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Investigating the Determinants of Local Scale Distribution of Ruellia tweediana (synonym R. brittoniana) in Natural Areas

Permanent Link: http://ufdc.ufl.edu/UFE0021595/00001

Material Information

Title: Investigating the Determinants of Local Scale Distribution of Ruellia tweediana (synonym R. brittoniana) in Natural Areas
Physical Description: 1 online resource (108 p.)
Language: english
Creator: Hupp, Karen V Shepherd
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: Agronomy -- Dissertations, Academic -- UF
Genre: Agronomy thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Ruellia tweediana is a non-native plant introduced to Florida prior to 1940 that has naturalized in disturbed uplands and wetlands of 8 states, the Virgin Islands, and Puerto Rico. A popular horticultural plant, R. tweediana is available in many nurseries across Florida, being very attractive to the homeowner because of its ability to grow in many environmental conditions, from a water plant to growing in well-drained, almost xeric conditions. Observations in natural areas suggest that the field distribution of R. tweediana was limited to a narrow band along the banks of waterways (a Ruellia zone between Upland and Submerged zones). The principle goal of this research was to determine why the natural area distribution of R. tweediana was limited to a narrow zone along the banks of waterways rather than in a broader distribution that includes both wetter and drier adjacent habitats. This research investigated whether 1): the observed distribution of R. tweediana could be explained by limitations due to environmental conditions: soil moisture, light, and soil temperature, and/or by the biotic interactions of competition with native and non-native plants; and 2) at which life-stage, seeds or seedlings, these limitations were most influential. In the seed burial study, the possibility of a persistent seed bank was found in the Upland and Ruellia zones while a seed limitation was found in the Submerged zone due to quick reduction in seed viability over the study period. In the seed germination study, R. tweediana seeds germinated in all tested soil types at very high percentages. In the field, germination studies provided differing results. In the first two experiments the highest germination percentages occurred in Ruellia plots where competition did not have an effect, and in the third experiment zone (Upland or Ruellia) did not matter while competition did, with cleared plots having higher germination. When R. tweediana seedlings were transplanted, they survived once established, irrespective of competition or zone treatment. The highest survival occurred in the cleared plots regardless of seedling size or location of zone. From these experiments it was concluded that there was a seed and microhabitat limitation in the Upland and Ruellia Zones, that once seeds are present they are capable of germinating and becoming juvenile seedlings, and when seedlings are present they are capable of surviving once established.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Karen V Shepherd Hupp.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Fox, Alison M.
Local: Co-adviser: Stocker, Randall K.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021595:00001

Permanent Link: http://ufdc.ufl.edu/UFE0021595/00001

Material Information

Title: Investigating the Determinants of Local Scale Distribution of Ruellia tweediana (synonym R. brittoniana) in Natural Areas
Physical Description: 1 online resource (108 p.)
Language: english
Creator: Hupp, Karen V Shepherd
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2007

Subjects

Subjects / Keywords: Agronomy -- Dissertations, Academic -- UF
Genre: Agronomy thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Ruellia tweediana is a non-native plant introduced to Florida prior to 1940 that has naturalized in disturbed uplands and wetlands of 8 states, the Virgin Islands, and Puerto Rico. A popular horticultural plant, R. tweediana is available in many nurseries across Florida, being very attractive to the homeowner because of its ability to grow in many environmental conditions, from a water plant to growing in well-drained, almost xeric conditions. Observations in natural areas suggest that the field distribution of R. tweediana was limited to a narrow band along the banks of waterways (a Ruellia zone between Upland and Submerged zones). The principle goal of this research was to determine why the natural area distribution of R. tweediana was limited to a narrow zone along the banks of waterways rather than in a broader distribution that includes both wetter and drier adjacent habitats. This research investigated whether 1): the observed distribution of R. tweediana could be explained by limitations due to environmental conditions: soil moisture, light, and soil temperature, and/or by the biotic interactions of competition with native and non-native plants; and 2) at which life-stage, seeds or seedlings, these limitations were most influential. In the seed burial study, the possibility of a persistent seed bank was found in the Upland and Ruellia zones while a seed limitation was found in the Submerged zone due to quick reduction in seed viability over the study period. In the seed germination study, R. tweediana seeds germinated in all tested soil types at very high percentages. In the field, germination studies provided differing results. In the first two experiments the highest germination percentages occurred in Ruellia plots where competition did not have an effect, and in the third experiment zone (Upland or Ruellia) did not matter while competition did, with cleared plots having higher germination. When R. tweediana seedlings were transplanted, they survived once established, irrespective of competition or zone treatment. The highest survival occurred in the cleared plots regardless of seedling size or location of zone. From these experiments it was concluded that there was a seed and microhabitat limitation in the Upland and Ruellia Zones, that once seeds are present they are capable of germinating and becoming juvenile seedlings, and when seedlings are present they are capable of surviving once established.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility: by Karen V Shepherd Hupp.
Thesis: Thesis (M.S.)--University of Florida, 2007.
Local: Adviser: Fox, Alison M.
Local: Co-adviser: Stocker, Randall K.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2007
System ID: UFE0021595:00001


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1e019b9c9d4f22f5bf40384dd953134cd2f2ee37







INVESTIGATING THE DETERMINANTS OF LOCAL SCALE DISTRIBUTION OF
RUELLIA TWEEDIANA (SYNONYM R. BRITTONIANA) IN NATURAL AREAS.




















By

KAREN VICTORIA SHEPHERD HIUPP


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2007

































O 2007 Karen Victoria Shepherd Hupp





























To my parents who always knew I had it in me and
to my husband who was there through the stressful times of this degree.









ACKNOWLEDGMENTS

I thank the members of my supervisory committee for their ideas, time, and mentoring. I

thank Alison Fox, for encouraging me to take this path in my life and making it actually possible

for me, and for help every step of the way. I thank Randall Stocker, for being extremely available

for questions and meetings and opinions. I thank Dr. Ramon Littell, for the many helpful

meetings to help me understand the statistical aspects of my thesis. I thank Sandy Wilson, for the

interest she carries with this species. I thank my family for their support, which kept me going to

complete this degree. To Jenn and Denise who were my cheerleaders. And this degree would not

have been possible without the help and friendship of Lisa Huey.












TABLE OF CONTENTS


page

ACKNOWLEDGMENTS .............. ...............4.....


LIST OF TABLES ................ ...............7............ ....


LIST OF FIGURES .............. ...............8.....


AB S TRAC T ............._. .......... ..............._ 1 1..


CHAPTER


1 INTRODUCTION ................. ...............13.......... ......


Study Site and Z ones .............. ...............17....
Research Questions............... ...............1


2 SEED VIABILITY INT SUBMERGED CONDITIONS AND SOIL EFFECTS ON
SEED GERMINATION ................. ...............27.......... .....


Introducti on ................. ...............27.................

Hypotheses............... ...... .. .. .. .......2
Seed Viability in Submerged Conditions .............. ...............28....
Soil .................. ........... ...............28.......
Materials and Methods ................ .. ....... ..............2

Seed Viability in Submerged Conditions .............. ...............29....
Soil ................ .......... ........... .............2

Soil effects on seed germination .............. ...............29....
S oil character sti cs............... ............3

Statistical Analyses............... ...............30
R e sults................... ............ .. .. ... ...... ............3

Seed Viability in Submerged Conditions ................. ...............31........... ...
Soil ................ .......... ........... .............3

Soil effects on seed germination .............. ...............31....
Soil characteristics............... ............3
D discussion ................ ...... ....... .. ........... .............3
Seed Viability in Submerged Conditions .............. ...............32....
Soil ................ .......... ........... .............3

Soil effects on seed germination .............. ...............33....
Soil characteristics............... ............3


3 SEED BURIAL .............. ...............41....


Introducti on ................. ...............41.................

H ypotheses................. ..............4
Materials and Methods .............. ...............43....












Study Site and Zone Characterization .............. ...............43....
Seed Burial Study ................. ...............45................
Statistical Analyses............... ...............46
Re sults............... .. ........... ...............46.......
Zone Characterization .............. ...............46....
Seed Burial Study ................. ...............47................
Discussion ................. ............ ...............48.......
Zone Characterization .............. ...............48....

Seed Burial Study ................. ...............49................


4 FIELD STUDY ................. ...............61........... ....


Introducti on ................. ...............61.................

Hypotheses............... .. ... ...............6
Seed Germination and Survival Experiments............... ..............6
Seedling Transplant Experiment .............. ...............62....
Material and Methods ................. ................. .. ...............63.....

Study Site and General Plot Establishment ................. ...............63...............
Seed Germination and Survival Experiments............... ..............6
Seed germination................ ...............6
Seed germination and seedling survival ................. ...............64........... ...
Seedling Transplant Experiment .............. ...............65....
Statistical Analyses............... ...............66
R e sults................ ... .. ....... .......... .. ..........6

Seed Germination and Survival Experiments............... ..............6
Seed germination................ ...............6
Seed germination and seedling survival ................. ...............68........... ...
Seedling Transplant Experiment .............. ...............68....
Discussion ................. ...............71.................
Conclusions............... ..............7


APPENDIX


A SPECIES LIST .............. ...............91....


B SPECIES BIOMASS ............. ...... ...............101..


LIST OF REFERENCES ............_...... ...............104...


BIOGRAPHICAL SKETCH ............_...... ...............108...










LIST OF TABLES


Table page

1-1 Florida counties in which Ruellia tweedian2a occurs .............. ...............20....

1-2 List of reported natural areas in which Ruellia tweediana occurs Florida. .....................21

1-3 List of conservation areas in which Ruellia tweediana occurs in South Florida .............22

1-4 Dry above ground biomass data for 1 m2 plOt, Ruellia weight is R. tweediana and
Others weight all other species in plot. Ruellia count number of R. tweediana
stems found in 1 m2 plOt. ............. ...............26.....

2-1 Soil analyses results for 10 replicates from Upland, Ruellia, and Submerged zones........37

2-2 Soil survey results of Ponoma Sand, depressional from Natural Resources
Conservation Service and United States Department of Agriculture .............. .... ........._..40

3-1 ANOVA table for percent missing of seed total. .............. ...............55....

3-2 ANOVA table for percent dead of seed total ................. ...............56........... .

3-3 ANOVA table for percent germinated of seed total ................ .......... ................. 57

3-4 ANOVA table for percent dormant of seed total ................ ...............58........... .

4-1 Means of covariates for Seed germination in field experiments ................. ................ ..83

4-2 ANOVA table for seedling Survival over 3.5 months in Seedling Transplant Study .......86

4-3 ANOVA table for Increase in Height over 3.5 months in Seedling Transplant Study......87

4-4 ANOVA table for Shoot Biomass in Seedling Transplant Study ................. ..................90

4-5 ANOVA table for Root Biomass in Seedling Transplant Study ................. ................ ..90










LIST OF FIGURES


Figure page

1-1 Map showing Gainesville creeks. Dark circle in the enlarged insert is the study site in
Payne' s Prairie Preserve State Park ................. ...............23...............

1-2 Photo on left is aligned with diagram on right to indicate the labeling of the three
zones, Upland, Ruellia, and Submerged, in the Hield taken from the Submerged zone.....24

1-3 The edge between the Upland zone and Ruellia zone, from the perspective of the
Upland zone, at Payne' s Prairie Preserve State Park, (January 2006). .............. ..... ..........25

2-1 Percent germination and percent viability of seeds over time in the Seed Viability in
Submerged Conditions study. The percent germinated of seeds remaining at each
time is represented in black and the percent viable of seeds tested each time is
represented in grey. .............. ...............35....

2-2 Number of germinated seeds in the "Soil effect on seed germination" experiments.
Experiment 1 seed germination is presented in the hash marked bars. Experiment 2
is presented with black for seeds on top of the soil and dark grey for seeds mixed in
the soil............... ...............36..

2-3 Soil parameters with different letters indicated significant differences among zones
within parameters (a of 0.05), n=10. ............. ...............38.....

2-4 Natural Resources Conservation Service and United States Department of
Agriculture map of soil series superimposed over an aerial photograph of a section of
Payne's Prairie Preserve State Park. Soil type 25 indicates Pomona sand,
depressional the area in which this Hield research was conducted .................. ...............39

3-1 Nylon mesh seeds bags attached to the bottom of surveyor' s flags with zip ties. .............52

3-2 Seed burial plot, closest flags in Submerged zone, center flags in Ruellia zone,
background flags in Upland zone. ............. ...............53.....

3-3 Average percent soil moisture for each zone February 2006 to March 2007. There
were significant differences between zones at each sampling interval and averaged
across tim e. ............. ...............54.....

3-4 Percent missing of seed total in each zone. U= Upland zone, R= Ruellia zone, and S=
Submerged zone. Different letters indicate significant difference between months
after burial (a of 0.05) for % missing averaged over all zones. Line fitted to averages
with an R2 = 0.93. ............. ...............55.....

3-5 Percent dead of seed total in each zone. U= Upland zone, R= Ruellia zone, and S=
Submerged zone. Percent dead were significantly higher at each month after burial in
the Submerged zone compared to the Upland and Ruellia zones. .................. ...............56










3-6 Percent germinated of seed total in each zone. U= Upland zone, R= Ruellia zone, and
S= Submerged zone. ............. ...............57.....

3-7 Percentages of total seed (50) that were recovered from the field that were 1)
missing, 2) dead, 3) germinated, or 4) viable but presumed dormant. The Zones are
represented by U= Upland R= Ruellia, S= Submerged. .................. ................5

3-8 Mean percent viable seed for 10 replicate samples per zone ................. ......._._.......60

4-1 Plot layout map for Seed Germination and Survival Experiments and Seedling
Transplant Experiment at Paynes Prairie Preserve State Park ................. ........._._.....75

4-2 Example of one set of paired plots in the Seedling Transplant Experiment with
competition factor applied. Left side of photograph = Not-Cleared treatment, right
side = Cleared treatment. This is within the Ruellia treatment, planted with a Small
seedling treatment. .............. ...............76....

4-4 Example of a Seed Germination Experiment plot within a Ruellia treatment and with
the Cleared treatment applied. ............. ...............78.....

4-5 Example of one replicate site plot layout for the Seedling Transplant Experiment. In
this split-split plot experimental design. Zone is the main-plot factor, Competition is
the sub-plot factor, and Size is the sub-sub-plot factor. ................ ....___ ..............79

4-6 Example of a plot from the Seedling Transplant Experiment with Upland, Cleared,
and Large seedling treatments. ............. ...............80.....

4-7 Example of a plot from the Seedling Transplant Experiment with Upland, Not-
cleared, and Small seedling treatments. .............. ...............81....

4-8 Percent germination rates for the first experiment of the Seed Germination Study. ....._...82

4-9 Percent germination rates for the second experiment of the Seed Germination Study. ....83

4-10 Percent seed germination rates over three months in the Seed Germination and
Seedling Survival experiment. .............. ...............84....

4-11 Average percent survival over three months in the Seed Germination and Seedling
Survival experiment. .............. ...............85....

4-12 Seedling Transplant Experiment two-factor interactions of seedling Survival. ................86

4-13 Transplant study two-factor interactions of seedling change in height over 3.5
m onth s ................ ...............87........... ....

4-14 Seedling Transplant Experiment average shoot biomass data. .............. ....................8

4-15 Seedling Transplant Experiment average root biomass data. .............. ....................8









LIST OF ABBREVIATIONS


U Upland Zone

R Ruellia Zone

S Submerged Zone



L Large plant treatment

SM Small plant treatment

SE Seed germination and survival experiments



C Cleared plot treatment

NC Not Cleared plot treatment









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of
Master of Science

INVESTIGATING THE DETERMINANTS OF LOCAL SCALE DISTRIBUTION OF
RUELLIA TWEEDIANA (SYNONYM R. BRITTONIANA) IN NATURAL AREAS.

