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Graduate Student Theses, Dissertations, & Professional Papers Graduate School
1988
Resource allocation to prey capture tissue in the aquatic carnivorous plant Utricularia vulgaris in northwestern Montana waters
Susan J. Trull The University of Montana
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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Resource Allocation to Prey Capture Tissue in the
Aquatic Carnivorous Plant, Utricularia vulgaris,
in Northwestern Montana Waters
By
Susan J . Trull
B.A., Carleton College, 1982
Presented in partial fulfillment of the
requirements for the degree of
Master of Arts
University of Montana
1988
Approved by:
Chairman, Board of Examiners
Dean, Graduate School /
/^ 7 fb Date
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Trull, Susan J., M.A., June 1988 Botany
Resource Allocation to Prey Capture Tissue in the Aquatic Carnivorous Plant, Utricularia vulgaris, in Northwestern Montana Waters (226 pp.)
Director: Vicki J. Watson '/ijv/"
Utricularia vulgaris L. plants were collected in northwestern Montana, from sites assumed to differ in dissolved inorganic nutrient availability. Plants were found to vary in their extent of prey capture tissue (bladders). To test the hypothesis that waters of low nutrient availabilities Induce greater development of prey capture tissue than do waters of higher nutrient availabilities, U^. vulgaris plants were raised under controlled laboratory conditions. A common garden experiment was conducted, and U^. vulgari s plants were found to retain site-specific characteristics of bladder production and other morphological traits. It was concluded that these traits are under the control of genotype and/or the environment of the turion-formîng plants rather than under the control of the environment under which turions develop into plants. Several experiments were conducted in which turions from the same site were exposed to different concentrations of nutrients and prey. Feeding regime and nutrient solution strength did not significantly affect morphological measurements used as indicators of prey capture tissue development. Duration of dormancy (i.e. length of time before experiments were begun) did affect morphological measurements. Development of prey capture tissue observed in lab-grown plants was primarily explained by their sites of origin. Plants originating from sites thought to have lower nutrient levels exhibited more prey capture tissue than did plants originating from sites thought to be rich in nutrients.
I I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Acknow1edgements
I thank the members of my committee for their advice and
encouragement: Dr. Vicki Watson (who read and commented on innumerable
drafts). Dr. Dick Fritz-Sheridan (who asked imaginative questions), and
Dr. Andy Sheldon (who saw this project through from the first, summer
afternoon graph). I also acknowledge the support and initial advising
of Or. James Habeck. Dr. Susan Knight provided invaluable culturing
suggestion from her research with Utricularia, for which I am most
grateful. Dr. Jack Stanford provided information on sites and facilitated
use of Biological Station equipment. Dr. Dave Patterson helped with data
analysis, and Mr. Dick Lane with preparation of computer files.
I thank Dr. Dave Bilderback for facilitating growth chamber use, and
Mr- A1 Johnson for assistance in obtaining equipment, culturing Daphnia,
and use of the greenhouse. Michael Matranga and Ed Keller of ChemStores
cheerfully let me take liters of distilled water; the Botany Department
provided technical and logistic support.
I owe thanks to Lynell Morrison and Nora Leetch for turion-
collecting prowess, and to Dr. Ernie Schulyer for other field assistance.
Lastly, I thank my family and friends for moral support, suggestions,
and other aid, in particular: Mark Bjorlie, Helena Chambers, Frank Dugan,
Karen Haagenson, Peter Lesica, Denise Mott, Lisa Schassberger, Jeff
Strachan, and Phil Tourangeau. Lisa Campbell has been a constant source
of encouragement, ideas, and companionship during long hours on the
third floor, and 1 thank her wholeheartedly.
I I I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Table of Contents
page
Abstract •<
Acknowledgements...... îii
Table of Contents...... iv
List of Tables...... vt
List of Figures...... viî
Chapter
1. Introduction...... 1
2. Study Organism...... 10
3- Literature Review...... 19
I. Feeding Experiments...... 20
I I . Constraints and Confounding Factors...... 32
111. Evolutionary Aspects...... 35
4. Study Sites...... 39
5- Methods and Materials...... 47
I. Summer Field Collection...... 47
11. Common Garden Experiment...!...... 51
III. Diet Experiment...... 60
6. Results...... 67
I. Summer Field Collection...... 67
II. Growth under Laboratory Conditions...... 67
111. Common Garden Experiment...... 69
IV. Diet Experiment...... 71
7 . Discussion...... 83
1. Summer Field Collection...... 83
I V
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. I I. Common Garden Experiment...... 89
III. Diet Experiment...... 101
Summary...... 114
Literature Cited...... 116
Appendix
A. Data Summaries, Boxplots, and Complete MANOVA Results for U. vulgaris Plants Collected in Summer...... 125
B. Data Summaries, Boxplots, Complete MANOVA Results, and Profile Plots for jJ. vulgaris Plants Raised from Turions in a Common Garden Experiment...... 141
C. Data Summaries, Boxplots, Complete MANOVA Results, and Profile Plots for JJ. vulgar i s Plants Raised from Turions in a Diet Experiment...... 168
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. List of Tables
Table page
1. Significance of Collection Site on Morphological Characters in ]J. vulgaris Plants Collected in Summer...... 75
2. Significance of Collection Site, Replicate Experiment, and the Interaction between These, on Morphological Characters in L[. vulgar i s Plants Raised from Turions in a Common Garden Experiment...... 76
3. Influence of Collection Site and Replicate Experiment on Formation of New Turions by L[. vulgar i s Plants Raised from Turions in a Common Garden Experiment -...... 77
4. Significance of Collection Site, Nutrient Solution Strength, Feeding Regime, and Interactions between these Factors, on Morphological Characters in vulgaris Plants Raised from Turions in a Diet Experiment...... 78
5- Significance of Experimental Season, Nutrient Solution Strength, Feeding Regime, and Interactions between these Factors, on Morphological Characters in jj. vulgaris Plants Raised from Turions in a Diet Experiment...... 79
6. Influence of Nutrient Solution and Feeding Treatments on Formation of New Turions by jj. vulgaris Plants Raised from Turions in a Diet Experiment...... 80
7- Statistics Showing the Association between Formation of New Turions and Treatments for jj. vulgar i s Plants Raised from Turions in a Diet Experiment...... 82
V I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. List of Figures
Figure page
1. Hypothetical Changes in Costs and Benefits of Bladder Production in Utricularia vulgaris as Nutrient Availabilities Increase...... 4
2. General Morphology of Utricularia vulgaris Showing Energy and Material Source and Sink Compartments-...... 6
3- Study Site Locations...... 40
4. Experimental Design for the Common Garden Experiment...... 56
5. Experimental Design for the Diet Experiment, Showing Treatment Number...... 6l
VI I
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter One
Introduction
Carnivorous plants have been studied for years, both
as objects of popular curiosity and as subjects for
physiological research, because of the paradox they
represent as "heterotrophic autotrophs" (Liittge, 1983).
While the plants are now known to absorb nutrients from
captured prey, the necessity of these supplements for
plant survival and reproduction is still a matter of
debate (Liittge, 1983; Sorenson and Jackson, 1968). Some
species are obligate carnivores while others seem to be
facultatively carnivorous (Skutch, 1928), growing
indefinitely without ancillary resources, although
perhaps not as vigorously as with prey inputs, or not to
reproductive stages (Arber, 1920; Dore Swamy and Mohan
Ram, 1971; Sorenson and Jackson, 1968).
The costs and potential benefits of carnivory are
also not clear, and seem to vary with the fertility of
particular habitats (Benzing, 1987). Prey-trapping
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. structures are thought to be modified leaves (Juniper,
1986), so that ontogenetic costs of production are
perhaps not high. However, traps are less suitable for
photosynthesis than are leaves, especially the more
advanced trapping mechanisms (Benzing, 1987), so that
a high return in terms of nutrients is necessary to
compensate for the resource outlay.
Resources shunted to trap production may diminish
the pool available for flowering, fruiting, and general
growth (Bloom, Chapin, and Mooney, 1985), but the ability
to supplement inorganic nutrient sources may allow these
plants to survive in areas where non-carnivorous plants
are poor competitors. These areas are most likely to be
those which are limited by nutrients and not by light,
water or some other factor, since the carnivorous habit
can only supplement nutrient supplies (Benzing, 1987).
In short, the costs must be outweighed by the benefits in
areas where carnivorous plant species are common and
populations are large (Benzing, 1987; Heslop-Harrison,
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1978).
The development of a strategy to mitigate substrate
nutrient paucity raises questions of resource allocation
in these plants; how does a plant divide its resources
between formation of prey capture tissue (PCT) and other
tissues under varying environmental conditions, and how
flexible is this resource allocation?
In a nutrient-poor environment, PCT development
requires no more materials than it does in a nutrient-
rich environment. But, energy costs to obtain these
materials might be greater due to their relative
dilution. However, the benefits of ancillary resources
should be greater in the extreme environment than in an
environment where nutrients are readily available. That
is, both the costs and benefits of trap possession
probably decrease as nutrient availability increases.
Figure 1 shows these cost-benefit relationships, and
suggests that carnivory is only adaptive in habitats with
low to moderate nutrient availabilities. A carnivorous
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 1. Hypothetical Changes in Costs and Benefits of Bladder Product ion in Utricularia vulgari s as Nutrient Availabilities Increase
carni vory not competi ti ve
carnivory carnivory adaptive maladaptive costs or benef i ts
costs= - benef i ts nutrient availability
Legend :
Material costs
Material and energy costs —. —.—.—
Gross benefits (i.e. relative importance of nutrients gained from carnivory) ......
Net benefits (i.e. gross benefits - material and energy costs) —
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. plant species would be expected to allocate more
resources to PCT development in its nutrient-poor
habitats, and more to non-PCT when growing in nutrient-
rich sites. Indeed, Givnish et al. (1984) report that
some carnivorous plants produce only non-PCT, or tissue
with reduced carnivorous function, during seasons when
nutrient availability is not the limiting factor to
growth.
My research centered on this allocation question,
with respect to the aquatic carnivorous plant,
Utricularia vulgaris L. : do U. vulgaris plants
growing in waters of low nutrient availability allocate
more of their carbon resources to PCT production
(bladders) than do U . vulgaris plants growing in
waters of greater nutrient availability? Potential
sources and sinks of energy and materials in
U , vulgaris are diagrammed in Figure 2.
To investigate the above question, one needs to know
the nutrient and prey levels to which a plant is exposed
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ■DCD O Q. C g Q. Figure 2. General Morphology of Utricularia vulgaris Showing Energy and Material Source and Sink Compartments ■D CD Sources : Sinks : C/)W 3o' O 3 CD f 1owers 8 frui ts
CD ^ turions 3. 3" CD ■DCD O Q. foliage ^ C a o 3 ■D O f meri steins* CD Q. stem bladders
■D CD primary bladder's (/)C/) secondary bladders bladders*
-'production, photo symbiont N fixers respiration, maintenance Since field sampling for nutrient levels is fairly
ambiguous, and for prey levels even less conclusive, due
to patchiness in space and time (Wetzel, 1983), I raised
Ü. vuïgaris under controlled laboratory conditions.
Bladderworts can be raised under controlled
conditions from turions, embryonic plants that arise
vegetatively on the "parent" plant in fall, are dormant
in winter, and develop into mature plants in spring.
Turions are protected by a layer of mucilage, thus
bladders are axenic. The dormancy of turions can be
broken experimentally by high temperatures (Winston and
Gorham, 1979a.)
Because turions already possess bladders, a
preliminary question must be addressed before the above
prediction can be examined, namely, is bladder production
controlled genetically, controlled by the conditions to
which the tur ion-forming "generation" was exposed, or
controlled by the conditions in which the plant develops
from the turion? Part of my research focused on this
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 8
problem of control of bladder production.
While much work has been done on carnivorous plant
taxonomy, morphology, and trapping physiology, only a few
studies have examined the effects of prey nutrients on
growth in carnivorous plants, and even fewer have
addressed resource allocation (Bosserman, 1983; Luttge,
1983; Pringsheim and Pringsheim, 1962). In this study,
I used laboratory and field experimentation to compare
resource allocation to PCT by Utricularia vulgaris
plants under differing prey and nutrient regimes.
My study consisted of three parts:
1. Summer field collections of mature U . vulgaris
plants to compare morphological characters and extent
of PCT tissue in plants from different sites.
2. A common garden experiment to compare development
of PCT and other tissues in plants grown from turions
collected from two different sites, when these
turions were grown under the same dissolved nutrient
conditions, without prey supplements.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3. A "diet" experiment to compare the effects of
different dissolved nutrient levels and feeding
regimes on development of PCT and other tissues in
plants grown from turions collected from two
different sites.
These three experiments were designed to examine the
degree of development of PCT under nutrient-poor and
nutrient-rich conditions, i.e. to test my resource
allocation prediction that a greater proportion of
resources are allocated to PCT under nutrient-poor
conditions. The experiments also would show whether
U , vulgaris plants respond to ambient conditions
despite past history. The experiments would not
distinguish whether any influence of past environment
was due to "parental" genotype, to "parental"
developmental environment, or both.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Two
Study Organism
Utricularia vulgaris, the greater or common
bladderwort, is an aquatic, floating, rootless,
carnivorous plant (Hitchcock and Cronquist, 1973). It
is a member of the family Lentibulariaceae, which
includes other carnivorous plant genera. The genus
Utricularia holds about 250 species, including other
floating aquatics like U, vulgaris, semi-ter res trial
and terrestrial forms, and epiphytes. The genus is
thought to have evolved by the Pliocene era (Muller,
1981), and fossil turions of U . vulgaris have been
found from the German interglacial period (Jung, 1976).
The chromosome number of CJ. vuigaris is n = 18 to 20
(Kondo, 1969).
U . vulgaris is circumboreally distributed in
slow-moving streams, lakes, ponds, boggy areas, and wet
meadows, and is fairly common (Ceska and Bell, 1973;
Meyers and Strickler, 1979; Rossbach, 1939). Its
10
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habitats range from acidic through neutral to alkaline,
and vary in temperature, nutrient availability, light
availability, and associated species (Ceska and Bell,
1973; Rossbach, 1939; this investigation).
The growth form of U. vulgaris is stoloniferous,
with side branches at a distance from the apical
meristem(s). Leaves arise alternately, are highly
dissected and generally two-parted at the base, with a
few teeth on some segments (Ceska and Bell, 1973; Crow
and Hellquist, 1985; Fig. 2.) However, the terms "stem"
and "leaf" are used mainly for convenience since organ
morphology in U. vulgaris is not readily homologous
with other vascular plants. The vegetative plant has
been suggested to be entirely a root system, a stem
system, or a single, much divided leaf, as well as a stem
and leaf combination (Arber, 1920; Sculthorpe, 1957).
U . vulgaris grows at branch tips and decays behind,
often fragmenting into several pieces, each of which can
survive and propagate. Indeed, the vegetative
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propagation capablities of U . vulgaris are incredible:
almost any cell, whether of flower, leaf, stem, or turion
leaf (see below), can become meristematic, dividing to
form a new stem and eventually populating an entire pond
(Arber, 1920; Goebel, 1904 and Gluck, 1906 in Sculthorpe,
1967) .
U . vulgaris also reproduces sexually, through
the formation of yellow, bilaterally symmetric flowers
which are supported on a scape above the water surface
(Hitchcock and Cronquist, 1973). Flowers may be self
fertilizing (Winston and Gorham, 1979a) or chasmogamous.
Pollen is stephanocolporate (Kapp, 1969; Thanikaimoni,
1966) and fruits are capsules with many small,
endospermless seeds (Ceska and Bell, 1973; Hitchcock,
Cronquist, and Ownbey, 1959).
U . vulgaris is perennial, overwintering in cold
climates through the formation of turions, or winter buds
(Rossbach, 1939 ; Sculthorpe, 1967). Turions are clusters
of leaves with condensed internodes that are formed in
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late summer, or when the plant experiences cold, nutrient
or water stress (Maier, 1979; Sculthorpe, 1967). Turions
are covered by coarser leaves, turn brownish when mature,
and possess a thick coating of mucilage. These
structures usually remain attached to the senescing
parental stem (which by autumn is only a tough vascular
strand within a spongy cortical layer, most leaves having
abscised), and are pulled downward by this stem, so that
they overwinter below the ice (Arber, 1920; this
investigation). With further decay of the old plant, the
turions are released and float to the surface. As
temperatures warm in the spring these turions reflex,
green up and elongate into new plants (Winston and
Gorham, 1979a). Turions may fragment and single coarse
leaves may each give rise to several plantlets. Turions
are high in starch and stored materials, and therefore
can withstand drying and freezing well (Maier, 1973;
Winston and Gorham, 1979a).
Utricularia vulgaris has a reduced vascular system.
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consisting mainly of a few, poorly developed tracheids
and small groups of phloem elements. There is an
endodermis, and the cortex has lacunae and some fibrous
ground tissue (Sculthorpe, 1967). It follows the C-3
photosynthetic pathway, and bladders are photosynthetic
(Luttge, 1983). Photosynthetic rates are probably low
relative to terrestrial plants (Thai, Haller, and Bowes,
1976).
The common bladderwort captures crustaceans, insect
larvae, fish fry, rotifers and other aquatic organisms,
sometimes including algae and the vascular plant Wolffia,
in small bladders, or utricles, by suction (Botta, 1976;
Hegner, 1925; Roberts, 1972). The utricles are of three
types: stem bladders, occurring in clusters of zero to
five at petiole bases; primary bladders, usually larger,
0.5 to 5 mm long, occurring singly near the principal
bifurcations of each leaflet ; and secondary bladders,
smaller, and occurring at distal leaf divisions
(Hitchcock and Cronquist, 1982; Wallace, 1977, 1978;
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this investigation; Fig. 2). These three types are all
functional and capture prey of different sizes (Wallace,
1977) .
Bladders vary in size and number per leaflet, and
are formed while the leaflet is still part of the
bunched, meristematic tip. They seem to arise from both
leaf and stem tissue, but exact homology is under
contention (Arber, 1920; Heywood, 1978). Utricles are
light to bright green when young, sometimes with a
reddish, anthocyanin shading, becoming violet to black
with age and use (Botta, 1976; Lloyd, 1933; this
investigation).
Utricles consist of a thin layer of cells, nearly
transparent, with many two- and four-armed glands that
secrete digestive enzymes (acid phosphatase, protease,
esterase) and absorb digestive products (Slack, 1979).
Utricles are attached to the stem and the leaflet arms
by short stalks and easily abscise, with age as well as
poor growing conditions, algal infections, and excessive
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movement or handling (Lloyd, 1933; Sorenson and Jackson,
1968; this investigation). Bladders have a trap door
that seals tightly, and two types of hair-like organs,
called antennae and bristles (Darwin, 1897), project from
this area. Antennae are long, branched processes which
may help to guide prey to the traps, or to protect the
tripping mechanism from too large aquatic visitors
(Johnson, 1987; Meyers and Strickler, 1979). Bristles
are non-branched, shorter organs which function as
triggers to open the door, allowing an inrush of water
and prey (Hegner, 1925; Luttge, 1983; Sydenham and
Findlay, 1973). The bladder cells actively transport
chloride anions outward, and sodium and potassium cations
inward, resetting the bladder, and forcing it to assume a
concave shape when ready to fire. There may also be an
electrical, excitatory step in trap firing (Sydenham and
Findlay, 1973).
Prey live for varying periods of time within the
bladders, and are digested gradually (Arber, 1920). Some
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algal species and protozoans not only survive, but grow
and multiply within the utricles. Some of these algae
are cyanophytes, and fix nitrogen which is then released
to the plant (Botta, 1976; Wagner and Mshigeni, 1986).
Some large prey, such as mosquito larvae and fish fry,
are caught by head or tail and ingested in segments
(Hegner, 1925).
Utricularia does not have major economic value,
although it is eaten by a few fish and provides shade
and shelter for them, and is used as fodder for pigs
and cattle (Sculthorpe, 1967). The plants also help
control mosquito larvae populations (Jha, Jha, and
Kumari, 1978; Schwartz, 1974; Skutch, 1928), help
control the spread of schistosomiasis in the Caribbean
by capturing flukes (Gibson and Warren, 1970), and may
help control the spread of radioactive waste (Deksbakh,
1964). U* vulgaris may also have some value as a "green
fertilizer" in rice fields because of the cyanophyte
nitrogen fixers associated with it (Wagner and Mshigeni,
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1986; Woelkering, 1976). U . vulgaris is occasionally
a nuisance weed in waterways (Heywood, 1978).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Three
Literature Review
The phenomenom of carnivory in plants has prompted
investigations into why it might be adaptive, and what
selection pressures might have led to its evolution. The
general consensus is that carnivory fills macro- and
micronutrient needs for plants in low nutrient
environments (Folkerts, 1982; Heslop-Harrison, 1978).
Carnivorous plants seem to have adequate photosynthetic
pigments for securing carbon (Pringsheim and Pringsheim,
1962), and many carnivorous plants have reduced root
systems suggesting prey inputs are compensatory
(Schmucker and Linnemann, 1959, in Aldenius, Carlsson
and Karlsson, 1983).
Heslop-Harrison (1978) reported that nitrogen and
phosphorous are important gains from carnivory, but which
is more important depends on the habitat (Benzing, 1987).
On the other hand, Folkerts (1982) suggested that the
acid nature of many carnivorous plant habitats decreases
19
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micronutrient availability, hence carnivory may give its
greatest benefit in securing these elements. He also
felt that carnivory may only be important when the
habitats are under nutrient stress (e.g. when Gulf Coast
pitcher plant bogs have not been fire-swept in a long
time.)
I. Feeding Experiments
Feeding experiments have been performed on several
species of carnivorous plants, and differing results have
been found. Aldenius et al. (1983) grew Pinguicula
vulgaris L. on local soil and enriched local soil, with
and without insect supplements. They found that both
watering with a complete nutrient solution and addition
of insects caused increased dry weights, increased
numbers and lengths of leaves, and increased nitrogen
and phosphorous tissue concentrations. They concluded
that P. vulgaris was using nitrogen and phosphorous
from the captured insects, as well as some other
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substance that helped the roots take up nitrogen,
perhaps iron or molybdate ions. Their experimental
plants came from two sites that differed in nutrient
richness, and the benefits of the insect enhanced diet
were greatest in the plants from the richer site. This
result does not agree with the general hypothesis that
the most benefits of carnivory are realized by plants
from the poorest sites (Chandler and Anderson, 1976;
Givnish et al., 1984; Sorenson and Jackson, 1968).
Aldenius et al. suggested phenological variation between
the two sites might be a confounding variable, but also
noted that, if a prey input increased root uptake of
minerals, then richer soils would lead to better growth.
Karlsson and Carlsson (1984) also worked with
P. vulgaris, simulating insect capture by applying
blocks of agar containing nitrogen, phosphorous, or
microelements to the leaves. They found that phosporous
blocks induced biomass increases, and concluded that
phosphorous was the most important supplement gained by
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carnivory in the common butterwort. They also reported
that application of nitrogenous agar caused an increase
in root to leaf weight ratios.
For another species of butterwort, Pinguicula
lusitanica L ., Harder and Zemlin (1967c) found increased
leaf development, increased chlorophyll, and more flowers
on plants grown on nitrogen- and phosphorous-deficient
inorganic media that were given Drosophila, egg yolk, or
ammonium phosphate. Untreated plants did not flower.
Harder and Zemlin (1968) also found that Pinus pollen
given to P. lusitanica leaves caused an increase in
number of leaves and diameter of rosettes, as well as
slowing aging, promoting flowering, and deepening the
plants' green color. Nonetheless, these researchers
(1967c) felt that unambiguous proof for enhancement of
plant development by captured prey inputs had only been
given for Drosera and Utricularia,
Chandler and Anderson (1976) similarly experimented
with species of Drosera, growing plants in sand cultures
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deficient in nitrogen, sulfur, phosphorous, or the micro
elements, and feeding with Drosophila. Using D. binata
Labill., they found that optimal growth occurred with
insect supplements and a nitrogen-deficient medium, while
added nitrate inhibited growth. Chandler and Anderson
found increased growth in D. w hittakeri Planch, when
flies were given to plants lacking root access to any
nutrients or to inorganic sources of nitrogen and sulfur,
but not in plants denied phosphorous or microelements.
However, Drosophila supplementation did cause increased
phosphorous tissue concentrations on phosphorous
deficient and complete media.
For another tuberous species of sundew, Drosera
erythrorhiza Lindl. , Pate and Dixon (1978) found
Drosophila to be an effective source of nitrogen and
phosphorous.
Fabian-Galan and Salageanu (1969) observed
translocation of carbohydrates and amino acids from prey
to the plant in Drosera capensis L. , and from prey in
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mature traps to growing points in Aldrovanda vesiculosa
L., but not from young traps.
