctoral dissertation, Louisiana State University, Baton Rouge. 83 pp.
Jensen, R. L. and L. D. Newsom. 1972. Unpublished data on file with Department of Entomology, Louisiana State University, Baton Rouge.
Jensen, R. L., J. W. Thomas, Jr., and L. D. Newsom. 1970. Unpub lished data on file with Department of Entomology, Louisiana State University, Baton Rouge.
Johnson, R. H. 1910. Determinate evolution in the color pattern of the lady b e e tle s . Carnegie In s t. Wash. Publ. 122:1-104.
Kogan, M. 1973. In litt.
Landis, B. J., and H. C. Mason. i938. Variant elytral markings °£ Epilachna varivestis. Entomol. News 49:181-184.
Lutz, F. E. 1908. Notes on the inheritance of variations in the color pattern of Crioceris asparagi. Psyche 15:50-52,
McConnell, W. R. 1915. A unique type of insect injury. J. Econ. Entomol. 8:261-266.
McEwen, F. L., and C. M. Splittstoesser. 1964. A genetic factor controlling color and its association with DDT sensitivity in the cabbage looper. J. Econ. Entomol. 57:197-199.
Merrell, D. J., and C. F. Rodell. 1968. Seasonal selection in the leopard frog, Rana pipiens. Evolution 22:284-288.
Metcalf, C. L., and W. P. Flint. 1962. Destructive and useful in se c ts. McGraw-Hill Book Company, In c .: New York. 1087 pp.
Metcalf, Z. P. 1962. General catalogue of the Homoptera. VIII. Cercopoidea. Part 3. Aphrophoridae. North Carolina State College Press: R aleigh. 600 pp.
Milliron, H. E. 1958. Economic insect and allied pests of Delaware. Delaware Agr. Exp. Sta. Bull. 321:1-87,
M otsinger, R. E ., J . L. Bagent, S. D. Hensley, N. L. Horn, and L. D. Newsom. 1967. Soybean diseases and insects of Louisiana. Louisiana Coop. Ext. Serv. Publ. 1558:1-24.
Newsom, L. D. 1973. Personal communication.
Norman, A. G. 1963. The soybean: genetics, breeding, physiology, nutrition, management. Academic Press: New York. 239 pp. 116
Patel, K. K., and J. W. Apple, 1966, Chlorinated hydrocarbon resistant northern corn rootworm in Wisconsin. J. Econ. Entomol. 59:522-525.
Peterson, A. 1960. Larvae of the insects. Part II. Edwards Bros., In c .: Ann Arbor. 416 pp.
Petty, H, B., and T. L. Wainscott. 1961. Soybean insects. Succ. Farm. 59(5):48-51.
Pliske, T. E. 1972. Sexual selection and dimorphism in female tiger swallowtails, Papilio glaucus L. (Lepidoptera: Papilionidae): a reappraisal. Ann. Entomol. Soc, Amer. 65:1267-1270.
Popenoe, E. A. 1877. A list of Kansas Coleoptera. Trans. Kansas Acad. Sci. 5:21-40.
Popenoe, E. A. 1889. Some insects injurious to bean. Kansas Agr. Exp. Sta. Ann. Rep. 2:206-212.
Reichmuth, W. 1967a. Nachveis von Vitamin A in Larven des Kartoffelkafers Leptinotarsa decemlineata Say. Z, Naturforschung 22:1232-1233.
Reichmuth, W. 1967b. Uber Beziechungen zwischen Pigmentierung und Empfindlichkeit beim Kartoffel Kafer Leptinotarsa decemlineata Say: Ein Beitrag zur physiologischen Bedeutung von Carotinoid- Pigmenten mit dem Nachweis von Vitamin A bei Insekten. Z. angew. Zool. 54:285-296.
Richards, 0. W. 1961. Insect polymorphism: an introduction to the study of polymorphism in insects. Symp. Roy. Entomol. Soc. London 1:1-10.
Ruppel, R. F. 1973. In l i t t .
Schmelter, H. 1879. Coleoptera of the neighborhood of New York, Bull. Brooklyn Entomol. Soc. 1:55.
Sheppard, P. M. 1952. A note on non-random mating in the moth Panaxla dominula L. Heredity 6:239-241.
Shimer, H. 1871. Additional notes on the striped squash beetle. American Nat. 5:217-220.
Simpson, G. S., A. Roe, and R. C. Lewontin. 1960. Quantitative Zoology. H arcourt, Brace and Company, In c .: New York. 440pp.
Smith, S. G. 1950. Cytotaxonomy of the Coleoptera. Can. Entomol. 82:58-68.
Sweetman, H. L. 1958. The principles of biological control. Wm. C. Brown Company: Dubuque, Iowa. 560 pp. 117
Tan, C. 1949. Seasonal variations of color patterns in Harmonia axyr id is. Proc. 8th Int. Cong. Genetics 1:669-670.
Thompson, W, R. 1943. A catalogue of the parasites and predators of insect pests. Section 1. Parasite host catalogue. Part 1. Parasites of the Arachnids and Coleoptera. Imp. Agr, Sur,, Inst, Entomol., Parasite Ser.: Belleville, Ontario. 151 pp.
Timofeeff-Ressovsky, N. E. 1940. Mutations and geographical vari a tio n . pp. 73-136. In: Huxley, J . E ., Ed. (The new system- atics. Oxford University Press: New York, 583 pp„)
Tower, W. L, 1906, An investigation of evolution in chrysomelid beetles of the genus Leptinotarsa. Carnegie Inst, Wash, Publ. 48:1-302,
Tower, W. L, 1910. The mechanism of evolution in Leptinotarsa. Carnegie Inst, Wash. Publ, 263:1-384.
Townsend, C, H, T. 1885. A list of Coleoptera collected in Louisiana, on or south of parallel 30°. Can. Entomol. 17:66-73.
Walters, H, J. 1970. Bean pod mottle virus disease of soybeans. Arkansas Farm Res. 19(2):8.
Webster, F. K. 1888. Report on the season’s observations, and especially upon corn insects. US Comm. Agr. Rept. 1887:147-154.
Webster, F. M. 1894, Insects of the year. USDA Div, Entomol., Insect Life 7:202-207.
Webster, F. M. 1895. On the probable origin, development, and diffusion of North American species of the genus Diabrotlca. J, New York Entomol. Soc. 3:158-166.
Webster, F. M. 1896, On the probable origin and diffusion of North American species of the genus Diabrotica. J. New York Entomol. Soc. 4:67-68.
Webster, F. M. 1909. T!e trend of insect diffusion in North America, Entomol. Soc, Ontario Rept. 32:63-67.
Webster, F. M. 1902. The diffusion of insects in North America. Psyche 10:47-58.
Wickham, H. F. 1897. The C oleoptera o f Canada. XIX. The Chrysomelidae of Ontario and Quebec (continued). Tribe IX Galerucini. Can. Entomol, 29:7-12.
Wilcox, J. A. 1954. Leaf beetles of Ohio (Chrysomelidae: Coleoptera). Ohio Biol. Sur. Bull. 43:353-506. 118
Wuensche, A. L. 1970. Unpublished data on file with Department of Entomology, Louisiana State University, Baton Rouge.
Zappe, M. P. 1919. The bean leaf beetle, Cerotoma trifurcata (Forst.). Connecticut Agr. Exp. Sta. Bull. 211:327-329.
Zulueta, A. de. 1932. Heredity in the X and T chromosomes of Phvtodecta. Proc. 6 th Int. Cong. Genetics 1:389 APPENDIX Table 1. Least-squares analysis of variance for mating frequency of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Source o f Degrees o f Mean v a r ia tio n freedom square '
Total 2545 ** Generation 3 8.03
9 Color 5 0.37nS
? Harking 3 0 .04nS d Color 4 0.46n® d Marking 3 0.31« ~ . n s
9 Color x 9 Marking 15 0.43 d Color x d Marking 12 0 . 16ns
9 Color x d Color 20 0 . 2 0 °®
9 Marking x d Marking 9 0 . 2 2 n8
Residual 2471 0 .2 1
1)p.oi - 2 .06 c 15 & 1000 degrees of freedom 121
Table 2. Mean mating frequencies of reciprocal color x color pairings of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Female Mating Female Mating x Male Frequency x Male Frequency
Ye X Be .477 Be X Ye .389
Ye X Pi .295 P i X Ye .311
Ye X Sa .445 Sa X Ye .454
Ye X Or .390 Or X Ye .530
Be X Pi .214 Pi X Be .292
Be X Sa .334 Sa X Be .383
Be X Or .217 Or X Be .339
Pi X Sa .228 Sa X Pi .286
Pi X Or .226 Or X Pi .394
Sa X Or .469 Or X Sa .418
r = .661*1)
^Analysis of correlation shown in Appendix Table 3. Table 3. Analysis of correlation between mating frequencies of reciprocal color x color pairings of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Source o f Degrees o f Mean v a ria tio n freedom square '
T otal 9 tm u
Correlation 1 .023*
R esidual 8 .004
1>f.05 * 5-32 « 1 & 8 degrees of freedom Table 4, Mean mating frequencies of reciprocal marking x marking pairings of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Female Mating Female Mating x Male Frequency x Male Frequency
No x Vi .347 Vi x No .313
No x Co .432 Co x No .435
No x Su .236 Su x No .311
Vi x Co .398 Co x Vi .368
Vi x Su .370 Su x Vi .471
Co x Su .217 Su x Co .375
r = .509nsl)
^Analysis of correlation shown in Appendix Table 5. Table 5. Analysis of correlation between mating frequencies of reciprocal marking x marking pairings of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana, 1970-71.