By

Karen Victoria Shepherd Hupp

December 2007

Chair: Alison Fox
Major: Agronomy

Ruellia tweedian2a is a non-native plant introduced to Florida prior to 1940 that has

naturalized in disturbed uplands and wetlands of 8 states, the Virgin Islands, and Puerto Rico. A

popular horticultural plant, R. tweediana is available in many nurseries across Florida, being very

attractive to the homeowner because of its ability to grow in many environmental conditions,

from a water plant to growing in well-drained, almost xeric conditions. Observations in natural

areas suggest that the field distribution ofR. tweediana was limited to a narrow band along the

banks of waterways (a Ruellia zone between Upland and Submerged zones). The principle goal

of this research was to determine why the natural area distribution of R. tweediana was limited to

a narrow zone along the banks of waterways rather than in a broader distribution that includes

both wetter and drier adj acent habitats. This research investigated whether 1): the observed

distribution of R. tweediana could be explained by limitations due to environmental conditions:

soil moisture, light, and soil temperature, and/or by the biotic interactions of competition with

native and non-native plants; and 2) at which life-stage, seeds or seedlings, these limitations were

most influential. In the seed burial study, the possibility of a persistent seed bank was found in

the Upland and Ruellia zones while a seed limitation was found in the Submerged zone due to










quick reduction in seed viability over the study period. In the seed germination study, R.

tweediana seeds germinated in all tested soil types at very high percentages. In the field,

germination studies provided differing results. In the first two experiments the highest

germination percentages occurred in Ruellia plots where competition did not have an effect, and

in the third experiment zone (Upland or Ruellia) did not matter while competition did, with

cleared plots having higher germination. When R. tweediana seedlings were transplanted, they

survived once established, irrespective of competition or zone treatment. The highest survival

occurred in the cleared plots regardless of seedling size or location of zone. From these

experiments it was concluded that there was a seed and microhabitat limitation in the Upland and

Ruellia Zones, that once seeds are present they are capable of germinating and becoming

juvenile seedlings, and when seedlings are present they are capable of surviving once

established.









CHAPTER 1
INTTRODUCTION

The movement of plant species by humans from one continent to another has increased

over time with the expansion of transportation and commerce (D'Antonio and Vitousek 1992).

Both the frequency and number of species moved has increased (Mack et al. 2000). As these

plant species are brought to new areas they may have characteristics that help them displace

native species: effective reproduction and dispersal mechanisms needed for the new area,

superior competitive ability, few to no herbivores or pathogens, ability to occupy vacant niches,

and capability of altering the invaded habitats (Gordon 1998).

The issue of invasive species in natural areas has received increasing attention for several

reasons. Threatened or endangered species protected under the Endangered Species Act are at

risk due to interference from, and predation by, invasive species (Pimentel et al. 2000). In

addition, the cost of invasive species control has negatively affected the budgets of many state

and federal agencies, and private landholders. In the state of Florida, more than $300 million has

been spent to control invasive plant species between 1980 and 2006 (Ferriter et al. 2006), and the

state of Florida is currently spending $30 million annually to control invasive species (D.

Schmitz, DEP, pers. comm.).

There are approximately 17,000 native plant species in the United States and about 2,500

species native to Florida (Pimentel et al. 2000; Pimentel et al. 2005). In contrast, about 25,000

non-native plant species have been introduced for cultivation in Florida alone (Frank and McCoy

1995). Of these species introduced to Florida, more than 925 have escaped and become

established in surrounding natural areas (Gordon 1998). Driving the importation of new non-

native species is a $15.2 billon environmental horticultural industry in Florida (Hodges and










Haydu 2005). And finally, about 85% of all plants imported into the United States arrive

thorough the Miami International Airport (Frank and McCoy 1995).

Ruellia tweedian2a is a non-native species that was introduced to Florida sometime before

1940 from Mexico. Its native distribution in Mexico is San Luis Potosi, Tamaulipas, Hidalgo,

Puebla, and Veracruz (GRIN 2004). It also extends into South America with a distribution in

western Bolivia, Paraguay, Uruguay, and northeastern Argentina, limited areas of southern

Brazil, and in open areas of eastern Chaco, located in wet or periodically flooded, sunny places,

such as ditches, streams, riverside lawns, or temporarily inundated areas (Ezcurra 1993).

Since its introduction it has naturalized in Alabama, Florida, Georgia, Hawaii, Louisiana,

Mississippi, South Carolina, Texas, the Virgin Islands, and Puerto Rico (USDA 2007), in USDA

hardiness zones 8 through 11 (Gillman 1999). The popularity of R. tweediana in the horticultural

trade appears to have increased in recent years and it is available for purchase in many nurseries

across Florida. This species holds great attraction to the homeowner because of its ability to

grow in many environmental conditions, from a water plant to growing in well-drained, almost

xeric conditions.

The genus Ruellia is in the family Acanthaceae which contains approximately 256 genera

(Zomlefer 1994). Ruellia is a large genus containing about 250 species found mostly in the

tropics and subtropics (Long and Lakela 1976). Five native species (R. caroliniensis, R. ciliosa,

R. noctiflora, R. pedunculata subsp. pinetorum, and R. succulents) and three naturalized non-

natives (R. malacosperma, R. tweediana, and R. ciliatiflora) occur in Florida (Wunderlin and

Hansen 2007). There are seven synonyms for Ruellia tweediana: Arrhostoxylon microphyllum,

Cryphiacanthus angustifolius, Ruellia brittoniana, R. coerulea (R. caerulea), R. ignorantiae, R.

microphylla, and R. spectabilis (Wunderlin and Hansen 2007) and eight cultivars: 'Chi Chi',









'Katie Pink', Katie Purple', 'Katie Variegated', 'Katie White', 'Morado Chi', 'Purple Showers',

and 'Snow White' (Wilson and Mecca 2003). A new cultivar 'Oh What a Feeling,' is presumed

to be a hybrid of R. tweediana x R. caroliniensis (S. Wilson, UF, pers. comm.) and is being sold

by http ://www.plantdelights. com.

Ruellia tweedian2a is an erect herbaceous perennial with one to many stems which are

either green or purple depending on the light conditions in which it grew (pers. obs.). Stems have

prominent nodal swelling and are documented to grow up to 1 m tall (Tobe et al. 1998), but

plants have been observed that are over 1.5 m tall (pers. obs.). The leaves are opposite, linear to

lanceolate and up to 30 cm long and less than 2 cm wide. The flowers are solitary or in few-

flowered cymes on axillary peduncles with a lavender or blue tubular corolla 2 to 4 cm long and

five-lobed. Cleistogamy (self-fertilization without flower opening) is known to occur and is

characterized by small, tubular greenish brown corollas (Long and Lakela 1976). Plants bloom

throughout the year in Florida (Tobe et al. 1998). Capsules are 2 to 2.5 cm long, cylindrical, with

suborbicular seeds about 2 mm wide (Long and Lakela 1976). Ruellia tweedian2a produces an

average of 20.6 seeds per capsule, each weighing an average of 1.78mg (Wilson et al. 2004).

Explosive dehiscence of the seed capsules results in seed dispersal distances from the parent

plant of 2.5-3 m (Witztum and Schulgasser 1995). A common family trait, the seed coat of R.

tweediana becomes mucilaginous when moistened (Zomlefer 1994). Some seed predation has

been noted in Florida by the M~elanagromyza ruelliae fly (Huey et al. 2007).

Populations of R. tweediana have appeared in natural areas across the state of Florida in

recent years. Wunderlin and Hansen (2007) reported R. tweediana in twenty-eight Florida

counties (Table 1-1). The Florida Exotic Pest Plant Council Distribution Database (FLEPPC

2007a), reported R. tweediana in 19 different natural areas (Table 1-2) in five different









community types: 1) pine flatwoods, prairies, 2) hardwood (hammocks, tree islands, etc.), 3)

freshwater marshes, 4) rivers, springs and 5) salt marsh. In 2001, FLEPPC upgraded R.

tweediana from a Category II (potential problem) to Category I due to "altering native plant

communities by displacing native species, changing community structures or ecological

functions, or hybridizing with natives" and its status has not changed since (FLEPPC 2007b).

The Institute for Regional Conservation's database, "The Floristic Inventory of South

Florida Database Online" contains distribution records of more than 2,400 species of plants in

South Florida conservation areas. Ruellia tweediana is listed in 18 conservation areas in South

Florida (Table 1-3) (Gann et al. 2007). Ruellia tweediana has been observed in other Florida

natural areas which have not been reported to these databases (pers. obs.).

According to the "Institute for Food and Agricultural Science (IFAS) Assessment of the

Status of Non-Native Plants in Florida' s Natural Areas," R. tweediana is concluded to be

invasive and not recommended by University of Florida-IFAS (UF-IFAS) faculty, in the north

and central parts of Florida. In southern Florida this species may be recommended by UF-IFAS

with "caution but should be managed to prevent its escape" (Fox et al. 2005). Experts questioned

for the IFAS Assessment process indicated that in several locations R. tweediana coverage

constitutes 50% of the infested stratum and that it is changing community structure by adding a

new stratum or increasing plant density in the stratum by 5-fold. It is also likely that it is altering

the hydrology within a community (C. Gantz, UF, pers. comm.).

At Blackwater Creek in Hillsborough County the population of R. tweediana is a very

dense monoculture in the swamp next to the creek. While it is not growing in the permanent

waterways, it is growing on exposed sandbars in the flowing water (A. Fox, UF, pers. comm.). It

is commonly found growing in narrow zones along waterways, for example at Frog Creek in










Manatee County (L. Huey, UF, pers. comm.), Hogtown Creek and Payne's Prairie Preserve State

Park in Alachua County (pers. obs.), Ocklawaha River in Marion County (L. Huey, UF, pers.

comm.), and The North Fork Buffer Preserve in St. Lucie County (A. Fox, UF, pers. comm.). At

Fakahatchee Strand State Park in Collier County, R. tweediana is growing down to the swamp

edge along the side of well-traveled, dry lime-rock roads. Where the swamp has flooded into the

road, R. tweediana can be found growing in the middle of the road (pers. obs.). At Tradewinds

Park in Broward County, a population ofR. tweedian2a is growing along the side of a horse trail

in an old dry stream bed (pers. obs.). At Long Key Natural Area in Broward County, it is

persisting under the canopy of trees of an abandoned homestead (pers. obs.). In Seminole County

at Lake Jessup, R. tweediana has formed a large monoculture under the cabbage palm hammock

even with grazing and trampling by cows (pers. obs.). These observations suggest most R.

tweediana populations in natural areas are associated with water, and upland populations are

likely to have persisted after deliberate planting and cultivation, or from populations that were

established under wetter conditions.

The native species R. caroliniensis grows in all but seventeen of Florida' s counties

(Wunderlin and Hansen 2007). Despite having this widespread distribution, it is typically found

as only a few scattered plants in any one place, not in dense monocultures. Ruellia caroliniensis

does not exhibit the narrow zonal water-related distribution exhibited by R. tweediana, although

under cultivation the two species exhibit similar habitat tolerances (pers. obs.).

Study Site and Zones

Field research on R. tweediana was conducted near Gainesville, Florida (Alachua County)

in small tributaries of Sweetwater Branch Creek on approximately 1.33 hectares in Payne's

Prairie Preserve State Park (Figure 1-1). Study sites were selected so the survival of R. tweediana

within the zone of R. tweediana monocultures could be compared with that in adj oining









elevation zones in which R. tweediana is not found. The adj oining zones were either more

upland with a diversity of native plants, or at a lower elevation and frequently/constantly

submerged. These three zones will be called "Ruellia", "Upland", and "Submerged" respectively

(Figures 1-2, 1-3). The Ruellia and Submerged zones were typically of 2-4 m wide while the

Upland zone varied in size but was generally wider than 10 m. On average (n = 60 per zone), the

number of species in a 1 m2 plOt of the Upland zone was 15.5 while in the Ruellia zone it was

6.7. The aboveground dry biomass for the Upland zone for R. tweediana was 0.0 g and other

species was 62.8 g and for the Ruellia zone for R. tweediana was 321.2 g and other species was

12.4 g. The average number of R. tweediana stems per plot in the Upland zone was 0.3 and the

250.8 in the Ruellia zone (Table 1-4, Appendix Table A-1 and Table B-1).

Research Questions

Considering that R. tweediana is found in a range of horticultural conditions, from aquatic

to almost xeric, the principle goal of this research was to determine why the natural area

distribution of R. tweediana was limited to a narrow zone along the banks of waterways rather

than in a broader distribution that includes both wetter and drier adj acent habitats. This research

investigated whether 1): the observed distribution of R. tweedian2a could be explained by

limitations due to environmental conditions: soil moisture, light, and soil temperature, and/or by

the biotic interactions of competition with native and non-native plants; and 2) at which life-

stage, seeds or seedlings, these limitations were most influential.

There were six experimental components: 1) a growth chamber experiment Seed

Viability in Submerged Conditions' and 2) a greenhouse experiment Soil Effects on Seed

Germination' are both described in Chapter 2. Four of the experiments were conducted in the

field, 3) a 'Seed Burial' experiment to assess the longevity of seeds in the soil, is described in










Chapter 3; 4) a Seed Germination' experiment, 5) a Survival' experiment' and 6) a Seedling

Transplant' experiment, are described in Chapter 4, to address the primary question in the field.










Table 1-1. Florida counties in which Ruellia tweedian2a occurs, (Wunderlin and Hansen's online
database, Atlas of Florida Vascular Plants, 2007).


Counties
Alachua
Brevard
Broward
Charlotte
Collier
Escambia
Franklin
Hendry
Highlands
Hillsborough
Indian River
Lake
Lee
Leon


Levy
Manatee
Marion
Miami-Dade
Monroe Mainland
Orange
Palm Beach
Pinellas
Putnam
Sarasota
Seminole
St. Lucie
Sumter
Volusia










Table 1-2. List of reported natural areas in which Ruellia tweedian2a occurs Florida, (The Florida
Exotic Pest Plant Council Distribution Database, 2007).
Natural Areas County
Tradewinds Park Broward
Forman Wilderness Preserve Broward
Myrtle Slough @ CR 74 Charlotte
Big Cypress National Park Collier
Fakahatchee Strand State Park Collier
Blackwater Creek Hillsborough
Marsh Branch Hillsborough
East Side Canal Hillsborough
Anclote River Hillsborough


Ocklawaha River
Long Key State Park
Lake Park Scrub Natural Area
No. Jupiter Flatwoods Natural Area
Anclote River
Alligator Creek Conservation Area
Cameron Property (SJRWMD)
Fort Mose State Park
No. Fork Buffer Preserve
Center Hill


Marion
Monroe
Palm Beach
Palm Beach
Pasco
Pinellas
Seminole
St. Johns
St. Lucie
Sumter










Table 1-3. List of conservation areas in which Ruellia tweediana occurs in South Florida, (The
Floristic Inventory of South Florida Database Online, Gann et al. 2007).
Conservation Area Counties
Arch Creek Park Miami-Dade
Big Cypress National Preserve Collier, Miami-Dade, Monroe
Caloosahatchee Regional Park Lee
Camp Owaissa Bauer Miami-Dade
Castellow Hammock Park Miami-Dade
Collier-Seminole State Park Collier
Deering Estate at Cutler Miami-Dade
Dry Tortugas National Park Monroe
Dupuis Reserve Martin, Palm Beach
Enchanted Forest Park Miami-Dade
Everglades National Park Collier, Miami-Dade, Monroe
Fred C. Babcock-Cecil M. Webb Wildlife Management Area Charlotte
Greynolds Park Miami-Dade
Hugh Taylor Birch State Park Broward
Little Hamaca Park Monroe
Long Key/Flamingo Road Natural Area Broward
Secret Woods Buffer and Nature Center Broward
Simpson Park Miami-Dade

















QI






-- ,








Pd
my
Qo




OE


-o

Ob
=0
'aM
O~c~



















Upland Zone


Figure 1-2. Photo on left is aligned with diagram on right to indicate the labeling of the three
zones, Upland, Ruellia, and Submerged, in the field taken from the Submerged zone.



















SRuellia Zone






SUpland Zone






Figure 1-3. The edge between the Upland zone and Ruellia zone, from the perspective of the
Upland zone, at Payne's Prairie Preserve State Park, (January 2006).