Christensen (1976) found that Sarracenia flava L.
plants deprived of soil nutrients and prey were fairly
small and showed some chlorosis. Plants grown on poor
soil and fed insects had increased tissue concentrations
of nitrogen and phosphorous, but not of calcium,
magnesium, or potassium. Plants grown with abundant
fertilizer and given insects did not show increased
tissue concentrations relative to plants grown with
fertilizer but not given insects. Christensen
hypothesized that insectivory may interfere with nutrient
uptake when nutrients are abundant.
Hermann, Platt, and von Ende (1987, pers. comm.)
found increases in growth and clone numbers in S. flava
plants that were fed. They did not observe effects
until a year or more had elapsed but suggested the
impacts of withholding prey could be ameliorated by the
underground storage organ this species possesses.
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Plummer and Kethley (1964) observed absorption of
amino acids, peptides, and other nutrients from prey by
leaves of S. fla v a . They decided that the gains from
carnivory may be greater for immature plants than for
adults.
Roberts and Oosting (1958) reported that various
previous experiments on Dionaea muscipula Ellis were
inconclusive, and found that fed plants in their study
showed more vigorous vegetative growth, but that unfed
plants watered with distilled water did better than unfed
plants watered with an inorganic nutrient solution
(excessive concentrations or wrong proportions of
nutrients were suggested as a possible cause.)
Observers of Utricularia, like those of terrestrial
carnivorous plants, have noted more vigorous growth in
plants which capture prey. Skutch (1928) reported that
putting asparagin, albumen or flesh extract into bladders
with pipettes resulted in increased chlorophyll in
bladder antennae and larger bladders (including "giant”
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bladders in U. vulgaris that measured 6.2 mm in length).
Also, several adventitious shoots arose from the leaves
bearing treated utricles. Another consequence of the
artificial feeding was formation of two bladders per
stalk and stimulation of leaf apices to form bladders.
Skutch recorded the results of Busgen's (1888) feeding
trials as well: treated Utricularia vulgaris plants were
longer and developed more leaves than unfed plants, and
in one series of experiments, untreated plants formed
unseasonal turions while fed plants grew well.
One of the classic feeding studies of Utricularia was
undertaken by Pringsheim and Pringsheim (1962). Their
U. exoleta R. Braun plants showed good vegetative growth
in inorganic nutrient solutions but only flowered if
organic compounds were added (peptone and meat extract).
Pringsheim and Pringsheim (1967, in Sorenson and Jackson,
1968) further found peptone and beef extract necessary
for good vegetative growth in U , minor L. and
U, ochroleuca R. Hartman, but could not induce flowering
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with organic additives.
Sorenson and Jackson (1968) experimented with
U , gibba L. in magnesium- and potassium-deficient and
complete media, and fed paramecia to some plants. They
discovered that feeding did not cause a growth increase
in plants in complete media and only caused a small
increase in plants in the magnesium-deficient media.
However, fed plants in their potassium study, in both
complete and incomplete media, did elongate significantly
more than unfed plants. Paramecia treatments also
increased the number of internodes, and allowed formation
of more bladders, but the latter result was confounded
by differing intensities of algal infection which caused
bladder abscission. Sorenson and Jackson’s experiments
supplied live prey, so that utricles were activated in
the study, which was not the case in other research
reviewed here.
Dore Swamy and Mohan Ram (1969, 1971) grew U , inflexa
Forsk. axenically and tried adding beef extract, casein
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hydrolysate, peptone, tryptone, and yeast extract. All
of these organic nitrogen sources enhanced vegetative
growth, but depressed flowering, and yeast extract
completely inhibited flowering. Dore Swamy and Mohan Ram
observed flowering with and without glycine in the
medium, and concluded that animal protein is not
necessary for flowering in U. inflexa. They also found
that beef extract, casein, and tryptone delayed bladder
abscission, while high light (6000 lux) promoted bladder
reddening and abscission. Mohan Ram, Harada, and Nitsch
(1972) confirmed that U . inflexa could use nitrate as
its nitrogen source.
Harder (1963), raising U . exoleta in a mineral
nutrient solution and treating some plants with
autoclaved D a f ^ ia infusions, found that untreated
plants became dormant, while supplemented plants
flowered. Peptone extract also induced flowering. He
inferred that natural carnivory is not a strategy for
nutrient assistance, but rather a source for
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reproductive cycle requirements. Harder and Zemlin
(1967a), however, induced flowering in U. stellaris L.
without animal supplements.
Harder (1970a) tried various proportions of nutrient
solution and autoclaved Daphnia (ranging from 18 to 4500
Daphnia per 100 ml of solution) on five species of
Utricularia, and found increases in dry weights of
U, minor, U, exoleta, and U. ochroleuca when 300 or more
Daphnia were administered. Daphnia decoctions induced
flowering in U, exoleta, but no flowering was observed
for U, vulgaris, U. minor, or U. ochroleuca whether or
not these species were "fed".
Johnson (1987) noted that absorbed nutrients were
rapidly moved to the growing points of Utricularia, and
Coleman, Dollar, and Boyd (1971) found quick movement of
phosphorous from bladders to stems and leaves when
U. inflata Walt, plants were exposed to radioactively
labelled ostracods. They also stated that the
carnivorous absorption pathway is probably more
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important than the foliar absorption route only in
infertile waters.
Harder and Zemlin (1967c) addressed the possibility
of carnivorous plants using carbon from captured prey.
They found that growth in U. stellaris, U. exoleta,
U. minor, U. ochroleuca, and U. vulgaris was
enhanced by saccharose and glucose, and to a lesser
extent by fructose, maltose and cellobiose. Flowers were
more abundant on plants grown in solutions with added
sugar. These effects were seen in plants grown under
light and dark conditions, but were more apparent in the
light-cultured plants. Harder (1970a) determined that
sugar added to nutrient solutions had a greater effect
on growth and flowering in these five species than did
Daphnia decoctions. Harder and Zemlin (1967b) found
growth promotion by sugar in a non-carnivorous submerged
aquatic, Apogoneton distachius, as well. Harder (1970b)
observed dry weight increases in U , minor plants grown
in an inorganic solution and beef extract when sugar and
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acetate were added. The extent of the increase depended
on concentrations of the sugar and acetate: more acetate
allowed lower sucrose concentration for maximum effect
and vice versa. With sucrose supplementation, he also
recorded growth in plants held in darkness, albeit less
growth than in plants exposed to light.
Similarly, Dore Swamy and Mohan Ram (1969) found that
increased sucrose levels in growth media for U . inflexa
resulted in development of lateral branches by release of
apical dominance, and that higher levels of sucrose
induced morphological change: 6% caused bushy plants
with short internodes, and 8% caused bushy, dark green
plants with small pigmented bladders and reduced
flowering. On the other hand, they (1971) recorded poor
growth in plants grown in darkness on a medium including
sucrose and glycine: small leaves, light green
coloration, and elongated stems. Bladders, however,
were no different from those of light-grown plants.
For DroserOf Chandler and Anderson (1976) determined
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by low level light experimentation that insects were not
an important carbon source.
In summary, the literature on plant feeding research
suggests that carnivorous plants derive nitrogen,
phosphorous, sulfur, and some micronutrients from their
prey, but that the actual nutrient of greatest importance
depends on the species and the environment. Further, the
necessity of ancillary resources for completion of the
life cycle also depends on species and habitat. Lastly,
some carnivorous plants can use the carbon skeletons of
prey, but none have been found to survive without
photosynthesis.
II. Constraints and Confounding Factors
In the literature, there has also been some mention
of constraints on the benefits of carnivory, and of
factors confounding the demarcation of such gains.
Moeller (1978) noted that carbon may be limiting for
Utricularia purpurea Walt., which does not use
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bicarbonate. Thus, experiments which do not provide
sufficient inorganic carbon in a form the experimental
species can use may not find the growth increases
expected from insect dietary additions.
Botta (1976) listed species of cyanophytes that
survive indefinitely in bladders of Utricularia obtusa
Sw. , U. platensis Speg., and U , foliosa L ., and
Bosserman (1983) mentioned nitrogen fixation by
periphyton associated with U, purpurea, U. juncea Vahl,
and U. inflata, Wagner and Mshigeni (1986) measured
nitrogen fixation by epiphytes and bladder-dwelling algae
of U . inflexa and suggested the process is intensive
enough to give the association potential as a
biofertilizer. They also raised the possibility of
nitrogen contributions to Utricularia by heterotrophic
bacteria. Consequently, Utricularia and other aquatic or
phytotelm (water-holding) carnivorous plants may have a
third nitrogen source.
Rossbach (1939) reported that Utricularia species of
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northern regions which form tarions usually develop fewer
flowers than their more southerly conspecifics, and only
infrequently produce fruit. Thus turions may be resource
sinks not considered in feeding experiments. Similarly,
Skutch (1928) explained that the turion food supply may
compensate for prey inputs in recently sprouted plants,
another factor to be considered in analysis of feeding
experiment results. Tubers as nutrient sources and sinks
in Drosera erythrorhiza were noted by Pate and Dixon
(1978).
The photosynthetic contribution of trapping organs
(Hegner, 1925; Luttge, 1983) also confounds cost/benefit
analyses.
Lastly, Moeller (1980) discussed the effects of
temperature on growth in U . purpurea, and Maier (1979)
showed effects of light intensity on production in
U, vulgaris. These and other environmental factors
interact with substrate fertility and prey nutrient
inputs to produce observed growth and development of
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plants.
III. Evolutionary Aspects
Some recent papers have explored models for the
evolution of carnivory and attempted to conduct
cost/benefit analyses. Thompson (1981) compared
insectivory with myrmecophily (ant-fed plants),
suggesting both nutrient supplementation strategies
evolved in response to similar ecological conditions.
He noted that insectivory is advantageous in moist,
low nutrient habitats, since such species use water
freely in glandular secretion and absorption processes,
while epiphytic myrmecophily is the workable design for
open canopy forest sites where dryness prohibits
insectivory.
Benzing (1986) agreed that ecological factors limit
carnivorous plants to moist, exposed habitats, where
photosynthesis is not limited and costs for secretory
lures and other trapping implementia are not excessive.
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He further stated that myrmecophily is a less costly
strategy and therefore found in more stressful habitats.
Juniper (1986) postulated that the origins of
carnivory are polyphyletic, and present convergence
represents a limited number of techniques to compensate
for habitat sterility. He noted that elements of the
suite of carnivory characters are found in many other
plants, and only in situations where carnivory would be
advantageous did the entire syndrome evolve.
Bloom et al. (1985) applied economic theory to
calculate how plants should develop in order to maximize
growth in their environments, and pointed out the
necessity of considering nutrients and water as currency
as well as carbon— the usual base for analysis. They
deduced that carnivory occurs when stocks of water and
nutrients are imbalanced. Growth in nutrient-poor
environments must be slow, but plants of these habitats
adjust for their supply levels and are less flexible in
allocation patterns than are plants of more fertile
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habitats. Nutrient-starved plants were also predicted to
show less sequestering of resources for sexual
reproduction.
Givnish et al. (1984) suggested the main advantage of
carnivory is increased photosynthesis via mineral
nutrient supplementation allowing increased
photosynthetic rates and/or increased numbers of
photosynthetic units. They hypothesized that
photosynthetic benefits will level out when factors other
than nutrients become limiting. This supports the rule
of thumb mentioned earlier, that prey capture is most
advantageous in the least rich sites. Givnish et al.
concluded that the greatest benefits of carnivory will
accrue to plants in moist, sunny, low nutrient habitats,
while in dry or shaded low nutrient habitats, benefits
will be less and level out sooner.
Benzing (1987) added that carnivory is rare because
capture and absorption of prey is not an economical way
to supplement nutrient uptake in most environments.
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These "generally unfavorable energetics" also prevent
carnivorous plants from being vast, dominant communities
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Four
Study Sites
Plants and turions were collected in Flathead and
Lake Counties of Montana. Sites varied in size y water
chemistry, plant species composition, and size and vigor
of Utricularia vulgaris populations. Tur ions for
laboratory experimentation were collected from three
sites: East Bay of Flathead Lake, McWenneger's Slough,
and Tykeson Pond; plant materials from these as well as
three other sites were examined: Daphnia Pond, Loon
Lake, and "Tykeson*s Kettle". Locations of these sites
are shown in Figure 3.
East Bay, Flathead Lake, is a shallow, marshy area
with numerous aquatic plant species including Hippurus
vulgaris L . , Myriophyllum spicatum L . , Potamogeton spp. ,
Ranunculus sp., Typha latifolia L . , Utricularia minor,
and U. vulgaris (nomenclature follows Dorn, 1984).
Water in East Bay is clear beyond the depths at which
Utricularia is found. Flathead Lake is classed as
39
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 3; Study Site Locations 40
East Bay, Flathead Lake Lake County T22N R19W Sec. 5-6
McWenneger's Slough Flathead County T28N R20W Sec. 6
Tykeson Pond Lake County T26N R19W Sec. 16 64 Daphnia Pond 4 5 Lake County T26N RI9W Sec. 1
Loon Lake Lake County T26N R19W Sec. 10
"Tykeson's Kettle" Lake County T26N R19W Sec. 15
Flathead Lake
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oligo-mesotrophic (Stanford, Stuart, and Ellis, 1983),
but East Bay itself is probably mesotrophic. (This
classification, and the trophic levels for the other
sites, are based on one-time water chemistry
measurements, including conductivity, pH, alkalinity,and
dissolved oxygen, as well as on water color and
hydrophyte species composition.) The substrate is sandy
in open water, silty under Typha stands. U, vulgaris
is found in several locationswithin East Bay: among the
Typha stands, in open, shallow water near Typha, and
in windrows with other floating macrophytes and detritus.
U. vulgaris is not a dominant plant in this site
(cover 5 to 25%), but is found in small patches.
McWenneger's Slough is a large shallow slough, wooded
on the southeast, open to the north and west, which
contains a luxuriant, diverse plant assemblage. Besides
U . minor and I/, vulgaris, species found in the slough
include: Carex spp. , Ceratophyllum demersum L . , Chara
sp. , Elodea nuttallii (Planch.) St. John, Lemna trisulca
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L., L. turionifera h ., Myriophyllum spicatum, Nuphar
lutea L .f Polygonum amphibium L., Potamogeton spp.,
Sagittaria sp. , Scirpus sp. , Spirodela polyrhiza (L.)
Schleiden, Typha latifolia and Wolf fia columbiana
Karsten.
The summer Secchi disk transparency is 0.5 m, and the
water is meso- to eutrophic. The substrate is coarse
and sandy in some places, a thick silt in others.
U. vulgaris grows at various depths in the Slough, among
the other macrophytes. Plants are large and healthy,
growing in patches, varying from 25 to 50% cover. Of the
sites described, and others in Lake County that were
visited, McWenneger's Slough is the mother lode for
U. vulgaris— plants are by far the largest, longest, and
most vigorous there.
Tykeson Pond is a small, shallow, somewhat dystrophic
pond surrounded by open forest to the south and east and
by logging roads to the north and west. It is dominated
by Menyanthes trifoliata L . , and also hosts Carex spp.,
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Lemna turioniferaf Nuphar lutea, Phalaris arundinacea L. ,
Potamogeton natans L. and other Potamogeton spp.,
Potentilla palustris (L.) Scop., and U, vulgaris. The
water is soft and dark, with a summer Secchi depth of
0.25 m. Sediments are peaty, brown, and coarse.
U. vulgaris grows densely with 50 to 7 5% cover.
Daphnia Pond is also small and shallow, a late
successional pond dominated by the emergent plants
Phalaris arundinacea, Scirpus sp. , and Typha latifolia.
Other macrophytes in Daphnia Pond include: Ceratophyllum
demersum, Lemna turionif era, Myriophyllum spicatum,
Nuphar lutea, Polygonum amphibium, Potamogeton natans L .
and other Potamogeton spp., and U. vulgaris. The summer
Secchi depth in the dark, dystrophic water is 0.18 m.
At the steep bank edges, the substrate is gravelly, but
in most areas there is an organic, mucky bottom.
U. vulgaris seems localized in distribution in
Daphnia Pond, contrary to reports from earlier years
(Sheldon, 1987, pers. commun.). During 1987, U. vulgaris
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occurred in the south end of the pond, among the Typha
stems and in openings in the emergent stands. Plants
grew singly in the cattails and in small groups in the
openings, with cover increasing from 5 to 25% to 25 to
50% respectively.
Loon Lake is a forested lake with an extensive mat
developing on the northwest side. Aquatic vegetation is
characterized by Carex spp. , Chara sp. , Elodea canadensis
Michx. , My r to f^y Hum spicatum, N ajas flexilis (Willd.)
Rost, and Schmidt, Nuphar lutea, Polygonum amphibium,
Potentilla palustris, Scirpus sp. , Typha latifolia and
U, vulgaris (with U. minor as well, on the mat). The
water is clear to the depths at which U . vulgaris is
found, about 0.80 m, and the lake is mesotrophic. The
substrate is silty and marl is present. U. vulgaris
grows singly, and is scattered, with a cover ranking of
1 to 5%.
"Tykeson's Kettle" is an unnamed, more or less
mesotrophic, sunken pond in the forest adjacent to
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Tykeson Pond. Much of its surface is covered thickly by
Lemna turionif e r a other macrophytes present are Chara
s p ,, Myriophyllum spicatum, Nuphar lutea, Phalaris
arundinacea, Potamogeton spp,, and U, vulgaris. Where
there is no Lem na, the summer Secchi depth is 0,5 m. The
sediments are brown, organic and silty. U. vulgaris
grows in small patches, 5 to 25% cover.
All sites described, except McWenneger's Slough, have
been created or affected by glacial activity; the Slough
is an artifact of meanders of the Flathead River (Alt and
Hyndman, 1986). All six areas are subject to ice
formation in the winter, and East Bay is also subject to
major water level fluctuations due to drawdown for
hydroelectric purposes. East Bay further differs from
the other sites in being the only one which is influenced
by water inputs other than ground water and
precipitation, i.e. the others have no inlets or outlets.
Climate is similar for all sites, a "modified North
Pacific Coast type", with one-half the annual
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precipitation falling between May and July (NOAA, 1985).
The mean yearly maximum temperature is 55.8 the
yearly minimum is 33.0 ®F. Precipitation averages
15.36 inches per year, excluding an average 50.8 inches
of snow. Annually, about 71 days are clear, 80 partly
cloudy, and 214 cloudy (Kalispell, Montana averages;
NOAA, 1985).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Five
Methods and Materials
I. Summer Field Collection
I gathered intact plants of Utricularia vulgaris
at six sites (Fig. 3) in late July and August of 1987 y
and transported these samples in local water. Within a
few days of collection, I determined the following for
the sample plants : stem diameter at one representative
point, length of plant or shoot section (some plants were
broken during handling), number of leaves (leaflet pairs)
per plant, and number of primary bladders per leaflet
for at least 13 leaflets on each plant. Although I
counted bladders for the current year's growth of each
plant, with handling, some bladders abscissed.
Consequently, I estimated the potential number of
bladders per leaflet based on a growth pattern discerned
from previous observations. I did not count stem
bladder or secondary bladder positions since their
47
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presence and number are more variable than those of
primary bladders, and since some of them had abscissed
also.
I computed summary statistics for the variables
measured and for indices computed from them: leaflet
pairs per cm of stem, number of bladders (or positions)
per plant (average bladders per leaflet x two leaflets
per leaf x total leaves) and bladders (or positions) per
cm of stem (total bladders/length). I drew boxplots for
these variables for each site, and conducted a
multivariate analysis of variance (MANOVA).
My data were quite variable and may have violated the
MANOVA assumptions, depending how conservatively one
follows the assumption guidelines (Ott, 1984; Patterson,
1988, pers. comm.). Specifically, the variance was not
common to the different sites, and this inequality was
made worse by the small and unequal sample sizes.
Samples were neither random nor independent. The
assumption of normal distribution for each variable in
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the population was probably adequately met. Because the
assumptions of parametric statistics may not have been
met, I also conducted a nonparametric MANOVA.
Because of the small sample sizes, I could not test
for differences among populations for all variables
measured (Patterson, 1988, pers. comm.). I chose four
variables that would roughly indicate development of PCT
versus other tissues and allow testing of my allocation
hypothesis: bladders per leaflet, per plant and per cm,
and leaflet pairs per cm. In my laboratory experiments,
I used slightly different, more appropriate, indicator
variables for PCT development, but I did not have the
necessary data to compute these indicators in the Summer
Field Collection.
I assumed that dissolved inorganic nutrient levels
regulate allocation of resources to PCT, while
carnivorous nutrient inputs control the amount of growth
within the allocation framework. Plants receiving many
nutrients via the carnivorous pathway might be expected
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to decrease allocation of resources to PCT if overall
(i.e. foliar and carnivorous) nutrient uptake was the
regulating mechanism. However, the limited lifespan of
traps suggests this is not the case. Bladders, for
example, darken and decay with use, and Dionaea leaves
may only catch three insects before becoming inactive
(Slack, 1979).
A plant stimulated by low substrate nutrient
availability to produce many traps will continue to be
stimulated to allocate resources to PCT. The quantity of
nutrients reaped by these traps will affect the amount of
growth the plant can complete, and whether or not it can
flower. Thus, carnivorous plants growing in nutrient-
poor sites which have large populations of potential prey
should exhibit high levels of PCT, but also be large
sized.
For Utricularia vulgaris, and other aquatic
carnivorous plants, sites with few dissolved nutrients
usually also have low prey populations. Accordingly,
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based on my allocation hypothesis, I expected the
greatest numbers of bladders per cm to occur in plants
collected from the poorest sites. I expected bladders
per plant to be highest in the largest plants, i.e. those
from the richest site. Numbers of leaflet pairs per cm
and bladders per leaflet should be intermediate for
plants from both nutrient-poor and nutrient-rich sites,
since good growth increases numbers of leaflet pairs and
bladders per leaflet, while allocation to PCT does so as
well.
Anomalous sites with low dissolved nutrient levels
but high potential prey populations should produce
U. vulgaris plants with fairly high values for all four
indices. I did not examine prey availability at any of
the sites.
II. Common Garden Experiment
In October, 1987, I collected Utricularia vulgaris
turions from East Bay and McWenneger's Slough, by
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severing the persistent stem of the summer growth 2-3 cm
from the turion base. I selected plants haphazardly and
took one (occasionally two) turions per plant. I
transported the turions in local water in an insulated
container.
I later rinsed the turions in tap water to remove
algae, detritus, and mucilage, and placed them into
aquaria filled with distilled water (separated by
site). I put the aquaria into a controlled temperature
room (CTR) where turions were exposed to about 450 ft-c
fluorescent and incandescent light (General Electric
light meter type 214) on a 14 hour photoperiod, with the
temperature set for 30 ®C. The dormancy-breaking
procedure closely followed Winston and Gorham (1979a).
I added distilled water as needed to keep turions
submerged. Turions from McWenneger's Slough developed
rampant algal coverings which I removed by rubbing and
rinsing.
Due to temperature control difficulties, in one week
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I moved the turions to a growth chamber (Percival model
PT80) set for the same conditions, but with 850 ft-c of
fluorescent and incandescent light.
When sufficient turions had begun to sprout
(approximately 2 weeks after collection), I began the
Common Garden Experiment. I placed nine sprouted turions
from East Bay and nine from McWenneger's Slough into two
trays (33.5 x 26 x 8.5 cm), one for each site, filled
with nutrient solution.
I used Pringsheim and Pringsheim's (1962) nutrient
solution throughout the growing period for this
experiment, diluted to one-half strength to slow algal
growth (Knight, 1987, pers. comm.). The nutrient
solution formula by weight, in distilled water, is:
KNOg 0.02% (NHi,)2 HP0 4 0.002% MgSOji.yHgO 0.001% CaSOij (saturated) 2 ml/100 Minor element solution 1 ml/100
Minor element solution :
EDTA 0.0 2%
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 54
FeSOfi.VHgO 0.07% ZnS0i|.7H20 0.001% MnSOi|. IHgO 0.0002% CuSOi|. 5 H2O 0.0000005 H^BOii 0.001% Co(NOg)2.6H200.0001% Na2MoOi|. 2 H 2O 0.0001%.
Pringsheim and Pringsheim's solution uses MnSOij.4 H2 O;
I substituted MnSOi|. IH2 O because it was more readily
available. Approximate pH values for the nutrient
solution were: 6.4 for one-half strength, (Orion
Research Digital Ionalyzer/501), 6.3 for full strength,
and 6.7 for one-tenth strength (the latter strengths were
used in the Diet Experiment, described below).
The turion trays were kept in the CTR, set at 24
during the day, and 18 °C at night, under about 400 ft-c
of fluorescent and incandescent light. I later
transferred the plants to larger containers (53 x 23 x
14 cm).