Source of Degrees of Mean variation freedom square
T otal 5
Correlation 1 ,005ns
Residual 4 .004 125
Table 6. Mean mating frequencies of male and female color- marking pattern types in laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
C olor- Female Mating Male Mating Marking Pattern Frequency Frequency
YeNo .321 .403 YeVi .360 .498 YeCo .561 .513 YeSu .451 .478
BeNo .348 .373 BeVi .295 .410 BeCo .200 .363 BeSu .353 .354
PiNo .379 .277 PiVi .340 .331 PiCo .000 .345 PiSu .377 .173
SaNo .315 .400 SaVi .318 .398 SaCo .371 .323 SaSu .421 .314
OrNo .373 .304 OrVi .369 .229 Or Co .450 .600 OrSu .396 .145
r = .186nsl)
^Analysis of correlation shown in Appendix Table 7. Table 7. Analysis of correlation between mating frequencies of male and female color-marking pattern types in laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Mean variation freedom square
Total 19
Correlation 1 .008ns
Residual 18 .013 127
Table 8 . Least-squares analysis of variance for average number of males required before impregnation of laboratory- reared female bean leaf beetles was achieved. Baton Rouge, Louisiana. 1970-71. '
Source of Degrees of Mean v a ria tio n freedom square
T otal 819 —
G eneration 3 0.61ns
Color 5 0.13ns
Marking 3 0.04ns
Color x Marking 15 0.17ns
Regression Preoviposition Period Linear 1 133.04**
Residual 792 0.32
^Generation x color x marking pattern means shown in Appendix Table 9. 128
Table 9. Mean number of males required before impregnation of laboratory-reared bean leaf beetle, females was achieved. Baton Rouge, Louisiana. 1970-71. '
Color- Generation Marking P attern I I I I I I IV
YeNo 1.53 1,22 1.00 l.OC YeVi 1.18 1.21 1.00 1.17 YeCo l.OC 1.00 1.00 l.OC YeSu 1.38 1.00 1.14 1.31
BcNo 1.64 1.17 1.10 1.26 3cVi 1.56 1.29 1.33 1.38 B eC o l.OC 1.17 1.00 BeSu 1.75 1.20 1.00 1.25
PiNo l.OC 1.00 1.10 1.C0 PiVi 1.50- 1.88 1.00 1.75 PiCo 1.00 -- PiSu -- 2.00 l.OC 1.00
SaNn 1.73 1<39 1.06 2.25 SaVi 1.63 1.27 1.19 1.00 SaCo — 1.20 1.60 1.00 SaSu 1.11 1.00 1.00 1.17
OrNo 1.25 1.-53 1.23 1.00 OrVi 1.42 1.22 1.25 1.20 OrC o 1.00 1.00 1.00 1.50 OrSu 1.50 1.00 1.00 --
CrNo 1.00 1.00 1.00 1.00 CrVi 1.38 1.09 1.17 2 .CO CrCo 1.00 1.00 — CrGu •> ~ 1.00
^Number of observations making up each mean shewn in Appendix Table 10. Table 10. Number of reproductively active female bean leaf beetles making up generation x color x marking p a tte rn means of variables obtained in the laboratory rearing experiment. Baton Rouge, Louisiana. 1970-71.
C olor- G eneration Marking P attern I I I I I I IV
YeNo 17 18 10 10 YeVi 17 14 9 6 YeCo 3 1 5 3 Ye3u 8 4 7 13
BeNo 25 36 31 27 BeVi 25 31 20 26 BeCo 0 2 6 2 BeSu 4 5 8 12
PiNo 5 10 10 5 FiVi 8 8 4 PiCo 0 1 0 0 PiSu 0 2 4 iA.
SaNo 15 13 17 4 n SaVi o 15 IS 8 SaCo 0 5 5 3 SaSu 9 10 11 6
OrNo 20 19 26 3 Or Vi 12 27 20 10 OrCo 2 2 5 4 OrSu 6 1 S 0
CrNo 4 7 1 1 CrVi 8 11 6 1 CrCo 1 1 0 0 CrSu 0 0 1 c Table 11. Least-squares analyses of variance for preoviposit ion. oviposition, and postoviposition periods o^ laboratory-reared female bean leaf b eetles. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Preovipos it ion Oviposition Pos t ov ipos i t ion var iat ion freedom Period Per iod Period mean square mean square mean square
Total 819 —— —
Generation 3 10485.11-** 615.77** 3574.33**
Color 5 229.78ns 56.96ns 382.38ns
Marking 3 1003.19ns 46„77ns 95.88ns
Color x Marking 15 445.09ns 131,35ns 423,84ns
Residual 793 411.57 194.40 246.72
^Generation x color x marking pattern means are shown in Appendix Table 12. Table 12. Mean preoviposition, oviposition, and postoviposition periods of four generations of laboratory-reared bean le a f b eetles. Baton Rouge, Louisiana. 1970-71. ^
Preoviposit ion Period Oviposition Period Postoviposition Period Color- P attern i II I I I IV I I I I I I r 7 I I I I I I IV
YeNo 30.6 22.2 13.6 13.5 16.2 9.6 19.8 14.4 12.2 7.9 5.0 5.2 YeVi 24.2 21.6 14.1 15.3 11.4 7.6 13.0 17.3 17,7 5.9 12.6 6.7 YeCo 15.7 14.0 15.2 13.0 17.7 25.0 15.4 8.3 4.3 5.0 3.4 3.0 YeSu 38.0 13.0 15.1 22,S 15.0 10.0 7.6 16.7 4.9 4.8 6,4 5.6 BeNo 41.2 17.0 21.5 18.0 13.0 14.6 10.4 11.9 17.4 4.9 8.9 5.1 BeVi 46.4 23.1 25.3 21.1 16.0 11.8 8.1 12.5 8.3 6.1 4.9 4.6 BeCo — 15.0 14.0 14.0 -- 8.0 28.3 1.0 -- 21.0 7.7 74.0 BeSu 31.0 14.8 15.0 15.6 22.3 7.3 7.5 11.5 33.3 3.4 4.0 4.3 FiNo 27.6 15.5 18.2 12.2 13.2 15.8 10.0 11.4 5.0 9.2 11.8 5.0 PiVi 84.0 33.3 22.3 29.3 9.5 10.0 11.8 15.0 13.5 5.3 8.3 3.5 PiCo -- 16.0 — — 19.0 -- -- — 5.0 —— PiSu 51.5 12.5 17.C — 8.0 15.5 1.0 -- 2.0 3.5 ' 29.0 SaNo 35.3 20.9 13.1 27.3 18.7 9.9 10.6 5.5 20,5 3.8 12.6 10.5 SaVi 43.0 18.5 15.3 15.9 20.9 12.1 15.9 8.0 10.8 5.0 4.6 6.0 SaCo -- 22.6 27.8 13.0 — 4.6 9.8 15.0 12.6 4.6 7.C SaSu 19.2 18.4 15.7 16.2 20.0 13.0 14.6 9.7 8.7 15.6 3.5 4.8 OrNo 20.2 22.4 15.7 15.3 14.3 11.9 11.4 3.7 18.6 4.8 6.5 1.0 OrVi 31.5 18.8 15.3 19.7 17.3 15.7 14.7 16.8 19.7 5.8 5.2 5.0 Or Co 18.0 14.5 13.6 28.5 28.5 20.C 16.6 13.5 6.5 4.0 5.8 5.0 OrSu 31.8 21.0 13.2 14.2 3.0 11.7 -- 6.5 3.0 10.0 — CrNo 14.3 14.3 18.0 13.0 11.3 8.4 9.0 13.0 15.0 5.3 3.0 1.0 CrVi 27.9 17.8 23.5 14.0 11.6 12.8 7.0 1.0 39.9 5.4 24.3 1.0 CrCo 15.0 16.0 — — 1.0 13.0 -- __ 6.0 7,0 --— Ci*Su — “ 16.0 "• •• 13.0 ■* ■* • •* 3.0 mrm
^Number of observations making up each mean shown in Appendix Table 10. Table 13. Least-squares analyses of variance for proportions of adult life spent in preovi posit ion, oviposition, and postoviposition periods in laboratory-reared female bean leaf beetles. Baton Rouge, Louisiana, 1970-71. '
Source of Degrees of Preoviposit ion Oviposit ion Postoviposition v a ria tio n freedom Period Period Period mean square mean square mean square
T otal 819
Generation 3 0.034ns 0.081ns 0.026ns
Color 5 0.025ns 0,015ns 0. 001ns
Marking •5 0.007ns 0,004ns 0,013ns
Color x Marking 15 0. 011ns 0.007ns 0.009ns
Regression Appropriate Period Linear 1 25.780** 30.973** 16.808**
Regression Longevity 1 19.171** 3.238** 1.984**