Table 1-4. Dry above ground biomass data for 1 m2 plOt, Ruellia weight is R. tweediana and
Others weight = all other species in plot. Ruellia count = number of R. tweediana
stems found in 1 m2 plOt (Raw data in Appendix Table B-1).
Ruellia Others Ruellia
Factor Treatment n weight weight Units count
Zone
Ruellia 60 321.2 12.4 g 250.8
Upland 60 0 62.8 g 0.4
Size
Large seedlings 40 160.4 45.2 g 118
Small seedlings 40 154.4 36 g 124.8
Seed experiment 40 167.2 31.6 g 130.8
Competition
Cleared (Aug. 06) 60 179.2 43.6 g 123.6
Not-Cleared (Nov. 06) 60 142 31.6 g 125.6









CHAPTER 2
SEED VIABILITY IN SUBMERGED CONDITIONS AND SOIL EFFECTS ON SEED
GERMINATION

Introduction

In horticultural situations, Ruellia tweediana can grow in a range of sites from an aquatic state

to well-drained, almost xeric conditions. Mature plants have also been shown to survive for

several months growing in submerged conditions in laboratory trials (A. Fox, UF, unpublished

data). But observations from sites across Florida have shown that this species is not found in the

waterways but alongside them. A factor that might be causing this is a family trait of the

Acanthaceae, a mucilaginous gel that forms around some species seeds when moistened.

Gutterman et al. (1973) indicate that the gel aids long-distance dispersal of the seeds and

adhesion of seeds to the soil surface. A previous germination trial on R. tweediana seeds, with

and without the mucilaginous gel present, indicated that there was no difference in percent

germination (L. Huey, UF, unpublished). But Gutterman et al. (1973) also indicated that this gel

can inhibit germination when seeds are in excess water certain species within this family. Ruella

tweediana seeds produce a mucilaginous gel (pers. obs), so it is possible that continuous

submersion of seeds covered with this gel would inhibit germination in this species. There are no

reports in the literature that this possibility has been examined for this species.

Seeds can be in different physiological states after leaving the parent plant (Booth et al.

2003). Ruellia tweediana seeds do not exhibit primary dormancy but are instead non-dormant

after leaving the parent plant (Wilson and Mecca 2003). After dispersal, seeds that become

buried can have a forced quiescence due to environmental conditions in the soil that are

unfavorable for germination. It is not known whether R. tweediana seeds buried in unfavorable

environmental conditions exhibit secondary dormancy, where they still do not germinate even if

placed in optimal germination conditions (Booth et al. 2003).










For successful reproduction, germination of quiescent, non-dormant seeds has to occur at

the appropriate season and soil depth. Other factors that can affect germination include soil

structure, which determines the distribution and availability of water, solutes and gases, and the

chemical effects of minerals in the soil, although this is a poorly understood relationship.

Although most inorganic ions in soils do not have any specific effect on seed germination, nitrate

ions have been shown to stimulate germination of some seeds (Hilhorst and Karssen 2000).

A submersion experiment was conducted to determine if submerged R. Aveediana seeds

were capable of germination. If not, this could be limiting the distribution of R. Aveediana in the

constantly submersed sites. A second experiment tested the effects of several soil types including

those from the three Hield site zones on R. Aveediana seed germination. Chemical and physical

characteristics of the field site soils were also measured to provide information on the site

conditions and for comparison if there were significant differences in seed germination from soil

samples from the different zones.

Hypotheses

Seed Viability in Submerged Conditions

1. Null Hypothesis: The percentage of submerged seeds that germinate or survive during 50 days
in deionized water will not be different then zero.
Alternative Hypothesis: The percentage of submerged seeds that germinate or survive during 50
days in deionized water will be significantly different then zero.

Soil

2. Null Hypothesis: Seeds will have the same percentage germination in each tested soil type.
Alternative Hypothesis: Seeds will not have the same percentage germination in each tested soil
type.

3. Null Hypothesis: If there are any significant differences in seed germination in the soils from
different sources, this will correlate significantly with differences in soil characteristics.
Alternative Hypothesis: If there are any significant differences in seed germination in the soils
from different sources, this will not correlate with differences in soil characteristics.









Materials and Methods

Seed Viability in Submerged Conditions

Ruellia tweedian2a seeds were collected November 4, 2005 from Payne's Prairie Preserve

State Park. All seeds were collected on the same day, mixed thoroughly, and stored together in a

covered plastic container under refrigerated conditions. At beginning of the experiment

(3/10/06), these seeds had a viability of 95% when tested with tetrazolium dye (Peters 2000 as

described below).

Ten seeds were submerged in oxygenated deionized water in each of 10 replicate petri

dishes, which were sealed with parafilm. Petri dishes were incubated at 20/30 oC (12h/12h cycle)

and a 12 hour day/night cycle. Every ten days (3/10/06 4/20/06), all germinated seeds were

removed from the Petri dishes and placed on Farfard soil (Conrad Fafard, Inc.) in a greenhouse.

Two of the replicate Petri dishes were then removed from the experiment and all remaining

ungerminated seeds were tested with tetrazolium to see if they were still viable but dormant. For

this viability test, seeds were moved to a new petri dish lined with filter paper, they were cut

laterally and stained with 1.0% concentration of tetrazolium and placed in a dark, 30 oC

incubator for 12 hours, following the Peters (2000) Tetrazolium Testing Handbook. An entirely

stained embryo indicated that the seed was viable but dormant.

Soil

Soil effects on seed germination

Five replicates of soil slabs of approximately 30 cm x 30 cm x 6 cm depth were collected

from Payne's Prairie Preserve State Park from all three zones. Soil slabs and Fafard soil (Conrad

Fafard, Inc.) (used as a control soil) were placed in trays and air dried for ten days. Fifty seeds

(from the same batch of seeds used in the Seed Viability in Submerged Conditions'

experiment), were placed on top of the soil in each of four replications per zone. The 5th SOil










replicate had no R. tweediana seeds added to test for any germination from the field seed bank.

Trays were moistened, covered with plastic and re-watered as needed. During the 30 day period,

trays were monitored at least weekly, and seedlings were counted and removed as they were

observed.

This experiment was repeated using the previously described methods, but was expanded

to include a soil mixing treatment. For the seed mixing treatment, the soil was mixed to

redistribute seeds to depths ranging from 1 cm to approximately 3 cm. The factor effects tested

in this experiment were soil source and seed position in soil.

Soil characteristics

Ten replicate sites were chosen at Payne' s Prairie Preserve State Park based on the

presence of the Upland, Ruellia, and Submerged zones. A 1-liter volume of soil was collected

from each zone for each of the ten sites and was analyzed for organic matter (OM), cation

exchange capacity (CEC), soil pH, estimated nitrogen release, NO3-N (nitrates), readily

available phosphorus, total inorganic phosphorus, water soluble phosphorus, exchangeable

calcium, exchangeable hydrogen, exchangeable magnesium, exchangeable potassium, and

exchangeable sodium. A second set of soil samples was collected from each zone in a random

selection of 5 of the 10 replicate sites and was analyzed for texture. All analyses were conducted

by A & L Southern Agricultural Laboratories, Inc, Pompano Beach, FL (http://www.al-labs-

plains. com).

Statistical Analyses

Analyses of variance (ANOVA) and Tukey's multiple comparison tests were performed to

investigate differences in seed germination between the different soil sources and seed position

in the soil, and to compare soil characteristics from the different zones. Differences between









means will be reported with an alpha (a) value and ANOVA results will be reported with a p-

value. All statistical analyses were done with SAS Version 9.1 software (2007).

Results

Seed Viability in Submerged Conditions

Out of the 100 seeds that were submerged in deionized water, 42 seeds were tested for

viability of which 41 were viable. The other 58 seeds had germinated in the chamber during the

50 day study and had been placed on soil (Figure 2-1). These seedlings were followed for 30

days after the last seed germinated, and only 4 of these seedlings died.

Soil

Soil effects on seed germination

In the first soil slab experiment, a total of 7 seeds from the field seed bank germinated, 2 of

which were in soil to which no seeds had been added and 5 germinated from a visible capsule in

one replication of the Ruellia zone soil to which 50 seeds were added (this was known because

the seedling count was greater than 50). In the Upland, Ruellia, Submerged, and Control soil on

average 91.0%, 94.5%, 95.5%, 88.6% of added seeds (assumes no resident seed germination)

germinated respectively. These values are not significantly different from one another (p = 0.2)

(Figure 2-2).

In the second experiment, 77%, 73.5%, 78%, 88% of seeds germinated (assumes no

resident seed germination) respectively in the Upland, Ruellia, Submerged and Control soil and

these values were not significantly different from one another at an a of 0.05. Although the soil

source in this experiment was not a significant factor, the soil mixing treatment was significant

(p = 0.0001), with seeds on top of the soil having a higher average germination (88.8 %) than

mixed seeds (69.2 %) (Figure 2-2).









Soil characteristics

Soil pH, estimated nitrogen release, total inorganic phosphorus, water soluble phosphorus,

exchangeable hydrogen, exchangeable magnesium, and exchangeable potassium were not

significantly different between zones (Table 2-1). While OM, CEC, NO3-N (nitrates), readily

available phosphorus, exchangeable calcium, and exchangeable sodium were different among

zones (Table 2-1, Figures 2-3). Specifically, cation exchange capacity was highest in the Ruellia

zone with lower, and similar values in the Upland and Submerged zone (Figure 2-3). Compared

to optimal concentrations for crop growth, NO3-N (nitrates) rates were Medium, Medium, and

Low in the Upland, Ruellia, Submerged zones respectively (Table 2-1 and Figure 2-3). Readily

available phosphorus and exchangeable calcium were above the optimum rate for crops yield in

all zones (Table 2-1). Total inorganic phosphorus levels were all higher than expected in crop

soils while water available phosphorus was very low for all zones (A & L Southern Agricultural

Laboratories). High levels of salinity can be a limiting factor on plant growth (Warrence et al.

2002). The sodium gradient on these transects had higher concentrations in the submerged zone

and decreases to the Upland zone (Figure 2-3).

Soil texture was not significantly different among zones (a of 0.05). The soil classification

is sand with an average composition of 91.5% sand, 5.6% silt, and 2.9% clay.

Discussion

Seed Viability in Submerged Conditions

The maximum time that seed submersion was tested was only 41 days instead of the expected

50 days, because all seeds had germinated by day 41. Therefore the null hypothesis, the

percentage of submerged seeds that germinate or survive during 50 days in deionized water will

not be different then zero, has to be rej ected.









Soil

Soil effects on seed germination

It was excepted that there would be different germination rates among the zonal soil samples

and the control soil, but that was not the case. Flooding and soil type effects on seed germination

do not appear to be responsible for limiting the distribution ofR. tweediana.

Soil characteristics

Given that there were no differences in seed germination on the soil from different zones, it

can be concluded that differences in soil characteristics are not significantly influencing short-

term seed survival and seed germination. The NRCS (2007) soil series for the region within

Payne's Prairie Preserve State Park where the field research was conducted is Pomona sand,

depressional (Figure 2-4). This soil type has native vegetation composed primarily of cypress

(Taxoium~ distichum (L.) L.C. Rich.), swamp maple (Acer rubrum L.), tupelo

(Nyssa sylvatica Marshall var. biflora (Walter) Sarg.), bay, and some scattered pond pine (Pinus

serotina Michx.), and typical soil parameters as identified in Table 2-2 (NRCS 2007). Aerial

photographs of the study site from 1982 showed that it was completely surrounded by farmlands.

This compares to the 2005 aerial photograph in Figure 2-4.

A particularly interesting result in the soil analyses was that the Submerged zone had the

lowest value for percent organic matter and that all zones were above the NRCS expected range.

The soil pH was also higher than expected for this soil type (NRCS 2007) but horticulturists have

reported that R. tweediana can grow within the found range (Dave's Garden 2007). While there

were significant differences between zones for cation exchange capacity, all values were

considered within normal range for crop growth but were higher than recorded for this soil type

by NRCS (Table 2-2). Even with some of the soil parameters not being in the normal crop or









NRCS soil survey ranges or having significant differences in the soil characteristics among

zones, it appears that this did not affect seed survival and germination in the two experiments.





20 3
Days Submerged


Figure 2-1. Percent germination and percent viability of seeds over time in the Seed Viability in
Submerged Conditions study. The percent germinated of seeds remaining at each time
is represented in black and the percent viable of seeds tested each time is represented
mn grey.























S30-
















Control Upland Ruelha Submerged
Soil Type



Figure 2-2. Number of germinated seeds in the Soil effect on seed germination" experiments.
Experiment 1 seed germination is presented in the hash marked bars. Experiment 2 is
presented with black for seeds on top of the soil and dark grey for seeds mixed in the
soil.











Table 2-1. Soil analyses results for 10 replicates from Upland, Ruellia, and Submerged zones.
Analysis conducted by A & L Southern Agricultural Laboratories, Inc. on May 15,
2006.


Optimum
crop
nutrient
rangeb


Zone

Upland Ruellia Submerged Units


A&L ratings


Soil Parameters


% Sig.a
meq/100g Sig.
NS
Lbs./A NS
ppm Sig.
ppm Sig.
ppm NS
ppm NS


Organic matter
Cation exchange capacity
Soil pH
Estimated nitrogen release
NO3-N (nitrates)
Readily available phosphorus
Total inorganic phosphorus
Water soluble phosphorus
Exchangeable
Calcium
Hydrogen
Magnesium
Potassium
Sodium


8.22
18.56
7.6
208.3
13
76.8
147.45
0.44

3269.5
0.29
169.9
40.3
35.25


8.69
24.53
7.17
217.7
7.55
39.3
145.7
0.45

4322
0.44
185.2
40.85
100.4


6.9
15.31
6.76
197
5.35
58.1
135.15
0.27

2513.5
0.63
228.4
27.25
111.5


5 to 35
6.0-7.0 H. VH, H
VH, VH, VH
M, M, L
20-30 VH, H, VH
40-60 VH, VH, VH
VL, VL, VL

VH, VH, H

30-70 M M. M
90-125 VL, VL, VL
L, H, H


ppm
meq/100g
ppm
ppm
ppm


"Sig. = Significant (p=0.05), NS = Not Significant between zones (Figure 2-1)
bMOst soil test readings on the report are given a rating of very low (VL), low (L), medium (M), high (H), or very high
(VH). The purpose of these readings is to provide a guideline for determining optimum nutrient levels for crop growth.











a b ab


ba a


12 Upland
O Ruellia
11 Submerged


100-




80-



60-




40-
bab
a ab b

20 ab a b
b ab



% Organic matter Cation exchange NO3-N ppm Readily Available Calcium ppb Sodium ppm
capacity meq/100g Phosphorus ppm


Figure 2-3. Soil parameters with different letters indicated significant differences among zones
within parameters (a of 0.05), n=10.























































40 80 180 240


N 40
A.


liMs im esucs
~ijCmunervationSevie


Web Soil Survey 2.0
National Cooperative Soil Survey


8110/2007
Page 1 of 3


Figure 2-4. Natural Resources Conservation Service and United States Department of
Agriculture map of soil series superimposed over an aerial photograph of a section of
Payne's Prairie Preserve State Park. Soil type 25 indicates Pomona sand, depressional
the area in which this field research was conducted.










Table 2-2. Soil survey results of Ponoma Sand, depressional from Natural Resources
Conservation Service and United States Department of Agriculture October 2006
(http:.//websoilsurvey .nrcs.usda.gov)
Soil Parameters Units
Effective cation-exchange capacity .3-3.5 Meq/100g
Soil reaction pH 3.5-5.5 pH
Salinity 0-2 mmhos/cm
Sodium adsorption ratio 0-4 --
USDA texture sand --
Sand -a
Silt 0-15 %
Clay 1-6 %
Moist bulk density 1.2-1.5 g/cc
Saturated hydraulic conductivity 42.34-141.14 Micro m/sec
Available water capacity .05-.10 In/In
Organic mater 1-3 %
aValue not provided









CHAPTER 3
SEED BURIAL

Introduction

A soil seed bank is the reserve of viable seeds present in the soil and on its surface

(Roberts 1981). A transient seed bank is characterized by a lack of viable seeds after a year under

field conditions; whereas a persistent seed bank has viable seeds after a year (Thompson and

Grime 1979). Once the mature plant population is controlled, the management of an invasive

species will be much easier if it has a transient seed bank. Re-inspection and re-treatment is only

needed for two years to ensure that none of the germinating seeds in the soil develop to produce

seeds and replenish the seed bank (Magda et al. 2004). Conversely, the management of a species

with a persistent seed bank will be more long-term and depend directly on the longevity of the

seed bank (Gardener et al. 2003; Swanton and Booth 2004).