I started a replicate of this experiment one week
later, when more turions had sprouted, using 14 East Bay
plants and 14 McWenneger's Slough plants in 46 x 24 x 15
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 55
cm aquaria. Complete experimental design is shown in
Figure 4.
Nutrient solution was added whenever needed to
maintain levels, and nutrient solution was changed
completely once in the Replicate 1 aquaria to control
algal growth. No prey were offered to plants in this
experiment.
After four weeks of growth I took a series of
measurements on each plant: stem diameter at one
representative point on each branch longer than 3 cm, or
at three points along the stem if single-stemmed; length
for each segment ; number of leaflet pairs per segment;
length of ten healthy, mature leaflets; number of
bladders on 15 representative leaflets. Bladders tended
to fall off with algal infection and handling, so I
estimated the potential number of primary bladders per
leaflet as before, again not counting stem or secondary
bladder positions. I recorded wet (blotted) weights for
each plant (Ohaus model B300 electronic digital scale)
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 56 Figure 4. Experimental Design for the Common Garden Experiment
SI te of Turion CollectIon East Bay McWenneger's Slough
ReplI cate ExperIment ran from 10/28-11/27 9 turions 9 turions
Replicate Exper iment ran from 11/4-12/5 14 turions 14 turions
Modified Pringsheim and Pringsheim (1962) nutrient solution at one-half strength and no prey for all treatments.
All treatments were exposed to summer conditions: 24 °C day, 18 °C night, 14 hour photoperiod, approximately 400 ft-c for four weeks.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 57
and noted presence or absence of new turions.
I measured each plant in a large white enamel tray,
with the plants in shallow water. Measurement precision
was lowered by the water and by the flimsy nature of the
plants. I dried all plants for about 24 hours at 35 ®C
and recorded dry weights.
I repeated the measurement process for plants in the
second replicate experiment after they had grown for four
weeks.
For both replicate experiments, I calculated summary
statistics by site. I used the variables of the Summer
Field Collection, plus leaflet length and bladders per
gm (total bladders/dry weight). I also drew boxplots
for the variables, for each site and each replicate. I
analyzed the effects of independent variables (collection
site, replicate experiment) on dependent variables
(morphological measurements) using MANOVA, in order to
consider overall variation simultaneously (Patterson,
1988, pers. comm.).
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Since my data base was small, I again chose four key
variables for the MANOVA: leaflet length, leaflet pairs
per cm, and bladders per cm and per gm. If developing
turions respond primarily to ambient conditions,
there should be no significant differences between sites
for these variables. However, if developing plants
retain site-specific traits, it would be more difficult
to test PCT predictions. Based on my allocation
hypothesis, I expected plants grown from turions from
sites thought to be low in nutrients to have higher
numbers of bladders per cm and per gm (standardized
measures of PCT development). I also expected these
plants to have intermediate numbers of leaflet pairs per
cm (leaflet pairs per cm can indicate good growth and/or
PCT development) and shorter leaflets (leaflet length
indicates general good growth). I expected turions from
sites thought to be rich in nutrients to grow into
plants with long leaflets, intermediate numbers of
leaflet pairs per cm, and few bladders per cm and per gm.
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Where the MANOVA showed an interaction between
variables, I drew profile plots, using the means for each
site and replicate experiment for the variables in
question. Once again, my data and sample sizes were not
in strict adherence to MANOVA assumptions, so I conducted
nonparametric tests.
I used the categorical variable, presence or absence
of new turions, to further verify the plants' capacity
for response to ambient conditions. Presence or absence
of new turions would also indicate the suitability of the
growing conditions.
For this variable, I computed a chi-square test for
replicate experiments separately and in combination. I
also calculated Cramer's V (same as phi for the combined
replicates), and the contingency coefficient. When the
four treatments were considered separately, 25% of the
cells had expected frequencies less than five, which
makes chi-square accuracy, and that of the related
statistics, borderline. Accordingly, I computed lambda.
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a statistic based on proportional reduction of error
(NoruSis r 1986).
III. Diet Experiment
In late October, 1987, I collected Utricularia
vulgaris turions from Tykeson Pond and McWenneger’s
Slough. Some turions were allowed to germinate in
distilled water under high temperatures, following
methods outlined in the Common Garden Experiment. I
rinsed the rest and refrigerated them at 1-3 ®C.
After two weeks, many McWenneger's turions had
sprouted and I began the Diet Experiment. Germination
of Tykeson turions was sporadic and unsuccessful, so
these turions were not used. I placed nine McWenneger's
sprouts in each of six treatment trays (33.5 x 26 x 8.5
cm), dividing turions so that each tray had plants of
approximately equal size and developmental stage.
These trays were Treatments 1-6 in the experimental
design shown in Figure 5. Trays contained about 5 1 of
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 5- Experimental Design for the Diet Experiment, Showing Treatment Number
Nutrient Solution Strength
ful1 one-tenth
1 15 3 18
7 21 10 24
5 14 6 17 10 9 20 12 23
2 13 4 16 100 8 19 11 22
fall winter fall winter
Experimental Season
All trays were exposed to summer conditions: 24°C day, 18°C night, 14 hour photoperiod, approximately 850 ft-c for four weeks, in a variety of growth chambers and controlled temperature rooms.
Treatments 5~12 started one week after Treatments 1-4.
Treatments 13~24 started eleven weeks after Treatments 1-4.
Treatments 13-18 were for turions collected from Tykeson Pond, all other treatments were for turions collected from McWenneger's Slough
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 62
one-tenth or full strength concentrations of the nutrient
solution used in the Common Garden Experiment (Pringsheim
and Pringsheim, 1962). One week later, I started
replicate treatments, again using nine McWenneger's
Slough turions per tray. These trays were Treatments
7- 12.
I added nutrient solution when needed to Trays 1-12
and fed them weekly, by adding the appropriate number of
D a f ^ ia to each tray and briefly stirring the tray
contents. Plants slated to receive no prey were treated
identically to plants offered prey.
Treatment Trays 1-6 had to be moved several times
from one growth facility to another. In addition, my
initial feeding method (modelled after Sorenson and
Jackson, 1968) proved to be too stressful for the fragile
plants. Hence, I did not use data from plants in these
trays in statistical tests.
Algal growth in all 12 trays was problematic, and
when necessary, I changed nutrient solutions entirely
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and scrubbed trays. I also removed algae by floating
paper towelling on the solution surface and discarding
the towelling with adhered algae. The solution of some
trays was filtered with a fish aquarium system.
After four weeks of growth, I terminated the
experiment and took measurements as I did in the Common
Garden Experiment. I did not weigh individual plants,
but recorded wet and dry weights for each treatment
tray. I divided the tray dry weights by the number of
experimental plants per tray to estimate individual plant
weight, and divided that weight into the tray mean for
bladders per plant to calculate bladders per gm.
Nine weeks after Treatments 7-12 were begun, I
removed McWenneger's Slough and Tykeson Pond turions
from refrigeration and put them under high temperatures
to break dormancy. After about one week, I used these
sprouts for Treatments 13-24 in Figure 5. I used eight
Tykeson sprouts per tray for Treatments 13-18, and nine
McWenneger's sprouts per tray for Treatments 19-24.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 64
After four weeks, I ended this portion of the experiment
and recorded data as described in the Common Garden
Experiment. I again computed summary statistics and
drew boxplots for the variables.
I used Trays 7-12 and 19-24 to compare differences
in season of turion germination as well as nutrient
solution and feeding treatment differences. I used Trays
13-24 to compare the effects of collection site and
treatments, in effect, a second, more complex common
garden experiment. The four key variables I chose for
MANOVA were the same as for the Common Garden Experiment,
except that I did not use bladders per gm since I did not
have weights for individual plants. Instead I used
bladders per leaflet. I ran MANOVA on Treatment Trays
7-24 for the four variables, and on the ranks for these
variables, and plotted profiles using tray means where
interactions between factors were indicated.
For the trays acting as a common garden experiment, I
expected variables to have high, medium, or low values
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as described in the Common Garden Experiment section.
For trays used to examine effects of nutrient
solution strength and feeding regime, I expected turions
grown in one-tenth strength solution to develop as plants
from nutrient-poor sites: short leaflets, intermediate
numbers of bladders per leaflet and leaflet pairs per cm,
and many bladders per cm. For turions grown in full
strength nutrient solution, I expected long leaflets,
intermediate numbers of bladders per leaflet and leaflet
pairs per cm, and few bladders per cm. I expected
increasing prey availability to increase growth, but not
to change the relative extent of the indicator variables,
as explained in the Summer Field Collection section.
I calculated the chi-square statistic for presence or
absence of new turions in Trays 1-24, as well as
Cramer's V and the contingency coefficient. Since a
majority of the cells had expected frequencies less than
five, I also calculated the lambda statistic.
In this study, I intended for light, temperature and
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photoperiod conditions to be identical for all treatment
trays. However, due to the necessity of using several
controlled environment facilities, conditions were not
identical, and the extent of fluctuation varied also.
The variation between treatment conditions confounds
interpretation of experimental results.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Six
Results
I. Summer Field Collection
Utricularia vulgaris plants from the six sites had
definite differences in growth forms and size. The
effect of site of origin was significant at p = 0.000 for
each of the four key variables: bladders (or positions)
per leaflet, leaflet pairs per cm, bladders per plant,
and bladders per cm. Summarized results of the
multivariate analysis of variance, and of the MANOVA on
the ranked variables, are shown in Table 1.
Summary statistics for the variables measured are
shown in Tables A1-A3, and boxplots for each variable.
Figures A1-A7, show the extent of variation between and
within sites (Appendix A) . The complete MANOVA results
are also given in Appendix A, in Tables A4 and A5.
II. Growth under Laboratory Conditions
67
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The Utricularia vulgaris plants did not grow
particularly well under laboratory conditions. Treatment
trays and aquaria were susceptible to algal infections
(mostly Chlamydomonas and Euglena types), which seemed
to increase fragmentation in the plants. Some trays
developed unidentified fungal and/or rotifer or small
invertebrate scums. Growth periods longer than four
weeks were desirable, but plant health was prohibitive.
Growth forms varied from long, delicate, hair-like
leaves to stunted, flatter, thicker, coarser leaves along
the same stem. Shoots grew from the tips of turion
leaves as well as in one to several directions from the
old stem axis. Plant color varied from light to dark
green, with McWenneger's Slough plants darker than those
from Tykeson Pond and East Bay. McWenneger's plants
were also more massive and more mucilaginous than plants
from the other sites.
Bladders originated on all plants but did not persist
and did not develop to maturity in many cases. The
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degree of algal infection seemed positively correlated to
bladder abscission, as Sorenson and Jackson (1968) noted.
Leaves that had been part of the turions extended and
were clearly distinguishable from newly developed leaves.
These old turion leaves did possess some bladders. The
old turion end of many plants darkened and decayed into
fragile, slimy pieces.
Germination of collected turions was quicker in
January than in November. Plants from all three
collecting locations formed new turions during the
experimental period.
III. Common Garden Experiment
When grown under common conditions, the Utricularia
vulgaris plants from East Bay and McWenneger's Slough
showed differences in morphology similar to those
observed in mature plants collected from the field. For
some of the variables measured, differences seemed to be
increased or reduced relative to differences observed in
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field-collected plants, but this was not tested
statistically.
Summarized results from parametric and nonparametric
MANOVA on the four key variables (leaflet length,
bladders per cm, leaflet pairs per cm and bladders per
gm) are shown in Table 2. There were no significant
differences between the two replicate experiments for
any of the variables, while the effect of collection site
was significant for all variables in parametric and
nonparametrie tests except bladders per cm. In addition,
a significant interaction occurred between the site and
replicate factors, which confounds the magnitude of the
site effect. The interaction was only important (i.e.
intersecting rather than off-parallel profiles) for the
bladders per cm variable. The profile plot for this
interaction is shown in Figure B12 (Appendix B).
According to the ranked MANOVA, the interaction was also
fairly important for the variables leaves per cm and
bladders per gm (0.05 < p < 0.10).
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 71
For the one categorical variable measured, presence
or absence of new tarions, there was no relationship
between site of origin and development of tarions when
replicates for each site were analyzed together. When
the four sets of plants were examined separately, the
chi-square value was significant, and the other
statistics of association supported this. Statistics
for this variable are given in Table 3.
Summary statistics for variables for each site are
given in Tables B1-B4 (Appendix B). Boxplots for these
variables. Figures Bl-Bll (Appendix B), display the
extent of variation between sites and between replicates
of the experiment. Tables B5-B10 in Appendix B show the
complete MANOVA results.
The correlation between wet and dry final weights for
each plant was 0.89.
IV. Diet Experiment
In the Diet Experiment, the greatest differences in
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measurements were due to plants originating from
different sites. Like the mature plants collected from
the field, plants grown from turions from Tykeson
Pond were smaller in leaflet length, and had fewer
bladders per leaflet, but had more leaflet pairs and
bladders per cm than the McWenneger's Slough turions
(bladders per cm was not significantly different in field
plants; Tables C1-C3, Figs. C2, C3, C6, C8, Appendix
C). Again the degree of difference between sites seemed
to change for lab-grown plants relative to field-
collected plants.
The site effect was statistically significant, as
summarized parametric and nonparametrie MANOVA results
show for Trays 13-18 (Tykeson Pond) and Trays 19-24
(McWenneger's Slough; Table 4). An interaction
occurred between feeding regime and collection site for
these 12 treatment trays, but was only important for the
bladders per cm variable in the full strength solution
treatment. The profile plot for this interaction is
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shown in Figure C9 (Appendix C). For the other three
key variables, all the interaction profiles showed only
slight departures from parallel, as may be seen on the
boxplots (Figs. C2, C3, and C6, Appendix C).
For plants grown from turions collected only from
McWenneger's Slough, bladders per leaflet and per cm
differed significantly between experimental seasons.
Feeding regime and nutrient solution strength did not
have statistically significant effects on any of the
variables used as indicators of growth and PCT
development. Summarized results of parametric and
nonparametric MANOVA for Treatment Trays 7-12 and 19-24
are shown in Table 5.
The experimental season factor interacted with the
feeding regime factor, and with the solution strength
factor. These interactions were important, as the
profile plots show (Figs. CIO and Cll, Appendix C).
Solution strength and feeding regime also interacted
(Fig. C12), more so in the winter than in the fall
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experimental period.
For the categorical variable, presence or absence of
new turions, there was a significant correlation with
season of turion germination. New turions were rarely
formed in the fall, but were common in the winter
treatments. Counts, chi-square and related statistics,
and the lambda value are given in Tables 6 and 7.
Summary statistics for all 24 trays are given in
Tables C1-C4, and boxplots for each variable are shown
in Figures C1-C8 (Appendix C). Final weights per tray
are shown in Table C5; wet and dry weights correlated
well : r = 0.98.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 75 Table 1. Significance of Collection Site on Morphological Characters in U. vulgaris Plants Collected in Summer*
Significance of Univariate F Statistic Parametr i c (Nonpa rametric)
Variable
Bladders Leaflet Bladders Bladders per pairs per per leaflet per cm plant cm
Factor
Collection 0.000 0.000 0.000 0.000 Site (0.000) (0.000) (O.OOO) (O.OOO)
^Summary of Tables A4 and A5, Appendix A
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 76 Table 2. Significance of Collection Site, Replicate Experiment, and the Interaction between These, on Morphological Characters in IJ. vulgaris Plants Raised from Turions in a Common Garden Experiment*
Significance of Univariate F Statistic Parametric (Nonpa rametric)
Variable
Leaflet Leaflet Bladders Bladders length pairs per per in cm per cm cm gm
Factor
Collection 0.033 0.001 0.637 0.011 Site (0.023) (O.OOO) (0.557) (0.020)
Replicate Not significant in multivariate analysis Experiment
Interact ion
Site- 0.813 0.229 0.000 0.238 Replicate (0.761) (0.085) (0.000) (0.066)
'Summary of Tables B5"B10, Appendix B
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 77 Table 3. Influence of Collection Site and Replicate Experiment on Formation of New Turions by vu 1 gar i s Plants Raised from Turions in a Common Garden Experiment
Count Expected Value Column %
Pond New Turions Replicate Absent Present
East Bay 1 n 0 6.5 4.5 100% 0%
East Bay 2 2 13 8.8 6.2 13.3% 86.7%
McWenneger ' s 1 11 0 6.5 4.5 100% 0%
McWenneger's 2 6 8 8.2 5 . 8 42.9% 57.1%
For replicate experiments separately/together:
Chi-square 26.69/1.70
Degrees of freedom 3/ 1
Significance level 0.00/0.19
Cramer's V (Phi) 0.76/0.18
Contingency coefficient 0.61/0.18 Lambda 0.62/0.00
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 78 Table 4. Significance of Collection Site, Nutrient Solution Strength, Feeding Regime, and Interactions between these Factors, on Morphological Characters in |J. vulgar i s Plants Raised from Turions in a Diet Experiment*
Significance of Univariate F Statistic Parametri c (Nonparametric)
Variable
Leaflet Bladders Leaflet Bladders length per pairs per in cm leaflet per cm cm
Factor
Collection 0.000 0.000 0.000 0.000 Site (0.000) (0.000) (O.OOO) (O.OOO)
Feeding O.OBO 0.370 0.002 0.040 Regime (0.177) (0.124) (0.049) (0.096)
Solution Not significant in multivariate analysis Strength
Interact ion
Site- 0-375 0.067 0.031 0.011 Feeding (0.501) (0.l84) (0.459) (0.026)
Site- Not significant in multivariate analysis Solution
Feeding significant in multivariate analysis Solution
S i te- Solution- Not significant in multivariate analysis Feed i ng
*Based on Trays 13-24; Summary of Tables C6-C19, Appendix C
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 79 Table S- Significance of Experimental Season, Nutrient Solution Strength, Feeding Regime, and Interactions between these Factors, on Morphological Characters in jj. vulgaris Plants Raised from Turions in a Diet Experiment*
Significance of Univariate F Statistic Parametric (Nonparametr i c)
Variable
Leaflet Bladders Leaf let Bladders 1ength per pa i rs per i n cm leaflet per cm cm
Factor
Expérimenta 1 0.163 0.000 0.847 0.022 Season (0.135) (0.000) (0.823) (0.051)
Feed Î ng Not significant in multivariate analys i s Reg ime
Solution 0.111 0.055 0.198 0.994 Strength (Not significant in multivariate analysis)
Interaction
Season- 0.842 0.001 0.625 0.626 Feed i ng (0.892) (0.000) (0.538) (0.619)
Season- 0.720 0.338 0.001 0.030 Solution (0.707) (0.275) (0.000) (0.003)
Feed i ng- 0.553 0.031 0.266 0-387 Solution (Not significant in multivariate analysis,
Season- Solution- Not significant in multivariate analysts Feed i ng
*Based on Trays 7-12 and 19-24; Summary of Tables C20-C33, Appendix C
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 7) "OCD O Q. c s g Table 6. Influence of Nutrient Solution and Feeding Treatments on Formation of New Turions g vulgaris Plants Raised from Turions in a Diet Experiment T3 CD
WC/) 3o' 0 Count Tray** 5 Expected Value* CD Column % 1 2 3 4 8 5 6 New turions (§■3" 1 3 absent 13 9 5 2 CD 9 13 7.8 5.4 4.8 4.2 5.4 7.8 100.0% 100.0% 62.5% 28.6% 100.0% 100.0% 3. 3* CD present 0 0 3 5 0 0 "OCD 5.2 O 3.6 3.2 2.8 3.6 5.2 Q. 0.0% 0.0% 37.5% 71.4% 0.0% 0.0% O 3 ■D O 7 8 9 10 11 12 CD Q. absent 10 13 12 9 9 9 6.0 7.8 7.2 5.4 5.4 5.4 ■D 100.0% CD 100.0% 100.0% 100.0% 100.0% 100.0% 3 (/) present 0 0 0 0 0 0 o' 4.0 5.2 4.8 3.6 3.6 3.6 0.0% 0.0% 0.0% 0.0% 0.0% 0.0%
*Based on no association between new turion formation and treatment tray **These treatments were conducted during the fall experimental season; complete treatment details are given in Fig. 5- 0 0 o :x) CD ■o O Q.C 8 a Table 6. Influence of Nutrient Solution and Feeding Treatments on Formation of New Turions by tJ. vulgaris Plants Raised from Turions in a Diet Experiment (cont'd.) % (/) § Count Tray** O Expected Value* CD Column % 13 14 15 16 17 18 8
ë' New turions O 3 absent 0 0 0 0 0 1 CD 2.4 4.2 3.6 6,0 4,2 4.8 C 0.0% 0.0% 0.0% 0.0% 0.0% 12.5% p. 3" CD present 4 7 6 10 7 CD 7 ■o 1.6 2.8 2.4 4.0 2.8 3.2 O Q.C 100.0% 100.0% 100.0% 100.0% 100.0% 87.5% a o 3 ■a o 19 20 21 22 23 24
CD Q. absent 6 2 7 7 1 1 4.8 6.6 6.6 7.2 7.2 7.2 75.0% 18.2% 63.6% 58.3% 8.3% 8.3% ■o CD present 2 9 4 5 11 11 C/) (/) 3.2 4.4 4.4 4.8 4.8 4.8 25.0% 81.8% 36.4% 41.7% 91.7% 91.7% *Based on no association between new turion formation and treatment tray **These treatments were conducted during the winter experimental season; complete treatment details are g iven in Fig. 5•
00 82 Table 7* Statistics Showing the Association between Formation of New Turions and Treatments for vulgaris Plants Raised from Turions in a Diet Experiment*
Chi-square 167-99
Degrees of freedom 23
Significance level 0-00
Cramer's V 0.86
Contingency coefficient 0.65
Lambda 0.77
*Based on all Trays
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Chapter Seven
Discussion
I. Summer Field Collection
As Tables A1-A3 and Figures A1-A7 (Appendix A) show,
individual variation in plant measurements within sites
was often large; indeed, morphological variation along
individual plants was extensive in all three experiments.
The samples for these measurements were not random or
large, accounting for much of the non-normal distribution
patterns. However, the multiple analysis of variance and
the MANOVA on ranks (Table 1) both show that the pond in
which Utricularia vulgaris plants grew had a
significant effect on their size and development, and on
PCT allocation based on four indices.
However, PCT indices (leaflet length, bladders per
leaflet, leaflet pairs per cm, bladders per cm) were not
significantly different between all sites; sites whose
plants exhibit similar morphologies probably possess
83
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similar nutrient availabilities. For example, plants
collected from East Bay did not differ significantly from
McWenneger's Slough plants for bladders per leaflet,
bladders per cm, or leaflet pairs per cm. Nutrients are
probably almost as available in the Bay as they are in
the Slough. Bladders per plant did differ between the
two sites— no doubt a reflection of the greater length of
the Slough plants.
The significant site effect revealed by the MANOVA
is, of course, expected: better conditions induce more
growth than do poor conditions. Not having extensive
water quality data, I cannot definitively rank the six
sites by trophic status. One-time water chemistry
measurements and vascular hydrophyte species composition
(Schuyler, 1987, pers. comm.) suggest that McWenneger's
Slough is by far the richest site. As expected,
McWenneger's plants were the largest and most vigorous,
exhibiting the highest values for bladders per leaflet
and bladders per plant. Having a high number of bladders
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per leaflet can lead to a high number of bladders per
plant, but plants which increase their number of
leaflets per unit length, or dissection of leaflets, also
increase total bladder number.
McWenneger's plants did not have the greatest values
for the other two variables used as indicators of growth
and PCT allocation— leaflet pairs per cm, and bladders
per cm. The higher values for these variables were found
in plants from sites thought to be more nutrient-poor,
based on water chemistry and hydrophyte diversity
(Schuyler, 1987, pers. comm.; Wetzel, 1983). The higher
values indicate a shunt of resources towards PCT, since
increased bladders per cm values are not as linked to
good growth as bladders per plant values are, as
explained in Materials and Methods. Increased leaflet
pairs per cm could also indicate an allocation of
resources towards PCT, but that shunt is confounded by
the other roles of leaflets.
Leaflet pairs per cm and bladders per cm were highest
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at sites thought to be least fertile— Tykeson Pond and
"Tykeson's Kettle". Tykeson Pond is dystrophic, and
nutrients may well be unavailable. "Tykeson's Kettle"
seemed to be mesotrophic, but it may be that the abundant
Lemna plants take up most of the available nutrients
as well as much of the incident light.
PCT levels can probably be adjusted in different
ways, by adding leaflet pairs, by adding bladders per
leaflet, or both. Pond chemistry and light availability
may influence which of these strategies plants will
follow, in accordance with allowable strategies dictated
by genotype.