Residual 791 0.017 0.012 0.010
^Generation x color x marking pattern means are shown in Appendix Table 14. Table 14. Mean proportions of total adult life spent in preoviposition, oviposition, and post oviposition periods by four generations of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.^'
Preoviposit ion Period Oviposition Period Postoviposition Period Color- P a tte rn I I I III IV I II III IV I II III IV
YeNo .539 .610 .407 .450 .268 .256 .504 .427 .212 .176 .131 .165 YeVi .488 .652 .429 .423 .212 .242 .368 .422 .324 .159 .237 .180 YeCo .580 .320 .496 .583 .353 .590 .432 .347 ,113 .110 .110 ,130 YeSu .651 .505 .577 .533 .285 .358 .260 .362 .083 .190 .217 .137 BeNo .579 .574 .574 .531 .208 .324 .270 .350 .226 .147 .199 .159 BeVi .621 .607 .645 .588 .260 .264 .266 .318 .128 .176 .144 .134 BeCo — .495 .410 .160 -- .165 .467 .005 .375 .153 .825 BeSu .408 .616 .673 .583 .350 .296 .223 .320 . 243 .144 .159 .156 PiNo .616 .501 .482 .554 .238 .317 .302 .312 .176 .244 .254 .196 PiVi .615 .641 .519 .603 .135 .264 .293 .353 ,250 .115 .224 .073 rico -- .410 — — .490 ____ .120 -- PiSu -- .770 .448 .370 — .230 .480 .010 — .030 .123 .630 SaNo .489 .635 .485 .615 .266 .291 .299 ,165 .251 .117 .262 .255 SaVi ,510 .573 .524 .595 .315 .347 .386 .255 .175 .115 .132 .200 SaCo -- .628 .564 .380 — .144 .332 .443 .266 .138 .213 SaSu .502 .485 .587 .568 .323 .318 .348 .300 .210 .226 .120 .163 GrNo .477 .593 .554 .870 .264 .297 .316 .170 .278 ,148 .179 .053 Or Vi .465 .537 .536 .510 .278 .349 .361 .382 .258 .151 .149 .141 ChrCo .390 .415 .422 ,618 .455 .495 .424 .295 .215 .120 .186 .113 OrSu .565 .830 .505 — .315 .110 .312 — .143 ,110 .232 — CrNo .433 ,576 .630 .510 .268 .270 .310 .510 ,330 .211 .100 .030 CrVi .496 .531 .500 .980 ,185 .364 .207 .060 .331 .143 .323 .060 CrCo ,740 .460 — __ .040 .370 -- .290 .200 ---- CrSu ---- .520 — -- — .420 — -- -- .090 --
-^Number of observations making up each mean shown in Appendix Table 10. 134
Table 15. Least-squares analysis of variance for mean longevity of reproductive and nonreproductivc laboratory-reared female bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Heaj v a ria tio n freedom square ' '
T otal 168 __
Generation 3 5363.39**
Product ivity 1 956.76*
Color 5 375.64ns
Marking 3 8 7 .84ns r Productivity x Color j 525.80*
Productivity x Marking 3 186.61ns
Color x Marking 15 272.C4ns
Regression Number Linear 1 35,34ns
Res idual 131 182.92
1) F a0<; * 3.91 c 1 & 125 degrees of freedom
F = 2.27 c 5 & 125 degrees of freedom
^Generation x color x marking p a tte rn means are shown in Appendix Table 16. 135
Table 16. Mean longevity of four generations of reproductive and nonreproductive laboratory-reared bean leaf beetle females. Baton Rouge, Louisiana. 1970-71.
Reproductive Females
C olor- I II I I I IV Marking — _ _ P attern x n x n x n x n
YeNo 57.0 17 37.6 18 36.1 10 31,1 10 YeVi 51.2 17 33.0 14 37.7 9 37.7 6 YeCo 35.7 3 42.0 1 32,0 5 22.3 3 YeSu 55.9 8 25.8 4 27.1 7 43.2 13
BeNo 69.9 25 33.4 36 38.8 31 33.0 27 BeVi 68.8 25 39.0 31 36.6 20 36.3 26 BeCo -- 0 42.0 2 48.0 6 88.0 2 BeSu 84.5 4 24,0 5 24.5 8 29.4 12
PiNo 43.8 5 38.5 10 38.0 10 26.6 5 PiVi 105.0 2 46,5 8 40.3 8 45.8 4 PiCo 0 38.C 1 0 — 0 PiSu -- 0 59,5 2 29.5 4 45.0 1
SaNo 72.5 15 32.6 18 34.4 17 41.3 4 SaVi 72.6 8 33.5 f r 33.8 16 27.9 8 iaCo 0 37.8 5 40.2 5 32.0 3 SaSu 45.9 9 45.0 10 31.9 11 28.8 6
OrNo 51.1 20 37.1 19 31.6 26 18.0 3 Or Vi 66.6 12 38.3 27 33.2 20 39.5 10 Or Co 5 1 .C 2 36,5 2 34.0 5 45.0 4 OrSu 50.5 5 25.0 1 32.8 6 -- 0
CrNo 38.5 4 26.0 n/ 28.0 1 25,0 1 CrVi 78.0 8 34.0 l i 52.8 6 14.0 1 CrCo 20.0 1 34.0 i 0 — 0 CrSu __ 0 -- 0 30.0 1 -- 0 Table 16 (continued).
Nomreproductive Females
C olor- III I I I IV
Marking mmm mmm P attern x n x n x n x n
YeNo 47.3 43 31.0 12 50.0 4 26.0 2 YeVi 54.9 36 32.2 9 21.3 5 20.3 3 YeCo 35.8 4 0 -- 0 20.0 2 YeSu 45.3 7 21.3 4 17.5 4 31.0 6
BeNo 48.0 44 33.1 36 31.8 20 23.4 13 BeVi 52.3 62 36.2 36 25.3 19 18.9 16 3eCo 16.4 5 26.2 5 25.5 4 15.0 4 BeSu 34.9 10 31,0 7 31.6 15 22.3 3
PiNo 44.1 11 28.4 1C 34.6 8 16.0 2 PIVi 42.5 6 31.5 8 24.1 8 20.0 1 PiCo ... 0 20.7 3 26.0 7 33.7 3 PiSu 50.0 1 46.5 2 22.0 1 0
SaNo 60.1 14 36.8 25 26.6 18 29,0 6 SaVl 44.4 9 30.5 13 42.7 20 27.2 o SaCo 128.0 1 82.0 1 -- 0 29.7 7 SaSu 67.6 11 23.4 7 30.2 6 46.7 6
OrNo 52.2 27 33.5 20 30.9 17 42.5 5 Or VI 56.6 21 53.3 21 23.0 27 21.8 9 Or Co 40.8 5 86.0 1 52.0 1 23.3 3 OrSu 37.8 5 20.0 2 23.2 5 33.5 2
CrNo 60.2 14 42.6 11 14.0 1 0 CrVi 60.8 16 44.4 _/ 28.1 8 20.C 2 CrCo — 0 0 ... 0 0 CrSu 77.7 3 0 0 15,0 1 Table 17. Relationship between mean longevity and mating frequency of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Color Longevity Mating Form in Days Frequency
Ye 37.72 .423
Be 48.25 .299
Pi 47.05 .260
Sa 42.58 .356
Or 38.10 .397
Cr 32.02 .434
r « -.938**1)
^Analysis of correlation shown in Appendix Table 18, Table 18. Analysis of correlation between mean longevity and mating frequency of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Mean var iat ion freedom square '
T otal 5 --
Correlation 1 . 022**
Residual 4 .0007
^ 0 1 = 16.3 c 16.5 degrees of freedom 139
Table 19. Least-squares analyses of variance for egg-laying days and proportion of the oviposition period in egg-laying days for laboratory-reared bean leaf b eetles. Baton Rouge, Louisiana. 1970-71. s'
Source of Degrees of Egg-laying Oviposition variation freedom Days Proportion mean square mean square
Total 819 — —
Generation 3 1.53ns 0.09
Color 5 46.87nS o.oins
Marking 3 31.12ns o.oins
Color x Marking 15 15.73ns 0 . 01ns
Regression Oviposition -Mr-4- ** Period Linear 1 40210.94 13.47
Regression Egg-laying l-l Days Linear 1 5.27
Residual 791 31.00 0 .0 1
^Generation x color x marking pattern means shown in Appendix Tables 20 and 21. 140
Table 20. Mean number of egg-laying days per female of laboratory- reared be^m leaf beetle.' Baton Rouge, Louisiana. 1970-71.