There is limited knowledge of the importance seed banks should have in invasive species

management in natural areas (Swanton and Booth 2004). This is because there are not many

studies on seed banks in natural areas, but there is a large body of literature dealing with seed

banks in agricultural systems due to the economic importance of agricultural weeds and the fact

that many of them reproduce only by seeds (Roberts 1981). These studies can be reviewed to

help understand what might be occurring in natural systems.

In agriculture it is common to apply the short-term management of removing the above

ground biomass of weeds that are competing with crops and limiting the overall yield (Swanton

and Booth 2004). In the management of the above ground vegetation of agricultural weeds, the

crop is protected and all other plants are removed. This is achieved by selective herbicides that

do not kill the crop and by altering of the genes of the crop to make them resistant to, and hence

permit the use of, certain herbicides (e.g. Roundup Ready crops). This differs from natural









ecosystems where only the invasive species need to be removed and all other plants need to be

protected. This difference makes the use of herbicides in natural areas difficult and the altering of

genes not an appropriate concept. But Jordan et al. (1995) showed that seed mortality had greater

effect on weed populations in crops than plant mortality or decrease in seed production per plant.

The management of invasive species in natural ecosystems should include the management of

the species' seed bank (Swanton and Booth 2004).

For invasive species management it is important to have an understanding of the seed

viability, dormancy, and longevity of each invasive species (Gardener et al. 2003; Lutman et al.

2002; Mennan and Zandstra 2006). Understanding seed longevity helps to predict the rate of loss

of seeds from the seed bank (Baskin and Baskin 2006). Dormancy is important because it is the

reason that seeds can remain viable in the seed bank for extended lengths of time (Booth et al.

2003).

As summarized by Baskin and Baskin (2006), one manner of determining longevity is

based on inferring the seeds' age based on site information. This can be done in archeological

sites where viable seeds can be dated based on the layer of earth in which the seeds are found.

Age can also be inferred in agricultural fields where the field has been sown in one crop for an

extended period of time and the seeds that lie beneath can be assumed to be older than the sown

crop. Another method to determine longevity of a seed bank is to bury seeds in the soil, wait

various periods of time, exhume the samples, and check the seeds for viability. Experiments of

this nature have been conducted since the late nineteenth century.

In a previous study it was determined that fewer than 2% ofRuellia tweedian2a seeds were

viable after 13 months of burial in field conditions (Barnett, unpublished data). This suggested

that R. tweediana essentially has a transient seed bank and that long-term management needs










would be reduced. The first component of the present study was to replicate this previous work,

with seeds of R. tweedian2a buried in the Hield in areas that contained mature populations of R.

tweediana to determine their survival for at least fifteen months. The second component of this

study was to expand the area of interest to include adj oining zones in the Hield that did not

contain mature R. tweediana plants, to determine if seeds have different rates of germination or

mortality in these zones and if these zones could maintain transient or persistent seed banks. The

overall obj ective of this study was to determine if the Hield distribution of R. tweediana was

limited by seed bank dynamics.

Hypotheses

1. Null Hypothesis: Plots will have the same mean physical characteristics such as: Gap
Fraction, Light, Soil Moisture, and/or Soil Temperature in the Upland, Ruellia, and Submerged
zones.
Alternative Hypothesis: Plots will NOT have the same mean physical characteristics such as:
Gap Fraction, Light, Soil Moisture, and/or Soil Temperature in the Upland, Ruellia, and
Submerged zones.

2. Null Hypothesis: Buried R. tweediana seeds will NOT survive for more than a year.
Alternative Hypothesis: Buried R. tweediana seeds will survive for more than a year.

3. Null Hypothesis: Buried R. tweediana seeds will have the same number in these categories:
Missing, Dead, Total Germinated, Dormant, and Total Viable in the Upland, Ruellia, and
Submerged zones.
Alternative Hypothesis: Buried R. tweediana seeds will NOT have the same number in these
categories: Missing, Dead, Total Germinated, Dormant, and Total Viable in the Upland, Ruellia,
and Submerged zones.


Materials and Methods

Study Site and Zone Characterization

Ten replicate sites were located at Payne's Prairie Preserve State Park on December 15,

2005 based on the presence of adj acent Upland, Ruellia, and Submerged zones. A transect was

run from the center of the Submerged zone, through the Ruellia zone, and 5m into the Upland

zone at each of the ten replicate sites. Three 0.25 m2 plOts along these transects were created, one









in the center of each zone. The experimental design was Zone as the factor with three treatment

levels Upland, Ruellia, and Submerged.

Canopy photos and light measurements. Digital canopy photographs were taken above

every plot with a Nikon Coolpix 4500 digital camera on a tripod with a fish-eye lens attachment.

The camera was leveled with the top facing north for each photograph on a non-sunny day.

Photographs were analyzed with the HemiView 2. 1 (Delta-T Devices, Cambridge, UK)

computer program to classify each pixel in each photograph as either 'sky' or 'non-sky' to

calculate canopy closure, also known as 'Gap Fraction.' Gap Fraction is a value that ranges from

0 (a closed canopy) to 1 (an open sky). HemiView's settings were calibrated to site conditions at

Payne's Prairie Preserve State Park based on the latitude, longitude, declination, and altitude.

Photos were imported into the program, sized to include the whole image, aligned to have north

in the photo match north in the program, and the threshold (brightness of photograph) was

specified.

Light measurements were taken with a LI-COR LI-250 (LI-COR, Lincoln Nebraska) light

meter. Readings were taken at ground/water level in the center of each plot by averaging the

light readings over a 30 second period of time.

Soil moisture and temperature measurements. Soil moisture measurements 6 cm deep

were taken periodically during the study with a Dynamax Theta Probe type ML2x (Delta-T

Devices, Cambridge, UK) attached to a Delta-T Moisture Meter. The probe was calibrated to the

site conditions following the manufacturer's instructions. The measurements were recorded as a

dimensionless parameter that is expressed as percent volume. Thus 0 % corresponds to a

completely dry soil, and pure water gives a reading of 100 %. Soil temperature measurements









were taken at 8 cm depth with a VersaTuff Plus 396 Atkins (techniCAL Systems, Hamilton,

Ontario) temptec probe (oC).

Seed Burial Study

Ruellia tweedian2a seeds were collected November 4, 2005 from Payne's Prairie Preserve

State Park. All seeds were collected on the same day, mixed thoroughly, and stored together in a

covered plastic container under refrigerated conditions, and at the beginning of the experiment

(March 2006) had a viability of 95% when tested with tetrazolium dye (Peters 2000 as described

below). On March 9-10, 2006, fifty seeds were mixed with ~4 g of sand (Quikrete Premium Play

Sand which was sterilized at 100 oC for five days), then rolled tightly in a 15.2 cm by 15.2 cm

section of nylon mesh and attached to a surveyors' flag with a zip tie (Figure 3-1). Sand was

mixed with the seeds to decrease the spread of disease and fungus from one seed to another (Van

Mourik et al. 2005).

On March 13, 2006, one seed bag was buried 10 cm deep in the ground at the center of

each plot and the other nine bags were buried in a circle surrounding this bag. The area that the

bags were buried in was approximately 0.25 m2 (Figure 3-2). Based on high rates of seed death in

the Submerged zone in a preliminary study (not reported), the number of sampling bags placed in

the Submerged treatment was limited to five.

One bag was removed from each plot after 1, 3, 6, 9, 12, and 15 months (March 2006 to

June 2007). Bags were brought to the lab, zip ties were cut off and the sand was washed from the

seed bags. The number of seeds recovered was recorded, as was the number of seeds that had

germinated in the ground. Recovered, ungerminated seeds were placed in a petri dish lined with

filter paper, moistened with deionized water, and sealed with parafilm. Petri dishes were

incubated at 20/30 oC (12h/12h cycle) and a 12 hour day/night cycle for 30 days. These

conditions have been shown to promote successful germination (Wilson and Mecca 2003).









Germination was then recorded and all remaining ungerminated seeds were tested with the

vitality stain tetrazolium to see if they were still viable but dormant. Seeds were moved to a new

petri dish lined with filter paper, they were cut laterally and stained with 1.0 % concentration of

tetrazolium and placed in a dark, 30 oC incubator for 12 hours, following the Peters (2000)

Tetrazolium Testing Handbook. The seeds were evaluated for an entirely stained embryo which

indicated that the seed was viable but dormant.

Statistical Analyses

Dependent variables related to the seeds were the percent missing, percent germination,

and percent dead. "Total viable seeds" was the sum of the numbers of seeds germinating in soil,

capable of germinating, and dormant. Analyses of variance (ANOVA) and Tukey's multiple

comparison tests were performed to investigate changes in the dependent seed variables over

time and the differences between zones in: canopy photos, light measurements, soil moisture,

soil temperature, and dependent seed variables. Differences between means will be reported with

an alpha value and ANOVA results will be reported with a p-value. All statistical analyses were

done with SAS Version 9.1 software (SAS 2007).

Results

Zone Characterization

Canopy photos and light measurements. Canopy photographs revealed mean (+ standard

deviation) Gap Fractions of 0. 1147 (+0.007), 0. 115 (+0.009), 0. 1153 (+0.0010), for the Upland,

Ruellia, and Submerged treatments respectively. These indicated an almost closed canopy with

no significant difference between zones (p= 0.99). This was supported by the mean (+_standard

deviation) measurements of light in the understory, 67.9 Cpmols sl m-2 (+~50.5), 60.3 Cpmols sl m-2

(+28.3), and 50.2 Cpmols sl m-2 (+15.1) for the Upland, Ruellia, and Submerged treatments

respectively, which also showed no difference between the zones (p=.52). Full sunlight at solar









noon on a clear summer day is 2224 Cpmols sl m-2 (+_90) in Miami, Florida (Lee and Downum

1991).

Soil moisture and temperature measurements. Percent soil moisture was significantly

different between each zone at each sampling interval and when averaged across the study period

(ae = 0.05) Figure 3-3. Soil moistures, averaged over all sampling intervals, were 32 %, 52 %,

and 91 %, in the Upland, Ruellia, and Submerged treatments, respectively.

Average soil temperature in the Upland treatment (21.4 Co) was significantly different

from the Submerged (20.9 Co) treatment (a of 0.05), while the Ruellia treatment (21.1 Co) was

not different from the Upland or Submerged treatments.

Seed Burial Study

Samples were exhumed from the Hield April 14, 2006 (Month 1), June 16, 2006 (Month

3), September 13, 2006 (Month 6), December 16, 2006 (Month 9), March 15, 2007 (Month 12)

and June 19, 2007 (Month 15), (the last sampling time only included the Upland and Ruellia

treatments because of the reduced number of sample bags in the Submerged zone). Overall, a

low percentage of seeds were missing within each zone and there was no significant difference

between zones for this variable. However, there was an increase in the number missing over the

15 month period of burial (Figure 3-4, Table 3-1).

The percentage of seeds that were recovered but were dead varied significantly with burial

time and were significantly higher each month in the Submerged treatment compared to the

Upland and Ruellia treatments (Figure 3-5). There was a two-way interaction between treatment

and burial time with a p-value of 0.058 (Table 3-2) because Upland and Ruellia zones were only

different during some months. The same two-way interaction (p = 0.088) was found in the

percentage of seeds that germinated (Table 3-3 and Figure 3-6) because Upland and Ruellia

zones were similar during some months. Seeds that were viable but presumed dormant consisted










of 3.2 % of the total seed. There was a two-way interaction between treatment and burial time for

dormant seeds (p = 0.003) (Figure 3-7, Table 3-4), again because differences between Upland

and Ruellia zones only occurred at some sampling times.

The percentage of total viable seeds (total viable seeds included seeds germinating in soil,

capable of germinating, and dormant) averaged over burial times were 74.9 % for the Upland,

66.3 % for the Ruellia, 8.9 % for the Submerged (Figures 3-7 and 3-8). During the first 12

months the percentages of viable seeds for the Upland and Ruellia treatments were not

significantly different (a of 0.05), while the Submerged treatment was different from them both.

There was a significant difference between the Upland and Ruellia treatments during month 15

(the Submerged treatment was not sampled), with higher viability in the Upland treatment. In the

Upland and Ruellia treatments, by nine months after burial, percent total viable was significantly

reduced from its highest percentage (three months after burial in the Upland and one month after

burial in the Ruellia treatment Figure 3-8).

Discussion

Zone Characterization

Canopy photos and light measurements were taken for site and zone characterization and

to determine if differences in light could explain the field distribution of R. tweediana. All

habitats, however are under the same cypress canopy and no differences in Gap Fraction and

light measurements were found. Thus, the first null hypothesis has to be accepted for these two

variables.

Soil moisture decreased and soil temperature increased moving from the Submerged zone

to the Upland zone. This was expected due to the slight elevational increase along this transect,

and the thermal stability of water and the higher specific heat of water compared to land. These









data lead to the rej section of the null hypotheses that soil moisture and soil temperature in the

Upland, Ruellia, and Submerged zones would be the same.

Seed Burial Study

A previous study (E. Barnett, unpublished) indicated that R. tweediana had a transient

seed bank within the Ruellia zone but results here suggest otherwise. The high (34.4 % (SD + 9. 1

%)) total seed viability in the Ruellia zone after 15 months of burial indicated that the seed bank

is persistent. This difference in results between those two studies could be due to many factors.

Ruellia tweedian2a viability was sensitivity to storage conditions. Seeds had a rapid decline in

viability (~95% down to 0% 8%) when stored under ambient laboratory conditions when

containers are completely sealed or completely open for approximately 4 months. Higher

viability rates (95%) were found when seeds were stored in refrigerated conditions and covered

but not completely sealed. The previous study did not test the viability of the seeds at time of

burial. Another factor that was different was the addition of sand to the seed burial bags. This

buffer between buried seeds could have reduced disease spread and fungal growth during the

experiment.

Soil moisture, previously thought to be an indicator of where R. tweediana could survive,

was lower in the previous study than the current one (soil moisture of the Ruellia zone in the

previous study was similar to that in the Upland zone the current study). Thus, this variable is

unlikely to explain differences in seed viability between the two studies. The results of the

current study falsified the hypothesis that R. tweediana would not survive over a year, and this

species has a persistent seed bank.

The expansion of the experiment to include the seed survival in the adj oining zones

resulted in unanticipated outcomes. It was expected that there would be statistical differences in

viability between the Upland and Ruellia treatments because of the absence of R. tweediana










plants and the decreased level of moisture in the Upland zone. Instead, the percent viability was

not different between zones during the first 12 months and seed survival was actually greater in

the Upland zone after 15 months of burial. This suggests that under the conditions of this study,

seed survival is unlikely to be the factor limiting R. tweediana in the Upland zone.

Because of the results of the two experiments "Soil effects on Seed Greenhouse" and

"Seed Viability in Submergence Conditions" (Chapter 2), the high percentages of dead seeds

recovered from the Submerged zone were also unexpected. Reasons why this occurred could

include the anaerobic conditions in the field compared to the oxygenated deionized water used in

the "Seed Viability in Submergence Conditions" experiment. It also could have been due to

some toxicant or adverse biochemical conditions in the water that caused the seeds to die. The

tributaries of Sweetwater Branch Creek in which the research was conducted are directly

downstream of a sewage plant and a large homeless community that uses the stream. Additional

studies would be needed to determine what conditions in the Submerged zone caused the death

of the R. tweediana seeds. For example an analysis of water quality, including redox potential

might be instructive.

In this burial experiment, seeds were collected from may different plants, and were almost

certainly in different physiological states. It was not believed that the seeds had a primary

dormancy (Wilson and Mecca 2003) and once seeds were collected from parent plants many

seeds would be non-dormant and consequently could germinate in the bags in the soil. Some of

the buried seeds did not germinate in the soil but germinated once placed in the incubation

chamber. It is likely that these seeds had forced quiescent due to environmental conditions in the

soil unsuitable for germination. Other seeds did not germinate even after placement in the









germination chamber and those seeds exhibited induced secondary dormancy (called dormant

but viable in this text).

This experiment indicates that soil moisture and light are not the primary factors that

exclude R. tweediana from the Upland zone. It also demonstrated that there is potential for a

persistent seed bank to exist in the Upland and Ruellia zones. The viability of buried seeds

decreased over time but not as quickly as had been expected, and seeds did survive burial for a

year in each of the zones, although at very low percentages in the Submerged zone.






