Since high PCT levels can be achieved in several
ways, I cannot differentiate nutrient-rich sites from
nutrient-poor ones based on my limited data. The rich
site plants may not be allocating any extra resources to
PCT, but their growth pattern forms bladders as it forms
leaves. The poor site plants meanwhile may allocate
resources to PCT, sacrificing general growth, resulting
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in small-sized plants with many leaflet pairs per cm and
bladders per leaflet. Increasing the number of leaflets
increases PCT as well as increasing foliar absorption and
photosynthetic capabilities, so the plants may be relying
on more than one strategy for survival. I would expect
that rich site plants which automatically have much PCT
do not depend on carnivorous inputs to the extent that
plants do which specifically allocate resources to PCT
development. To verify this expectation, it would be
necessary to discern whether a given site's plants used
the foliar or carnivorous pathway as the main nutrient
input route, perhaps through the use of labelled
nutrients.
Sources of error in the summer data are many, and
interpretation of the meager results cannot be extensive.
Measurements depended on which part of the plant was
measured, whether in a young area with unextended
internodes, a mature area, or a senescing portion which
had already lost bladders and leaves.
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Moreover, the ponds did not differ solely in nutrient
availabilities, but were of different sizes and depths,
so that temperatures, lengths of ice-free seasons, etc.
varied greatly. For example, Tykeson Pond and East Bay
turions were mature earlier in the fall than were those
of McWenneger's Slough. Certainly plant developmental
stage also varied during the summer. Developmental stage
and length of growing season probably have an effect on
bladder number : the "leaves" of U, vulgaris may
actually be branching systems (Arber, 1920; Sculthorpe,
1967), and conditions which encourage continued growth
may increase the number of bladders per leaflet.
Nonetheless, trapping capacity and PCT allocation
vary significantly between sites. These could be genetic
differences, with each pond's population an ecotype,
where characteristic genes dominate because of natural
selection, where gene combinations have been fixed by
genetic drift/founder effect, or where each population
is clonal. On the other hand, the differences between
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sites could be entirely environmental. That would mean a
high degree of plasticity, not unusual for aquatic plants
(Sculthorpe, 1967), but unusual for plants adapted for
life in resource-poor environments (Bloom et al., 1985).
Thirdly, and most likely, the differences could be due
to an interaction of genetic and environmental factors.
The summer data show that there is a site effect on
resource allocation in C/. vulgaris, but do not reveal
the underlying cause.
II. Common Garden Experiment
A common garden experiment allows one to distinguish
between effects of current environment and effects of
genotypes or past environments. Within a common garden,
the Utricularia vulgaris plants grown from turions
from two sites did not grow to similar size, but
retained some of their apparently site-specific traits.
For example, the McWenneger's plants were larger in the
turion stage, and finished the experiment with
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significantly longer leaflets than the East Bay plants.
Bladders per gm and leaflet pairs per cm were also
significantly different between sites in the Common
Garden Experiment. I do not know if the variable
bladders per gm was significantly different among mature
plants collected from East Bay and McWenneger's Slough,
because I have no weight data for the Summer Field
Collection.
Surprisingly, leaflet pairs per cm was not
significantly different between those two sites for
plants collected in the field. Perhaps with more time,
the differences seen in the lab would have disappeared,
as the East Bay plants compensated for their initial
smaller size as turions. On the other hand, differences
between East Bay and McWenneger's Slough plants might
increase with time. If development could be followed
longer, an increasing or decreasing trend in differences
might suggest the relative importance of "parental"
environmental conditions versus current environmental
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conditions in controlling morphological variables like
PCT. The observation of a tendency toward change in the
extent of morphological differences between plants grown
from tarions from the two sites suggests plants may
indeed respond to current conditions.
The significant effect of site (MANOVA, Table 2) on
morphology in these plants suggests that natural
variation is not solely due to ambient environmental
factors. Differences could be genetic, although the
extent of variation within treatment cells and that
between replicate experiments for plants grown from
turions from the same site, seems excessive if each
population is ecotypic. Or, differences could have been
pre-set in the tarions by the "generation" that formed
the tarions. Growing these plants through another
"generation" would discriminate between these
alternatives, since genetic differences would persist
while any "parental" environment effect should disappear
since the new tarions would have been formed under the
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common garden conditions. I did not examine the newly
formed tur ions to any extent, so cannot say if any
differences between Tykeson Pond and McWenneger's Slough
turions seemed likely to persist into the next
"generation".
Turion formation is^ in itself, a resource sink.
Variations in the tendency to produce turions, or the
rapidity of formation, could affect other growth in
U. vulgaris. Tur ion formation is induced by cold,
drought, nutrient or light stresses (Maier, 1973;
Winston and Gorham, 1979a); probably by light or
crowding effects in this case.
As the chi-square and related statistics and the
lambda value show (Table 3), turion formation was not
correlated with collection site, therefore that response
was probably largely controlled by current environmental
conditions in the common garden. As the results of the
separate replicate analysis show, however, turion
formation was correlated with replicate experiment.
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No plants formed new turions in the first replicate. The
difference may be due to more crowded conditions in the
second replicate (more plants, smaller aquaria), or to
other differences in growing conditions.
The plants in the secondreplicate may have been
more likely to produce turions and go dormant because
they had recently been in the dormant stage. Moreover,
these plants grew from turions which left the dormant
stage less readily than the plants in Replicate 1 (the
quick sprouters). Winston and Gorham (1979a) found that
Utricularia vulgaris turions collected in Alberta,
Canada, when the "parental" plants were senescing, were
in a state of innate dormancy, from which turions
induced to sprout soon reformed turions. The
McWenneger's turions were in this stage when I collected
them; the East Bay and Tykeson turions had probably
passed into the imposed dormancy stage since their
"parental" stems were dead. Turions induced to sprout
from imposed dormancy did not reform turions in Winston
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and Gorham's study. Their growing conditions were
probably more favorable than mine. Winston and Gorham
(1979b) found dormancy to be hormonally controlled.
U. vulgaris plants observed in the field at Nimrod
Warm Spring (Montana, Granite County, T12N R15W Sec. 14)
had not formed turions in November, 1987, while plants at
Tykeson Pond, East Bay, and McWenneger's Slough had done
so. The Nimrod plants also did not form turions under
conditions of cold, low light, and low nutrients (tap
water), although they eventually died, but under the same
conditions East Bay, Tykeson, and McWenneger's plants did
form turions. Thus, the species does not seem to have an
endogenous rhythm that dictates turion formation, but
rather responds to various environmental cues in a manner
controlled by its physiological state. Of course, the
Nimrod plants could be a warm spring ecotype, where the
endogenous rhythm has a different setpoint.
In any event, if turions are formed by plants of
differing size and vigor, as was the case for the
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McWenneger's and East Bay plants, those turions will
start the next "generation" with a built-in size
difference. During my experimental period, and perhaps
during the relatively short growing season of many
northwestern Montana bodies of water, this initial
handicap may never be overcome. For example, if a
"parent" plant grows in poor conditions and produces a
tiny turion, that turion, even if placed in optimal
conditions, may never grow as vigorously as another
turion formed under better conditions but placed into a
suboptimal habitat. Several "generations" of turion
formation under good conditions, by plants originating
from different sites, are probably necessary before the
effect of vigorous "parent" plants could definitely be
ruled out.
A lag effect could also be due to preconditioning.
The environment that influences the "parent" plant's
health and indirectly affects the turion also directly
affects the early development of the turion. Bud and
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seed formation are thought to be developmental stages
that are particularly sensitive to preconditioning
(Rowe, 1964). During turion formation and maturation,
environmental conditions may induce changes that affect
later gene expression. Conditions during turion
maturation may also cause changes in growth factor
proportions in the turion, as Gutterman, Thomas, and
Heydecker (1975) found for Lactuca scariola seeds.
Accordingly, even if plants from different sites had
identical genotypes, preconditioning of turions could
cause observable differences in morphology. The turions
I collected were in different stages of maturation at
different sites, as mentioned earlier, so preconditioning
effects would certainly be possible in my experiments.
Furthermore, Utricularia vulgaris turions are not
sexually produced propagules, but rather a perennating
extension of the "parent" plant. Consequently, any
acclimation a U . vulgaris plant may have undergone
could be retained in the overwintering plant to be
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expressed when dormancy is broken.
Therefore, morphological differences that I observed
between plants from different sites in the Common Garden
Experiment have several, not mutually exclusive,
explanations :
1. Genetic differences, i.e. ecotypes at each site;
2. Differences in "parental" plant vigor and
ability to endow turions ("parental" environment
indirectly affects turion);
3. Differences in the turions' early environments
("parental" environment directly affects turion);
4. Differences in environmental conditions of
"parent" plants, causing acclimation that was not
lost during dieback to the turion phases.
My data do not allow me to determine which explanation,
or what combination of explanations, is correct.
Nonetheless, these data can be examined for agreement
with my hypothesis that plants from poorer waters will
develop more PCT.
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Plants grown from turions from East Bay had more
leaflet pairs per cm and bladders per gm than did plants
grown from turions from McWenneger's Slough. This
suggests that the lab-grown McWenneger's plants do not
need to invest as much of their carbon supply in PCT.
In the lab experiment, they may have had more nutrients
than the East Bay plants to begin with, because of their
larger turions, and they may have received more nutrients
by foliar absorption through their longer leaflets. In
the field, McWenneger's plants probably get more
nutrients by foliar absorption from the water than they
get from prey. The McWenneger's plants might have
benefitted from developing more PCT under the common
garden conditions (assuming nutrients were less available
in the nutrient solution than in the field), but may not
have done so due to preconditioning and acclimation
effects. Again, following development for several
"generations" in the common garden environment would help
to clarify what is occurring.
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Analyses of representative turions for starch and
nutrient levels, and tissue analyses of experimental
plants would allow comparison of resource allocation
efficiency and success.
The interaction effect between replicate experiment
and collection site for bladders per cm has a number of
possible explanations. The replicates were started one
week apart, had different numbers of plants (9 or 14) in
slightly different-sized aquaria, and had minimally
different growth regimes. A crowding effect,
differences in degree of algal infection, or other
factors could have caused the opposite response levels
of bladders per cm in the second replicate. That other
variables measured were not also affected is perplexing,
but the nonparametric MANOVA (Table 2) does indicate
some interaction (0.05 < p < 0.10) between site and
replicate for the variables leaflet pairs per cm and
bladders per gm. Neither MANOVA shows an interaction for
leaflet length; some morphological characters may be more
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responsive than others to conditions during development
from turion to plant.
The Common Garden Experiment ignored the effect of
prey on Utricularia vulgaris, in order to remove as many
confounding factors as possible. In the field, plants
from one site may respond differently to prey
than plants from another site, especially if such plants
differ in their allocation to PCT, as discussed in
Materials and Methods. Accordingly, my results need to
be field checked, and further experimentation would also
enhance the tentative conclusions that I have drawn.
(Part of the Diet Experiment, however, acts as a common
garden experiment with prey.)
Without information on nutrient saturation levels for
Utricularia vulgaris growth, I cannot say whether the
nutrient solution used is a good, mediocre, or sub-
optimal medium. Hence it is not clear if the plants
were under sufficient nutrient stress to maximize their
production of PCT.
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III. Diet Experiment
In the Diet Experiment, plants did not respond to
their treatment conditions in a statistically significant
manner. Observed tendencies suggest plants may respond
minimally to ambient conditions; with longer
experimental periods, this response might increase. That
there were interactions between treatment factors was not
surprising, most organisms respond to a suite of factors
in their environments so that they optimize survival,
growth, development and reproduction. The interactions
make the results harder to interpret; nonetheless,
several trends are apparent.
Plants grown from Tykeson turions had shorter
leaflets than those grown from McWenneger's turions,
when grown under common garden conditions. (I noted
that field-collected Tykeson plants appeared to have
shorter leaflets also, but I have no data to support this
observation.) Leaflet length is related to turion size,
since many leaflets are developed to primordial stages
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when the turion is formed. Having fewer nutrient/energy
reserves in their small turions could also have affected
leaflet length in lab-grown Tykeson plants, since leaves
seem to be indeterminate systems, as mentioned earlier.
The number of bladders (or positions) per leaflet was
less for plants grown from Tykeson Pond turions than for
those from McWenneger's turions grown under the same
conditions, as it was for field-collected plants from the
two sites. Few bladders per leaflet correlates with
having shorter leaflets. However, the difference between
plants from the two sites was not so dramatic for this
variable as for some of the others. This suggests the
lab-grown Tykeson plants may compensate for shorter
leaflets by increasing the degree of dissection of
leaflets, which increases bladder number because primary
bladders generally occur near the points of bifurcation.
Further, field-collected and lab-grown Tykeson
plants had more leaflet pairs per cm than field and lab
McWenneger's plants. This can be another developmental
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Strategy to increase trap number, provided that leaflets
are highly dissected, as I noted above. However, the
increase in leaflet number also increases foliar
absorption potential (and the need to do this concurs
with the probable nutrient-deficient state of the
organism), and enhances photosynthetic capabilities.
The stimulus for increasing leaflet number is not
discernible from my experiment.
The higher bladders per cm values for lab-grown
Tykeson plants relative to lab-grown McWenneger's plants
suggest that the increase in leaflet pairs per cm was
indeed due to the necessity of increasing nutrient
inputs from carnivory. Perhaps the plants from large
turions made at a rich site (McWenneger's) do not need
the nutrient inputs from carnivory, while the plants from
small turions made at a poor site (Tykeson) do. In the
Summer Field Collection, bladders per cm was not
significantly different between the two sites; with time
the lab difference might have disappeared.
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These between site differences seen in the lab
experiments agree with my hypothesis of increased
allocation of resources to PCT in less fertile sites.
Interestingly, Tykeson Pond plants increased the number
of bladders per cm with increasing feeding levels, while
McWenneger's Slough plants did not. This response agrees
with my assumption (see Materials and Methods) that
dissolved nutrient levels regulate PCT allocation, while
prey inputs affect growth. This increase also shows a
response by the plants to ambient conditions.
The data from the Diet Experiment also seem to agree
with my conclusions from the Common Garden Experiment,
that past history of the individual plant and/or the
early environment of the turion play a role in "progeny"
development. Treatments 13-24 acted as a common garden
experiment between plants from more disparate sites.
Again the common conditions did not induce similar growth
or PCT development in turions from different sites, and
significant differences for morphological measurements
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seen in field-collected plants were also found under lab
conditions. Differences in morphology were greater for
this common garden experiment, comparing Tykeson Pond
with McWenneger’s Slough, than for the comparison of East
Bay with McWenneger's. Since East Bay seems to be more
similar to McWenneger's Slough in terms of nutrient
availabilies, this result was expected. As I stated in
the Common Garden Experiment section, continued
experimentation, perhaps including genotype studies (e.g.
electrophoresis), would be necessary to determine the
cause of morphological differences between if* vulgaris
plants from different sites.
Looking at plants grown from turions only from
McWenneger's Slough, there was a significant effect of
experimental season on the variables bladders per leaflet
and bladders per cm. Generally there were more bladders
per leaflet in plants grown during the winter. This may
be related to depletion of starch and other reserves in
the turions by the plants while they remained dormant
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(Maier, 1973; Winston and Gorham, 1979a.) Thus the
developing plants would have a greater need to supplement
nitrogen, phosphorous and other minerals by carnivory
than plants grown in the fall from less depleted buds.
In nature, by spring, turion reserves would be very low,
and developing plants in nutrient-poor environments
would probably have to develop more PCT more quickly
than would plants developing from turions held only
until winter and raised under laboratory conditions.
The bladders per cm response also may be related to
turion reserve depletion, but the interaction for this
variable between experimental season and nutrient
solution strength makes explanation difficult.
Nutrient solution strength and feeding regime did
not, by themselves, have significant effects on any of
the morphological measurements analyzed by MANOVA.
Bladders per leaflet, leaflet pairs per cm, and bladders
per cm were affected by interactions of these factor
levels with each other and with experimental season, so
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that main effects cannot be determined. Obviously, the
PCT stimulation process is complicated. An effect of
nutrient solution strength or feeding regime might have
shown if development had been followed for a longer
time, as past "generation" influences decreased.
Unlike the other three key variables, leaflet length
for lab-grown plants did not vary significantly with
season, prey or nutrient level. This morphological trait
seems to be tied to the turion, and all turions from the
same site were approximately the same size.
The development of new turions occurred more often
in the winter treatment trays than in the fall ones, and
more often on Tykeson Pond plants than on McWenneger's
Slough ones. This may be explained by the turion
reserve drop over time, so that new plants in the winter
experimental trays were more sensitive to growing
conditions. Tykeson Pond turions were smaller than
McWenneger's turions, so reserve depletion may have been
more serious for them.
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New turion formation was not greatly correlated with
feeding level or nutrient solution strength, although
turion formation did seem to be more common in trays
providing few dissolved nutrients. It may be that none
of the experimental conditions were suitable for good
growth in the plants, but rather were stressful,
inducing reversion to the dormant phase. New turion
formation could also be tied to hormone levels in the
old turion which increase the probability of turion
formation (Winston and Gorham, 1979b) and act to prevent
plant growth during fleeting warm periods in winter.
Using the tray weights to compute bladders per gm
gives data which support my hypothesis: Tykeson plants
had many more bladders per gm than did McWenneger's
plants.
Overall, I think the variable bladders per gm is the
best indicator for development of PCT stimulated by the
need to increase nutrient uptake. High bladders per
leaflet values can be due to good growth, without
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reflecting an increasing reliance on carnivory. Any
increase in trapping capacity due to increased leaflet
pairs per cm is confounded with leaflet roles in
photosynthesis and nutrient absorption. Bladders per
cm values do standardize trapping potential, but do not
show any shunting of resources from general growth to
PCT development. But high bladders per gm values mark
the plant that allocates resources to PCT.
Unfortunately, I have no weight data for the Summer
Field Collection of Utricularia vulgaris, and the
weight data for the Diet Experiment is not for
individual plants. Thus I must mainly interpret
resource allocation in these two experiments using the
less appropriate indices of leaflet pairs per cm,
bladders per leaflet, and bladders per cm.
I did not collect data on the occurrence of stem and
secondary bladders. Yet plants from nutrient deficient
habitats may boost PCT through these traps instead of,
or in addition to, increasing numbers of primary
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bladders. Of course, increasing the number of leaflet
pairs and the degree of leaf dissection, which boosts
primary bladder numbers, would also increase numbers of
stem and secondary bladders.
With regard to the constraints mentioned in the
Literature Review, my experimental conditions probably
were not restrictive. Temperatures were equal to summer
temperatures in Utricularia vulgaris habitats. Light
availability in the lab was at least as great as in the
darker-watered study sites. Carbon dioxide should have
been sufficiently available, since the trays were open
to diffusion, and the nutrient solution was buffered.
Although U. purpurea was found to be unable to use
bicarbonate (Moeller, 1978), U. vulgaris probably
can. Moeller states that the aquatic vascular plants
dependent on free carbon dioxide are usually rosette
plants, which U. vulgaris is not. It is more
similar to Elodea and Ceratophyllum r which do use
bicarbonate (Moeller, 1978).
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However, sources of error in this study were many.
One such source is the problem of not being able to
insure that Dapfmia were trapped and digested
rather than dying and adding to the nutrient solution.
My initial feeding method would have taken care of this
problem, but would also have killed the experimental
plants. The loss of bladders and incomplete maturation
of bladders, due to algal growth, handling, and other
laboratory effects, decreases the likelihood that all or
even most of the Daptmia were captured. Bladder
abscission may also be related to nutrient
availabilities: Bloom et al. (1985) noted that leaves
senesce when the carbon budget becomes negative.
Bladders are modified leaves, and the growing conditions
may not have provided a positive budget.
Elimination of variation in temperature and light
regimes would be advantageous, as would be better control
of algal growth. Larger growing containers and a flow
through nutrient solution system would probably allow
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the experimental period to continue for longer than four
weeks (Knight, 1987, pers. comm). Larger sample sizes
would improve statistical inference. Data defining
nutrient and prey availabilities for each site would
eliminate doubt from my assumptions regarding trophic
status and the mechanism regulating resource allocation
to PCT. Most importantly, related field work is
necessary to validate the conclusions I have drawn.
Nevertheless, overall my data support my research
hypothesis, that Utricularia vulgaris plants grown to
maturity in waters of lower nutrient availabilities
allocate more of their carbon resources to PCT
development than do U . vulgaris plants in richer
waters (based on water chemistry and hydrophyte
diversity). My data also suggest that ambient conditions
in which turions develop into plants exert little control
over bladder production, at least during initial weeks of
growth. Rather, the plants' genotypes and/or past
environments (of past growing phases as well as turions'
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early history) seem to exert more control over bladder
production.
Carnivory, as an adaptation, allows plants to
colonize areas they otherwise could not. It is
reasonable that, along with the evolution of carnivory,
came mechanisms to regulate the extent of its use.
Carnivory apparently does not allow plants to adapt to
new conditions very rapidly, which may contribute to the
relative rarity of the carnivorous habit.
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Summary and Conclusions
The following hypothesis was stated for the carnivorous plant
Utricularia vulgaris; plants growing in sites relatively low in
dissolved, inorganic nutrients will exhibit less vigorous growth but
will allocate more resources to prey capture tissue (PCT) than plants
growing in nutrient"rich sites.
Based on collections of mature plants from a series of sites
thought to possess a range of nutrient levels, plants from poorer sites
do exiiibit less vigorous growth, while producing more bladders per cm
of stem.
Based on allowing turions from these sites to develop for four
weeks under common garden conditions, plants from poorer sites do
exhibit less vigorous growth while producing more bladders per cm of
stem and per gram of plant.
The common garden experiments also showed that allocation of
resources to PCT appeared to be controlled largely by genotype and/or
the environment under which the turion was formed (the field environment)
rather than the environment in which the turion developed into a plant
(the lab environment). Periods of development longer than four weeks
might have reduced (or increased) differences in plants from
different sites. Following plants over several vegetative “generations"
in a common garden would be required to determine if genotype or a
lagged environment effect has greatest control over PCT development.
As a consequence of the genetic/lagged environment effect,
experiments exposing the developing turions to different prey levels
114
Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. and inorganic nutrient levels for four weeks showed little effect of
these treatments on PCT development/allocation. If PCT allocation
is not solely genetically controlled, but requires several turion
"generations" before plants respond to lab conditions, these treatments
may eventually be found to affect PCT allocation.
115
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Appendix A. Data Summaries, Complete MANOVA Results, and Boxplots for
Utricularia vulgaris Plants Collected in Summer
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Table Al. Data Sufitfiiar i es For Uiriculacia uulsaris Plants Collected in S u ni mer: Stem Diameter Plant Lenathf and Leaflet Pairs per Plant b y Collection Site
Collection Site Stem Total Leaflet Diameter L e n 31 h Pairs i n mm in cm per Plant-
T ■/ K e s o n
Mean 1 .0 3 9. 18 1 9 . 33 Std. Deviation .0 5 2 .3 1 5 .2 8 Mini mum 1 .0 0 5 .6 0 12 Maximum 1 . 10 1 1 .6 0 27 Samp le Size 6 6 6
Daphn i a
Mean . 38 1 8 .9 8 31 . 75 Std. Deviation .0 5 6. 62 6. 85 Mini m u (ti . 90 1 0. 00 22 Maxi mum 1 .0 0 2 5 .2 0 37 Sample Size 4 4 4
Loon
Mean 1 . 67 10. 03 14 . 67 Std. Deviation . 32 2 . 72 8 ,3 3 M i n i m u r« 1 . 30 7. 50 8 Max imum 1 .9 0 1 2 .9 0 24 Samp le Size 3 3 3
East Bax
Mean 1 .1 7 1 9 .2 3 20. 75 Std. Deviation . 42 3 .3 6 3 . 3 0 Mini m um . 90 15. 40 17 Maximum 1 .8 0 2 3 .6 0 25 Samp le Size 4 4 4
Kettle
Mean .9 2 2 5 .8 2 GO . 60 Std. Deviation . 1 3 5. 60 8 .0 8 Minimum . 70 1 B . 30 50 Max i mum 1 . 00 3 3 . 70 72 Samp le Size 5 5 5
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Tab 1e A l . Data Summaries for Uicicularia uulsaris Plants Collected in Summer : Stem Diameter, Plant Lenathf and Leaflet Pairs per Plant b y Collection Site (coni'd.)