Color- Generation Marking P attern I I I I I I IV
YeNo 8.5 8.2 18.1 11.8 YeVi 7.8 6.8 11.0 13.7 YeCo 13.0 24.0 14.2 7.3 YeSu 11.8 8.0 7.4 8.8
BeNo 9.2 9.4 8.2 1C.2 BeVi 12.9 8.2 7.6 9.9 BeCo .... 7.0 19,2 1.0 Be3u 22.0 7.2 6.9 11.2
FiNo 10.2 9.9 9.5 9.4 FiVi 8.0 8.1 9.3 14.0 PiCo 18.0 — PiSu 8.0 12.8 1.0
SaNo 12,6 8.3 7.8 5.0 SaVi 14.8 10.9 10.5 6.9 SaCo 4.2 9.0 12.7 SaSu 10.3 11.6 9.5 6.7
OrNo 11.1 11.1 9.1 3.3 OrVi 14.5 11.8 11.5 12.2 OrCo 25.5 19.5 15.6 13.3 OrSu 13.0 3.0 10.0 --
CrNo 7.8 7.3 6.0 1.0 CrVi 9.5 11.3 6.3 1.0 CrCo 1.0 13.0 --— CrSu •« — m — 12.0
^Number of observations making up each mean shown in Appendix Table 10.
o 141
Table 21. Mean proportion of the oviposition period in actual egg-laying days of laboratory-reared bean leaf beetle. Baton Rouge, Louisiana. 1970-71.-)
C olor- Generation Marking P attern x I I I I I IV
YeNo .872 .929 .928 .880 YeVi .065 .932 .854 .845 YeCo . 890 .960 .908 .910 YeSu .911 .863 .993 .772
BeNo .867 .909 .950 .873 BeVi .897 .935 .972 .857 BeCo .920 .817 1.000 BeSu .813 .934 .974 .963
PiNo .820 .843 .971 .924 PiVi .835 .905 .918 .950 PiCo ... .950 — — FiSu — 1.000 .865 1.000
SaNo .781 .932 .879 .923 SaVi .881 .910 .848 .784 SaCo .956 .954 .847 SaSu .799 .910 . 844 .817
OrNo .861 .965 .899 .953 Or Vi .839 .904 .910 .853 OrCo .940 .945 .920 .988 OrSu .927 1.000 .883 --
CrNo .773 .946 .670 1.000 CrVi .870 .924 .928 1.000 GrCo 1.000 1.000 -- CrSu - “ .920 " ••
^Number of observations making up each mean shown in Appendix Table 10. 142
Table 22. Least-squares analysis of variance for egg production of laboratory-reared bean leaf beetles. Baton Rouge, Lcuisi.ana. 1970-71.
Source of Degrees of Mean v a ria tio n freedom square
T otal 819 —
Generat ion 3 66844.01**
Color 5 19509.67ns
Marking 3 11228.10ns
Color x Marking 15 9593.69ns
Regression Oviposition Period Linear 1 7886775.02**
Residual 792 13124.41
^G en eratio n x color x marking p a tte rn means shown in Appendix Table 23. Table 23. Hean egg production of laboratory-reared bean leaf b eetles. Baton Rouge, Louisiana. 1970-71. '
Color - Generation Marking P a tte rn I I I I I I IV
YeNo 85.8 106.9 290.8 189.1 YeVi 89.2 86.6 186.9 121.0 YeCo 69.3 369.0 250.4 176.3 YeSu 174.6 111.5 102.9 113.7
BeNo 114.0 148.1 117.5 159.7 BeVi 152.0 125.7 101. C 147.3 BeCo -- 81.5 264.7 19.0 BeSu 195.3 112.2 91.4 164.7
PiNo 138.6 122.2 116.0 162.2 PiVi 59.0 129.1 104.0 161.3 PiCo 266.0 -- -- PiSu -- 182.5 111.8 5.0
SaNo 141.1 132.6 102.5 57.0 SaVi 155.1 136.9 149.6 84.8 SaCo -- 41.2 102.0 107.3 SaSu 103.4 174.8 123.8 132.8
Or No 109.6 176.4 112.8 74.0 OrVi 181.0 154.1 191.6 187.0 Or Co 332.0 197.5 327.2 226.8 OrSu 108.2 16.0 121.2 --
CrNo 49.0 91.1 69.0 70.0 CrVi 131.6 152.5 7S.0 9.0 CrCo 3.0 199.0 — --
m . mm ■» ~ CrSu •" *• 111.0
Number of observations making up each mean shown in Appendix Table 10. Table 24. Least-squares analyses of variance for eggs per laying day, eggs per day in the oviposition period, and eggs per day through lif e for laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. ^
Source of Degrees of Eggs per Egg- Eggs per Day in Eggs p e r Day variation freedom laying Day Oviposition through Life mean square Period mean square mean square
Total 819 -- • • --
Generat ion 3 172.98* 377.44** 23.00
Color 5 I5.24ns 24.56ns 2 . 02ns
Harking 3 66.30ns 88.64ns 0 . 10nS
Color x Marking 15 32.25nS 31.68ns 1 .20nS
Regression Egg Production JLJU Linear 1 21689.52 13167.72 7577.92**
Regression Egg-Laying Days Linear 1 15043.24** — --
Regression Oviposition Period Linear 1 -- 10574.05** --
Regression Longevity Linear 1 — — 509.38
Residual 791 62.13 64.04 1.85
^Generation x color x marking pattern means shown in Appendix Table 25. Table 25. Mean number of eggs per laying-day, eggs per day in the oviposition period, and eggs per day through li f e of four generations of laboratory-reared bean leaf beetle females. Baton Rouge, Louisiana. 1970-71.*'
Eggs per Eggs per Day in Eggs per Day Color- Laying Day Oviposition Period through Life Marking Pattern IIIIII IV I II III IV I II III IV
YeNo 9.6 15.1 14.7 15.1 8.6 14.5 13.7 13.4 1.9 3.4 7.5 5.8 YeVi 8.4 13.3 23.7 8 .8 7.2 12.7 22 .2 7.6 1.6 2 .8 5.3 3.1 YeCo 5.9 15.4 13.2 20.5 5.4 14.7 12.1 19.3 1.6 8 .8 6 .6 7.5 YeSu 12.1 12.6 1 2 .0 12.4 10.6 11.0 11.9 10.0 3.7 4.3 3.8 3.2 BeNo 9.8 14.8 13.3 15.8 8.2 13.8 12.5 12.6 2 .1 4.3 3.6 4.8 BeVi 12.6 14.3 13.2 12.5 12 .0 13.6 12 .8 11.0 2 .6 3.9 3.9 3.8 BeCo -- 9.7 15.5 19.0 — 8.7 13.8 19.0 -- 1.6 5.6 0.3 BeSu 9.3 15.1 12.1 15.7 10.7 14.1 11.7 15.3 3.7 4.5 2 .8 4.7 PiNo 8.9 14.6 12.3 24.2 8.3 12.6 12.0 23.1 1.9 2.7 3.6 5.7 PiVi 6.9 11.0 10.6 12.8 6.0 9.9 9.8 12.2 1.2 3.5 3.2 4.4 PiCo -- 14.8 ------14.0 ------7.0 •— — PiSu — 22.9 9.3 5.0 — 22.9 8 .1 5.0 -- 5.5 3.8 0 .1 SaNo 8.3 16.2 16.6 9.9 7.0 15.5 15.0 9.1 1.7 4.3 3.5 1.6 SaVi 7.4 14.5 13.1 12.1 6 .8 12.2 11.8 1 0.0 2.5 4.0 4.5 2.9 SaCo -- 10.4 12.4 7.2 -- 10.3 12.1 6 .0 — 1.5 3.3 3.0 SaSu 8.7 14.2 10.5 24.1 6.5 12.7 8.7 14.0 1.9 4.5 3.3 4.4 OrNo 11.6 16.6 12.2 29.7 10.6 16.0 11.7 29.0 2 .1 4.4 3.7 4.5 OrVi 12.6 16.5 13.7 14.4 11.5 14.6 13.4 12.7 3.2 4.0 5.2 4.4 OrCo 10.6 10.3 16.6 18.1 9.8 9.7 15.1 17.9 5.2 4.9 8 .1 5.2 OrSu 7.8 5.3 12 .8 — 7.3 5.3 11 .0 -- 2.7 0 .6 3.4 -- CrNo 11.1 15.0 11.5 70.0 9.9 14.3 7.7 5.4 1.4 3.2 2.5 2 .8 CrVi 10 .0 1 1.8 1 0.0 9.0 9.1 11.0 9.3 9.0 2 .2 4.6 2.5 0 .6 CrCo 3.0 15.3 —-- 3.0 15.3 ---- 0 .2 5.9 ---- CrSu ---- 9.3 ------8.5 ——-- 3.7 --
**Number of observations making up each mean shown In Appendix Table 10. 146
Table 26. Least-squares analysis of variance for mean dally oviposition of laboratory-reared bean leaf beetles through four 10-day Intervals beginning at day 11 after emergence. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Mean variation freedom square * /
Total 289 -«
-tr-fl- Generation 3 368.40
Time 3 40.37nS
Color 5 51.05*
Harking 3 63.65**
Time x Color 15 13.75nS
Time x Harking 9 23.5ln®
Color x Harking 15 20.29nS
Regression Number Linear 1 0.08ns
Residual 235 19.15
*^F oi “ 3.88 c 3 & 200 degrees of freedom
F Qj =» 2.26 c 5 & 200 degrees of freedom 2) Generation x color x marking pattern means shown In Appendix Table 27. 147
Table 27. Mean dally oviposition of laboratory-reared bean leaf beetles through four 10-day Intervals beginning at day 11 after emergence. Baton Rouge, Louisiana. 1970-71.