Figure 3-1. Nylon mesh seeds bags attached to the bottom of surveyor' s flags with zip ties.


















































Figure 3-2. Seed burial plot, closest flags in Submerged zone, center flags in Ruellia zone,
background flags in Upland zone.












53









































F M A M J J A S O N
-A Upland -X- Ruellia -5- Submerged


D J F M


Figure 3-3. Average percent soil moisture for each zone February 2006 to March 2007. There
were significant differences between zones at each sampling interval and averaged
across time.












bc


30 -b




bb


XX



10 -

~a xa


'Oi O
aa


oio

0 2 4 6 8 10 12 14 16
Months After Burial



Figure 3-4. Percent missing of seed total in each zone. U= Upland zone, R= Ruellia zone, and S=
Submerged zone. Different letters indicate significant difference between months
after burial (a of 0.05) for % missing averaged over all zones. Line fitted to averages
with an R2 = 0.93. (Submerged zone not sampled at 15 months.)

Table 3-1 ANOVA table for percent missing of seed total.
Source of Variation Degree of Freedom F Value P Value
Zone 2 0.29 0.7497
Month 5 25.99 0.0001
Zone*Month 9 0.94 0.4952
































10


8
Months After Burial


10 12


Figure 3-5.


Percent dead of seed total in each zone. U= Upland zone, R= Ruellia zone, and S=
Submerged zone. Percent dead were significantly higher at each month after burial in
the Submerged zone compared to the Upland and Ruellia zones. (Submerged zone not
sampled at 15 months.)


Table 3-2 ANOVA table for percent dead of seed total.
Source of Variation Degree of Freedom F Value
Zone 2 212.7
Month 5 3.71
Zone*Month 9 1.88


P Value
0.0001
0.0034
0.0588















































Table 3-3 ANOVA table for percent germinated of seed total.
Source of Variation Degree of Freedom F Value P Value


100.0


90.0 Z1


80.0


50.0


8
Months After Burial


10 12


Figure 3-6. Percent germinated of seed total in each zone. U= Upland zone, R=
S= Submerged zone. (Submerged zone not sampled at 15 months.)


:Ruellia zone, and


126.85
10.66
1.73


0.0001
0.0001
0.0875


Zone
Month
Zone*Month










Table 3-4 ANOVA table for percent dormant of seed total.
Source of Variation Degree of Freedom F Value P Value
Zone 2 4.91 0.0086
Month 5 8.04 0.0001
Zone*Month 9 2.97 0.0028






















1~11~1 111111 11111


100%


-o~


20%


IT-1 I-3 IT-6 IT-9 11-12 11-15


R-1 R-3 R-6 R-9 R-12 R-15
Zone-Month


S-1 S-3 S-6 S-9 S-12


O # Missing # Dead El Total # Germinated


S# Dormant


Figure 3-7. Percentages of total seed (50) that were recovered from the field that were 1)
missing, 2) dead, 3) germinated, or 4) viable but presumed dormant. The Zones are
represented by U= Upland R= Ruellia, S= Submerged. The numbers represent the
time buried, e.g. 9=nine months buried.













10 -a
ab ab
ab
90 -1 ab


80-
b b
ab
70 -1 bc

5 Month 1
S60 Month 3
I FF I I Month 6
~n I 1 [OMonth 9
50 -o I c 5 Month 12
.H Month 15

S40-


30-a

ab



20




Upland Ruellia Submerged



Figure 3-8. Mean percent viable seed for 10 replicate samples per zone. (Percent Viable included
seeds that germinated in bag/chamber and dormant seeds.) Different letters indicate
significant differences at ac of 0.05, within a zone over time.









CHAPTER 4
FIELD STUDY

Introduction

Persistence and spread of most plant species is commonly limited by a combination of seed

and microhabitat availability (Eriksson and Ehrlen 1992; Scherff et al. 1994; Kollmann et al.

2007). Scherff et al. (1994) proposed that all it would take to overcome these limitations and

expand a species' distribution is successful seed dispersal coupled with opportunistic seedling

growth in a new habitat. Understanding what limits persistence and spread is of great importance

for understanding distribution of invasive non-native species. Currently, however, the specific

seed and microhabitat limitations are unknown for most non-native plants (Turnbull et al. 2000;

Kollmann et al. 2007).

Initial establishment processes are thought to be of great importance in determining the

distribution of plants. Several studies (cited by Foster 1999) indicate that it is in the early

establishment stages of life history when a plant may be most sensitive to competition. Studies of

colonization potential beyond the boundaries of an existing plant population are typically

achieved by comparing the establishment of seedlings from seeds transplanted into foreign

microsites to that of controls planted into the home habitat (Scherff et al. 1994).

The overall objective of the Hield study was to investigate whether the observed Hield

distribution of R. tweediana could be explained 1) by limitations due to environmental

conditions: soil moisture, light, and soil temperature, and/or by the biotic interactions of

competition with native plants and 2) at which life-stage, seeds or seedlings, these limitations

were most influential. The first component of the present study was to determine if seeds of R.

tweediana could germinate in the Upland or Ruellia zone. If they germinated, the next

component was to determine if the seedlings could survive over a three month period of time.










The third component was to determine if transplanted seedlings could survive in the Upland or

Ruellia zone.

Hypotheses

Seed Germination and Survival Experiments

1. Null Hypothesis: Plots will have the same mean Gap Fraction at both levels of Zone and/or
Competition.
Alternative Hypothesis: Plots will NOT have the same mean Gap Fraction at both levels of
Zone and/or Competition.

2. Null Hypothesis: R. tweediana seeds will have the same percentage germination at both
levels of Zone and/or Competition.
Alternative Hypothesis: R. tweediana seeds will NOT have the same percentage
germination at both levels of Zone and/or Competition.
If Alternative Hypothesis is accepted then:
0 Null Hypothesis: Values of Light, Soil moisture, and/or Soil temperature will be
the same for at both levels of Zone and/or Competition
Alternative Hypothesis: Values of Light, Soil moisture, and/or Soil temperature
will NOT be the same at both levels of Zone and/or Competition.

3. Null Hypothesis: Germinated R. tweediana seedlings will have the same percentage survival
at both levels of at both levels of Zone and/or Competition.
Alternative Hypothesis: Germinated R. tweediana seedlings will NOT have the same
percentage survival at both levels of Zone and/or Competition.
If Alternative Hypothesis is accepted then:
0 Null Hypothesis: Values of Light, Soil moisture, and/or Soil temperature will be
the same for at both levels of Zone and/or Competition.
Alternative Hypothesis: Values of Light, Soil moisture, and/or Soil temperature
will NOT be the same at both levels of Zone and/or Competition.

Seedling Transplant Experiment

4. Null Hypothesis: Plots will have the same mean Gap Fraction at both levels of Zone, Size,
and/or Competition.
Alternative Hypothesis: Plots will NOT have the same mean Gap Fraction at both levels
of Zone, Size and/or Competition.

5. Null Hypothesis: Transplanted R. tweediana seedlings will have the same percentage
Survival, Growth, Root biomass, and/or Shoot biomass at both levels of Zone, Size and/or
Competition.
Alternative Hypothesis: R. tweediana seedlings will NOT have the same percentage Survival,
Growth, Root biomass, and/or Shoot biomass at both levels of Zone, Size and/or
Competition.
If Alternative Hypothesis is accepted then:









o Null Hypothesis: Values of Light, Soil moisture, and/or Soil temperature will be
the same for at both levels of Zone, Size and/or Competition
Alternative Hypothesis: Values of Light, Soil moisture, and/or Soil temperature
will NOT be the same at both levels of Zone, Size and/or Competition.

Material and Methods

Study Site and General Plot Establishment

Research was conducted on small tributaries of Sweetwater Branch Creek located in

Payne's Prairie Preserve State Park, Alachua County, Florida. Ten replicate sites were located

during May 2006 based on the presence of the Upland, Ruellia, and Submerged zones. A transect

was established from the center of the Submerged zone, through the Ruellia zone, and 5 m into

the Upland zone at each of the ten replicate sites. In the experimental design, Zone was the main-

plot factor with two treatments: Upland and Ruellia (Submerged zone was excluded from these

experiments). Within each replicate site, six 1 m2 plOts were laid out on a transect parallel to the

zone orientation with a 0.5 m wide buffer surrounding each plot as seen in Figure 4-1. These

plots were paired (2 x 1 m2 plOts) for the sub-plot factor, which was Competition, with two

treatments: Cleared and Not-Cleared (Figure 4-2). In the Cleared treatment all vegetation was

removed with a gas-powered weed trimmer to ground level. One week later all regrowth and

remaining vegetation was removed by hand, resulting in a vegetation-free plot, and this condition

was maintained throughout the study. Vegetation was not disturbed in the Not-Cleared treatment.

Prior to vegetation removal within the Cleared treatment, a 0.25 m2 area was sampled where

species were recorded and collected for above ground biomass. At the end of the study it was

repeated in the Not-Cleared treatment (Table 1-4, Appendix Table A-1 and Table B-1).

This created a set of three paired plots, with the members of the each pair immediately next

to one another while the three sets of pairs were not always adj oining one another (Figure 4-1).

Treatments were located randomly (Figures 4-1 and 4-3). Light intensity at the soil surface, soil









moisture, and soil temperature were recorded at each plot every two weeks during the study, for

use as covariables. Canopy photos were taken in the plots at the beginning of the Seed

Germination and Survival Experiments and Seedling Transplant Experiment for site

characterization. For details about the methods and instruments used see Chapter 3.

Seed Germination and Survival Experiments

Seed germination

A seedling germination study was conducted using the Hyve of the 10 replicate sites on the

eastern side of the Prairie study area. This was a split-plot experimental design with Zone (main

plot) and Competition (sub-plot) as factors. A 3 x 2 grid was created using string on a 1 m2 PVC

frame and was placed over one member of each paired plot. Six empty 10 cm diameter plastic

flower-pots, with bottoms removed, were inserted into the ground with one pot in each section of

the grid (Figure 4-4). Ten seeds were placed on the soil surface in five of the pots, leaving one

for a check to see if there was a seed bank already in the soil or if new seeds fell into the pots.

This study was conducted the first time from August 13 through October 4, 2006 with each plot

watered with 7.5 liters of creek water applied with a watering can on rain-free days for the first

two weeks. The study was conducted a second time October 4 through November 20, 2006 but

the plots were not watered. Seedlings were counted as they emerged and were removed from the

pots.

Seed germination and seedling survival

An experiment identical to the first (August 13-October 4) Seed Germination study was

conducted in the remaining fiye replicate sites on the western side of the Prairie study site from

August 13 through November 16, 2006. Pots were watered with 7.5 liters on rain-free days for

the first two weeks. Seedlings were counted bi-weekly but not removed.









Seedling Transplant Experiment

Soil from the Upland and Ruellia zones was collected May 23, 2006 and June 2, 2006. The

soil was air dried and sifted. Four hundred R. tweediana seeds were placed in seedling trays of

each soil type on each of June 1 and June 30, 2006 to create two age groups for the Size factor,

Large (15 cm tall) and Small (3 cm tall) seedlings. On June 30, Large seedlings (June 1 group)

were repotted into 10 cm wide, square pots and seedlings were transplanted to the field in

August.

A split-split plot experimental design was used with Zone as the main-plot factor with

Upland and Ruellia as treatments. The sub-plot factor was Competition with Cleared and Not-

Cleared treatments, and the sub-sub plot factor was Size with Large and Small seedlings. The

two sets of paired plots in each zone that were not used for the germination studies were used for

this study. A 4 x 3 grid was created using string on a 1 m2 PVC frame which was placed over

each plot (Figure 4-5). In one set of paired plots, 10 Large seedlings were randomly transplanted

into the grid squares and in the other set of paired plots the 10 Small seedlings were randomly

transplanted into the grid squares. Thus a total of 80 seedlings, 40 Large and 40 Small, were

planted in each replicate site (Figure 4-6 and 4-7). The five eastern replicate sites were planted

the week of August 1 and the western five replicate sites were planted the week of August 7,

2006. Seedling plots were watered with 7.5 liters of creek water on rain-free days for the first

two weeks to minimize transplant shock. This experiment was conducted over 3.5 months ending

on November 16. During this period, survival of seedlings was noted and biweekly

measurements of seedling height were recorded. On November 16, 2006 all surviving plants

were collected, separated into shoots and roots and dried before weighing for above-ground and

below-ground biomass.









Statistical Analyses

ANOVA of canopy photo measurements was used to investigate differences in Gap

Fraction between zones. A mixed-model ANOVA was used for hypothesis testing in the Seed

Germination and Survival Experiments and the Seedling Transplant Experiment. This was done

with SAS Mixed procedure for all experiments except the Seedling Survival Data, which was

analyzed as binary data, 1=1ived 0=dead, with the SAS Glimmix macro. Light intensity, soil

moisture, and soil temperature measurements were averaged over time and analyzed as

covariables with Proc Mixed. All statistical analyses were done with SAS Version 9.1 software

(SAS 2007). The Zone factor was analyzed as a one-sided test because of the preexisting

knowledge that the Ruellia treatment (the zone in which vast populations of mature R. tweediana

occur) would have high rates of seed germination and seedling survival.

Results

Seed Germination and Survival Experiments

Canopy photos. Canopy photographs revealed mean (+ SD) Gap Fractions of 0. 113

(+0.013) and 0. 112 (+0.005), respectively for the Zone treatments Upland and Ruellia, and 0. 113

(+0.009) and 0. 112 (+0.01), res ectively for the Com petition treatment Not-Cleared and Cleared.

This indicated an almost closed canopy with no significant difference between zone treatments

(p= 0.7) or competition treatments (p=0.9).

Checks. The purpose of the checks was to determine if there was germination from a seed

bank or if seeds were being deposited and germinating during the experiments. Only one

seedling was removed from one check pot during the first (August) germination trial. Given this

extremely rare occurrence, the checks were removed from all datasets before the analyses were

conducted.









Seed germination

During the first germination experiment, 1 of the 100 pots had germination over 100 % (11

seeds instead of 10 indicating the presence of an "outside" seed). During the course of the

experiments, a few pots were disturbed by armadillos, 3 in the first experiment and 1 in the

second. Data from these pots were removed from the analyses because there was a high number

of replicate pots and no pattern was found in the disturbance.

In the ANOVA for the first (watered) experiment, Zone (Upland, Ruellia) had a p-value of

0.06, with germination rates of 67.8 % (Ruellia) and 57.9 % (Upland) (Figure 4-8). In the second

experiment, Zone had a p-value of 0.04, with the Ruellia treatment having a higher percentage

germination rate (3 5.4 %) than the Upland treatment (21.2 %), Figure 4-9. The p-value for the

Competition factor in the first experiment was 0.37 and 0.56 for the second experiment and the

Zone x Competition interaction was 0.8 and 0.12, respectively for either the first or second

experiment.

In the Proc Mixed analysis all three covariables, light intensity, soil moisture, and soil

temperature, had an influence on the Zone factor in the first experiment, while only soil moisture

had an influence on the second experiment. If these covariables could be set to an equal level in

all treatments, the average germination would be the same between the zones in the first

experiment. The means for all the covariables light, soil moisture, and soil temperature are

presented in Table 4-1. The overall lower percent germination for the second experiment (note

different scales on y axis in Figures 4-8 and 4-9) was possibly because the seeds were not

watered, but could have been due to other factors. The analysis of the covariables supports the

suggestion that soil moisture was influential.









Seed germination and seedling survival

During the course of this experiment, 6 pots were disturbed by armadillos. For these pots,

no further data were collected and the data collected until that point were removed from the

analyses.

During the germination part of this experiment, the Competition factor had a p-value of

0.08, with the Cleared treatment having a higher percentage germination (65.4 %) than the Not-

Cleared treatment (55.0 %) (Figure 4-10). The factor of Zone, and the Zone x Competition

interaction had p-values of 0. 17 and 0.2, respectively. These results were unexpected since this

experiment was very similar to the first seed germination experiment except that it lasted one

month longer. The covariables that were analyzed using Proc Mixed did not have an influence on

this experiment.

At the end of the three months, 49 pots (out of the 100 pot total) had no surviving

seedlings. These 49 pots were excluded from the statistical analysis so the relationship between

factor level and survival could be assessed, (inclusion of the 49 pots would have created a zero-

inflated dataset, which is very complex to analyze).