Collection Site Stem T o t a l L e a f 1 e t D ia m e t e r L e n g th P a ir s
i n film in cm Per P la n t
McNenneser's
Mean 2 . 43 40. 70 4 0 .3 3 Std. Deviation .5 2 4 .8 7 7 .3 1 M inim um 1 . SO 3 4 .2 0 38 Maximum 3 . 0 0 47. 50 GO Sampie Size 6 B G
TOTAL
Mean 1 .3 9 21 . 83 34 . 39 Std. Deviation .66 12 . 33 1 8 .1 4 M i n i fii u m . 70 5. GO 8 Maximum 3. 00 4 7 .5 0 72 Sample Size 28 28 28
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Table A2 Data Sufluiiaries for U lnicularia iiulaanxs Plants Collected in Summer: Leaflet Pairs per cmr Bladders/Positions per Leaflet» and Bladders/Positions per Plant by Collection Site
Collection Site L e a f l e t Bladders/ T o t a l Pa i r s P o s i t i o n s B l a d d e r s / p e r cm per Leaflet P o s i t i o n s p e r P la n t
T y K e s o n
Mean 2. 1 1 8 . SB 3 3 0 .7 8 Std. Deviation . 2 G 2 .0 5 6 9 .5 2 M in i mum 1 . 72 6 21G Maximum 2 . 42 12 40G Sample Size 6 G G
D a p h n i a
Mean 1 . 76 7. 79 5 0 1 .6 1 Std. Deviation . 32 .67 1 4 1 .3 2 M i n i m u fii 1 . 47 7 300 Maximum 2 . 2 0 8 600 Sample Size 4 4 4
Loon
Mean 1 . 39 1 0 .7 3 3 1 2 .5 5 Std. Deviation . 42 1 . 23 1 6 6 .3 8 M in i mum 1 .0 7 10 158 M axi mum 1 .86 12 489 Samp le S iz e 3 3 3
E a s t B ay
Mean 1 .0 8 13 20 5 4 9 .0 5 Std. Deviation .0 3 2 0 1 3 2 .4 5 M i n i r>i u m 1 . 05 1 1 426 Maximum 1 . 1 2 IB 674 Samp le S iz e 4 4 4
Kettle
Mean 2 . 38 10 67 1 3 0 1 .5 4 Std. Deviation . 23 1 13 2 8 3 .3 8 992 M in i mum 2 . 1 4 10 Maximum 2 . 73 12 1763 Sample Size 5 5 5
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Table A2. Data Sufiiftiar i e s For U tric u la ria uulaaris Plants Collected in Summe rI Leaflet Pairs per CM» Bladders/Positions per Leaflet» and Bladders/Positions Per Plant by Collection Site ( c on t ' d . )
Collection Site Leaflet Bladders/ T o t a l P a i r s P o s i t i o n s B l a d d e r s / p e r CM per Leaflet P o s i t i o n s p e r P la n t
McWenneaer ' s
Mean 1 .1 8 1 5 .2 4 1 4 8 0 .8 3 Std. Deviation . OS 1 . 16 2 8 4 .1 4 M i n i M u M 1.11 13 1017 Max i MUM 1 .2 6 16 1936 Sample Size 6 6 G
TOTAL
Mean 1 . 69 1 1 . 22 8 0 4 .2 0 Std. Deviation .5 4 3 .0 2 5 3 1 .4 6 M i n i M u M 1 .0 5 6 158 Max imum 2 . 7 3 16 1936 Samp le Size 28 28 28
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Table A3. Data Summiar i es for Utricularia uulaacis Plants Collected in Summer I Bladders/Positions per cm b r Collection Site
Collection Site Bladders/ Positions per cm
T X K e s o n
Mean 3 G .79 Std. Deviation G .59 Minimum 29.63 Maximum 48.42 Samp le Size G
Daphn i a
Mean 27.07 Std. Deviation 2.5G M i n i fri u m 23.81 Maximum 30.00 Sample Size 4
Loon
Mean 29.G5 Std. Deviation 8.42 Minimum 21.05 Maximum 37.87 Sample Size 3
East Bay
Mean 28.G5 Std. Deviation 5.56 Minimum 22.31 Maximum 35.82 Samp le Size 4
Kettle
Mean 50.64 Std. Deviation 4.22 Mini mum 43.38 Maximufii 54.22 Sample Size 5
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Table A3. Data S u ni ni a r i es for Utriculacia uulsaris Plants Collected in Summer I Bladders/Positions Per cm by Collection Site (cont'd.)
Collection Site Bladders/ Positions Per cm
McWenneaer's
Mean 3G.1G Std. Deviation 3.98 Minimum 29.72 Maximum 40.7G Sample Size 6
TOTAL
Mean 35.81 Std. Deviation 9.33 Mini mum 21.05 Maximum 54.22 Sample Size 28
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Table A5. Nonparametric MANOVA Showina the Effect of Collection Site on Four Variables (Bladders/Positions per Leaflet» Bladders/Positions per cn Plant» Bladders/Positions per cm» Leaflet Pairs per cm) Measured cn on Utri cularia vulaaris Plants Collected in Summer o'3
M ultivariate Tests of Sianif icance (S = 4»M = 0 »N = 8 1/2) 8
c5' Test Name Value Approx, F H/poth. OF Error DF Sis. of F
Pi 1lais 2.27353 5.79454 2 0 . 0 0 8 8 . 0 0 . 0 0 0 Hotel 1 inas 12.84090 11.23573 2 0 . 0 0 70.00 . 0 0 0 WilKs .00988 9.67839 2 0 . 0 0 83.37 . 0 0 0 3- 3" Ro vs .87018 CD
CD ■o O cO. Uni varia te F -test s with (5»22) D. F. g. o 3 ■o Variab1e Hypoth, SS Error SS Hvpoth. MS Error MS F Sia. of F o Bladders 8. per leaflet 1444.30000 382.70000 288.88000 17.39545 18.80549 . 0 0 0 O C Bladders ■O per cm 1240.70000 586.30000 248.14000 28.65000 3,31107 CD . 0 0 0
en en Bladders o' 3 PL': Plant 1531.71887 295.28333 308.34333 13.42197 22.82402 . 0 0 0
Leaf 1e t pairs per cm 1501.83333 325.36887 300.32667 14.78939 20.30889 VjO . 0 0 0 W 134
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K » ÜJ X) I '4 + <-> M c m ro 01 '—( + — — + -H Q- I I 3 + t + 3 U I I Oi + — — + "0 CL 4 tn q + + •H ■i-t I I 3 ns * + — Q y OL I I 4 + + q 4 Of 3 4 Ü. + — + I I q m o Of + * + X I \ Ü. + — + tn 4-> o t- Of 0. 5 o Of X 5 in t- Ot o 3 c c Of Qj cn M I •H k JC (- on LU T3 X 01 tJ jJ ■1-4 Of ITf 3 X en 't E C'J IO LÜ Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 138 c + —« I + TJ I Hi + 11 o *» 11 — 1 cn + + I I a c + * u o I I + + UI ♦J o c #—4 + + 1—1 I I CL O + w U I I ui + + •H X W JO m m S + + H o I I 3 + + 3 Î- I I OI + + 0. « 4 w t- + — — + H • w I I 3 ro + + + Cl 3 CL I I •H + U 1* 3 U. + + f- ro I I o u + + C k I I + + Ul • • o {. ,— f (U Q. 1 3 X 5 1 i n o {- 01 o 3 1 C f x _J 01 c CO cn 1 ro ' x •i-t 01 n > • 1 • n - JZ «— « t- in 11 UI 1 ID 01 4-> *-> o x r 1 1 > i n - w 3 X — « cn in OJ 1 r OJ XO ÜJ Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 139 OJ U) + c I -w C w + LU O I TJ + 4J U U ai a* + + I I o o LU * + LU U O I I + + u> >* *-> -a c ru t» + — + - I C I I O. m + * + lU I I u1 Q- + — 4- -4 (-1 î- tl 01 en B. 4- — — 4- I I 3 4- ^ 4- 3 I 1 4- — — 4- m H <-> tl nj u» 4- H o I 3 Q. + + X Q tl ^ I I H W 4- 4- H !- H 01 3 X) + f- m I o <—< 4- tt- 00 I 4- 1/1 o f- —t 01 OL c 1 3 X S 1 in o u 01 o 3 1 c 01 5 00 en 1 tc Of to 1 -•-» - SL u r~ lU n LU1 T3 01 m 4- > cn 1 OI (A •—1 3 X W lO 1 TZ N X O LU tn 1 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 140 c 0> + — + *> t I + + — X "OUI I I 01 + — + c u o Oi *—4 4-> + + o I I o 0/ » + LU u F—y I 1 + + VI o «-> U c ro X + x> I I Û. + * + LU 5 I I o + H W u tn a> 010. + + H I I 3 1/1 + + 3 c I I o + + m ■ w H H w m v> + — + ( o I I 3 Q. * + U I I H tn + — + W L- w 3 n "O TJ + — + Î- m I I o I—1 + » + r*.CÜ 1 i + — + 1/1 4-> • ■ 1 o u # ■ 4 m a. s X 3 X s in o L- Of o 3 c 0/ E V. 01 m 01 nj w 1. -C (. in OJ C'I o 4 J N LU ~ o X 01 *> X l i t in X h- r Cl % o tu CO Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 141 Appendix B. Data Summaries, Complete MANOVA Results, Boxplots, and Profile Plots for Utricularia vulgaris Plants Raised from Jurions in a Common Garden Experiment Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1 4 2 T a b le B i . Data Summar i es For U lricuiaria uulsarls Plants Raised From Jurions in a Common Garden Experiment I Stem Diameter» LeaFIet Length and Plant Length b y Collection Site and Replicate Site Stem L e a F I e t T o t a l Re p 1 i c a t e D i ame t e r Length Length i n mm in cm in cm E a s t Bay 1 Mean .71 1 .96 30.62 Std. Deviation . 13 .45 13.12 M in i mum .50 1 . 45 7. 60 Ma X i mum .88 2. 78 51 . 20 Samp le Size 1 1 1 1 1 1 E a s t Bay Mean 56 1 .71 27.36 Std. Deviation 19 . 43 15.13 M in i mum 13 1.11 7.20 Maximum 83 2. 46 60. 80 Sample Size 14 12 14 M c W e n n e 3 e r ' s 1 Mean . 98 2 .25 27. 30 Std. Deviation . 17 .46 7. 72 M in i III u m . 70 1 .53 13.30 Max i mum 1 .27 3.10 39.90 Sample Size 10 10 10 M c W e n n e 3 e r ' s Mean . 86 1 .94 28.52 Std. Deviation . 09 . 27 12. 43 Minimum . 70 1 . 58 7.70 M axi mum 1 . 00 2.64 51.10 Sample Size 13 13 13 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1 4 3 T a b 1 e B 1 . Data Summaries for Utricularia uulaaris Plants Raised From Jurions in a Common Garden Experiment: Stem Diameter, Leaflet Length and Plant Length by Collection Site and Replicate (cont'd.> Site Stem Leaflet Total Re p1icate Diameter Length Length i n mm in cm in cm TOTAL Mean 7G 1 . 95 28.41 Std. Deviation . 43 12.35 Minimum 13 1.11 7.20 Maximum 27 3,10 GO. 80 Samp le Size 48 4G 48 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1 44 Table B2 Data Summaries for U tricularia uulsacis Plants Raised from Turions in a Common Garden ExperimentI Leaflet Pairs per Plant, Leaflet Pairs per cm, and Bladders/Positions per Leaflet by Collection Site and Rcr-licate Site L e a f l e t L e a f 1e t B la d d e r s / R e p 1 i c a t e P a ir s Pa i r s P o s i t i o n s p e r P l a n t p e r cm per Leaf let East Bay 1 Mean 1 1 0 .7 3 3 .5 6 10. 16 S t d . D e v i a t i on 5 3 .8 4 .4 7 1 . 15 Mini mum 24 2 . 9 9 0 Maximum 219 4 .3 1 12 Sample Size 1 1 1 1 11 E a s t Bay Mean 94 . 07 3. 74 7 .9 6 Std. Deviation 4 2 .5 8 . 86 1 .31 M inim um 39 2 . 95 6 M axi mu m 181 5. 42 10 Sample Size 14 14 14 McWenneaer's 1 Mean 71 . 90 2 .6 5 1 1 .0 5 Std. Deviation 1 8 .8 5 . 33 1 .56 M in im u fii 31 2 .2 5 8 13 Max i mum 97 3 .1 7 Sample Size 10 10 10 McWenneser's 1 1 .35 Mean 81 . 23 3 . 09 1 .62 Std. Deviation 2 7 . 2 2 . 79 8 M in i mum 28 2 . 1 6 Maximum 134 4 . 83 14 13 Samp le Size 13 13 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 145 Tab 1e B2. Data Summar i e s For U tricularia uulsacis Plants Raised From Turions in a Common Garden Experiment : LeaFIet Pairs per Plant, LeaFIet Pairs per cm» and Bladders/Positions per Leaflet by Collection Site and Reelicate ( c o n t ' d . ) Site LeaFIet L e a F I e t B l a d d e r s / Re p1 i c a t e Pa i r s Pa i r s P o s i t i o n s p e r P l a n t per cm Per LeaFIet TOTAL Mean 8 9 . 7 9 3 .3 0 1 0 .0 3 Std. Deviation 3 9 .6 5 . 78 1 .97 M in i mum 24 2 .1 6 6 M axi mum 219 5 .4 2 14 Sample Size 48 48 48 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 1 46 Table B3 Data SuAimar i e s For U lricu la ria uulsacis Plants Raised From Jurions in a Common Garden Experiment: Bladders/Positions per Plant» Bladders/Positions per cm» and Bladders/ Positions per sm fay Collection Site and Rep 1i ca t e Site T o t a l Bladders/ B 1 a d d e r s / R e a l i c a t e B l a d d e r s / P o s i t i o n s P o s i t i o n s P o s i t i o n s p e r c m p e r am P e r P la n t Dr Y We i ah t E a s t B ay 1 Mean 2 2 2 0 .3 0 71 . 79 1 0 7 6 3 .8 5 Std. Deviation 1 0 3 6 .4 7 7. 59 7 0 6 3 .2 7 M i n i m u m 570 5 7 . 73 5 1 7 8 . IS M a X i m u m 4000 8 4. 47 2 8 7 9 0 .6 7 Samp le Size 1 1 1 1 1 1 E a s t B ay Mean 1 5 3 2 .4 3 5 9 .0 5 1 4 0 3 2 .7 3 Std. Deviation 7 7 3 .3 5 1 4 .8 4 1 5 4 0 4 .8 2 M in i mum 437 36 . 92 4 2 0 0 .0 0 Maximum 2 872 8 8 .9 0 6 2 4 0 0 .0 0 Sample Size 14 14 14 McWenneaer's 1 Mean 1 6 1 4 .5 6 5 8 . 2 5 5 2 3 4 .6 7 Std. Deviation 5 5 0 .1 7 8 .6 1 1 4 1 5 .4 1 M i n i m u m 587 4 4. 13 3 0 0 5 .8 0 Max i mum 2535 68 . 05 6 8 5 2 .4 1 Samp le Size 10 10 10 McWenneaer ^ s Mean 1 8 9 4 .0 9 68 . 32 7 2 3 9 .2 4 Std. Deviation 8 2 6 .7 7 9 . 2 8 1 7 5 6 .1 5 M in i m um 605 5 2 .4 0 4 4 7 1 .8 8 Maximum 3627 81 .2 0 1 0 9 9 9 .4 2 Sample Size 13 13 13 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 147 Table B3. Data Buiiiniaries For Utcxcularia uulaacia P l a n t s R a is e d f r o m J u r i o n s in a Common G arden Experiment : Bladders/Positions per Plant, Bladders/Positions per cm, and Bladders/ Positions per am by Collection Site and Replicate (cont'd.) Site T o t a l B l a d d e r s / Bladders/ Re p 1 i c a t e B l a d d e r s / P o s i t i o n s P o s i t i o n s P o s i t i o n s per cm p e r am p e r P l a n t D ry W e is h t TOTAL Mean 1 8 0 5 .1 3 6 4 .3 1 9 6 1 0 ,7 8 Std. Deviation 8 3 5 .9 9 1 1 . 96 9 4 4 4 .0 6 M i n i m u rti 437 3 6 .9 2 3 0 0 5 .8 0 M axi mum 4000 88 . 90 6 2 4 0 0 ,0 0 Sample Size 48 48 48 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 148 Table B4. Data Summaries for Ulcicularla uulsacis P l a n t s R a is e d f r o m T u r io n s in a Common G arden Experiment I Final Wet and Dr x Weishts bx Collection Site and Reelicate S i t e Blotted Final Dr x R e a l i c a t e W e is h t W e is h t i n sm i n sm E a s t Bax 1 Mean 2 .6 4 4 253 Std. Deviation 1 .6 1 0 138 M in i mum . 39 .0 8 Maximum 6. 13 . 46 Sample Size 1 1 1 1 E a s t B ax Mean 1 .6 1 0 193 Std. Deviation 1 . 199 140 M i n i fii u m , 16 .01 Maximum 4 .2 0 .41 Samp le S iz e 15 15 McWenneser's 1 Mean 3 .8 3 7 334 Std. Deviation 2. 155 165 M inim um .8 0 . 10 Maximum 7 .3 7 .6 9 Samp le S iz e 1 1 1 1 McWenneser' s Mean 2 . 8 9 5 .3 0 3 S td . Dev i at i on 2. 100 . 182 M in i mum 1.11 .0 9 M a X i fii u m 8 . 6 6 . 82 Sample Size 14 14 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 149 Table B4. Data Summaries for U tricularia uulaaris P l a n t s R a is e d From T u r i o n s in a Common G arden Experiment % Final Wet and Dry Weishts by Collection Site and Replicate (cont'd.) S i t e Blotted F i n a l D ry Rep 1 i c a t e W e is h t W e ig h t in sm in sm TOTAL Mean 2 . GGG .266 Std. Deviation 1 .9 1 0 . 162 M inim um . 16 . 01 Maximum S . GG . 82 Samp le Size 51 51 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 73 ■oCD I Table B5. Parametric MANOVA Showina the Effect of Collection Site on Four I Variables (Leaflet Lenathf Leaflet Pairs per cm, Bladders/Positions per cm. Bladders/Positions per am) Measured on U. vulaaris Plants Raised from Jurions in a Common Garden Experiment (/)t/) 3o' Multivariate Tests of Significance (S = 1. M = 1 . N = 18 1/2) 3" Test Name Value Exact F H/POth. DF Error DF Sia. of F CD 8 Pillais .38487 5 . 10042 4.00 39.00 .001 â ’ Hate 11inas .62568 G . 10042 4.00 39.00 .001 WilKs .61513 6.10042 4.00 39.00 .001 Ro /s .38487 Note.. F statistics are exact. 3- 3" CD Univariate F-tests with (1.42) D. F. "OCD O D. C Variab1e H ’/poth. SS Error SS H/Poth. MS Error MS F Sia. of F a o 3 ■o Leaf 1e t o Lenath .78372 6.80126 .78372 .16193 4.83974 .033 & Leaf let Pairs per cm 5.27800 16.72252 5.27800 .39816 13.25612 .001 co % Bladders per cm 22.83592 4242.31304 22.83592 101.00745 .22608 .637 c/)CO o' 3 Bladders per am 166280190 975634277 166280190 23229387.6 7.15818 .011 ■DCD 0 Q. 1 Table 66. Parametric MANOVA Showina the Effect of Reelicate on Four Variables (Leaflet Lenath, Leaflet Pairs per cm, Bladders/Positions per cm, ■o CD Bladders/Positions per am) Measured on U . vulaaris Plants Raised from Turions in a Common Garden Experiment C/) (/i o' =3 Multivariate Tests of Sisnificance (S = 1, M - 1 , N = 18 1/2) Test Name Value Exact F H/poth. DF Error DF Sia. of F 8 Pillais c5' .15486 1.78661 4.00 39.00 .151 Note 11inas .18324 1.78661 4.00 39.00 .151 WilKs .84514 3 1.78661 4.00 39.00 . 151 CD Ro ys .15486 Note.. F statistics are exact C p. 3" CD ■oCD Univariate F-tests with (1,42) D. F. O Q.C a Variab1e H'/Poth. SS Error SS H /P o th . MS Error MS F Sia. of F o 3 ■a o Leaf let lenath .86397 6.80126 .86397 .16193 5.37233 .025 CD Q. Leaf let o pairs per cm .52260 16.72252 .52260 .39816 1.31256 .258 c ■o CD Bladders C/) per cm 47.68313 4242.31304 47.68313 101.00745 .47208 C/) .496 Bladders per am 998820.692 975634277 998820.632 23229387.6 .04300 .837 VI ■o I I Table B7, Parametric MANOVA Showins the Effect of the Interaction Between Collection Site and Reelicate on Four Variables (Leaflet Lenathf % Leaflet Pairs per cm» B1adders/Positions per cm» Bladders/Positions per am) Measured on jj. vulaaris Plants Raised from Turions in a (/) Common Garden Experiment o'3 Multivariate Tests of Sisnificance (S = 1» M = 1 » N = 18 1/2) CD 8 Test Name Val ue Exact F Hypoth. DF Error DF Sis. of F c5' Pillais .36898 5.70116 4.00 39.00 .001 Hote11inas .58473 5.70116 4.00 39.00 .001 Wilks .63102 5.70116 4.00 39.00 .001 Ro ys .36898 a 3" Note.. F statistics are exact. CD TDCD O Q. Univariate F-tests with (1,42) D. F . C a o 3 Variable Hypoth. SS Error SS Hypoth. MS Error MS “O F Sia. of F o Leaf let CD Q. lenath .00914 6.80126 .00914 .16193 .05646 .813 Oc Leaf let ■D pairs per cm .59246 16.72252 .59248 .39816 1.48802 CD ,229 % Bladders O 3 per cm 1672.76537 4242.31304 1672.76537 101,00745 16.56081 .000 Bladders per am 33246863.9 975634277 33246863.9 23229387.6 1.43124 .238 vnN> ■o o Q. C 8 Q. Table B8. Nonparawetric MANOVA Showina the EfFect oF Collection Site on Four Variables (LeaFIet Lenath, LeaFIet Pairs per cm, Bladders/Positions ■D Per cm, Bladders/Positions per am) For U, vulaaris Plants Raised CD From Turions in a Common Garden Experiment en en Multivariate Tests oF SianiF icance (S = 1, M = 1 , N = 18 1/2) Test Name Val ue Exact F Hypoth. DF Error DF Sia. oF F 8 Pillais .53841 11.37275 4.00 39.00 .000 Hotel Iinas 1.16644 11.37275 4.00 39.00 . 000 Wi Iks .46159 11.37275 4.00 33.00 .000 Ro '/s .53841 Note.. F statistics are exact. 3. 