Generation I
Color- 11 - 20 21 - 30 31 - 40 41 - 50 Marking Pattern * n1^ x n x n x n
YeNo 5.1 26 7.0 64 7.5 52 7.9 38 YeVi 5.1 46 5.7 56 5.9 46 4.7 22 YeCo 4.7 6 4.3 18 6.4 10 2.3 10 YeSu 7.8 10 14.8 26 8 .2 18 0 .0 10
BeNo 5.2 26 7.8 72 10.4 68 7.3 34 BeVi 5.5 38 9.3 96 9.9 72 7.8 64 BeCo -- 0 -- 0 0 — 0 BeSu 9.0 2 4.0 22 0.5 20 9.7 20
PiNo 5.0 8 0.3 4 4.1 10 1.7 10 PiVi 6.7 6 8.3 6 — 0 -- 0 PiCo -- 0 -- 0 -- 0 -- 0 PiSu -- 0 -- 0 — 0 — 0
SaNo 2 .8 18 3.3 74 4.0 46 4.0 26 SaVi 9.7 16 8.9 32 5.7 30 1 2 .8 28 SaCo -- 0 -- 0 -- 0 -- 0 00 SaSu 6 .0 26 * 44 0 .6 22 0 .1 28
OrNo 5.2 52 7.4 108 6 .6 60 9.5 34 OrVi 1 2 .8 22 9.3 78 7.2 42 7.4 20 OrCo 3.0 2 9.1 16 9.5 10 14.3 10 OrSu 4.9 14 9.6 36 8.3 18 2 .8 8
CrNo 7.4 18 0.5 14 2.5 10 6 .0 4 CrVi 8 .0 6 9.7 28 7.7 12 2.9 10 CrCo 1.5 2 -- 0 — 0 — 0 CrSu «• — 0 “ •* 0 0 0
*^n is the total number of female oviposition period days per subclass in each 10-day time interval. 148
Table 27 (con tin u ed ).
Generation II
Color- 11 - 20 21 - 30 31 - 40 41 - 50 Harking Pattern x n x n x n x n
YeNo 11.5 50 11.4 76 9.9 36 4.5 16 YeVi 11.4 50 10.3 44 8 .8 22 4.5 6 YeCo 20.5 6 17.4 10 9.9 10 0 YeSu 13.1 24 4.1 16 16.5 4 -- 0
BeNo 1 2 .8 134 ,1 2 .5 142 11.1 96 7.5 48 BeVi 15.1 86 13.1 110 9.7 56 10.5 20 BeCo 9.2 10 8.9 8 — 0 -- 0 BeSu 14.1 22v 12.3 18 8 .0 4 -- 0
PiNo 12 .1 34 7.7 46 7.1 40 2 .8 30 PiVi 7.3 18 15.7 30 11.9 30 8 .0 8 PiCo 11.5 4 16.5 10 9.2 6 0 PiSu 19.8 6 2 1 .0 4 -- 0 — 0
SaNo 11.9 56 13.6 84 10.6 36 14.6 8 SaVi 11.5 48 7.8 82 12.2 38 10 .8 18 SaCo 1 2 .0 10 5.4 10 -- 0 -- 0 Sa Su 12.5 42 1 2 .8 62 12.9 26 9.3 10
Or No 12.1 66 11.5 74 15.7 36 15.7 26 OrVi 11.5 100 12.2 132 9.1 88 7.8 32 OrCo 8.4 10 10 .6 14 10.2 10 7.6 8 OrSu 0 -- 0 -- 0 — 0
CrNo 11.3 26 1 0 .8 24 5.4 16 _ a. 0 CrVi 10.7 42 11.3 78 14.4 20 6.5 6 CrCo 26.0 4 9.5 10 -- 0 0 CrSu -- 0 -- 0 — 0 -- 0 149
Table 27 (continued).
Generation III
Color- 11 - 20 21 - 30 31 - 40 41 - 50 Marking Pattern x n x n x n x n
YeNo 14.6 54 13.1 72 15.9 56 19.5 10 YeVi 13.6 46 12.4 52 11.9 22 23.3 6 YeCo 12.5 22 12.9 36 20.7 18 23.3 6 YeSu 14.6 32 10.3 20 6 .0 8 -- 0
BeNo 1 2.2 120 1 1 .8 102 12.7 50 7.7 38 BeVi 11.5 78 1 2 .0 60 16.6 12 - - . 0 BeCo 13.9 30 10.1 46 11 .6 32 9.8 22 BeSu 11.4 28 1 2 .0 28 7.5 10 0.5 2
PlNo 9.1 44 11.3 34 13.3 12 9.4 10 PiVi 10.4 30 10.7 24 0.3 10 4.5 10 PiCo 0 -- 0 -- 0 -- 0 PlSu 7.2 24 6 .2 26 5.8 20 -- 0
SaNo 11.7 82 9.9 64 5.3 20 0 SaVi 11.9 68 10 .8 100 4.8 46 7.9 20 SaCo 7.4 30 10.2 18 2 0 .8 4 -- 0 SaSu 9.2 44 9.4 48 7.0 40 0.3 12
OrNo 10.4 110 10.4 118 8 .0 56 1.9 20 OrVi 12.2 86 1 2 .8 102 1 2 .1 70 19.0 30 OrCo 16.3 26 20.5 32 14.4 18 28.8 10 OrSu 11.3 30 9.3 34 6 .0 12 -- 0
CrNo 10.5 2 1 2 .0 8 0 0 CrVl 8 .1 16 16.8 10 11 .0 8 0 CrCo -- 0 -- 0 — 0 -- 0 CrSu 14.0 4 5.5 10 0 0 (continued)-
Generation IV
11 - 20 2 1 - 3 0 31 - 40
x n x n x n n
13.5 62 8.2 58 12.5 28 8 7.3 32 5.1 42 9.8 22 14 19.7 18 17.5 20 27.2 10 2 8 .6 52 5.3 70 8.5 30 20
14.3 160 11.1 154 13.7 62 4 9.5 98 11.4 116 13.8 70 26 6 .2 12 — 0 0 0 11.7 44 14.6 52 15.3 32 16
13.2 24 15.1 20 15.3 12 6 8.5 14 12.6 20 12.6 10 2 9.2 20 10.4 20 11.5 12 0 2.5 2 — 0 — 0 0
9.0 20 2.5 8 0 0 11.5 28 6 .6 32 17.0 10 2 6 .8 28 8.3 30 9.2 18 0 14.0 20 12 .0 38 10.7 6 0
23.3 6 11.5 8 0 0 8 .0 44 12.9 50 9.9 48 34 15.5 10 19.3 22 13.3 18 8 — 0 -- 0 -- 0 0
11.7 6 1.6 8 w — 0 0 4.5 2 — 0 ----- 0 0 0 -- 0 ----- 0 0 — 0 -- 0 -- 0 0 151
Table 28. Least-squares analysis of variance for mean dally oviposition of laboratory-reared bean leaf beetles through four 10-day intervals beginning at oviposition initiation. Baton Rouge, Louisiana. 1970-71.
Source of Degrees of Mean variation freedom square
Total 261 —
Generation 3 2 0 0 .0 1
Time 3 9.85ns
Color 5 25.24ns
Marking 3 47.95ns
Time x Color 15 24.94ns
Time x Marking 9 11.09nS
Color x Marking 15 19.56ns m C CM H 00
Regression Number Linear 1 •
Residual 207 21.25
^G eneration x co lo r x marking pattern means shown in Appendix Table 29. 152
Table 29. Mean daily oviposition of laboratory-reared bean leaf beetles through four 10-day periods beginning at ovi position initiation. Baton Rouge, Louisiana. 1970-71.
Generation I
C olor- 1 - 1 0 11 - 20 21 - 30 31 - 40 Marking 1) Pattern x n x n x n x n
YeNo 6.9 116 7.3 66 5.1 28 1 .8 12 YeVi 6 .6 102 4 .0 40 3 .9 30 0 .2 12 YeCo 5.3 20 4.7 10 3.1 10 lo 0 10 YeSu 11.0 66 9.2 34 10.7 12 18.6 10
BeNo 6.9 168 9.7 82 11.4 52 8 .1 30 BeVi 8.4 182 8 .8 100 8.9 72 9.1 36 BeCo — 0 -- 0 0 — 0 BeSu 4.7 34 3 .4 30 8.3 30 10.9 20
PiNo 6.5 32 16.3 20 10.0 16 _ m 0 PiVi 5.3 20 7.5 2 -- 0 — 0 PiCo -- 0 -- 0 0 — 0 PiSu — 0 -- 0 -- 0 0
SaNo 5 .8 128 6 .8 66 8 .0 40 11.5 30 SaVi 8.3 56 10 .0 40 8 .1 22 7.1 20 SaCo -- 0 -- 0 -- 0 0 SaSu 6 .1 66 5 .4 38 4 .8 30 4 .6 28
Or No 7.9 146 6.9 96 7.0 38 3.0 10 Or Vi 9.6 104 8.9 60 8.2 20 4.1 14 OrCo 8.4 18 9.5 10 14.3 10 16.8 10 OrSu 7.6 54 9.0 24 1 .6 10 4.0 2
CrNo 5.2 26 1.7 10 3.6 10 0 CrVi 9.0 50 9.5 18 2.9 10 19.6 10 CrCo 1.5 2 -- 0 -- 0 — 0 CrSu — “ 0 0 0 0
^n is total number of female oviposition period days per subclass in each 10-day time interval. 153
Table 29 (continued).