Zone for percent survival had a p-value of 0.36, while Competition was a highly

significant factor (p= 0.0003), the Cleared having a larger percent survival (85 %) than Not-

Cleared treatment (40 %) (Figure 4-11). The Zone x Competition interaction had a p-value of

0.35. The covariables analyzed for this experiment did not show any influence on the study.

Seedling Transplant Experiment

Canopy photos. Canopy photographs revealed mean Gap Fractions that did not vary for

any factors of Zone (p= 0.8), Competition (p= 0.85), or Size (p=0.59). The average Gap Fraction

of 0. 114 (+0.011 SD) indicated an almost closed canopy.









Survival and Height. Of the 800 seedlings transplanted, 662 seedlings survived (83%

survival). Zone and Competition had p-values of 0.055 and <.0001, respectively (Table 4-2). The

mean survival was higher in the Ruellia treatments (0.85 + 0.36 SD) than it was in the Upland

treatment (0.81+ 0.39 SD). The mean survival was higher in the Cleared treatments (0.93 + 0.26

SD) than it was in the Not-Cleared treatment (0.73 + 0.45 SD). The interaction between the Zone

and Competition factors had a p-value of 0.002 (Table 4-2). In the Cleared treatment the survival

was higher in the Ruellia zone and in the Not-cleared treatment the survival was higher in the

Upland zone (Figure 4-12:A). The final interaction was between the Size and Competition

factors with a p-value of 0.047 (Table 4-2). In the Cleared treatment the survival was nearly the

same for with Large and Small plants while in the Not-Cleared treatment the survival of Large

was greater than Small plants (Figure 4-12:B).

In the ANOVA of the change in seedling height over the 3.5 month study, Zone, Size, and

Competition were all significant (p<.0001) (Table 4-3). The mean change in height was greater

in the Ruellia treatments (1.63 + 1.8 SD) than it was in the U land treatment (0.59 + 0.65 SD .

The mean change in height was greater in the Large treatments (1.49 + 1.78 SD) than it was in

the Small treatment (0.73 + 0.95 SD The mean chan e in hei ht was greater in the Cleared

treatments (1.53 + 1.76 SD) than it was in the Not-Cleared treatment (0.69 + 0.95 SD). The

interaction between the Zone and Competition factors had a p-value of 0.0003 (Table 4-3). The

difference in change of height between the Ruellia and Upland zones was greater in the Cleared

than the Not-Cleared treatment (Figure 4-13:A). The final interaction was between the Zone and

Size factors with a p-value of 0.0095 (Table 4-3). The difference change in height between the

Ruellia and Upland zones was greater in the Large than the Small seedling size (Figure 4-13:B).









Biomass. For shoot biomass, Zone had a p-value of 0.051 (Table 4-4). The mean dry shoot

biomass was higher in the Ruellia treatments (0. 108 g + 0.114 SD) than it was in the Upland

treatment (0.079 g + 0.84 SD). Size and Competition both had a p-value of <.0001 (Table 4-4).

The mean dry shoot biomass was higher in the Large treatments (0. 144 g + 0. 113 SD) than it was

in the Small treatment (0.042 g + 0.019 SD). The mean dry shoot biomass was higher in the

Cleared treatments (0. 129 g + 0. 115 SD) than it was in the Not-Cleared treatment (0.058 g +

0.068 SD). There was an interaction between the Zone, Size, and Competition factors with a p-

value of 0.007 (Table 4-4, Figure 4-14). In the Cleared treatment the dry shoot biomass was

higher in the Ruellia zone and in the Not-cleared treatment the dry shoot biomass was higher in

the Upland zone (Figure 4-14).

For root biomass, Size and Competition both had a p-value of <.0001 (Table 4-5). The

mean dry root biomass was higher in the Large treatments (0. 072 g + 0.050 SD) than it was in

the Small treatment (0.019 + 0.019 SD The mean dr root biomass was hi her in the Cleared

treatments (0.059 g + 0.051 SD) than it was in the Not-Cleared treatment (0.032 g + 0.037 SD).

There a interaction between the Zone, Size, and Competition factors with a p-value of 0.008

(Table 4-5). In the Cleared treatment the dry shoot biomass was higher in the Ruellia zone and in

the Not-cleared treatment the dry shoot biomass was higher in the Upland zone (Figure 4-15).

Covariables. In the analysis of the covariables light, temperature, and moisture, only

moisture had a significant effect on the Seedling Transplant Study. For seedling growth,

moisture influenced the zone treatment. For shoot biomass, moisture influenced one, two-way

interaction, Ruellia, Not- cleared Upland, cleared; and one three-way interaction, Ruellia,

Small, Cleared Upland, Large, Cleared. For root biomass, moisture influenced one two-way









interaction, Ruellia, Not Cleared Upland, Not-Cleared. The extent to which the covariable of

moisture influenced the experiment warrants further investigation.

Discussion

In the Seed Germination and Seedling Survival Experiments, the absence of seedlings in

all but one check pot suggests that natural seed rain and seed bank of R. tweedian2a was low

during the study period, indicating either little seed production during the 3-month study period,

dormancy, or possibly a general limitation in seed availability. When seeds are sown in the field,

germination occurs and juvenile seedlings are found. However, it must be noted that seeds that

were mixed with the soil in the greenhouse study (Chapter 2) had decreased rates of germination

compared to when they were on the soil surface. If there are strong seed limitation factors such

as seed production, seed dispersal, and/or seed predation, they are possibly important for

controlling spread ofR. tweediana. If seeds were present in the soil, dormancy was not broken

under the conditions of these studies. A seed bank study in the Upland zone is needed to

determine if R. tweediana has a seed bank there. A seed production and dispersal study needs to

be conducted in the existing mature populations of R. tweediana. Both of these experiments

could further explain the seed limitation that is occurring in the Upland zone.

Another factor that may be involved is that seed predation by the M~elanagromyza ruelliae

fly causes a 25 % reduction in seed viability and also inhibits the seed capsules from explosively

dehiscing (Huey et al. 2007). How widely this is occurring and its effect on seed production and

dispersal or seedbanks warrants further investigation.

The Competition factor did not have an effect in the first or second Seed Germination

experiment but it did in the Seed Germination and Seedling Survival experiment. This positive

effect of the disturbance of clearing away competing vegetation in both Ruellia and Upland

zones indicates a microhabitat limitation and suggests competitive suppression by the resident










vegetation. This is also seen in the Seedling Transplant data, where the survival of transplanted

seedlings, change in height, and root and shoot biomass were always higher in cleared plots and

were more self-limited by R. tweedian2a than by native vegetation. Survival rates of the seedlings

indicated that the seedling size or zone in which seedlings were transplanted had little influence

if plots were cleared (Figure 4-12). While there was microhabitat limitation, it was not

completely limiting the seedlings and they were still able to survive and grow in the Not-Cleared

plots.

Environmental conditions that were measured during the experiment to determine if they

explained the zonation of R. tweediana did not reveal conditions that prevented seed germination

or seedling emergence beyond the current population edge. Moisture did influence some of the

results of the experiments, for example in the second Seed Germination experiment where

germination was probably reduced by the absence of watering at the beginning of the

experiment, but the germination percentages were still above 20%.

Conclusions

If there is an area that has not been invaded by R. tweediana but is otherwise similar to the

study site at the Prairie and seed reaches that site, it would likely become invaded in a similar

manner as the Prairie, and the invasion would start around the waterways. The seeds would

germinate even if there was some native plant competition, and would survive and grow. There

are environmental conditions around the waterways that differ from the Upland zone, that could

not be specifically characterized, resulting in higher germination, survival, growth, and biomass.

But even with these high values in the Ruellia zone and the seed and microhabitat limitations that

were identified in the Upland zone, there was still some germination and survival in the Upland

zone when seeds or plants were sown there with competition from native vegetation. It seems

that the full extent of the invasion is not yet realized at this site and R. tweediana plants will









move into the Upland if seed reaches it, especially if there is limited competition. During the

experiment the beginnings of this expansion into the Upland by seedlings (10-15cm tall) were

observed, a few of which appeared in Upland plots (Appendix Table 1-A).

Also R. tweediana extensively reproduces vegetatively with rhizomes and layering and

rooting at the nodes (pers. obs.). There was extensive expansion, by layering, of populations into

the waterways when the water was low during 2006 and this changed the course of the water, as

a once moving waterbody became two separate pools. Germinated seeds were also monitored in

one of the creek beds that had little water for a few months. The seedlings survived for over a

month until they were completely submerged (and 2 seedlings survived for 2 months

submerged), but in one of the plots there was a ~25 cm long R. tweediana stem that layered into

the plot and never died.

The existing monocultures of mature R. tweediana plants with the abrupt line between

zones can suggest the population is no longer expanding. However, during the course of these

studies, vegetative expansion was observed into the Upland zone. A future study to

experimentally monitor the rate at which the population is expanding at this local scale would

allow better predictions of the extent and rate of spread into the Upland zone once the habitat

closest to the waterways has been colonized.

Determining the mechanism by which the habitat closest to the waterway (at this stage of

invasion in this site, the habitat that became the Ruellia zone) was initially colonized, was not an

objective of this study. However, the presence of R. tweediana within the study site and creeks

across Gainesville, Florida, is likely due to the widespread use of this popular ornamental plant

(pers. obs.), which creates a source of propagules and seeds in residential areas upstream of the

invaded sites. The mucilaginous gel produced by these seeds when wetted, promotes dispersal of










this nature by allowing the seeds to stay buoyant in the water. Once the seed leaves the water, the

gel dries and glues the seed to the soil (Gutterman et al. 1973).

While R. tweediana plants grown from seed are easy to kill with herbicides in greenhouse

experiments (pers. obs.) this is not necessarily true once it becomes vegetatively established in

natural communities (R. Stocker, UF, pers. comm.). This understanding, along with the

knowledge of R. tweediana's ability to invade and expand once it has arrived in a natural

community, means that early detection and rapid removal of new populations in natural areas

should be a priority. A large scale study to look at common characteristics of invaded sites across

Florida would be needed to get a complete understanding of the ecological and propagule

pressure factors that influence the invasion of new sites. The results presented herein can

contribute to improved predictions of where R. tweediana is likely to invade on a state-wide

basis in the future. Importantly, these results provide more definitive information to guide local

level predications about how R. tweediana invasions will proceed once the species becomes

established within a site.











































Small
SSeed


I__


I


Figure 4-1. Plot layout map for Seed Germination and Survival Experiments and Seedling
Transplant Experiment at Paynes Prairie Preserve State Park.


N





































Figure 4-2. Example of one set of paired plots in the Seedling Transplant Experiment with
competition factor applied. Left side of photograph = Not-Cleared treatment, right
side = Cleared treatment. This is within the Ruellia treatment, planted with a Small
seedling treatment.











Ilii



12r3rl


Smal Seed ings


Large Seed~mgs


Seed


Seed


Large: Seedlings


Small Seedimgs


Figure 4-3. Example of the plot layout for one replicate site.


Upland Zone


Ruellia Zone


I Cleared
I No3t Cleared







































Figure 4-4. Example of a Seed Germination Experiment plot within a Ruellia treatment and with
the Cleared treatment applied.











Water


Not Clear Cleared
xxx x
xX xxx x
xxx xxx
x x xxx
Small Small



Not
Cleared Cleared
xx xx
x xx xx x
xxx xxx
xx xx

Large Large


Ruellia Zone













Upland Zone


Figure 4-5. Example of one replicate site plot layout for the Seedling Transplant Experiment. In
this split-split plot experimental design. Zone is the main-plot factor, Competition is
the sub-plot factor, and Size is the sub-sub-plot factor.


Not
Cleared Cleared
xxx xx
xX x xxx
xx xxx
xx xxx

Large Large



Not
Cleared Cleared
xxx xxx
x x x x
xx x x
xxx xxx
Small Small





































Figure 4-6. Example of a plot from the Seedling Transplant Experiment with Upland, Cleared,
and Large seedling treatments.





































Figure 4-7. Example of a plot from the Seedling Transplant Experiment with Upland, Not-
cleared, and Small seedling treatments.













p = 0.06 Ap = 0.37 B
70


60-


i 50-


~40-


S30-






20



Ruellia Upland Cleared Not Cleared


Figure 4-8. Percent germination rates for the first experiment of the Seed Germination Study. A)
The dark grey bars represent the data averaged over the two treatments (Upland and
Ruellia) of the Zone factor. B) The light grey bars represent the data averaged over
the two treatments (Cleared and Not-Cleared) of the Competition factor. Bars
represent standard error and within-factor p-values are at the top.













p = 0.56


40-









~20-


10


Ruellia


Upland


Cleared


Not Cleared


Figure 4-9. Percent germination rates for the second experiment of the Seed Germination Study.
A) The dark grey bars represent the data averaged over the two treatments (Upland
and Ruellia) of the Zone factor. B) The light grey bars represent the data averaged
over the two treatments (Cleared and Not-Cleared) of the Competition factor. Bars
represent standard error and within-factor p-values are at the top.


Table 4-1. Means of covariates for Seed germination in field experiments.


Ruellia
Cleared
19.27


Ruellia
Not-Cleared
19.38


Upland
Cleared
19.30


Upland Units
Not-Cleared
19.32 Co

24.39 Clmols s' m-2
24 %
23.61 Co

12.32 Clmols s' m-2
28 %


First
experiment


Second
experiment


Soil Temp

Light
Soil Moisture
Soil Temp

Light
Soil Moisture


24.34
24
23.70

23.48
60


24.29
24
23.77

14.28
54


24.53
23
23.60

12.95
32


p = 0.04





Ruellia


Upland


Cleared


Not Cleared


Figure 4-10. Percent seed germination rates over three months in the Seed Germination and
Seedling Survival experiment. A) The dark grey bars represent the data averaged over
the two treatments (Upland and Ruellia) of the Zone factor. B) The light grey bars
represent the data averaged over the two treatments (Cleared and Not-Cleared) of the
Competition factor. Bars represent standard error and within-factor p-values are at the
top.


p = 0.17


p = 0.08











p = 0.36


Ruellia Upland Cleared Not Cleared


Figure 4-11. Average percent survival over three months in the Seed Germination and Seedling
Survival experiment. A) The dark grey bars represent the data averaged over the two
treatments (Upland and Ruellia) of the Zone factor. B) The light grey bars represent
the data averaged over the two treatments (Cleared and Not-Cleared) of the
Competition factor. Bars represent standard error and within-factor p-values are
indicated.


p = 0.0003
















0.95 -
0.9 -

0.85 --Rellia
0.8 EB Upland
0.75 -
0.7 -

0.65 -
0.6
Cleared Not-Cleared p=0.002


0.95
0.9

0.85 -9Larg
0.8 El Sal

0.75
0.7

0.65
0.6
Cleared Not-Cleared p=0.05


Figure 4-12. Seedling Transplant Experiment two-factor interactions of seedling Survival. A)
Zone x Competition interaction with p=0.002. B) Size x Competition interaction with
p=0.05.

Table 4-2. ANOVA table for seedling Survival over 3.5 months in Seedlinn Transolant Studv


Source of Variation


Degree of Freedom


F Value Pr > F
4.84 0.0553
1.3 0.2698
20.52 <.0001
1.8 0.1964
10.79 0.0022
4.23 0.0468


Zone 9
Size 18

Competition 37
Zone*Size 18

Zone*Competition 37
Size*Competition 37
Note: Not capable of calculating three-way interaction











3- A 3- B

2.5 -1 2.5



-9- Ruellia 15- -Ruli
.9-G-Upland .6 --Upland


0.5 -0.5 -~

0 0
Cleared Not-Cleared p=0.003 Large Small p=0.0095



Figure 4-13. Transplant study two-factor interactions of seedling change in height over 3.5
months. A) Zone x Competition interaction with p=0.003 B) Zone x Size interaction
with p=0.0095.