3" CD "DCD Univariate F-tests with (1,42) D, F. O Q. C g. Variable Hypoth. SS Error SS Hypoth. MS Error MS F Sia. oF F 3o "O LeaF1 et o Length 883.44107 6679.51827 883.44107 159.03615 5.55497 .023 CD Q. LeaFlet pairs per cm 2661.97843 5286.78721 2661.97843 125.87589 21.14764 000 ■D CD Bladders cn per cm 46.81903 5606.92541 46.81903 133.49822 .35071 .557 cn o' 3 Bladders per am 918.28210 6538.78520 918.26210 155.68536 5.89819 .020 73 ■OCD I I Table B9. Nonparametric MANOVA Show ins the EfFect of Reelicate on Four Variables (LeaFIet Lenath» Leaf let Pairs per cm » Bladders/Positions per cm» B1adders/Positions per aw) for U. vulaari s Plants Raised from Turions in a Common Garden Experiment cnC/) o'3 Multivariate Tests of Sisnificance (S = 1 » M = 1 » N = 18 1/2) 3" CD Test Name Val ue Exact F H'/Poth. DF Error DF Sia. of F 8 (S' Pillais .15355 1.76874 4.00 39.00 . 155 3" Ho te 11inas .18141 1.76874 4.00 39.00 .155 Wilks ,84845 1.78874 39.00 .155 3 4.00 CD Ro YS .15355 Note.. F statistics are exact. 3. 3" CD T3CD Univariate F-tests with (1 »42) D, F . O Q.C 2r Variable H/poth. SS Error SS Hypoth. MS Error MS F Sia. of F o 3 "O o Leaf let Lenath 801.74177 8679.51827 601.74177 159.03615 3.78368 .058 CD Q. Leaf 1e t O pairs P e r cm 93.89712 5286.78721 93.89712 125.37589 .74595 .393 C ■D CD Bladders cn per cm 93.21797 5606.92541 93.21797 133.49822 .89827 cn . 408 o' 3 Bladders per am 349.35138 6538.78520 349.35138 155.68536 .24396 , 142 VI 73 ■OCD 0 Q. 1 Table 810. Nonparame t rie MANOVA Showina the EfFect oF the Interaction Between Collection Site and Replicate on Four Variables (LeaFIet Lenath, LeaFlet Pairs per cm, Bladders/Positions per cm. Bladders/ Positions per am) For (J. vulaaris Plants Raised From Turions in a (gC/) Common Garden Experiment o"3 Multivariate Tests oF Sian if icance (S = 1, M = 1 , N = 18 1/2) CD 8 Test Name Value Exact F Hypoth. DF Error DF Sia. oF F (S' 3" Pillais .3778G 5.92173 4.00 39.00 .001 Ho te 11inas .60736 5.92173 4.00 39.00 .001 3 CD Wilks .62214 5.92173 4.00 39.00 .001 Ro'/s .37786 3. Note.. F statistics are exact. 3" CD ■oCD O Univariate F-tests with (1,42) D. F. Q.C a o 3 Variable Hypoth. SS Error SS Hypoth. MS Error MS F Sia. oF F "O o LeaFlet CD Lenath 14.88830 6679.51827 14.88830 159.03615 .09362 .761 Q. LeaF1e t pairs per cm 392.13011 5286.78721 392.13011 125.87589 3.11521 "O .085 CD (/) Bladders o' per cm 2647.41644 5606.92541 2647.41644 133.498: 19.83110 . 000 3 Bladders per am 555.75562 6538.78520 555.75562 155.68536 3.56974 .066 U1 vn 7] "OCD O Q. C 8 Q. F i a u r e B1. Boxplots for Ulricularia uulaacia Plants Raised from Turions in a Common Garden Experiment! Stem Diameter by "D CD Collection Site and Replicate C/) C/) 1.27 E 0 8 KEY (O' > Median - 25%, 75% X Hiah/Low 4-— ■# —4- X 0 Outlier f 3. 3" E Extreme X + *--i+ —4- CD +- + -+ X 4--4-- + 4--4--4- ■DCD O mm I Q. C X a X E2 o3 "O o CD Q. 4 h —- ■D .37 X CD (/) o' 3 Site E M M Replicate 1 ui o\ 157 m o c u •u u - _J T) OJ 11 VI ••H a . m TO CK u V» ♦J c • ■ «3 4-> + — + r—V c I I a . l l lU + » + E r j I I «1 + — + f- W 11 m 0. m X 11 + + H LU 1 I =1 TO — — — + + => C O 1 I H «H + + nj X ? — I H L Q. W TO 11 nj a 0 : + + '—( I 1 3 C XJ + + w C'J a o c P \ H S TO + + w H Q 01 3 U + + U TO tn I I o X — + + w t i . c c I I — 1 o + + VI VI o C U o 0> (U 0. »—1 1 X 3 +j X f- rH 1 IT. o t- 11 TO o 3 o 1 c bv 01 O CÛ r- U 1 TO V. ■—1 ll w o > 1 -« «. -C I—* u s 1—1 11 1 1—1 lU 1 X> X TO 4-1 4J o 1—1 4-» 0. .• X: 1 1* ITi 3 X • ■—r ll N n 1 E r-j X O tu 1-1 tn IZ m 1 1 * 1 X O lU m u 5 m •H Ll Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 158 in c o H in 0 o u CL u. v> "O L 01 (U in T3 - w 1 3 OJ 0} m tt ,— 1 m CÛ u in ••H #—4 c • # Q. ra 01 + — — — + c q : 1 1 a . Of O X — — + * + X w t : c i c "O 1 1 «1 ■wt c + — — — + H u m » Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 159 N L> c to ,—1 o CL {- u . U- o •o lU -C in 4-> pi m rO C CC ÜJ > C % # ro ta •— + c I CL 01 O + + f) I I Ul w + — + -4 L M OJ ro 0. a t X tu + + "4 LU «-> ( I 3 to LU O + + LU r 3 c u I I ÜJ ■pf + + W ~o ■ t 4 u (L U to ÜJ n> 13 c n + + •4 I I LU t ' I 3 c r t O X — + 4" + — — — X 3 o c I I 4 10 + + W E 4 o m n U «-» •Pi t - 10 tn T ------i o — — + * + — '— — LU U- c c -p4 o ! ; in Pl ■*-» in o c o p—1 o m 01 0. p* f" 1 1 3 tri Q f - Ot ro X t - r—# 1 u o 3 o 1 c _J 01 5 -w 01 *-4 CÛ F- u 1 to (U •—* o > - 1 p JC ,— 1 L E u CÛ tL CD LU 1 TJ X in • •H OJ X: 1 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 160 ai Ol c (- o w f- TO u. Ql ■3 TO TO in •— t •irl (4- TO TO CK TO _J in «-> c • * TO TO »-> + — + .—t C TO I I O. TO TO W + * + W z : c j 5 I I ifl #—4 + — + ■H f- 0. W TO TO TO C. q : TO X + — + H lU T? I I 3 C + * + o 3 c TO I I TO + — + TO T5 TO ■H U W TO TO (5 cn + + H I I 3 c c + + TO o o I I H e + + W 5 o TO d U TO .—I + + TO #—4 I I UJ o TO + + I I (4. C U + + in X *-» in o C ,—1 o 4J 0. 'M c t 3 TO X u TO 1 ITi TO u TO o o 3 f—t 1 c r ' _J TO 5 ■v. -«H Qj t - CL 1 TO TO CD > 1 w «. s : 1 f- t'J 0. —I TO CJ 1 TO ITi w 3 X c n Ù1 in 1 n C'J X o ÜJ Qj 1 1 * 1 X a Ul 01 t. 3 01 W Ll Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 61 w 0. «n S P O M t . rd t* . CL T 5 * » + — + —« c I I o . 01 01 ÜJ LU + * + s: M S I I vi ^ m + — + H t - o t | Ol nj (L •—I m % 0. + — + ^ lU 11 I I 3 OE + * 3 C I I 01 T l + — + •Hnj TJu m C « m m 13 01 + — + —I Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 162 tn c o *4 •>4 s m o o u Q. Ü- V, tn TJ Ï- tu UJ TJ TJ m 10 a: mW 01 0Û 4J UJ ro u c • « w ro 4 J '—1 + — — + <— f c 0. I I O. 01 Ol UJ + * + E M £ q : I I ul w + — — ^ •H f - TJ t4 01 C fO IL re — + m X UJ Ol I I 3 4-> + * + o 3 c ••"4 1 I Ol tn + — — — + m Xf u. c » m D ITJ a - 4 + + H *» I t ÜJ M 3 c u + + 3 o Ol I I 4 S .-4 + + W H — 4 «4 o o 3 U U + + (_ X I I 10 ÜJ o JQ + + c*. c I I tJ + + UJ c 4 J in re o C m a Q. Q. — 4 1 3 1 r i o t- Ol ro X u L. u o 3 Ol 1 c _J 01 E 1 re — 4 Ol Qj i~ Q. CD fil f - 4 o p—4 f- 4-> C£) •U n. o UJ 1 TJ X tn 4-» Ü*i •4 fil o 3: I 01 IT» -4 3 X en û: r* 4- z: M I 3 U CÛ 1 * 1 X O UJ f- 3 01 Ü_ Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 163 c o E U» o o J- a . Q- Sv u> TJ Ï- (U 01 m ■o w tn m c : •-4 00 Ul *» 0) c nî m + c U I a . 01 «-I — — + * + — r ri £ r—t I Ul 0. + HI- OJ U 0> cn UJ 0. 01 % T? + + •H ÜJ c I I 3 to + + 3 c I I at Ot + + m z? H t . W to tn m C5 + — — + '—( c I f 3 c o w + * + ÜJ C i a o M I I •HE 4-> + — — + w E U *4 o Ot 3 U i—1 + —— + m o 1 1 o U % — + + — — W q . c -1 X + — — — + u> Ul o c ai •—1 o U 0. ••H 1 X 3 rC X t- f. 1 tn o t- 0; O o 3 01 1 c _j Ol E 00 h- a 1 to •iHI Ol o 1 X u— # 1- o eu CD w 1 "O m CD *> 0. Ol ITi *-4 3 X —« 01 1 U3 cn iv 00 00 1 r r 1 % O UJ m m 1 n 1 » 1 XO UJ 01 u 01 w U. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 164 w» c o 5 in o o u CL u . •V. Ul • o Î- (U c m • ro ra + + c o I 1 CL Ol — 1 UJ + + X z: C'J H .— 1 t I V* (L + + t- O j U 01 CK to (L 01 X 73 + - i UJ C I 3 UJ X * 3 c i HI Oil + m 73 «-> •H (. M m eft m a + + H c i I 3 c o UJ UJ + + X UJ C'J u o •pH I I ■H E + + » S O *4 o Ol n CJ #—4 + + {- M o I I o U UJ X — + » + w u . C X •f — ------+ in JO VJ in O c E eu F-* o 01 IL «-I 1 X 3 rd X L t- 1 ITi o u 0 1 o o 3 Oj 1 c |X _J Ol E oil 1 - 0. 1 10 'X •pH 0 1 p. p"H O 01 r-H o 1 • pH JC f~ 2 4p> CD a (X UJ 1 73 X 0 1 01 w Qj X: 1 01 in •pH 3 X ' O 1 E C'J XOUJ in en û: m o ou cn 1 1 y 1 XO W o 01 CO n t. C'J Ll Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 165 "O «u O c m O u Ü. 4-> O l T > 3 0 1 U l p i>4 «4 re m c (V Ü. Ul 0 1 c « * 4J> n3 4-> m c u + + CL O l -w I I E p— 41 ÜJ o + + z: C'J u l •4 CL I ■H C- 0 1 + U 0 1 o : m 0. 01 X 7 3 ÜJ c + + 3 td I I 3 c — + + — — Ol 0 1 I I iq 7 3 * > 4- + t - — 4 CJ ro U l CD - 1 c + — + 3 c o I I U o O X — + » + u M ■H 4J I I W E u + H o o > n U •— 4 t- m o + + o U I I ÜJ (4- c O + + — X \ 1 W 1 j O + + 4 ^ U l o tJ> c 1 o JC Ol 0. 0 1 4J X c - -4 1 X 3 1 ir . o t- Ol o 3 0 1 Q CÜ 1 - 3 1 c rx _ l 01 I m X. ■•H Ol Ol ■-H O 1 JC •—1 {- c o ÜJ 1 XJ X m 4-> 4-1 m cn Q. iC 1 Ol ir. 3 X n •4 01 cv n U l T-« 1 z: M % o til • 0: Q] r- 1 I » 1 X o W m t . 3 m Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 166 JC m s HI o 3 X U Q Ol V» I—4 —* 10 ffi c q: u_ in c m a #—4 c + + CL 0> N I 1 5 LÜ + + UJ r : N VI I I H L- Of + + U 01 «-> rd a 10 n X Ü H ID + + 3 I I a c OL + + 0* OJ I I It) X? cc + + •H L. LI 10 -o IdCD c •H ID + + 3 c I I U o Ol + + — ■H c I I LI S + + *4 o in 3 U c t- 10 o + + o I I u. c L> + + Ixl rl Ü 1 I tn 01 + + int—f o C «-a o O 01 (L rl u *-> X t- 3 os o 3 N in o u Ol u Oj F* XI c f'- Ol s 10 'X w 01 s: yW L- c O Ol o Q. ui LU X 01 «-> 01 4" CD 5(: Ol in -i 3 X o i n C E n C) X o tu ' m 1^ 1 X o U.I Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 167 Figure B12. Profile Plot for the Interaction of Collection Site and Replicate Experiment on Bladders per cm for L[. vulgaris Plants Raised from Tarions in a Common Garden Experiment Key: East Bay — McWenneger’s Slough bladders per cm \ Repli cate 1 2 Experiment Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 168 Appendix C- Data Summaries, Complete MANOVA Results, Boxplots, and Profile Plots for Utricularia vulgaris Plants Raised from Turions in a Diet Experiment Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 169 Tst'le Cl. Data Summaries for Utricularia vulgaris Plants Raised from Turions in a Diet ELxpe r i iTient Î Stem Diameter» Leaflet Lendthr and F'*lant Lendth by Treatment Tray Treatment Tray Stem L.eaflet Total D if!Ille t e t' L i?i id I h Lendth in Him in cm in cm Mean .94 1 . 9 5 16.95 S t Q , D e V i it t ion . J 1 .43 15.07 Minimum .76 1.46 6.30 Ma ; ; imum 1.13 2.99 64.90 Sample Size 13 11 13 Mean .85 2.13 16,33 Std. DeVicl Lion .16 .42 4.35 Minimum .53 1,56 9,30 Ma;; i mu m 1 .00 2 . 7 2 2 2 . 0 0 Sample Size 9 3 9 3 M t; a n .96 2 . 2 9 .1.3.78 Std, Deviation .06 .55 4 . 6 0 M i n .1. Ill u m . 34 1 .37 9 . 1 0 M a ; ; i m ij ni 1 . 0 5 2 . 9 6 2 0 . 6 0 Sample Size 8 6 8 Mean .92 2.42 16.13 Std ♦ Devi at i on .18 . 35 ^ M i n i m u in .60 1.91 ^ O Ma;: i mu m 1.20 2.94 33.20 Sample Size 7 6 7 Mean .90 2 . 1 9 44.98 Std. Deviation .12 .25 16.9^ Minimum .<^2! l.Svj 18.00 Maximum 1 * 00 2♦a 1 68./ SaiTiF'le Size 9 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 170 T a b l e Cl. Data Summarles for Utrxcularia vulaaris Plants Raised f rom Turions in a Diet Experiment I Stem Diameter » Leaflet Lendth y and Plant Lendth by Treatment Tray (cont'd.) Treatment Tray Stem L e f 1 e t T o t a 1 Diameter Lendth .endt h iII mm in cm in cm Mean .84 1.93 21 .60 Std. Deviation . 17 . 44 9.65 Min i mum .53 1.10 10.20 Maximum 1 .03 2.37 46. 10 Sample Size 13 13 13 7 Mean 88 1.99 42 75 Std. Deviation 19 . 29 19 56 Mi I I imurn 64 1 . 55 20. 80 Max i mum 24 2.40 71 .40 Sample Size 10 10 10 8 ne an .81 2.00 '■> ' ) Std. Devi ation . 15 * A- 12.00 M i n i m urn . 60 1 . 7 J. 16.10 Maximum 1.10 2.46 59.50 Sample Size 13 12 13 Mean .84 2.09 38. 31 Std. Deviation . 13 . 28 15.21 Minimum . 62 1 .59 16.20 63.60 Maximum 1 .02 2.40 Sample Size 12 12 10 37.43 Mean . 71 1 .87 Std. Deviation ♦ 11 .36 19.70 16.00 Mi ni mum . 53 1 .44 68.60 Maxi mum .84 2.41 9 Sample Size 9 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 171 T able Cl * Hata Summaries for Utriculacia vulaaris Plants Raised from 1urions in a Diet E>îper i ment î Stem Diameter y Leaflet Lendthf and Plant Lendth by Treatment Tray (cont'd.) T l'e a t m e? n t T »• a Stem Leaflet T o t a 1 Di amc'te r L e n d t h Lendth i n m iTi in cm in cm 11 Mean ♦ 82 1,87 35, 92 Std, Deviation . 19 ,26 24,03 Minimum ,54 1 , 26 15,00 Ma;c i mum 1 ,19 2,21 79,80 Sample Size 9 9 12 Mean ,81 2,02 37,47 Std, Deviation , 09 ,28 26 , 96 Minimurn ,60 1 ,65 15,00 Moximum ,93 2,57 101,80 Samp 1e Size 9 9 13 Mean 60 1,27 16.53 Std, Deviation 14 , 22 11 ,08 Minimum 40 , 98 5,90 Maximum 70 1 ,48 31 ,90 Sample Size 4 4 4 14 9.97 Mean ,33 1,08 Std, Deviation ,21 ,19 5,03 5,30 Minimum , 10 ,78 18,50 Max i mum ,53 1,31 7 Sample Size 7 7 1,17 1 5, 58 Mean ,44 8,11 Std, Deviation ,08 , 19 Minimum ,33 , 86 5 ,00 25,70 Max i mum .56 1 . 35 S a ni p 1 e Size 6 6 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 172 Table Cl. bata Summaries for Utricular ia vulsîaris P 1 a II t s lÂ’ a i sed f r o ly, T u r i o ii s i ri ei Diet Eixeer i ment I Stem Di a me te r y Leaflet Length r ai id Plant Lei lü tli by Treatment T ray Treaitment Tray S t e m Leaflet T otal Diameter Lei iF(t h L e ri rî 1- h in mm in cm i I I c III 16 Mean .41 1.51 22.70 Std. Deviation .22 .37 11.15 Minimum . 13 1.05 12. 30 M aXimum . 70 2.17 48. 90 Sample Size 1 0 10 10 17 Mean . 42 1 .42 18.10 Std. Deviation . 20 .19 11.16 Mi II i mum . 10 1 . 1 9 7.90 Maximum .67 1.64 38.50 Sample Size 7 7 7 18 Mean .48 1 . 13 9.44 S td , Deviation . 24 . 21 4.19 M i II i mu m . 10 .82 4.20 Max imum .80 1.42 16.30 Sample Size 8 8 8 19 Mean .95 87 19.28 Std. Deviation .22 46 9.20 Mini mum .80 1 15 7.40 Maxi mum 1 .40 60 33. 80 S a in p 1 e Size 8 8 8 20 20.43 lir a I I 89 1.98 Std. Deviation 13 . 3 J. 6.75 1 1.50 M i n i mum 70 1 .45 33.50 Maxi mum 13 2.44 Sample Size 11 11 11 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 173 Table C l , Ijata Suiiiiiiaries for U;L ricuiax^ia vulsiaris Plants Raised f rom Turions in a Diet E;-;pe r i ment î Stem Diameterr Leaflet Lendthf and Plant Length bw Treatment T ray (cont'd*) TrecJtment Tray t e m Leaf1 et Total Diameter Lenath Lerii^thi i n m m i n e m i n c iTi 21 Mean . 84 1 * 94 24 .74 Std* Deviation * 18 *23 7 *5 2 Minimum *63 1 *63 1 5 .0 0 Ma;; i mum 1 *1 7 2 *2 6 3 6 .0 0 Sample Size 11 11 11 Me a n 1 * 05 1*94 3 5 .2 1 Std * Devis; t ion * 21 . 19 15*37 Mi nimum *80 1 * 65 1 7 *40 Maximum 1 * vf) 2 .3 5 6 8 .3 0 Sample Size 12 12 12 Î3 Mean * 93 1 * 94 3 3 *3 3 Std* Deviation *21 * 38 1 9*6 9 Mini mum *68 1 . 30 1 4 *80 Maximum 1 *33 2 * 5 7 78* 70 Samp1e Size 12 12 12 24 Mes; n *85 1 * 72 2 5 *7 8 Std* Deviation *20 . 28 10*49 Mini mum *57 1*3 4 15.32 Maxi mum 1 *30 2 * 4 0 5 3 *4 0 Sample Eîize 12 12 12 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 174 Ta ble C 2 ♦ Data Sun,maries for Ut.ricularia vulsiacis Pl ants Raised f rom Turions in a Diet Experiment I Leaflet Pairs per Plant» Leaflet P Treatment T ras; Leaflet Leaflet B1adders/ F’ i r «i F'a i rs Positi ons per Plant per cm per Leaflet Mean 5 0 , 15 3 ,2 5 11 ,92 Std, Deviation 33, 12 ,81 ,95 Minimum 21 2,22 11 Maximum 150 5 ,2 0 13 Sample Size 13 13 12 Mean 47 78 2 . 9 7 11 , 28 Std, Deviation 12 44 ,42 1 ,1 7 Mini mum 30 2 ,3 1 10 Maximum 63 3 ,5 6 13 Sample Size 9 9 9 lie an 6 3 ,2 5 4 ,5 2 1 0 .0 8 Std, Deviation 2 4 .6 4 .51 2 ,6 9 MiniiTium 3 6 3 ,9 6 8 Maximum 97 5 ,2 4 16 Sample Size 8 8 8 11 ,10 Mean 67 . 43 4,68 Std, Deviation 3 0 . 6 7 1 ,41 1 ,6 5 9 Minimum 35 3 ,5 8 14 Max i mum 119 7 ,5 0 7 Sample Size 7 7 11 ,84 M e a n 110,00 2 ,6 1 1 ,09 Std, Deviation 2 7 ,0 1 ,59 10 Minimum 65 2,11 14 Maximum 148 3 ,6 1 9 Sample Size 9 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 175 Table C2» Bata Summaries for Utricular ia vulsfaris Plants Raised from Turions in a Diet Experiment I Leaflet Pairs per Plant ? Leaflet Pairs per cm y and Bladders/Positions per Leaflet by Treatment Tray (eont"d *) Treatment Tray L e £} f 1 e t Leaflet Ed adders/ P airs F’ a i r s Pos i t i ons per PI ant per cIII per Leaflet Mean 79,31 3 . 74 9,41 Std, Deviation 33,86 ,60 1,23 Minimum 4 6 2, 93 8 Maximum 165 4,80 12 Sample Size 13 13 13 Mean 121,70 2,87 11,24 Std, Eievi at ion 54,33 , 28 1 , 18 Mi n i mum 55 2,49 10 Maximum 198 3 , 26 13 Sample Size 10 10 10 8 Mean 94 ,46 2,91 10,44 Std, Deviation 37,97 , 45 1,11 Mini mum 43 2 , 26 8 Maximum 165 3, 72 12 Sample Size 13 13 13 Mean 105,92 2,83 11,41 Std, Deviation 38,50 ,28 1,00 Mini mum 52 2,32 9 13 Maximum 179 3,29 Sample Size 12 12 12 10 9.94 Mean 121,78 3, 44 1,46 Std, Deviation 53,01 , 56 7 Minimum 69 2, 65 12 Maximum 205 4 . 50 9 Samp' 1 e Sizt^ Q 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 176 Table C2. Data Summaries for Utricularia vulaaris Plants Raised from Turions in a Diet Experiment* Leaflet F'ai 1rs per Plant r Leaflet Pairs per cm f and B1adders/Pos i t i ons per Leafiest by Treatment Tray (cont'd.) T reatmen t Tra Leaflet l.eaf let DIadders/ F" £i i r s P a i r s Posi t i ons e r Plant per cm per Leaflet 11 Mean 112*89 3*35 10.07 Std. Deviation 62*32 *48 1 *04 Min i mum 56 2.77 9 Maximum 229 4.13 11 Sample Size 9 9 9 12 Mean 115*89 3*21 11 .46 Sîtd* Deviation 77*49 .51 .91 Mini mum 43 2*51 10 Max imum 302 4.21 13 Sample Size 9 9 9 13 Mean 58*00 4*21 8*90 Std * Deviation 18.17 1 . 44 1 *39 Minimum 35 2.48 8 Max imum 79 5.93 10 Sample Size 4 4 4 14 Mean 47. 14 5.30 9 47 07 Std * Deviation 12*32 1 *55 1 7 M i nimum 27 2*81 10 Max i mum 65 7* 64 7 Sample Size 7 7 15 9,40 M e 3 n 86 . 