Generation II H o H Color- S 11 - 20 21 - 30 31 - 40 Marking Pattern x n x n x n x n
YeNo 9.9 118 12.2 46 7.2 28 -- 0 YeVi 10 .1 80 14.2 22 6 .8 16 2.5 2 YeCo 18.7 10 15.3 10 9.3 6 -- 0 YeSu 10.9 32 8.1 12 -- 0 0
BeNo 12.7 252 12.3 120 7.1 64 2 .6 16 BeVi 12.4 224 11.7 76 4 .2 24 0 .9 12 BeCo 8.4 14 11.3 4 — 0 -- 0 BeSu 13.0 36 12.1 8 -- 0 -- 0
PiNo 10.9 62 7.5 40 4 .0 38 2 .6 14 PiVi 1 1 .8 52 11.2 30 13.8 6 -- 0 PiCo 12.9 10 13.7 10 — 0 -- 0 PiSu 20.3 18 — 0 — 0 -- 0
SaNo 12.2 136 13.2 48 9.4 10 .... 0 SaVi 9.9 124 10.1 58 11.0 14 — 0 SaCo 7.9 26 -- 0 -- 0 -- 0 SaSu 13.3 84 1 1 .8 42 9 .8 14 - - 0
OrNo 1 1 .8 146 15.7 58 17.4 28 16.6 14 Or Vi 10.3 196 13.0 110 8.4 56 9.9 14 Or Co 10 .0 20 9.0 10 9 .8 10 4 .0 2 OrSu 2 .0 4 ~- 0 -- 0 -- 0
CrNo 11.5 44 6.5 20 2 .0 2 0 CrVi 10.3 98 1 1 .8 44 14.1 10 — 0 CrCo 18.6 10 3.3 4 — 0 — 0 CrSu 0 0 -- 0 0 154
Table 29 (continued).
Generation III
C olor- 1 - 1 0 11 - 20 21 - 30 31 - 40 Marking Pattern x n x n x n x n
YeNo 12.2 88 14.7 64 2 0 .2 36 11.5 10 YeVi 13.5 72 1 0 .8 40 19.6 12 2 1 .0 2 YeCo 12.9 46 16.8 22 20.7 12 2 1 .0 2 YeSu 13.3 44 8.9 16 - - 0 -- 0
BeNo 1 1 .0 194 11.4 86 11.5 40 6.3 16 BeVi 1 1 .8 124 1 0.2 36 24.8 4 -- 0 BeCo 11.5 48 11.4 38 10 .6 30 8.2 20 BeSu 11.4 42 13.2 16 4.2 10 — 0
PiNo 9.7 66 1 2 .0 34 10.7 10 0 PiVi 7.8 58 10 .0 20 5.7 10 9.9 10 PiCo -- 0 -- 0 -- 0 -- 0 PiSu 6 .8 36 6.4 24 4 .7 10 -- 0
SaNo 11.3 110 7.5 36 5.3 20 0 SaVi 10.2 132 8 .6 64 4.9 20 7.2 20 SaCo 7.2 40 12.9 12 34.0 2 — 0 SaSu 8.7 76 9.1 40 4 .8 24 0 .0 10
OrNo 9.9 200 1 0 .0 86 4 .1 22 7.9 10 Or Vi 11.9 146 12.9 84 16.5 40 14.4 28 Or Co 15.7 40 2 1 .6 24 19.5 16 22.3 8 OrSu 1 0.2 44 1 0.1 22 5.7 10 — 0
CrNo 6.9 10 H 0m 0 ... 0 _ _ 0 CrVi 1 1.1 40 3 .3 8 -- 0 0 CrCo -- 0 — 0 — 0 — 0 CrSu 8.9 10 5.5 4 -- 0 .. 0 155
Table 29 (continued).
Generation IV
Color - 1 - 1 0 11 - 20 21 - 30 31 - 40 Marking Pattern x n x n x n x n
YeNo 11.5 92 9 .4 42 19.0 18 25.3 4 YeVi 7.0 56 3 .6 24 10.7 20 4.2 10 YeCo 17.7 32 25.3 16 9.5 2 -- 0 YeSu 7.3 108 7.3 42 6 .8 22 6 .0 20
BeNo 12 .0 240 14.5 110 11.8 42 0 BeVi 9.6 194 12.7 106 12.2 44 10.6 8 BeCo 6 .2 12 -- 0 -- 0 -- 0 BeSu 12.6 88 14.3 48 13.6 14 — 0
PiNo 13.0 30 16.2 20 6.9 12 —_ 0 PiVi 10.1 36 9.7 20 10.9 8 — 0 PiCo 9.0 28 1 1 .8 18 10.5 6 -- 0 PiSu 2.5 2 -- 0 -- 0 -- 0
SaNo 8 .6 36 4 .0 2 _— 0 _ _ 0 SaVi 9.4 58 11.3 14 -- 0 -- 0 SaCo 6 .2 38 10.4 28 7 .8 10 — 0 SaSu 11.5 48 15.4 16 -- 0 -- 0
OrNo 16.6 14 0 _ - 0 0 Or Vi 10.9 72 13.4 50 6.3 42 10.2 14 OrCo 14.2 26 2 0 .1 20 11.3 14 — 0 OrSu -- 0 -- 0 -- 0 -- 0
CrNo 7 .0 10 3 .3 4 0 _ _ 0 CrVi 4.5 2 -- 0 — 0 — 0 CrCo -- 0 — 0 — 0 — 0 CrSu 0 •* 0 0 —— 0 Table 30. Oviposition initiation, oviposition cessation, and mortality of laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71. Oviposition Oviposition Oviposition Day Initiation*' Cessation^) M ortality^ Cessation*) Mortality^)
2 »*> 166 130 27 4 -- 241 395 -- 68 6 317 572 — 151 8 3 388 638 -- 207 10 52 477 684 -- 267 12 166 497 702 24 319 14 352 547 718 89 380 16 521 578 729 148 434 18 598 616 742 211 477 20 632 641 753 265 509 22 674 669 761 319 553 24 691 689 766 369 572 26 702 706 769 429 609 28 712 722 771 462 629 30 721 747 775 504 658 32 726 763 777 528 679 34 730 771 782 553 701 36 734 777 784 584 713 38 736 781 785 612 725 40 738 786 788 633 734 42 741 - - . — 649 — 44 744 ---- 664 -- 46 747 ---- 679 -- 48 __ ---- 688 -- 50 ——— 695 — Observed 820 820 820 820 820
^Measured from adult emergence 2)Measured from oviposition initiation •^Measured from oviposition cessation 158
Table 31. Mortality of reproductive and nonreproductive laboratory-reared bean leaf beetles. Baton Rouge, Louisiana. 1970-71.
Days Reproductive Nonreproductive after Females Females Emergence
12 — 92 15 — 196 18 93 290 21 164 366 24 238 415 27 310 454 30 362 496 33 413 518 36 465 542 39 514 570 42 557 593 45 587 622 48 608 644 51 635 670 54 651 692 57 666 709 60 682 729 63 698 740 66 702 749 69 709 770 72 716 75 728
Observed 820 898 159
Table 32. Oviposition response of laboratory-reared bean leaf beetles, beginning at adult emergence and oviposition initiation. Baton Rouge, Louisiana. 1970-71.
Egg Production Egg Production Days from Adult ^ from Oviposition Emergence (xlO ) Initiation (xlO )
2 164 4 -- 295 6 -- 425 8 -- 530 10 — 620 12 — 707 14 53 773 16 153 836 18 228 891 20 325 933 22 414 970 24 500 997 26 574 1020 28 643 1043 30 709 1060 32 762 1072 34 813 1083 36 850 1093 38 883 1098 40 912 1104 42 934 -- 44 952 -- 46 968 -- 48 981 -- 50 993 -- 52 1002 — 2 Total Egg Production 1137 x 10 160
Table 33. Least-squares analysis of variance for mean weight of bean leaf beetles collected from St. Landry and Catahoula Parishes, Louisiana. 1971.
Source o f Degrees o f Mean )2 ) v a ria tio n freedom square
Total 118 —
Replication 4 31.26**
Color 5 1.89*
Marking 3 3.51**
Color x Marking 15 i . i o ns
Regression Number Linear 1 0 . 0002ns
Residual 90 0.81
4 & 90 degrees of freedom 1>F.01 - 3 -54 s 5 & 90 degrees of freedom F.05 " 2 ’32 '
F 0 i - 4 .0 1 i 3 & 90 degrees of freedom
^Replication x co lo r x marking pattern means shown in Appendix Table 34. Table 34. Mean weight in milligrams of bean leaf beetle color-marking pattern subclasses. St. Landry and Catahoula P arish es, Louisiana. 1971.