Table 4-3. ANOVA table for Increase in Height over 3.5 months in Seedling Transplant Study.
Source of Variation Degree of Freedom F Value Pr > F
Zone 18 26.05 <.0001
Size 54 19.23 <.0001
Competition 54 23.48 <.0001
Zone* Size 54 7.24 0.0095
Zone*Competition 54 14.68 0.0003
Size*Competition 54 1.28 0.2621
Zone* Size*Competition 54 2.75 0.1032












Ruellia


Upand


0.24

0.2

0.16


S0.12

0.04


0.24

0.2

-e Large 0.
S 9alla 0.12


0.04


-e Large
-9 Snall


Geared Nobt-Geared


Geared Nokt-Gleared


Feared


NJot-Geared


0.240

0.200

0. 160

0. 120

0.080

0.040

0.000


-0 Ruellia
-8 Upland


-e Ruellia

| 0 Upland|


Izy


Small


Isy


Small


Small


Large


0.240

0.200

0.160

0.120

0.080

0.040

0.000


0.240

0.200

0.160

0.120

0.080

0.040

0.000


SClea
-B-Not-Clear


SClea
-0-Not-Clear


~


Ruellia Upland


Ruellia Upland


Figure 4-14. Seedling Transplant Experiment average shoot biomass data. Three-way interaction:
A- separated by Zone factor; B- separated by Competition factor; C- separated by
Size factor.


0.240

S0.200


S0.120

S0.080

0.040

0.000













Raellia


QIl~and


A

0.1



S0.08



ooz




0


0.1


0.08


-9 Inige
-a sen al

002a

0~c


-9 Inigee
-- sI


n~emfier wnt-nema d


n~emfier wnt-nema


Feared


Not-Gleared


0.1




0.06




0.02


0


-9 Rxella
-0 UFpland a


-9 Rxella

| -9- pland|


I~nge


Snrall


Inige


Snrall




Sndll


Ins-e


0.08

0.06







0


0.1


0.08




0.02




0


-El Not-Gear |


-9 Clear
-8 Not-Glear


Rxella UFpland


Rxella UFpland


Figure 4-15. Seedling Transplant Experiment average root biomass data. Three-way interaction:
A- separated by Zone factor; B- separated by Competition factor; C- separated by
Size factor.










Table 4-4. ANOVA table for Shoot Biomass in Seedling Transplant Study
Source of Variation Degree of Freedom F Value Pr > F
Zone 9 5.02 0.0518
Size 18 87.82 <.0001
Competition 36 52.79 <.0001
Zone* Size 18 0.47 0.4999
Zone*Competition 36 32.47 <.0001
Size*Competition 36 6.00 0.0193
Zone* Size*Competition 36 8.11 0.0072

Table 4-5. ANOVA table for Root Biomass in Seedling Transplant Study
Source of Variation Degree of Freedom F Value Pr > F
Zone 9 0.00 0.9907
Size 18 158.30 <.0001
Competition 36 46.38 <.0001
Zone* Size 18 7.99 0.0112
Zone*Competition 36 24.70 <.0001
Size*Competition 36 5.77 0.0216
Zone* Size*Competition 36 7.87 0.0081













APPENDIX A
SPECIES LIST


Table A-1. Species recorded from a 0.25 m2 plOt from the Cleared (C) (Aug 2006) and Not-

Cleared (NC) (Nov 2006) treatments of the paired plots of the Seed Germination and Survival

Experiments and the Seedling Transplant Experiment. L = Large seedling treatment, SM = Small
seedling treatment, SE = Seed germination and survival experiments.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae

Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae
Bignoniaceae
Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
Oleaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae

Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae
Vitaceae


Upland
L SM SE
C NC C NC C NC


Ruellia
L SM
C NC C NC


SE
C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K. Jansen
var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. ex Bureau
Macfadyena unguis-cati (L.) A.H. Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis (L.)
R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar stvraciflua L.
Stachvs floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. var. nitidus
(Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.) Planch.
Vitis rotundifolia Michx.
species 1
species 2


x xx xx x




x


x xx xx x

x



xx xx x x


x x


xx x x


xX xxx x


x
x x xx x


xX xxx x


x x
x


x x












Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae

Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae

Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland Ruellia
L SM SE L SM SE
C NC C NC C NC C NC C NC C NC
Genus species
Ruellia tweediana Griseb.x x x x x x
Acer negundo L.x x
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen x x x x
Verbesina virginica L.x x x x x x
Youngia japonica (L.) DC.
Vernonia Schreb.x
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentryx x x xx x x


Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1
species 2


x x


x
xx x


x x












Table A-1. Continued.
Plot
Zone
Size
Competition
Family Genus i
Acanthaceae Ruellia
Aceraceae Acer ne
Anacardianceae Toxicoc
Apiaceae Hydroc
Araceae Colocar
Araliaceae Hedera
Asteraceae Acmell
Jansen
Asteraceae Verbesj
Asteraceae Youngi
Asteraceae Vernon
Asteraceae Elephar
Bignoniaceae Campsi
Bureau
Bignoniaceae Macfad
Gentry
Blechnaceae Woodw
Caprifoliaceae Sambuc
(L.) R.
Commelinaceae Comme
Cucurbitaceae Meloth
Dioscoreaceae Dioscol
Fagaceae Quercu
Fagaceae Querus
Hamamelidaceae Liquida
Lamiaceae Stachys
Lamiaceae Salvia I
01eaceae Ligustr
Oxalidaceae Oxalis
Phytolaccaceae Petiverj
Pinaceae Pinus L
Poaceae Dichan
Poaceae Grass #
Poaceae Grass #
Poaceae Grass #
Poaceae Grass #
Poaceae Grass #
Poaceae Oplism
Ranunculaceae Ranunc
nitidus
Rosaceae Prunus
Rosaceae Rubus i
Saururaceae Saururu
Smilacaceae Smilax
Ulmaceae Celtis o
Vitaceae Parthen
Planch.
Vitaceae Vitis ro
? ~species
? species


Upland
L SM
C NC C NC


Ruellia
SE L SM SE
C NC C NC C NC C NC


species
tweediana Griseb.
:gundo L.
dendron radicans (L.) Kuntze
otyle L.
sia sp. Schott
helix. L.
a oppositifolia (Lam.) R.K.
var. repens (Walt.) R.K. Jansen
ina virginica L.
a japonica (L.) DC.
ia Schreb.
ntopus elatus Bertol.
is radicans (L.) Seem. Ex

lyena unguis-cati (L.) A.H.

Jardia Sm.
cus nigra L. ssp. Canadensis
Colli
:lina sp. L.
ria pendula L.
rea bulbifera L.
s virginiana P. Mill.
nigra L.
rmbar styraciflua L.
Sfloridana Shuttlw. ex Benth.

um sinense Lour.

ia alliacea L.


thelium (A.S. Hitchc. & Chase)







enus hirtellus (L.) Beauv.
:ulus hispidus Michx. Var.
(Chapman) T. Duncan
serotina Ehrh.
argutus Link
Is cernuus L.
sp. L.
,ccidentalis L.
locissus quinquefolia (L.)

tundifolia Michx.


x xx xx x


xx x

x


xxxxxxxxxx


X X X X


X XX XX X


x x
xx xx xx x


x
X X X


X X












Table A-1. Continued.


Plot
Zone
Sze
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae


Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae


Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


4
Upland Ruellia
L SM SE L SM SE
C NC C NC C NC C NC C NC C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L. x
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L. x
Youngia japonica (L.) DC.
Vernonia Schreb. x
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry x
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Collix
Commelina sp. L. x
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)x
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan x
Prunus serotina Ehrh. x
Rubus argutus Link x
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L. x
Parthenocissus quinquefolia (L.)
Planch. x
Vitis rotundifolia Michx.
species 1 x
species 2


x xx xx x


x


xx xx x


x x





xx xx x


xx x x







x


x x x x


x xx x


xx xx x











x x












Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae

Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae

Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland
L SM
C NC C NC


Ruellia
L SM
C NC C NC


SE
C NC


SE
C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1
species 2


x x xx xx x




















x x



xX X x xxx x x xx x


x x x
x x x


x x
x


x x


x x x












Table A-1. Continued.
Plot
Zone
Size
Competition
Family Genus species
Acanthaceae Ruellia tweediana Griseb.
Aceraceae Acer negundo L.
Anacardianceae Toxicodendron radicans (L.) Kuntze
Apiaceae Hydrocotyle L.
Araceae Colocasia sp. Schott
Araliaceae Hedera helix. L.
Asteraceae Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Asteraceae Verbesina virginica L.
Asteraceae Youngia japonica (L.) DC.
Asteraceae Vernonia Schreb.
Asteraceae Elephantopus elatus Bertol.
Bignoniaceae Campsis radicans (L.) Seem. Ex
Bureau
Bignoniaceae Macfadyena unguis-cati (L.) A.H.
Gentry
Blechnaceae Woodwardia Sm.
Caprifoliaceae Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelinaceae Commelina sp. L.
Cucurbitaceae Melothria pendula L.
Dioscoreaceae Dioscorea bulbifera L.
Fagaceae Quercus virginiana P. Mill.
Fagaceae Querus nigra L.
Hamamelidaceae Liquidambar styraciflua L.
Lamiaceae Stachys floridana Shuttlw. ex Benth.
Lamiaceae Salvia L.
01eaceae Ligustrum sinense Lour.
Oxalidaceae Oxalis L.
Phytolaccaceae Petiveria alliacea L.
Pinaceae Pinus L.
Poaceae Dichanthelium (A.S. Hitchc. & Chase)
Poaceae Grass #1
Poaceae Grass #2
Poaceae Grass #3
Poaceae Grass #4
Poaceae Grass #5
Poaceae Oplismenus hirtellus (L.) Beauv.
Ranunculaceae Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Rosaceae Prunus serotina Ehrh.
Rosaceae Rubus argutus Link
Saururaceae Saururus cernuus L.
Smilacaceae Smilax sp. L.
Ulmaceae Celtis occidentalis L.
Vitaceae Parthenocissus quinquefolia (L.)
Planch.
Vitaceae Vitis rotundifolia Michx.
? species 1
? species 2


6
Upland Ruellia
L SM SE L SM SE
C NC C NC C NC C NC C NC C NC


x xx xx xx x
x


x x


x x


x x x


x x x


x xx x x


x x


x x x


x x x x


















x x
x xx xx xx x













Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae


Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae


Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland
L SM
C NC C NC


Ruellia
SE L SM
C NC C NC C NC


SE
C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1
species 2


x x
x x xX xx


x xx x


x x


x x
x


x x

X X x

xX x x


x x


x x
x x x


x x


x x x


xx xx x


xx xx x
x xx xx x

x x













Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae


Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae


Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland Ruellia
L SM SE L SM SE
C NC C NC C NC C NC C NC C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.


x x x







x x


x x x


x x x x


x x
x


x
x
x x
x


xX x xx xx x


x x


x x x
x x


Pinus L.
Dichanthelium (A.S. Hitchc. & Chase) x
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.x
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1 x
species 2


x
x xx x


x xx x


x xx x


x x
x


x x x


x
x


xx x












Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae


Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae


Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland
L SM
C NC C NC C


Ruellia
SE L SM
NC C NC C NC


SE
C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1
species 2


x xx xx x


xx x


x XX x x x x x x
x


x x


xx x x


x xx xx x


x x


xx x


x x












Table A-1. Continued.


Plot
Zone
Size
Competition
Family
Acanthaceae
Aceraceae
Anacardianceae
Apiaceae
Araceae
Araliaceae
Asteraceae


Asteraceae
Asteraceae
Asteraceae
Asteraceae
Bignoniaceae

Bignoniaceae

Blechnaceae
Caprifoliaceae

Commelinaceae
Cucurbitaceae
Dioscoreaceae
Fagaceae
Fagaceae
Hamamelidaceae
Lamiaceae
Lamiaceae
01eaceae
Oxalidaceae
Phytolaccaceae
Pinaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Ranunculaceae


Rosaceae
Rosaceae
Saururaceae
Smilacaceae
Ulmaceae
Vitaceae

Vitaceae


Upland Ruellia
L SM SE L SM SE
C NC C NC C NC C NC C NC C NC


Genus species
Ruellia tweediana Griseb.
Acer negundo L.
Toxicodendron radicans (L.) Kuntze
Hydrocotyle L.
Colocasia sp. Schott
Hedera helix. L.
Acmella oppositifolia (Lam.) R.K.
Jansen var. repens (Walt.) R.K. Jansen
Verbesina virginica L.
Youngia japonica (L.) DC.
Vernonia Schreb.
Elephantopus elatus Bertol.
Campsis radicans (L.) Seem. Ex
Bureau
Macfadyena unguis-cati (L.) A.H.
Gentry
Woodwardia Sm.
Sambucus nigra L. ssp. Canadensis
(L.) R. Colli
Commelina sp. L.
Melothria pendula L.
Dioscorea bulbifera L.
Quercus virginiana P. Mill.
Querus nigra L.
Liquidambar styraciflua L.
Stachys floridana Shuttlw. ex Benth.
Salvia L.
Ligustrum sinense Lour.
Oxalis L.
Petiveria alliacea L.
Pinus L.
Dichanthelium (A.S. Hitchc. & Chase)
Grass #1
Grass #2
Grass #3
Grass #4
Grass #5
Oplismenus hirtellus (L.) Beauv.
Ranunculus hispidus Michx. Var.
nitidus (Chapman) T. Duncan
Prunus serotina Ehrh.
Rubus argutus Link
Saururus cernuus L.
Smilax sp. L.
Celtis occidentalis L.
Parthenocissus quinquefolia (L.)
Planch.
Vitis rotundifolia Michx.
species 1
species 2


x xx xx x


x x


x xx xx xx x


x
x x x


x x
x




x


x x


x x


x
xx x











APPENDIX B
SPECIES BIOMASS

Table B-1i. Biomass and the number of Ruellia tweediana stems in a 0.25 m2 plOt cleared at setup
of experiment in Cleared treatment (Aug. 2006) and in the Not-Cleared at the end of the
experiment (Nov. 2006).
Plot Zone Size Competition Ruellia weight Others weight Ruellia count
1 U L C 0.0 18.0 0
1 U L NC 0.0 4.6 0
1U SM C 0.0 14.0 0
1U SM NC 0.0 16.4 0
1 U SE C 0.0 16.0 0
1 U SE NC 0.0 20.3 0
1R L C 102.0 14.0 68
1R L NC 51.4 3.0 47
1 R SM C 74.0 8.0 62
1R SM NC 8.4 0.1 19
1R SE C 58.0 2.0 49
1 R SE NC 47.3 0.1 43
2 U L C 0.0 10.0 0
2 U L NC 0.0 8.8 0
2 U SM C 0.0 8.0 0
2 U SM NC 0.0 3.5 0
2 U SE C 0.0 22.0 0
2 U SE NC 0.0 9.0 0
2 R L C 58.0 8.0 49
2 R L NC 46.3 6.4 34
2 R SM C 84.0 4.0 56
2 R SM NC 38.6 0.0 44
2 R SE C 58.0 0.0 57
2 R SE NC 26.3 5.5 38
3 U L C 0.0 28.0 0
3 U L NC 0.0 38.3 0
3 U SM C 0.0 26.0 0
3 U SM NC 0.0 33.6 0
3 U SE C 0.0 26.0 0
3 U SE NC 0.0 13.8 0
3 R L C 44.0 2.0 35
3 R L NC 80.7 0.9 89
3 R SM C 112.0 2.0 64
3 R SM NC 70.8 3.1 64
3 R SE C 92.0 2.0 57
3 R SE NC 56.5 0.4 64
4 U L C 0.0 46.0 0
4 U L NC 0.0 37.8 0
4 U SM C 0.0 30.0 0
4 U SM NC 0.0 17.1 0











Table B-1. Continued.
Plot Zone Size Competition Ruellia weight Others weight Ruellia count
4 U SE C 0.0 26.0 0
4 U SE NC 0.0 20.9 0
4 R L C 170.0 4.0 115
4 R L NC 143.7 0.0 125
4 R SM C 126.0 0.0 106
4 R SM NC 124.0 0.0 132
4 R SE C 224.0 2.0 169
4 R SE NC 132.2 0.1 116
5 U L C 0.0 12.0 0
5 U L NC 0.0 2.71
5 U SM C 0.0 4.0 0
5 U SM NC 0.0 13.9 0
5 U SE C 0.0 16.0 0
5 U SE NC 0.0 2.6 0
5 R L C 108.0 2.0 62
5 R L NC 63.2 0.0 60
5 R SM C 58.0 0.0 59
5 R SM NC 76.9 0.0 49
5 R SE C 76.0 2.0 56
5 R SE NC 53.6 0.1 50
6 U L C 0.0 0.0 0
6 U L NC 0.0 22.0 0
6 U SM C 0.0 2.0 0
6 U SM NC 0.0 11.8 0
6 U SE C 0.0 22.0 3
6 U SE NC 0.0 1.9 1
6 R L C 46.0 34.0 25
6 R L NC 73.3 0.1 56
6 R SM C 234.0 4.0 96
6 R SM NC 119.8 0.1 68
6 R SE C 102.0 0.0 missing
6 R SE NC 104.1 0.05 95
7 U L C 0.0 14.0 0
7 U L NC 0.0 9.6 0
7 U SM C 0.0 12.0 0
7 U SM NC 0.0 14.7 0
7 U SE C 0.0 11.0 0
7 U SE NC 0.0 13.2 0
7 R L C 76.0 4.0 49
7 R L NC 54.5 2.8 50