67 5*98 Std* Deviation 37*45 1 *09 *87 Minimum 36 4*76 8 Maximum 142 7*47 10 Sample Size 6 6 6 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 177 Table C2* Data Summaries for Utricularia vuldaris Plants Raised from Jurions in a Diet Experiment* Leaflet Pairs per Plant y Leaflet Pairs per cm r and Bladders/Positions per Leaflet by Treatment Tray Treatment Tray Leaflet Leaf let E< 1 adders/ Pai rs P airs Positions per Plant per c m per Leaflet 16 Mean 95.90 4.29 9.71 Std * Deviation 45.89 . 59 1 .33 Mini mum 51 3. 49 8 Max i mum 206 5, 46 11 Sample Size 10 10 10 17 Mean 83.29 4.74 9.72 Std. Deviation 47.06 .63 .67 Minimum 40 3.96 9 Maximum 165 5.95 11 Sample Size 7 7 18 Mean 47.50 5.20 9.66 Std. Deviation 18.59 1 .07 .50 M i n i mum 18 3 . 99 9 Maxi mum 79 6.6)4 10 Sample Size 8 8 8 19 Mean 60.63 3,33 12.37 Std. Deviation 24.57 .73 1 .40 Minimum 33 2.23 10 Maximum 92 4.46 14 Sample Size 8 & 8 20 11.31 Mean 63.91 3.19 .57 Std. Deviation 18,64 .43 1 9 Mini mum 42 2.56 13 M a X imum 95 3.98 Sample Size 11 11 11 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 178 Table C2. Data Su mm a ries for Utricularia wulj^aris Plants Raised from Turions in a Diet Experiment I Leaflet Pairs per Plant y Leaflet Pairs j-er cm» and E<1 adders/Positions per Leaflet by Treatment Tray (cont'd») T r eat ment 7 ra^f L e B f ]. E4 t L. e a; f ] e t E< 1 adde r s/ F’ ct i V s> F'ai rs Positions per F’lant per cm per Leaflet Mean 75* 64 3 * 1 5 11 *80 Std* Deviation 16*92 *42 1 *20 Minimum 48 2*43 10 Maximum 105 3*67 14 Sample Size 11 11 11 Mean 90 * 75 2*68 12*34 Std * Deviation 30.41 .38 .84 Minimum 50 2.08 11 Max i mum 142 3*40 14 Sample Size 12 12 12 23 Mean 96* 17 3*12 11 *68 Std* Deviation 44.38 * 77 1 *43 Mini mum 62 2*45 10 Maxi mum 202 5 * 1 4 14 Sample Size 12 12 12 24 Mean 78*75 3*24 10* 84 Std* Deviation 24*02 1 *07 *82 M i n i IÏI u m 50 2*11 9 Max X mum 133 6*40 13 SJaritple? Size 12 12 12 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 179 Table C3. Data Summa r i es for Utricularia vulda cis Plants Raised from Turions in a Diet Eîîper i ment î B1 adder s/Posi tions per Plant and Bladders/Positions per cm by Treatment Tray T reatment Tray Total B 1 adders/ B 1adders/ F osi t i ons Positions per cm per Plant ne an 1202.72 73,66 Std. Devi a*t ion 928.90 15.62 Mi ni muni 490 50.37 Maximum 3900 99. 46 Sample Size 12 12 Mean 1084.12 67.28 Std. Deviation 333.16 13.59 Minimum 694 44.92 Max imum 1571 8 8 . 75 Sample Size 9 9 Mean 1329.50 91 .51 Std. Deviation 755.11 27.92 Minimum 603 62.40 Maxi mum 2703 148.51 Sample Size 8 8 Mean 1543.34 103.24 Std. Deviation 875.29 30.39 Minimum 649 69.01 Max iiri'.im 3284 151.00 Sample S>ize 7 7 Mean 2633,87 61 .01 Std. Deviation 761.69 9.89 Miniiri'JUfi 1248 50.79 Maxi mum 3651 78. 14 Sample Size 9 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 180 Table C 3 ♦ Data Summaries for Utricularia vulSaris Plants Raised from Turions in a Diet Experiment ; B1adders/Pos i t ions per Plant and D 1 adders/F'ositions per cm by Treatment Tray < con t 'd *) Treatment Tray To ta 1 Bladders/ B 1adders/ Posi tions Positions per cm per Plant Mean 1456.91 70. 86 Std, Deviation 518.19 17.14 M i. ri i. m u m 724 46. 10 Maximum 2684 113.37 Sample Size 13 13 7 Mean 2752.99 64.33 Std. Deviation 1309.98 6 . 73 Miiii mum 1357 49. 79 Maximum 4794 75.49 Sample Size 10 10 8 Mean 1984.12 60 . 59 Std. Deviation 847.03 10.20 Minimum 936 40.70 Max imum 3454 74 . 99 Sample Size 13 13 Mean 2365.81 64.78 Std. Deviation 726.63 10.34 Mi ni mum 1241 4 9.46 Maxi mum 3723 80. 38 Sample Size 12 12 10 M e a n 2489.82 68.50 Std. Deviation 1279.76 14.80 Minimum 990 40.57 Maximum 4647 89 . 40 Sample Size 9 9 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 181 Table C3. Data Summaries for Utricularia vulSaris Plants Raised from Turions in a Diet Experiment» B1adders/Positions per Plant and Bladders/Positions per cm by Treatment Tray < cor it ' d . > Treatment Tr^ry Total B 1adde rs/ B 1adders/ Pos i t ions Pos i ti ons per cm per Plant 11 Mean 2241 » 75 67.63 Std » Deviation 1201.81 12.15 M i n imum 1262 47.62 Maximum 4672 84. 12 Sample Size 9 9 12 Mean 2655.17 73.45 Std. Deviation 1815.74 11 .48 Minimum 1124 55.27 Maximum 6926 91 . 44 Sample Size 9 9 13 Mean 1030.63 74.92 £>td. Deviation 393.74 30. 88 Minimum 705 50.19 Maxi mum 1601 119.44 Sample Size 4 4 14 Mean 903.39 98. 14 Std. Deviation 300.99 20.71 Minimum 511 56.22 MaXimum 1361 116.50 Sample Size 7 7 Me a n 1623.13 113.08 Std. Deviation 744.54 26.90 Minimum 710 83.06 Maximum 2878 147.34 Sample Size 6 6 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 182 Table C3. Data Eîuirimaries for Utricularia vuls^aris Plants T-aised from Turions in a Diet [ZMF-er i ment î Bladders/Posi t ions per Plant and Bladders/F'ositions per cm bs Treatment Tray (c ont'd * > T r e E) t m e n t T r a y T o t a 1 D 1adders/ E{ 1 Et d d e r s / Posilions f’o-ii tions per cm per Plant 16 Mean 1900.21 82.34 Std. Deviation 1065.90 7. 24 Minimum 1081 70.00 Maximum 4477 91 ,56 Sample Size 10 10 17 Mean 1659.66 92.26 Std. Deviation 1015.78 14.33 Minimum 688 68, 12 Maximum 3410 115.81 Sample Size 7 7 18 Mean 919.03 100.24 Std. Deviation 379.38 20.41 Minimum 372 79.22 Maximi.im 1580 135.02 Sample Size 8 8 19 Mean 1507.52 83.32 Std. Deviation 661.33 24.22 Mini mum 733 46.67 Maximum 2333 118.92 Sample Size 8 8 20 Me Bn 1428.45 72.47 Std. Deviation 418.22 16.36 Mini mum 877 52. 21 Maximum 235 2 95. 42 Sample Size 11 11 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 183 Table C3, Data Summaries for Utricularia vuldaris Plants Raised from Turions in a Diet Experiment I Dladders/F'osi tions per Plant and D 1adders/Positions per cm by Treatment Tray (c e n t "d ♦) Tre;Ît ment Tray Total B 1 adde rs/ B1adde rs/ Positions Posi t i ons per cm per F' 1 a n t 21 Mean 1788.75 74 .00 Std t Deviation 457.69 10.07 M i n i m u iti 1107 49.96 Maximum 2559 86.22 Sample Size 11 11 ri e a n 2214.69 66.55 Std. Deviation 679.81 12.62 Minimum 1:336 48. 79 Maxi mum 3332 90. 76 Sample Size 12 12 23 Mean 2210.07 73.04 Std. Deviation 982.60 20.29 Minimum 1471 50.34 Maximum 4848 124.61 Sample Size 12 12 24 Mean 1713.86 71 . 09 Std. Deviation 575.32 29.25 Mini mum 1100 46 . 41 Maximum 3050 160.35 Sample £>ize 12 12 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 184 Table C4 ♦ Data Eîunimaries for Ut ricularia vuis(aris Plants Raised from Jurions in a Diet Experiment I Bladders/F’ositions per Sm by Treatment Tray Treatnient Bladders/ Tray Positions per Sm 1 7478.05 2 6294.89 3 4748.21 4 3662.16 5 7140.01 6 7575.93 7 9493.07 8 9733.42 9 10103.10 10 9700.60 11 12933.17 12 11433.75 13 11778.63 14 31618.65 15 30433.69 ]6 33932.32 ]7 29780.77 18 33419.27 19 10963.78 20 10008.25 21 11642.75 22 8227.95 03 9404.55 24 9222.57 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 185 Table C 5 , Bata Sunriniaries for Utricularia vuldaris Plants Raised from Turions in a Diet Experirrient Î Final Wet (Blotted) and Dry Weights by Treatment Tray Treatment Traya y Blotted F i na 1 Weidht Dry Weight in dm in dm 1 34,66 1 . 93 2 27,20 1 ,55 3 38, 50 2,24 4 47,97 2.95 5 56.92 3,32 6 32, 12 2,50 7 54,23 2,90 8 44,05 2. 65 9 43,48 2,81 10 34,27 2,31 11 30,43 1 ,56 12 30,56 2,09 13 4,22 ,35 14 3,99 ,20 15 4,65 ,32 16 10,61 ,56 17 6 , 36 .39 18 3.58 .22 19 22,45 1 , 10 20 24,97 1 ,57 21 27. 16 1 ,69 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 186 Table C5« Data Summa r i es for Utricularia vulaaris Plants Raised from Turions in a Diet Experdmentî Final Wet (Blotted) and Dry We i a ht s; by Treatment Tray (cont'd.) T r e a t ment Tra y D1o11e d F i na1 Weidht Dr^» Weisht in dm in dm 22 49.09 3.23 23 40.69 2.82 24 32.38 2.23 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 187 o O O O o O 4^ O P o o O c 01 G u 4 - fS £- C? O OJ "U •fi li_ en c o -p > ü_ o O O Lu m un un X cu o c O X en r-! Oi o o O X X X 4 - = 1 G * * ♦ f i en X X ri f3 un iX X f i cn ar G r t tn X r? un c X M T-i X f4 f i o (. X? en t-i f4 ■H OJ G -P i>- 4 P ÜG o G f o o o en X un X Oi b1 tn G o o o z f i f i un c G E ?! » * * * r un f i X G G £- tn ?n m en f i T-i o *H Z £. Z es o- O' o o m X T"? CJ ■p G c •H (_ t—î X V y1 e % Lu LU en G G o u i X Ü_ X c- !l a in G G o o o U3 f i • Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 188 O -i rs O T N'. O O ÜJ o ^ ly s— sri J eu it! M O _ ! Û! ü_ ■ en -P e>- li") '=r es o Oi üi üi Ü-; CN CJ es V o o o c-i ü~: rs CD eu «p = 1 o o- (S T -i s ns U*3 S3 S3 eu c C-i tn O \ tn Ci Sp '0 ro % T-! ù: TD CD 01 U l tn Ol -P Lb. OO O CD es ÜO rs T i-i u* CU II O o o Z ? C4 C-J liO ro i= • • ♦ < - UD CJ SO Si o ■1; z s~ CD 4 : a— CD CJ Tp Tp Oj •J-! %: i- o CD CD GD G O ro rs Tp Ü. eu t- T-: T—i U. »« ‘i Ci. se. T -i rs 4 - in s \ l u lu CD O 0 cj LU ro Ù, -P \ -p H ■» o V! Cj Li_ OO O-P CD rs Sj ÜD ro 0! 3 : C i O o O CJ Z o CJ O o 4 - 0; C i * »» ro es c-j rs c-j "O ■Jt Sa ♦ CD CD CD y r * T -i ro UD LU rc r-j eù X r-j ro UD UD ro »» • T Ci r-4 •ïS i! O en o Tp 'T O- jC iXt r-f i -r-i ■X -X- -P CD 3 y. C-j ■n y aw X !0 * Tp o c ü . û_ o C i rc JCi CD ro O Ts 'C -fS tn \ CJ < - r-j ro T CD rs es CJ rs 3 VI c « O'*T r-i ro iZi o ro UJ C eu 3 m % 7—! CD üD t- C-J rs i~ _ j o es e> V G CJ ro » cn •fS >BS * • * en sC » • ■» ro -p •O 4 - iX -r-i s—i T-< s es CD rs Gj c < r cu rc C iX j Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 189 CO C l CD o is O S3 O r-j Oi c r-l 0; 4- rr, t. ro M di co CO o P c Ï- TO G « f «T T ü_ S3 un 9" iS o d j CU 5 e t e t C-J <- ro et et -Î-Î Û. s . •j-i < r 00 ro et rs V 3 (. o *•« et S3 T—; o 3 Ui w) Si es et ro S3 t-t •ro CL ■I-S * Q O % c t ro cn ■7-i : c w \ -I-i -P T—i 0 3 4 -) p c Ui £ Si SiT & c Ü •r4 U_ o O o cn es u3 rs *?• r l Û- a a o o o z *T et et Îi3 \ c . !! * «. « Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 190 ÜT rs rs. S3 M O 11 m m -p ». TT a îc ü cc 0! 33 (- m l3_ Ci r-o Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ■o I I Table CIO, Parametric MANOVA E)howins the E ffect of the In teractio n between Collection Site and Nutrient Solution Strength on Four Variables ■a CD (Leaflet Length y Bladders/Positions per Leaflet y Leaflet Pairs I per cm y B1adders/Positions per cm) Measured on U, vulsaris Plants C/) Fvaised from Tu rions in a Diet Experiment 3o' o Multivariate Tests of Sidnificance (S .1. y M = 1 y N 45 1/2) CD 8 Test Name Va 1 ue Exact F Hypoth, DF Error DF Sid, of F (S' P il l a i s ,07830 1.97507 4 . 0 0 93.00 ,105 3 Hotellinds ,08495 1.97507 4.00 93.00 ,105 CD Wil k s ,92170 1.97507 4.00 93.00 ,105 C Roys ,07830 p. Note,, F statistics are exact. ■DCD O Q. Univariate F - Les Is with (1 » 96) D, F, CQ ■o Variable potIt, E)E) E T' I'o r E>E> H poth, MS Erro r MS F Sid, of F O Leaflet CD Q. lendth ,36948 8,11073 ,36948 ,08449 4,37323 ,039 Bladders "O per le a fle t ,53029 127,22390 2,53029 CD 1,90930 170 C/) C/) Leaflet pairs per cm ,27918 68,37827 ,27918 ,71227 ,39195 ,533 Bladders per cm 40,92115 37163.1876 40,92115 387,11654 ,10571 ,746 192 0" S- LV U2 GJ O m Ù. C m û) 0) m O i Î-! « - u> 21 3 JQ •'t -p •1-; +) 1% m c t îï CD -iH û_ m £ L ü _ c o U 2 t 4 P 0 3 C i C i e s c o r n U 2 v G 'C T Ü 2 t -4 < r o r-t U l C i C-4 C 4 4 3 4 3 O - •T • H s- • i S ' G * •• IN r o o u o 4^ 3 m i - liO o o T-i Cl o & 1 2 C%l ♦ «. « m ü ! 2 1 X 2 1 C-4 t -4 w r-i G) c » Clj 4^ G -P > \ n C eu < r î—j ^ 1-4 ♦ O OO C O e s liO ÎN < r -P V il C o O O z < r C i C i u o ai m 12 • » ♦ < r ü 2 C i 4 3 jc c ai z : {- œ T SD u 0 3 C i t -4 ■p ai —j o G CD CD CO G O r o ÎN t 4 (~ >. P T — : T— i - «*T 4- -p t- 1 3 ü C 4 IX ï - T-i Î N o m ai O l G \ P i P CD ÛL t- 2 r 4 r o ■P C 3 H o o ■fii. II * û! -M c t v o O o P CO 0 3 U 2 O C i V -p c Q j X 3 3 o o O o O r 4 3 4 3 U 2 4 3 4 - 3 M X ■*- ■* ♦ r c ÎN r o 4 3 V ÜJ 1-4 -P U i * c o w œ X * T's O O o C-i x r G C i r o O 0 3 G CD m * * * ♦ JC o Ü G II o •31 G T—I e s -p 4-s a. *-î ■ X •i-i X c \ s - d t n 3 y 21 ai en 3 : • 2 ♦ p t- •1-t C- C L ! 2 1 3 p •!-i i- ai G » p C 4 M LO c o MO ÎN 4 3 3 •p "O 2 1 G r i T-i C i ♦ c o ■ N D- C i ÎN u 3 13 C' — î C » r v 1 0 O m s O r o i p j= 2: m o ■ 2 % r v ' C f v C i r s to 1-4 • e 4 Î/1 G c C l C i CJ e-v G T-i C i r o • 1 3 CCI ■ P •i4 *** c o 4 3 t - * ♦ * r o Reproduced with permission of the copyright owner. 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T — i T—i 1 — i u. o un O' f—i O !l no m m o Xv S3 rx O ü_ 4- CO CD CD T CO un o cn \ Î4» C O ** * CN CD o 14» C •f4 T-i -r-i •T c. o • -» * - • 04 14» r-4 01 1-i Oi •O P X. -r-i •H •O •fi O 1-4 cn (- no U» -M +) T"4 C- yn D iXi Û. <■ u_ o O o cn ■ un r-4 IX r-4 Z\ h - 5 o o o z ro un O' m >> 14» i! * ■» * to CD- r-4 <4- JZ £- £ (~ m 3 rt -CN 0 •p 04 C Z O cs i> O' o T—i 03 o to 01 (_ t_ •P c "3 £- s~ r-4 to 03 rx u 04 ro w LU o UI T-i o "C- o ro to r-4 a -Oi r-4 ct r-4 to 10 Oi OI to m m * o O' CN o O' 3 => r-4 X * * -» p o s~ o '0 m o Z «4- P LtJ Cj o- (- r-4 O' UD O'. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ■o I I Table C16, Nonparametric MANOVA Showing the Effect of the Interaction between Collection Site and Feeding Redime on Four Variables (Leaflet ■o CD Lendthr Bladders/Positions per Leaflet y Leaflet Pairs per cmy (/) Bladders/Positions per cm) Measured on vulsaris Plants Raised C/) from Turions in a Diet Experiment Multivariate Tests of Sidnificance (S M 1/2, N =: 45 1/2) 8 Test Name Value Approx* F Hypoth* DF Error DF Sid* of F ci- Pillais *24095 3*21904 8*00 188*00 *002 Hotellinds *29838 3*43140 8 . 0 0 184*00 *001 Wi Iks *76534 3*32637 8*00 186*00 *001 Roys 3- *21114 3" CD Note * * F statistic for WILK'S Lambda is exact* ■oCD O Q.C Univariate F-tests with (2 ,96) D* F * g. o3 ■o V a r i a hie Hypotli * S S E r r o r S S H y p oth * MS E r ro r M S F Sid. of F o Leaflet & lendth 2784*97734 192209*602 1392*48867 2002*18335 *69549 *501 3O" B1adders S- "8 per leaflet 8110*11958 225969*458 4055*05979 2353*84852 1 * 72274 .184 (/)CO Leaflet o' 3 pairs per cm 3642,39374 222530*110 1821*19687 2318*02197 *78567 *459 Bladders per cm 24267.0100 307909*663 12133*5050 3207*39232 3 * 70298 *026 t98 li. MO S3 S3 al Oi O O iX. 'T Oi UI il tH MO O O 3 0; * -ri r-t t. ifi i j to -fH P •H ro C u ÛL. to CD o nj rH ■I-t O P ü_ rs M m li. MO tH so MO OJ o o o MO SO ir-î rx Li ift 4- tH tH tH MO fx MO cx na 4- r4 O uD MO «- o <0 t- rx at Li. 0! ro ON tH -p -J Of CJ 01 C*J tH c c rH \ H MO i-! o 3 tH CD > Oi jC Qi IjD JC r—i V Le. O O O CD ÜD CJ fx Ci -p fn 4- = 1 1=1 o o o Z MO ÛD es fD L. ro i! MO CO iH Ci 4- 2i & i. FD M M c. CD Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 199 M ii1 D' r.! IX 01 04 Ci 04 Ù. ilT 3 + s 04 141 04 r— i £. 141 wi X i JCî •r-i •Ti r s nj C en c -I-i Û. 40 0 C. r— i •H ro p T - i O ' - O u. CN Ü0 05 CN - p 0 04 0 0-0 fO 05 'C T-i C4 r-i 141 4 - un Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 7J ■DCD O Q. C g Q. Table C19» Nonparametric MANOVA Sliowin*^ the Effect of the Interaction between Collection Sitev Feedinfl Redime and Nutrient Solution Strendth on ■D CD Four Variables (Leaflet Lend thy B1 adde r s /F'os i t i ons per Leaflet» Leaflet Pairs per cm » B1adde rs/Po s i t ions per cm) Measured on (/) (/) U* vuldaris Plants Raised from Jurions in a Diet Experiment Multivariate Tests of Sidnificance (S - 2» M - 1/2» N = 45 1/2) 8 Test Name Value Approx, F Hypoth, DF Error DF Sid, of F Pillais ,12819 1,60924 8.00 188,00 ,125 Hotellinds .13731 1,57910 8,00 184.00 ,134 Wilks .87578 1,59419 8,00 186.00 ,129 Roys ,07618 3 Note,, F statistic for WILK'S Lambda is exact. 3 " CD "O O Q. Univariate F-tests with (2»96) D, F, C a o 3 Vari able Hypoth. SS Error SS Hypoth, MS Error MS F Sid, of F "O o Leaflet CD Q. lendth 245,00191 192209.602 122,50095 2002,18335 ,06118 .941 Bladders ■o per leaflet 3527,65355 225969,458 1763,82678 2353,84852 ,74934 ,475 CD C/) Leaflet (/) pairs per cm 10085,5912 222530,110 5042,79560 2318,02197 2,17547 ,119 Bladde rs per cm 13400,7155 307909,663 6700,35775 3207,39232 2,08904 ,129 ONS o 201 n o rv rj o o 'T Ci o o c - p o OJ UI rH *t-4 c ^ o - p tn Q fi) m Ci -o 5 m _j (D £ O o O OJ ül 'T Ci u M o o o *Ï—i X <3 en c 3 «- o o o X X X ÜT Cl ai Cl Sù o M X < - i CL H3 0. o. o O M f i) Ci % -p ai S L U Dl o ÜT •H OJ Cl o ü ) £ -fi £ Cl •H 4-) fj M Ill î» ‘fH ÜT 3 O u; X 0 4 Ci Uî Q O 3 O 2 X X M Û. O w !! 03 -o ÜT X % ù_ a. OJ '•J 0 4 £. <- tn O U i X 2 C M X ai at •r-t i - T-Î *rH t - i L. ***« 4 — (- ». £. C. 0 4 O Cl o at LU U i M 3 ■r4 t H 0 4 P "O P G Ü i t } ■ H •H n Ci rO Ifi (_ o o O e n C C TÜTÜT p a G 3 a o o o 2 ÜT 0 4 ÜTO ü _ H- •* - ' f M 3 0 4 X U i »v X ». ♦ *T *T < r • X ÜT rH X JZ ai fH 2 JZ T-i X o CD C p (- O P P • - » *■ 2 t n G c II O GC 0 3 p c CL ‘X 0 4 * Cl I D m j t 31 •T Cl p rs "GXX T-i c r H G 'Î-Ï p a at cu u . 