Color- I II III IV \ Marking Pattern x n x n x n x n X n
YeNo 13.08 126 14.86 96 14.85 108 15.03 134 15.49 82 YeVi 13.29 95 15.68 96 15.48 152 15.60 123 15.61 75 YeCo 11.82 33 14.08 32 13.01 25 13.66 19 11.76 15 YeSu 12.78 52 15.46 48 14.65 44 15.16 43 15.02 29 BeNo 12.56 89 15.09 71 14.01 315 14.93 199 15.20 70 BeVi 12.60 61 15.23 69 14.55 364 15.52 154 15.67 74 BeCo 10.74 16 13.89 19 12.38 52 14.45 34 12.43 12 BeSu 11.76 24 14.70 18 13.81 73 15.83 38 14.48 20 PiNo 12.44 19 14.05 24 14.70 34 15.13 31 15.20 14 PiVi 12.53 16 14.82 17 14.49 38 13.94 17 14.53 18 PiCo 11.17 9 14.91 8 14.86 5 17.68 4 14.30 5 PiSu 11.59 7 15.90 6 14.98 6 15.51 15 16.65 2 SaNo 13.26 59 14.57 24 13.62 42 14.78 36 15.29 15 SaVi 12.33 27 15.04 64 14.57 46 15.02 33 14.67 26 SaCo 10.24 8 14.60 8 13.99 7 13.97 7 13.33 4 SaSu 13.82 14 15.15 17 13.30 25 15.03 12 13.65 10 Or No 12.66 53 15.13 34 13.65 60 14.79 46 15.44 26 Or Vi 13.48 32 14.82 29 14.04 67 15.65 39 15.59 35 Or Co 12.58 13 13.68 6 14.10 6 14.26 9 18.13 4 OrSu 13.02 22 14.40 18 14.50 14 15.01 10 15.76 5 CrNo 13.38 31 14.62 13 14.43 61 15.67 46 15.97 15 CrVi 13.33 21 14.61 17 14.95 . 63 15.55 52 17.13 26 CrCo 10.97 3 19.30 1 13.65 10 14.62 11 -- 0 CrSu 13.10 16 15.80 5 14.35 19 15.81 18 14.96 8 162
Table 35. Relationship between mean longevity of laboratory- reared bean leaf beetles and mean weight of field- collected populations. East Baton Rouge, St. Landry, and Catahoula Parishes, Louisiana. 1970-71.
Color Longevity Weight Form in Days in mg.
Ye 37.72 14.32
Be 48.25 13.99
Pi 47.05 14.47
Sa 42.58 14.01
Or 38.10 14.53
Cr 32.02 14.88
i) r - -.720 B
Analysis of correlation shown in Appendix Table 36. 163
Table 36. Analysis of correlation between mean longevity of laboratory-reared bean leaf beetles and mean weight of field-collected populations. East Baton Rouge, St. Landry, and Catahoula Parishes, Louisiana. 1970 71.
Source o f Degrees o f Mean va riation freedom square
Total 5 —
Correlation 1 9 9 .36ns
R esidual 4 22.53 Table 37. Mortality 48 hours following treatment of adult bean leaf beetles with topically-applied concentrations of methyl parathion. Baton Rouge, Louisiana. 1971.
Dosage in Number Killed/Number Tested u g ./b e e tle Ye Be Pi Sa Or Cr
May 4. 1971
Control 0 /1 1 1/20 0 / 6 0/15 0 /11 0 / 8 .005 1/11 0 /2 0 0 / 5 0/15 2/11 0 / 8 .01 0/11 0/20 0 / 6 1/15 0 /11 1/ 8 .02 1/11 5/20 1/ 6 3/15 1/11 5 / 8 .04 6 /11 19/20 5 / 6 13/15 11/11 6 / 8 .1 11/13 21/21 11/11 13/13 7 / 7 10/10
May 12. 1971
Control 12/60 2/39 0 / 9 0/19 0/25 1/13 .01 6/60 5/40 0 /1 0 2 /2 0 1/25 1/13 .02 16/45 24/35 6 /10 13/20 12/21 7/13 .04 43/46 33/34 10/10 20/2 0 21/21 13/13 .1 41/42 38/38 11/11 21/21 20/20 14/14
June 12. 1971
Control 4/45 2/30 0/10 1/20 1/15 0 / 6 .01 7/45 0/30 0 /1 0 0 /2 0 2/15 0 / 6 .02 3/45 5/30 3/10 5/20 3/15 0 / 6 .04 37/45 28/30 7 / 8 18/20 14/15 4 / 6 .1 48/50 44/45 6 / 6 28/28 23/23 10/10
June 30. 1971
Control 1/25 .2/25 1/25 2/25 3/25 4/25 .015 6/25 2/25 3/25 6/25 7/25 2/25 .02 4/25 11/25 12/25 15/25 17/25 12/25 .03 19/25 21/25 7/ 8 22/25 23/25 20/25 .04 24/25 24/25 - / o 17/17 24/25 22/25
July 24. 1971
Control 1/25 0/25 0/25 0/25 1/25 1/25 .015 4/25 5/25 4/25 5/25 6/25 0/25 .02 11/25 8/25 8/25 16/25 10/25 3/25 .03 12/25 20/25 13/25 20/25 10/25 16/25 .04 17/25 18/25 18/25 16/25 14/25 17/25 165
Table 38. Least-squares analysis of variance for overall percentage parasitization of the bean leaf beetle by Celatoria diabroticae (Shimer). St. Landry Parish, Louisiana. 1972.
Source o f Degrees o f Mean v a ria tio n Freedom square '
Total 23 —
Colors 5 7.34*
Markings 3 1.15nS
Regression Number Linear 1 0 . 33ns
Residual 14 1.65
n(. = 2.96 c 5 & 14 degrees of freedom • vJ 2) Figures used in the analysis are shown in Appendix Table 39. Table 39. Overall percentage parasitization of the bean leaf beetle by Celatoria diabroticae (Shimer). St. Landry Parish, Louisiana. 1972.
Color
Marking Ye Be Pi Sa Or Cr
% n % n % n % n % n % n
No 7.83 1175 3.95 1467 2.83 671 3.76 744 4.88 841 2.12 330
Vi 7.54 1167 4.37 1441 3.37 594 4.17 672 4.61 738 5.80 345
Co 4.46 157 4.78 230 2.02 99 5.69 123 5.26 114 4.76 42
Su 7.78 437 3.38 533 1.27 236 2.56 256 4.20 238 1.47 136 166 Table 40. Mortality of bean leaf beetles subjected to a constant temperature of 100° F. Baton Rouge, Louisiana. 1972.
Days Color Marking Pattern a fter I n itia tio n Ye Be Pi Sa Or Cr No Vi Co Su
1 48 14 9 20 35 1 54 50 1 22 2 376 320 121 156 202 65 544 443 58 195 3 428 402 153 201 234 75 629 540 77 247 4 432 438 173 227 256 79 668 589 88 260 5 439 450 179 232 265 -- 689 603 89 263 6 440 451 -- — -- — — — — 265
Observed 440 451 179 232 265 79 689 603 89 265 167 Table 41. Mortality of bean leaf beetles subjected to a constant temperature of 95° F. Baton Rouge, Louisiana. 1972.
Days Color Marking Pattern a fte r I n itia tio n Ye Be P i Sa Or Cr No Vi Co Su
1 24 17 10 6 10 9 18 43 6 9 2 64 66 22 22 36 19 76 97 22 34 3 91 89 45 50 69 35 137 167 27 48 4 117 108 50 60 73 37 162 194 31 58 5 235 279 139 116 172 85 396 428 62 140 7 323 389 183 153 208 108 525 570 77 192 8 379 504 234 187 250 135 666 694 88 241 9 406 529 254 201 266 147 710 741 94 258 10 425 571 302 238 293 159 798 800 103 287 11 431 586 317 255 309 170 832 838 108 290 12 434 592 323 263 320 174 849 852 a* a* 297 13 434 596 326 268 323 176 860 857 -- 298 14 440 598 328 269 324 -- 865 861 — 301 15 441 — ——— -- 866 — — —
Observed 441 598 328 269 324 176 866 861 108 301 Table 42. Mortality of bean leaf beetles subjected to a constant temperature of 90° F. Baton Rouge, Louisiana. 1972.
'■) Days Color e / Marking Pattern a fter I n itia tio n Ye Be Pi Sa Or Cr No Vi Co Su
2 106 64 17 21 36 5 121 89 13 26 4 156 113 43 44 69 21 194 180 29 44 6 176 155 73 60 89 44 267 231 41 59 8 198 171 78 66 99 49 298 253 44 67 10 222 201 94 88 128 64 371 292 54 81 12 237 286 132 151 171 83 507 377 67 110 14 310 393 159 184 208 94 632 472 94 151 16 331 439 189 203 228 112 702 540 97 164 18 342 482 203 216 235 118 741 572 104 179 20 359 514 214 229 252 123 782 610 111 188 22 364 530 225 230 265 125 805 631 112 191 24 365 549 235 237 272 129 820 652 116 199 26 368 559 241 244 276 132 830 657 118 206 28 372 573 242 252 285 133 844 679 119 213 30 376 584 247 255 288 134 853 688 121 216 32 377 593 251 258 294 861 700 122 219 34 378 598 252 261 298 -- 867 706 -- 224 36 381 601 253 263 299 -- 869 708 -- 224 38 — 606 254 264 300 -- 873 710 -- 227 40 — 608 — 265 301 — 875 713 -- 228
Observed 381 615 254 266 307 134 884 721 122 230 169 Table 43. Mortality of bean leaf beetles subjected to a constant temperature of 80° F. .Baton Rouge, Louisiana. 1972-73.
