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1988

Resource allocation to prey capture tissue in the aquatic carnivorous 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. 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

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problem of control of bladder production.

While much work has been done on carnivorous plant

, 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 , which

includes other carnivorous plant genera. The

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 , 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

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 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 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

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 (-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

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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%

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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

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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.

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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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Johnson, W.T. 1987. Bladderwort, Arizona'a carnivorous wildflower. Desert Plants 8(3):140-141.

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Perioden der Austrocknung. Flora 162:269-283

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Winston, R.D. and P.R. Gorham. 1979a. Tarions and dormancy states in Utricularia vulgaris. Can. J. Bot. 57:2740-2749.

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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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* 1 X o w 01 c. 3 <-c01 IL

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Appendix B. Data Summaries, Complete MANOVA Results, Boxplots, and

Profile Plots for Utricularia vulgaris Plants Raised from

Jurions in a Common Garden Experiment

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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

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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

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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

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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

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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

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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

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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

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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

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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

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Appendix C- Data Summaries, Complete MANOVA Results, Boxplots, and

Profile Plots for Utricularia vulgaris Plants Raised from

Turions in a Diet Experiment

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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

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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

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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

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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

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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

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Ta ble C 2 ♦ Data Sun,maries for Ut.ricularia vulsiacis Pl 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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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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

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 7 3 ■DCD O Q. C g Q. 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

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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

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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.