0 4 0 4 0 4 » OT 0 4 O < 3 < r 3 Cl o CO œ 0 3 w 0 3 0-4 O 0 4 o rs c p 0 4 0 4 0 4 ■ O < r r-T < - JZ p Ù: m o S 3 S 3 V 3 s O o T 0-4 v ) m G (J 0 4 0-4 0-4 * ÜT G 0 4 X ü T •» Cl at 'M n \ »•* 1 t c_ * ♦ •» T-i < E ( ~ P 4 - LU ÜT ÜTÜT G T 4 O «T 0 3 < r X G t-4 T-i GLU T-i ÜT r o 0 4 o CL rs X T-i O 2 ai r H G -- - M) < T Cl at £ L ’tH 2 c_ o r G X OX G JZ OT CX) Ü T *?H X 0'4 X c- CI) Ü T 0 4 ü T O Ü rs fi) •sH P r-4 0 4 ÜT X 0 4 G •rt < - M 3 0 4 X t4 -r-l LL. G G ÜT •q- < 0 ü T 3 » X ÜT X C- (- % X CO ÜT S 3 M at rH XO c o P IS-P Ui at * •» * • o ai - P ♦ - ♦ . 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Table C22» Parametric MANOVA Showing the Effect of Nutrient Solution Strength on Four Variables (Leaflet Lensithf B1adders/Positions per Leaflet» ■D CD Leaflet Pairs per cm» Bladders/Positions per cm) Measured on jJ. vuldaris Plants Raised from Turions in a Diet Experiment C/) C/) Multivariate Tests of Significance CD Test Name Value Exact F Hypo th * DF Error DF Sid * of F 8 Pillais * 10006 3*11331 4,00 112*00 *018 Hotellinds *1.1,119 3*11331 4*00 112*00 *018 Wi Iks *89994 3*11331 4*00 112*00 *018 Roys *10006 Note * * F statistics are exact* 3. 3" CD ■DCD O Univariate F-tests with (1 » 115) D* F. Q. C a Variable Hypotli* SSError SS Hypoth, MS Error MS F Sid* of F 3O "O O Leaflet lendth *22862 10*20612 *22862 *08875 2*57605 * 111 CD Q. Bladders per leaflet 5*06025 154*90420 5*06025 1*34699 3*75670 * 055 ■D CD Leaflet C/) pairs per cm C/) *56287 38*54306 .56287 *33516 1*67941 * 198 Bladders PC r cm *01279 30241*2424 *01279 262*96732 *00005 *994 ro o 204 c-j ü i o -P «T O r-j c-4 04 •c O c S3 r-4 04 01 =4- in 04 fO r. -r-4 01 3 Oi >C- -M -+4 —1 O ÛC CD 04 r - r JO (- m in y—i T-» Ol U. C-4 l i . r - 4 S 3 M 3 Oi ÛL c O o o l i l LO r s S 3 Η 1 ct} o <4- O o o CJ SS r H CM r-l JQ « 5 r - t o * ♦ * r s i > r s S 3 t - nj Û. H M 3 T T < T o -rJ * no c- m 91 r s i - O l . -H 01 > c. cn -P -P nj c. & 9 1 r - l i-t 3 o o o c n 111 CM S 3 C-4 O q - 3 il a o o o z f s CM M 3 04 u , rc > CD S 3 i i l r s Z f- S 3 c-4 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 205 O en O C-i ro o ro 03 fs ro o o !—Î Ul o c 4- r-i Of nc {- 03 ro G 3 _J Cl ■t4 en dr c. ». 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Further reproduction prohibited without permission. 212 li U~i o O IX o m ro IX r-j o m itî es 3 4 - H fi ro f- m ro ro Û1 jQ _i roi i-i 1—i Ü3 C t- 3 Q 3 fi > M O ro ü_ o o o o m cx I—i ♦ fi ü_ o o o if! cx X o V 31 fe- o o o lil -i-i ro c ro O o X un t- o ro c C4 X o ro f i J C 151 m Reproduced with permission of the copyright owner. Further reproduction prohiPited without perntission. 213 u_ m to o o c 0 0 un o 0 ll O 3 CC 0 0 0 1—4 m tn JD JC •I-I -p -r-i ro to c c n -r-i a. m O (- I—i ro ■p ù_ l i . £ > t o - 4 3 l l M 3 r v u n O 0 CK CN O CNCD r v r v Ü t—1 cn 4 - O o o r v m i V 0 4 to c- 4- •M O * ■* * S j 0 4 0 4 3 to (- v 3 0 4 0 4 C N 0 O 0 0 ti__ i Oi 01 t o T—! c r-i - r - i H-i c 3 c n o •!> > IG 0 0 i n JC jC • l i . O o o c n !V C N u n -4J 4- ZZ if o o o z 0 4 O' r v IV to 0 ** ♦ « r o C N 4- c 0 z S D 0 4 " T f - CN 1 - 4 M 3 u n O 0 o o C i 0 4 0 4 o 0 4 u n CO r v # - t . C i 0 4 0 4 im. • T -T-> c- "3 Î- t o C N N u n u CO 0 0 m UJÜJ 'C < T C N O' 0 0. u n M 3 0 4 4- 3 • r - i 0 4 0 4 0 4 t o 4- G w cn (_ ‘ Ui r4 0 Reproduced with permission of the copyright ownec Furthor reproduction prohibited without permission. ■DCD O Q. C g Q. Table C33« Nonpar «met rie MANOVA Showirr.-( the Effect of the Interaction between ■D Experimental Seaaon» Feeding Fîesiiine and Nutrient Solution Strength CD on Four Variables (Leaflet Lendthy Bladders/Posi tions per Leaflet y C/) C/) Leaflet Pairs per cm y Bladders/Positions per cm) Measured on U. vuldaris Plants Raised from Turions in a Diet Experiment Multivariate Tests of' S:i.dnii'i.cnce (S M 1/ 2 t N 55 ) 8 Test Name Value Approx. F Hypoth. DF Error DF Sid. of F (O' F' i 11 a i s .05602 ♦81406 8.00 226♦00 ♦ 591 Hotellinds .05782 .00232 8.00 222.00 ♦ 601 Wilks .94468 .80820 8.00 224.00 ♦ 596 3. Roys .03739 3" CD Note.. F statistic for WILK'S Lambda is exact. ■DCD O Q. C a Univariate F-tests with (2,115) D. F. 3o "O Variable Hypoth. SS Error SS Hypoth. MS Error MS F Sid. of F o CD Leaflet Q. lendth 3142.54624 294778.841 1571.27312 2563.29427 .61299 .543 Bladders "O CD per leaflet 790.58724 304693.706 395.29362 2649.51049 .14919 .862 (/) (/) Leaflet pairs per cm 3994.40196 264484.102 1997.20098 2299.86175 .86840 .422 Bladders per cm 2079.30169 367512.367 1039.65084 3195.75971 .32532 .723 ho 4-. ■DCD O Q. C g Q. Figure Cl. Boxplots for U, vu.lgaris Plants Raised from Turions in a Diet Experiment: Stem Diameter by treatment Tray ■D CD 1.40 X E C/) KEY C/) 1 ! $ Median + -+-+ - 25%, 75% X X I 1 F-+-+ X Hieh/Low 1 +-+-+ 1 1 0 O u tlie r 1 » I 1 + -+-+ 1 1 E2 1 1 t-t-t +-+-♦ 1 » 1 E E x tr*# * +-+- 8 1 * 1 t - * - + 1 * 1 X X 1 1 1 I X me 1 1 + ' + - f t - + -+ X 1 $ I 1 1 F-+-+ I 1 « 1 1 +-+-+ +-+-+ ci' 1 X X +-+-+ +“+-+ 1 1 1 E + - + -F 1 1 1 X 1 X ! 1 y 1 « 1 X I » 1 1 1 1 1 1 1 +-+-+ ; t 1 3 F -+ -+ 3" t .10 X CD X X CD ■D eOLSTft 1 1 1 1 1 1 1 O FEED 1 1 1 1 1 2 Q. 2 2 3 3 C SEASON 1 2 3 3 1 1 1 2 2 1 ■ 1 a TRAY 11 16 2 2 1 6 12 17 23 3 3O 10 18 24 2 "O 1.40 O KEY X X CD $ Median Q. I I 0 - 25%, 73% X + ■+- + t X X Hiah/Low I I I X I I 0 O u tlie r F-+- + I I ==*- I I +-+-+ I E Extreme I I * I I I I I ! * I ! * I + -+-T ■D ! * I I + “+-+ I * I Key: CD +-+- f I I I I + “ +■ + -+ - + X f - l - i I I I « I f-+- + X I » I +-+-♦ + -+ -+ Solitr • Nutrient Solution Stfe")Wt C/) X C/) ! I X X 1 • one-tenth strength + - + - + X 2 • ful\ strength + -+- +-+'i ! * I « I Feed • Feeding Regime +-+-F 1 • no prey X 2 • 10 prey .10 J • 100 prey + - + - F Season ■ Cxperlmtnte) Season 1 > fall BOLSTR 2 2 2 2 • winter FEED 1 1 2 2 2 2 2 2 2 2 ro SEASON 1 2 3 3 3 3 F-» I 2 2 TRAY e 13 IV 1 1 2 tn 9 14 20 1 2 7 15 21 ■o I I Figure C2. Boxplots for U. vulgaris Plants Raised from Turions in a Diet Experiment: Leaflet Length by Treatment Tray X 2.»» 1 X 4 -4 -4 ■o KEY I CD I I t - + “ t 4-4 X t I I • Median I t (/) I I t X œ - 25%, 75% I « t X I o' t-i“+ E + - + - t X 1 I * I 3 % Hiah/Low 4 -4 -4 4 -4 -4 0 ü u tU a r I X X X I + O I I I I E Extra## I +-♦-+ I I « I I I I I X X I « I I +-*-+ I + -*-+ I « I CD I « I 4 -4 -4 4 -4 ' +-+-+ I +-+-+ I I I c# I « t 1 8 +-+“+ X X I ! 4 -4 -4 I 4 -4 -4 I I 0 +-E- + Q + -+ ” + 4-4 -4 X (S' I * I 1 I » I I X I ! I t I I I 4 -4 -4 I I t + -+- + X +-+-+ X I « I 4 -4 -4 X I X ,78 a 3" CD 1 1 2 SOLSTR 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 2 2 3 3 3 3 CD FEED 1 "O SEASON 1 1 2 1 1 2 2 1 1 2 2 1 O TRAY 4 11 16 22 6 12 17 23 3 10 18 24 2 Q.C a o 3 2,99 "D O KEY X • Median 1 X CD I X X Û. - 25%, 75% X ! I I 4-*-4 4 -4 -4 I X X HlflM/Uflw 4 -4 -4 0 O u tlie r 4 -4 -4 4 -4 -4 I I I * i 4 -4 -4 4-4-4 Key: I I E Extreme I I I I I I I I « I X I « I O 4 -4-4 I I C i • i 1 t 4-4-4 4 -4-4 I I T I Solslr • Nutrient Solution Strenjtit ! « I "O c# 4 -4 -+ I * I X I I I I I I I 1 • onc'tenth ttrengch 4 -4 -4 CD X I I I 4 -4-4 I * I I I Î • full ttrength 4-4-4 X I 4 -4-4 4 -4-4 X C/Î 4 -4-4 X X Feed ■ Feeding Regime C/) X 4 -4 -4 I I 1 • no prey o' I $ I 3 ; » I 4 -4 -4 2 " 1 0 prey I I I I 4-4-4 3 • 100 B'ey X 4 -* -4 4-4-4 I . X Seaton « Experimental Seaton X .78 1 - fall 2 • winter S0L8TR 2 2 2 2 2 2 2 2 2 2 2 FEED 1 1 1 2 2 2 2 3 3 3 3 1 1 2 2 ro SEASON 1 2 2 1 1 2 2 I—> TkAY 8 13 19 5 9 14 20 1 7 15 21 a t ■o o Q. C 8 Figure C3, Boxplots for U. vulgaris Plants Raised from Turions in a Diet Experiment: 81 adders/Positions Q. per Leaflet by Treatment Tray ■D 16 CD KEY X C/) C/) X I • Median % X I I E I - 25%, 75% + -+-■+■ I X HidM/Lou 0 +-+“+ 0 O u tlie r I I I I +-+-+ I I E EMtrem* +“+“+ 0 I * I £ +-+-♦ I » I X X X I I 0 I I 8 +-+-+ ♦-+-+ I I I I * I + -+ -+ I « I I I 1 +-+"+ I « I +-+-•+ X +-+-+ X X I I t * I +-+-+ I » I I I ♦-»-+ +-*-+ X +- i-+ I I t * I I t +-+-+ X +-+-+ I +•+“+ I I + -+ - + I X X I +“ +•+ X I X CD I X X 3. 3" CD SOLSTR 1 1 1 1 1 1 1 1 1 1 1 1 2 CD "O FEED 1 1 1 1 2 2 2 2 3 3 3 3 1 O SEASON 1 1 2 1 1 2 2 1 1 2 2 1 Q. TRAY 4 11 16 22 6 12 17 23 3 10 IB 24 2 3o ■o 16 o KEY CD « Median Q. - 25%, 75% +'+■ X X X Hiwn/Low I « X +-+• + +-+-+ +-+“+ 0 O u tlie r X I +■+-+ I I « I X I I I * I Key: E Extreme I I I I * I I I ♦ -+ “ + t « I +-+-+ I « I I I + "+-■t Solslr Nutrient Solution StrenjUi I t +-+-+ X I I I I "8 1 ■ one*tenth strength ! « I X +-+-+ ♦-+“+ X + -+•+ 2 • full strength +-+- + +-+-+ I I * 1 i*+ +-*-+ (/) I I I X I I I I w Feed • Feeding Regime o' X I * I I I + •+ '+ I • no prey 3 I I + -+-+ I 1 ■ 10 prey 1 I 1 X +-+-+ X 3 " 100 prey Season - Experimental Season 1 • fall 2 ■ winter T SOLSTR 2 2 2 2 2 2 2 2 2 3 FEED 1 1 2 2 2 2 3 3 3 ro SEASON 2 2 1 1 2 2 1 1 2 2 r-* TRAY 13 17 5 9 14 20 1 7 IS 21 *»J :x) ■O(D O D. C Figure C4, Boxplots for U. vulgaris Plants Raised from Turions in a Diet Experiment: Length of Plant by g Q. Treatment Tray loi.eo titr (/) o' t Median 3 - 23%, 75% X M M h / L o u X 0 O u tlitr I E E N tr*** t +-+- 8 cm E 1 X 1 1 +-+-I +“+-+ 0 ! 1 1 S' 1 1 1 1 +-+-+ X 1 1 X 1 1 t * 1 ! ! 1 +-+-+ 1 1 X ! 1 1 X 1 1 X 1 * 1 1 1 1 1 a 1 +-+-+ 3 +-+-I 1 a 1 +-+“+ +“+-+ + -+-+ 1 1 +-+-+ 1 a 1 +-+-+ 1 a I CD 1 1 + -+“♦ I * I X 1 « 1 + -+-+ 1 1 +■+-+ f-+-+ 1 +-a-+ 1 « I X +-+-+ + -+- + X 1 « 1 X 1 a 1 X + -+-+ X +-+"+ 1 1 E C +-+-+ X +-T-+ +-+-+ a P- 4,20 +-+-+ 3 " CD 2 CD SOLSTR 1 1 1 1 1 1 1 1 1 1 1 1 ■o 1 1 2 2 2 2 3 3 3 3 1 O FEED 1 1 1 Q. SEASON I 1 2 2 1 1 2 2 1 1 2 2 c TRAY 4 11 16 22 6 12 17 23 3 10 18 24 2 Q. ■o lot.BO o KEY CD a Median Q. - 25%. 75% X Hlah/Low X X ! 0 O u tlie r 1 X £ Ktf. 4 -4 -4 E Extreme E 1 1 1 1 1 1 Solttr • Nutrient Solution Strenjtik T 3 1 1 CD cm X 4 -4 -4 4 -4 -4 1 • one-temth itrength 1 1 a 1 1 1 I 1 2 • full etremgth 1 1 1 1 a 1 1 a 1 C/) 4 -4 -4 (/) ♦ -+-F X 1 1 1 1 X t 1 Feed « Feeding Regime 1 a 1 X 1 4 -4 -4 1 1 4 -4 -4 1 1 1 1 1 « no prey 4 -4 -4 1 a 1 +-+-+ + -+-+ 4 -+ “ + 1 4 -4 -4 1 1 4 -4 -4 2 « 1 0 prey 1 1 1 1 1 1 a 1 X 1 X 1 a 1 4 -4 -4 X I 1 3 • 100 prey X 1 a 1 4 -4-4 X 4 -4 -4 4 -4 -4 1 a 1 1 a 1 4 -4 -4 +-+“+ X 1 a 1 X 4 -4 -4 4 -4 -4 Seejon » Experimental Seaion X 4.20 X 4 -4-4 X I « fall 1 « winter 2 2 2 SOLSTR 2 2 2 . 2 2 2 2 2 3 3 FEED 1 1 1 2 2 2 2 3 3 ro 1 2 2 SEASON 1 2 2 1 I 2 2 1 »—• 15 21 00 TRAY 8 13 19 5 9 14 20 1 7 i - 219 O) CL (O CL d> O X L « -I X I I CL X C41*1 n t n c o *r— -4 f4p, C 4 B I i "O X 4- » 4- 01 I I iO ûd I 4- X I I I * X «A ; I A3 CL i n A3 I I ^ C 4 C 4 4- * 4- — X cn| C4 I I 3 +-> > _ 0 = > | 5 ri ^ ri ^ â l r I O X 4^ » 4- X 4 2 ^ I I CL C I I X 1 M 3 X 3 in 1 a 9 V) O 9 CJ> 1 c -1 9 9 c Ps «J 9 # 1 m Nb 9 £ S 9 9 £ g N >- 1 X tA Q CA >• r* X W Q tA O UI 1 *o X m *# UI « « o "O X a -I UI 'T « B i 9 lA 3 X O UI u eg 9 l/ l X O UI UJ >K 3 1 X. C 4 X O w (A U. UI I- X ri X a Ui UI IJL tA h- 1 1 X o Ui # 1 X o UI Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 220 s_ 0) CL I/I s _ r . rtJ 1 Q_ ;L i + J 0 ) I l f I X 2 u M- c « Æ £ JÏ S. s ^ 1 1 1 _J ii £ 0 0 0 X + * '+' X 04 *4 04 g • • V — + 2 — 7 ■ ■ • III • • 4 ^ t C 1 4 ■o 1 X * ^ — X ^ r» 04 ♦ i i 1 CM S ♦ Hk 1 .r J s- û} I I _ CL -f. X X M Ci CD LU * ^ >e W r t rv in C O ir» S- X — 4- Z 5 e X X N M Ci o ^ CN r< fs. 04 -o O) I f < o ■ » CL: I I ( n -M i I c X 4- ♦ X fO 1 ( in 04 *4 11*! *f— I I s. w X 4' * 4. X «5 t I >>«3 3 5L. 04 «4 04 »■ > H“ X — 4- # • 4------y 4-* 3 |OJ c S- E 04 «4 04 n t f x - - f o 4^ X 4 - * X 4 - (O Q) V) S- + > #— O 04 -4 -N œ I CL_a »‘ î X o E CD u X 3 VIO Oi tf 1 M 3 C N _J » fi k* o 1 40 o &. • X X 9# m Ui O 1 C N • i % c k - o X b. >■ o ^ Ui « « \ o 1 m 'S " 4 « ( A A t n •O K «1 O UI w Û£ 1 » j : ^ _ i U I « « 3 X 04 O) u. UÜ e- CJ « -9» •*» & 40 1 *0 *4 o U I U I ÛC X 04 X O UI I & 4 0 ^ 3 X ( D U . u ) w fs. 1 r 04 X O m 1 X o ui §: 1 * I X o UI g . Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 73 CD "O 0 Boxplots for U. M b t H Plants Raised from Turions in , Diet Experiment-. Bladders/Positions Q. Figure C7 1 per Plant by Treatment Tray 6926 % hEt (/) t Median - 25%, ?5X X o" X Hi9h/L0M 3 I 0 O u tlie r E Extreme I 1 3- X CD X I I +-T-+ I 8 E- + -E I I ( +* + “ + X 5 + -E-E +-+-+ I I ! +-+-+ 1 $ +-■ + -+ I $ I I I â- I 1 I I * I I 3 +- + - + I I I I I + - + - + * I I I + -+ - + +- + -E I * I I I I 1 * 1 +-+ I « I X X +-+-T X + -*- + + - + “ + X t -*-E ! I 3 I +-+-+ +-T- 1 + -* - + CD X X X X 372 C 1 1 1 2 3. 1 1 1 I t 1 1 3 1 SOLSTR 1 3 3 3 2 2 2 2 1 1 1 1 2 2 1 FEED 1 2 2 1 CD 2 1 18 24 2 SEASON 1 17 23 3 10 -O 16 22 6 12 TRAY 11 IC a o 3 ■O O 6926 MEY CD a. * Median X - 25%, 75% I Kay: 3 X Hlflh/Low O 0 Outlier I Solstr ■ Nutrient Solution StrengO» E Extreme ■ D X X 1 • one-tenth strength } CD 2 ■ full strength + -+ -+ 1 C/î 1 I + -+ ” + X feed • Feeding Regime (g E-+-4 E X 1 * t 1 * 1 1 + -+ -+ 1 ■ no prey O* I I X 3 +-E- + +-+-+ 1 1 I * I 2 "1 0 prey +-+-+ +"*•6 : I + -+ - + +-+--V , 1 1 1 I I ) • 100 prey I » ! X X I « I + - + -+ 1 * 1 X X X +-+-+ + -* -+ +-+-+ t~+“ E X Season " Experimental Season X 1 * 1 +-+-+ X + -+ - + X 1 - fall X 372 2 • winter 2 ro 2 2 . 2 ro 2 2 2 3 SOLSTR 3 2 2 3 1 1 7 2 1 ;; 2 FEED 1 1 2 2 2 I 7 15 21 SEASON 1 o < 20 19 5 1 TRAY 8 13 ■o Figure C8. Boxplots for U, vulgaris Plants Raised from Turions in a Diet Experiment: Bladders/Positions I per cm by Treatment Tray I 160.39 E KEY E ■g * Median X - 25%, 75% 1 c/) X Hish/Low E 1 (g 0 + - f - 1 o' 0 O u tlie r 1 1 E 3 E Extreme 1 1 1 1 1 1 + -+ -+ 1 I 1 1 + - + -f 1 « 1 1 1 3" CD 1 « 1 +-+-+ X X 1 * ! f - + -+ X 1 « 1 X 1 1 X 1 * 1 1 X I + -+-+ X 1 +-F -+ 1 8 1 1 1 +-+-+ 1 + -+-F + - + -+ + -*-+ +-+-+ X X + -+ - + 5 +-+-+ +-*-+ X +-+-+ ! 1 : » I 1 1 E 1 * 1 I 1 c5' 1 1 1 4 1 1 « 1 I--+-1 0 1 1 +“+“ + 1 * 1 1 « I +-+-+ 1 I + * + -+ ! 1 ! ♦-+-+1 1 1 +-+-F 1 X +-+-+ 0 I +-+-F 3 X X XXX CD 40,57 E C p. SOLSTR I 1 1 1 1 1 I 1 1 1 I 1 2 3" FEED 1 1 1 1 2 2 2 2 3 3 3 3 CD 1 SEASON 1 1 2 2 1 1 2 2 1 1 2 2 I TRAY 4 ■OCD 11 16 22 6 12 17 23 3 10 18 24 2 O Q. C 160.35 a G KEY X 3 ■ D * Median O - 25%, 75% X Hldh/Lou X 0 O u tlie r t CD X +-+ Key: Q. E Extreme I I I I +-+- 1 1 Solstr • Nutrient Solution SirenjtJi + - * - + X X 1 I 1 • one-tenth strength + -+ -+ 1 ! ! 2 • full strength "O 1 t + -+ - + +“+-+ + - + CD X X 1 I 1 * I 0 1 * Feed ■ Feeding Regime + -+-+ +“+“ + + -+ -+ 1 1 1 1 + -+ 1 • no prey I I * I +-+■ Î 2 ■ 10 prey C/) 1 1 » 1 1 « 1 1 1 + - * - + X C/) + -»- + 1 t * 1 f - + - 1 +-+-+ +-+-+ + ~ t“ t 3 • 100 prey I + “ +” + ! E X X X E 0 Season • Experimental Season 40,57 1 ■ fall 2 • winter SOLSTR 2 2 2 2 2 2 2 2 2 2 FEED 1 1 1 2 2 2 2 3 3 % SEASON 1 2 PO 2 1 1 2 2 I I p ro TRAY B 13 IP 5 V 14 20 1 7 2 \ PO 223 Figure C9. Profile Plot for the Interaction of Collection Site and Feeding Regime on Bladders per cm for vulgaris Plants Raised from Turions in a Diet Experiment* K e y : Tykeson's Pond - McWenneger's Slough-— Bladders per cm 0.1 strength nutrient solution full strength nutrient solution 100 Feeding Reg ime *Based on Trays 13-24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 224 Figure CIO. Profile Plot for the Interaction of Experimental Season and Feeding Regime on Key Variables for U^. vulgar t s Plants Raised from Turions in a Diet Experiment* Key : fall experimental season - — -- winter experimental season ----- Variable: 0.1 ful 1 Nutrient Solution Strength Leaflet length Leaflet pairs per cm Bladders per leaf 1 et Bladders per cm 0 10 100 0 10 100 Feeding Reg ime *Based on Trays 7-12 and 19-24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 225 Figure Cil. Profile Plot for the Interaction of Experimental Season and Nutrient Solution Strength on Key Variables for U. vu 1ga r i s Plants Raised from Turions in a Diet Experiment* Key fall experimental season ------winter experimental season ---- 0 10 100 Feed i ng Variable: Regime Leaflet length Leaflet pairs per cm Bladders per 1eaf1 et Bladders per cm 0.1 full 0.1 full 0.1 full Nutrient Solution Strength *Based on Trays 7-12 and 19-24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 226 Figure C12 Profile Plot for the Interaction of Feeding Regime and Nutrient Solution Strength on Key Variables for IJ. vulgaris Plants Raised from Turions in a Diet Experiment* Key : 0.1 strength nutrient solution ------full strength nutrient solution ----- Variable; fall winter Experimental Season Leaflet length Leaflet pairs per cm Bladders per leaflet 4k / / \ \ / \ Bladders per cm / / 0 10 100 1 10 100 Feed i ng Reg ime *Based on Trays 7-12 and 19-24 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. m IL 01 01 X w + + '—1 ÜJ -# 1 I 3 cn X — + * + — 3 c 1 I U c + + nj ■3 o L #-# W Ti a u + + I 1 3 c X **“ — + + ÜJ C'4 a o I t H 5 o + + q u o 3 U X + + L 1 I o 4-> + * + — Lü u . c QU 1 I i-i + + in (*- 4_> in nj o c OJ eu o 3 0. * W 1 X 3 1 i n o [_ OJ nj X t - t - u o 3 OJ I c n . _J Oj 5 CL 1 m Oi n > 1 - x: *—t {. to tu I T-t Ü J 1 m *-» 0. ' W OJ , X: 1 OJ in X U 1 e t n t i : C'J X o W 0 0 i * 1 X o UJ 01 u 3 m