Days Color Marking Pattern a fte r I n itia tio n Ye Be P i Sa Or Cr No Vi Co Su
2 48 30 23 15 18 9 55 60 8 20 4 88 60 33 32 40 24 112 115 16 34 6 131 112 49 46 54 30 161 181 23 57 8 147 126 57 54 65 32 181 208 26 65 10 176 170 74 69 79 38 227 268 29 81 12 193 190 81 77 81 40 246 295 31 89 14 229 220 100 88 100 49 291 343 39 108 16 240 237 116 93 107 50 317 361 42 118 18 272 277 126 108 115 58 361 411 46 133 20 284 287 136 115 120 61 382 432 47 137 24 314 326 145 125 135 73 428 492 51 153 28 348 365 158 134 150 77 470 538 61 169 32 380 387 174 147 163 81 508 583 64 183 36 391 407 181 151 177 89 527 610 69 196 40 403 426 192 164 182 94 548 644 74 201 46 422 440 199 168 192 103 570 669 76 215 52 443 465 215 176 213 111 603 713 86 227 58 455 491 218 187 227 118 640 738 89 235 64 465 509 222 194 236 125 665 758 93 242 70 478 521 226 199 241 127 681 774 95 249
Observed 536 623 260 241 278 158 804 897 107 295 Table 44. Mortality of bean leaf beetles subjected to a constant temperature of 70° F. Baton Rouge, Louisiana. 1972-73.
Days Color Marking Pattern a fte r I n itia tio n Ye Be Pi Sa Or Cr No Vi Co Su
7 39 42 17 21 21 15 68 61 12 13 14 85 80 26 36 38 22 106 120 19 41 21 118 107 34 49 54 25 140 162 28 56 28 152 143 49 70 78 34 191 225 42 67 35 171 168 61 80 93 42 228 262 46 78 42 187 189 71 89 97 43 242 290 52 91 49 222 222 82 105 113 46 302 360 61 113 56 257 253 92 114 124 55 319 380 70 125 63 282 284 107 128 135 56 361 422 75 136 70 304 316 119 141 151 64 407 461 82 147 77 312 341 126 155 160 70 439 485 83 158 84 316 358 131 158 168 72 455 501 86 162 91 334 395 146 170 184 80 495 552 88 175 98 342 428 153 187 201 84 536 579 91 190 105 356 447 160 194 209 86 557 600 97 199 112 362 460 167 203 217 88 571 620 98 209 119 372 495 175 213 228 90 601 651 99 223 126 381 509 183 224 237 92 623 677 99 232 133 385 526 189 232 245 99 639 704 100 239 140 390 540 197 240 251 102 656 721 104 245
Observed 416 592 213 260 285 117 707 791 114 277 171 Table 45. Mortality of bean leaf beetles subjected to a constant temperature of 60° F. Baton Rouge, Louisiana. 1972-73.
Days Color Marking Pattern a fter I n itia tio n Ye Be Pi Sa Or Cr No Vi Co Su
7 16 12 5 8 7 3 22 19 6 5 14 53 33 16 31 20 10 73 55 12 24 21 94 54 34 43 37 13 115 94 21 46 28 131 71 46 61 43 17 144 133 28 65 35 161 91 55 87 66 24 193 175 36 81 42 198 121 69 111 90 31 258 225 40 98 49 235 153 82 136 114 46 312 286 49 120 56 267 170 99 158 130 54 352 332 54 140 63 294 195 112 176 144 58 386 376 60 157 70 324 216 127 192 168 63 442 410 67 171 77 334 228 130 200 179 66 466 437 68 176 84 357 242 137 211 187 70 491 459 72 182 91 371 258 143 217 194 72 514 473 76 192 98 388 275 147 224 204 74 544 488 80 198 105 412 281 152 229 218 80 571 511 82 208 112 423 294 155 237 226 82 593 527 82 215 119 425 301 156 241 230 85 607 531 82 218 126 427 312 157 246 232 86 615 541 83 221 133 428 321 159 248 237 87 622 545 85 228 140 431 324 163 252 240 90 633 548 86 233
Observed 440 347 165 267 242 96 663 557 94 243 Table 46. Mortality of bean leaf beetles subjected to a constant temperature of 50° F. Baton Rouge, Louisiana. 1972-73.
Days Color Marking Pattern a fter I n itia tio n Ye Be Pi Sa Or Cr No Vi Co Su
7 52 26 21 21 14 4 72 42 14 10 14 70 46 35 46 36 7 119 76 18 29 21 86 73 58 61 52 12 165 104 31 44 28 90 79 59 65 55 12 170 110 32 50 35 96 91 68 72 68 16 197 127 33 56 42 101 103 73 82 71 17 212 139 37 61 49 113 108 77 86 75 18 223 149 40 67 56 121 116 80 94 81 19 239 157 42 75 63 125 131 88 104 92 21 262 172 45 84 70 137 142 93 113 96 21 285 184 46 89 77 149 158 101 122 109 25 314 205 53 94 84 166 180 113 135 113 28 344 228 58 107 91 171 189 117 145 H 9 30 358 243 63 109 98 171 206 123 152 125 33 376 255 67 114 105 172 208 123 152 129 34 378 261 67 114 112 179 218 128 158 135 39 395 271 73 120 119 187 227 137 167 138 41 413 284 77 125 126 201 233 140 170 141 41 427 293 78 130 133 202 240 142 172 145 41 438 296 79 131 140 202 244 145 173 148 43 444 298 80 135
Observed 241 356 185 208 194 61 554 420 113 160 Table 47. Slope of log-time probit lines obtained in the bean^leaf beetle temperature tolerance study. Baton Rouge, Louisiana. 1972-73.
Temperature (° F.) o o o o O o 50° O' 00 o O 95° 100°
Color
Ye 1.44+0.12 3.13+0.18 2.28+0.11 1.76+0.06 2.45+0.24 4.06+J.50 6.33+0.83
Be 1.62+0.09 2.82+0.15 2.31+0.18 1.79+0.04 3.0240.27 4.51+0.72 6.18+0.62
Pi 1.56+0.07 3.23+0.17 2.34+0.19 1.82+0.07 3.19+3.23 4.40+0.87 5.69+0.59
Sa 1.79+0.12 2.92+0.11 2.22+0.17 1.64+0.05 3.23+0.26 4.0740.48 5.49+0.27
Or 1.65+0.07 3.40+0.26 2.12+0.14 1.71+0.08 2.75+0.19 3.98+3.43 5.01+0.52
Cr 1.69+0.14 2.90+0.13 1.84+0.13 1.58+0.07 3.54+0.23 3.82+0.57 8.30+1.75
Marking Pattern
No 1.58+0.10 2.93+0.16 2.26+0.18 1.69+0.05 2.94+0.24 4.44+0.56 5.74+0.76
Vi 1.51+D.09 3.33+0.21 2.18+0.13 1.7840.05 2.79+0.22 4.03+0.57 5.66+0.51
Co 1.43+0.09 2.51+0.12 2.18+0.10 1.75+0.08 3.06+0.30 3.39+0.14 6.80+0.57
Su 1.91+0.08 2.88+0.08 2.20+0.14 1.74+0.05 2.89+0.24 4.30+0.60 5.83+0.27
Computed from mortality data shown in Appendix Tables 40-46. 174 Table 48. Mortality of bean leaf beetles subjected for 7 days to constant temperatures ranging from 50 to 100° F. Baton Rouge, Louisiana. 1972.
Colors Number dead/Number Temperature observed ° F. Ye Be Pi Sa Or Cr
60 16/440 12/347 5/165 8/267 7/242 3 / 96 70 39/416 41/592 17/213 21/260 21/285 15/117 80 138/536 118/623 54/260 50/241 59/278 31/158 90 193/381 169/615 77/254 65/266 99/307 49/134 95 323/441 389/598 183/328 153/269 208/324 108/176 100 440/440 415/451 179/179 232/232 265/265 79/ 79
Markings
Temperature Number dead/Number observed ° F. No Vi Co Su
60 22/563 19/557 6/ 94 5/243 70 68/707 61/791 12/114 13/277 80 170/804 192/897 26/107 61/295 90 293/884 250/721 43/122 67/230 95 725/866 570/861 77/108 192/301 100 689/689 603/603 89/ 89 265/265 VITA
Donald Charles Herzog was born to Carl C h ristian and Leona Anna
Herzog at Lampassas, Texas, on April 3, 1944. He attended St. Paul
Lutheran Day School in Wilson, Texas, and graduated from Wilson High
School in May of 1962.
From September 1962 until May 1966, He attended Texas Techno logical College, Lubbock, Texas, where he received the Bachelor of
Science Degree in Entomology.
He married the former Sandra Jane Koslan of Wilson, Texas, in
May of 1966, and is the father of one daughter, Dawn Michelle.
In June of 1966, he entered the Graduate School of Louisiana
State University under an NDEA T itle IV Fellowship in the Department of Entomology. He received the Master of Science Degree in May of
1968.
In September of 1968, he was inducted into the U.S. Army. After serving a one-year’s tour of duty in Viet Nam with the 101st Airborne
Division he returned to the Graduate School of Louisiana State
University in June of 1970 under the NDEA Title IV Fellowship. With the expiration of the fellowship in September 1970, he was granted a
Graduate Assistantship. Since July 1971, he has been employed fu ll time as a Research Associate in the Department of Entomology. He is currently a candidate for the Doctor of Phiolosophy Degree.
176 EXAMINATION AND THESIS REPORT
Candidate: Donald Charles Herzog
Major Field: Entomology
Title of Thesis: Some Biological Implications of Polymorphism in the Bean Leaf Beetle, Cerotoma trifurcata (Forster) Approved:
Major Professor and Chairman
Dean of the Graduate School
EXAMINING COMMITTEE:
J l 2
Date of Examination:
April 25, 1973