Table B-1. Continued.
Plot Zone Size Competition Ruellia weight Others weight Ruellia count
7 R SM C 18.0 12.0 15
7 R SM NC 12.4 5.3 26
7 R SE C 44.0 4.0 44
7 R SE NC 52.3 0.8 73
8 U L C 0.0 10.0 0
8 U L NC 0.0 14.1 0
8 U SM C 0.0 22.0 0
8 U SM NC 0.0 0.1 0
8 U SE C 0.0 20.0 0
8 U SE NC 0.0 15.3 0
8 R L C 32.0 2.0 19
8 R L NC 57.8 0.1 52
8 R SM C 60.0 4.0 96
8 R SM NC 43.0 1.5 59
8 R SE C 54.0 12.0 28
8 R SE NC 89.6 0.4 49
9 U L C 0.0 6.0 0
9 U L NC 0.0 14.2 0
9 U SM C 0.0 28.0 0
9 U SM NC 0.0 11.2 0
9 U SE C 0.0 2.0 0
9 U SE NC 0.0 0.5 0
9 R L C 86.0 2.0 60
9 R L NC 87.7 2.6 81
9 R SM C 74.0 6.0 64
9 R SM NC 69.1 0.3 77
9 R SE C 62.0 0.0 62
9 R SE NC 71.8 0.9 57
10 U L C 0.0 28.0 0
10 U L NC 0.0 35.5 0
10 U SM C 0.0 10.0 0
10 U SM NC 0.0 22.2 0
10 U SE C 0.0 16.0 0
10 U SE NC 0.05 7.5 1
10 R L C 130.0 4.0 55
10 R L NC 91.8 2.4 48
10 R SM C 84.0 6.0 45
10 R SM NC 56.5 1.6 47
10 R SE C 142.0 2.0 96
10 R SE NC 128.1 0.05 67










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BIOGRAPHICAL SKETCH

Karen V. S. Hupp, the daughter of Stephen and Kathryn Shepherd was born on 1981, in

Chicago, Illinois. The third of four children who mostly grew up in Spartanburg, South Carolina,

she graduated from Paul M. Dorman High School in 1999. She received a BS degree in

Environmental Science from Catawba College in North Carolina in 2003. During that program

she worked on invasive plant removal at the Cowpens National Battle Field and conducted

research on the restoration of granite outcrops at Dunn's Mountain, NC. Since September 2003,

Karen has worked as an OPS assistant with Alison Fox's research team, providing excellent

support to many proj ects related to invasive plants. In Fall 2005, she started her MS program at

the University of Florida with the obj ective of further increasing her knowledge of invasive plant

ecology and management. Also that fall she married Jason R. Hupp on September 18.





PAGE 1

1 INVESTIGATING THE DETERMINANTS OF LOCAL SCALE DISTRIBUTION OF RUELLIA TWEEDIANA (SYNONYM R. BRI TTONIANA) IN NATURAL AREAS. By KAREN VICTORIA SHEPHERD HUPP A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2007

PAGE 2

2 2007 Karen Victoria Shepherd Hupp

PAGE 3

3 To my parents who always knew I had it in me and to my husband who was there through th e stressful times of this degree.

PAGE 4

4 ACKNOWLEDGMENTS I thank the members of my supervisory comm ittee for their ideas, time, and mentoring. I thank Alison Fox, for encouraging me to take th is path in my life and making it actually possible for me, and for help every step of the way. I th ank Randall Stocker, for being extremely available for questions and meetings and opinions. I thank Dr. Ramon Littell, for the many helpful meetings to help me understand the statistical aspects of my thesis. I thank Sandy Wilson, for the interest she carries with this sp ecies. I thank my family for thei r support, which kept me going to complete this degree. To Jenn and Denise who we re my cheerleaders. And this degree would not have been possible without the help and friendship of Lisa Huey.

PAGE 5

5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........7 LIST OF FIGURES................................................................................................................ .........8 ABSTRACT....................................................................................................................... ............11 CHAPTER 1 INTRODUCTION..................................................................................................................13 Study Site and Zones........................................................................................................... ...17 Research Questions............................................................................................................. ....18 2 SEED VIABILITY IN SUBMERGED CONDITIONS AND SOIL EFFECTS ON SEED GERMINATION.........................................................................................................27 Introduction................................................................................................................... ..........27 Hypotheses..................................................................................................................... .........28 Seed Viability in Submerged Conditions........................................................................28 Soil........................................................................................................................... ........28 Materials and Methods.......................................................................................................... .29 Seed Viability in Submerged Conditions........................................................................29 Soil........................................................................................................................... ........29 Soil effects on seed germination..............................................................................29 Soil characteristics....................................................................................................30 Statistical Analyses..........................................................................................................30 Results........................................................................................................................ .............31 Seed Viability in Submerged Conditions........................................................................31 Soil........................................................................................................................... ........31 Soil effects on seed germination..............................................................................31 Soil characteristics....................................................................................................32 Discussion..................................................................................................................... ..........32 Seed Viability in Submerged Conditions........................................................................32 Soil........................................................................................................................... ........33 Soil effects on seed germination..............................................................................33 Soil characteristics....................................................................................................33 3 SEED BURIAL.................................................................................................................... ..41 Introduction................................................................................................................... ..........41 Hypotheses..................................................................................................................... .........43 Materials and Methods.......................................................................................................... .43

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6 Study Site and Zone Characterization.............................................................................43 Seed Burial Study............................................................................................................45 Statistical Analyses..........................................................................................................46 Results........................................................................................................................ .............46 Zone Characterization.....................................................................................................46 Seed Burial Study............................................................................................................47 Discussion..................................................................................................................... ..........48 Zone Characterization.....................................................................................................48 Seed Burial Study............................................................................................................49 4 FIELD STUDY.................................................................................................................... ...61 Introduction................................................................................................................... ..........61 Hypotheses..................................................................................................................... .........62 Seed Germination and Survival Experiments..................................................................62 Seedling Transplant Experiment.....................................................................................62 Material and Methods........................................................................................................... ..63 Study Site and General Plot Establishment.....................................................................63 Seed Germination and Survival Experiments..................................................................64 Seed germination......................................................................................................64 Seed germination and seedling survival...................................................................64 Seedling Transplant Experiment.....................................................................................65 Statistical Analyses..........................................................................................................66 Results........................................................................................................................ .............66 Seed Germination and Survival Experiments..................................................................66 Seed germination......................................................................................................67 Seed germination and seedling survival...................................................................68 Seedling Transplant Experiment.....................................................................................68 Discussion..................................................................................................................... ..........71 Conclusions.................................................................................................................... .........72 APPENDIX A SPECIES LIST................................................................................................................... ....91 B SPECIES BIOMASS............................................................................................................101 LIST OF REFERENCES.............................................................................................................104 BIOGRAPHICAL SKETCH.......................................................................................................108

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7 LIST OF TABLES Table page 1-1 Florida counties in which Ruellia tweediana occurs.........................................................20 1-2 List of reported na tural areas in which Ruellia tweediana occurs Florida........................21 1-3 List of conservation areas in which Ruellia tweediana occurs in South Florida...............22 1-4 Dry above ground biomass data for 1 m2 plot, Ruellia weight is R. tweediana and Others weight = all other species in plot. Ruellia count = number of R. tweediana stems found in 1 m2 plot....................................................................................................26 2-1 Soil analyses results for 10 replicates fr om Upland, Ruellia, and Submerged zones........37 2-2 Soil survey results of Ponoma Sa nd, depressional from Natural Resources Conservation Service and United St ates Department of Agriculture................................40 3-1 ANOVA table for percent missing of seed total................................................................55 3-2 ANOVA table for percent dead of seed total.....................................................................56 3-3 ANOVA table for percent ge rminated of seed total..........................................................57 3-4 ANOVA table for percent dormant of seed total...............................................................58 4-1 Means of covariates for Seed germination in fi eld experiments........................................83 4-2 ANOVA table for seedling Survival over 3.5 months in Seedling Transplant Study.......86 4-3 ANOVA table for Increase in Height ove r 3.5 months in Seedling Transplant Study......87 4-4 ANOVA table for Shoot Biomass in Seedling Transplant Study......................................90 4-5 ANOVA table for Root Biomass in Seedling Transplant Study........................................90

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8 LIST OF FIGURES Figure page 1-1 Map showing Gainesville creeks. Dark circle in the enlarged insert is the study site in Paynes Prairie Pres erve State Park...................................................................................23 1-2 Photo on left is aligned w ith diagram on right to indica te the labeling of the three zones, Upland, Ruellia, and Submerged, in th e field taken from the Submerged zone.....24 1-3 The edge between the Upland zone and Ruellia zone, from the perspective of the Upland zone, at Paynes Prairie Pres erve State Park, (January 2006)...............................25 2-1 Percent germination and percent viability of seeds over time in the Seed Viability in Submerged Conditions study. The percent germinated of seeds remaining at each time is represented in black and the percen t viable of seeds tested each time is represented in grey............................................................................................................ .35 2-2 Number of germinated seeds in the So il effect on seed germination experiments. Experiment 1 seed germination is presented in the hash marked bars. Experiment 2 is presented with black for s eeds on top of the soil and dark grey for seeds mixed in the soil....................................................................................................................... .........36 2-3 Soil parameters with different letters i ndicated significant differences among zones within parameters ( of 0.05), n=10..................................................................................38 2-4 Natural Resources Conservation Servi ce and United States Department of Agriculture map of soil series superimposed over an aerial photograph of a section of Paynes Prairie Preserve State Park. Soil type 25 indicates Pomona sand, depressional the area in which this field research was conducted.....................................39 3-1 Nylon mesh seeds bags a ttached to the botto m of surveyors flags with zip ties..............52 3-2 Seed burial plot, closest flags in Subm erged zone, center flags in Ruellia zone, background flags in Upland zone......................................................................................53 3-3 Average percent soil moisture for each zone February 2006 to March 2007. There were significant differences between zones at each sampling interval and averaged across time.................................................................................................................... .....54 3-4 Percent missing of seed total in each z one. U= Upland zone, R= Ruellia zone, and S= Submerged zone. Different letters indicat e significant difference between months after burial ( of 0.05) for % missing averaged over all zones. Line fitted to averages with an R2 = 0.93...............................................................................................................55 3-5 Percent dead of seed total in each zone. U= Upland zone, R= Ruellia zone, and S= Submerged zone. Percent dead were significan tly higher at each month after burial in the Submerged zone compared to the Upland and Ruellia zones......................................56

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9 3-6 Percent germinated of seed total in each zone. U= Upland zone, R= Ruellia zone, and S= Submerged zone...........................................................................................................57 3-7 Percentages of total seed (50) that we re recovered from the field that were 1) missing, 2) dead, 3) germinated, or 4) viable but presumed dormant. The Zones are represented by U= Upland R= Ruellia, S= Submerged.....................................................59 3-8 Mean percent viable seed for 10 replicate samples per zone.............................................60 4-1 Plot layout map for Seed Germinati on and Survival Experiments and Seedling Transplant Experiment at Paynes Prairie Preserve State Park...........................................75 4-2 Example of one set of paired plots in the Seedling Transplant Experiment with competition factor applied. Le ft side of photograph = NotCleared treatment, right side = Cleared treatment. This is within the Ruellia treatment, planted with a Small seedling treatment............................................................................................................. .76 4-4 Example of a Seed Germination Experiment plot within a Ruellia treatment and with the Cleared treatment applied............................................................................................78 4-5 Example of one replicate site plot layout for the Seedling Transplant Experiment. In this split-split plot experime ntal design. Zone is the main -plot factor, Competition is the sub-plot factor, and Size is the sub-sub-plot factor......................................................79 4-6 Example of a plot from the Seedling Tr ansplant Experiment with Upland, Cleared, and Large seedling treatments...........................................................................................80 4-7 Example of a plot from the Seedling Transplant Experiment with Upland, Notcleared, and Small seedling treatments..............................................................................81 4-8 Percent germination rates for the first experiment of the Seed Germination Study..........82 4-9 Percent germination rates for the second e xperiment of the Seed Germination Study.....83 4-10 Percent seed germination rates over th ree months in the Seed Germination and Seedling Survival experiment............................................................................................84 4-11 Average percent survival over three months in the S eed Germination and Seedling Survival experiment...........................................................................................................85 4-12 Seedling Transplant Experiment two-fact or interactions of seedling Survival.................86 4-13 Transplant study two-factor interacti ons of seedling change in height over 3.5 months......................................................................................................................... .......87 4-14 Seedling Transplant Experiment average shoot biomass data...........................................88 4-15 Seedling Transplant Experiment average root biomass data.............................................89

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10 LIST OF ABBREVIATIONS U Upland Zone R Ruellia Zone S Submerged Zone L Large plant treatment SM Small plant treatment SE Seed germination and survival experiments C Cleared plot treatment NC Not Cleared plot treatment

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11 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science INVESTIGATING THE DETERMINANTS OF LOCAL SCALE DISTRIBUTION OF RUELLIA TWEEDIANA (SYNONYM R. BRI TTONIANA) IN NATURAL AREAS. By Karen Victoria Shepherd Hupp December 2007 Chair: Alison Fox Major: Agronomy Ruellia tweediana is a non-native plant introduced to Florida prior to 1940 that has naturalized in disturbed uplands and wetlands of 8 states, the Virg in Islands, and Puerto Rico. A popular horticultural plant, R. tweediana is available in many nurseries across Florida, being very attractive to the homeowner because of its abi lity to grow in many environmental conditions, from a water plant to growing in well-drained, almost xeric conditions. Ob servations in natural areas suggest that the field distribution of R. tweediana was limited to a narrow band along the banks of waterways (a Ruellia zone between Up land and Submerged zones). The principle goal of this research was to determine why the natural area distribution of R. tweediana was limited to a narrow zone along the banks of waterways rather than in a broader di stribution that includes both wetter and drier adjacent hab itats. This research investigated whether 1): the observed distribution of R. tweediana could be explained by limitations due to environmental conditions: soil moisture, light, and soil temperature, and/or by the biotic interactions of competition with native and non-native plants; and 2) at which life-stage, seeds or seedlings, these limitations were most influential. In the seed burial study, the possibility of a pe rsistent seed bank was found in the Upland and Ruellia zones while a seed limitation was found in the Submerged zone due to

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12 quick reduction in seed viability over th e study period. In the seed germination study, R. tweediana seeds germinated in all tested soil type s at very high percentages. In the field, germination studies provided differing results. In the first two experiments the highest germination percentages occurred in Ruellia plot s where competition did not have an effect, and in the third experiment zone (Upland or Ruel lia) did not matter while competition did, with cleared plots having high er germination. When R. tweediana seedlings were transplanted, they survived once established, irrespec tive of competition or zone tr eatment. The highest survival occurred in the cleared plots regardless of seedling size or location of zone. From these experiments it was concluded that there was a se ed and microhabitat limitation in the Upland and Ruellia Zones, that once seeds are present they are capable of germinating and becoming juvenile seedlings, and when seedlings are pr esent they are capable of surviving once established.

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13 CHAPTER 1 INTRODUCTION The movement of plant species by humans fr om one continent to another has increased over time with the expansion of transportati on and commerce (DAntonio and Vitousek 1992). Both the frequency and number of species move d has increased (Mack et al. 2000). As these plant species are brought to new areas they may have characteristics that help them displace native species: effective reproduction and disper sal mechanisms needed for the new area, superior competitive ability, few to no herbivores or pathogens, ability to occupy vacant niches, and capability of altering the invaded habitats (Gordon 1998). The issue of invasive species in natural areas has received increasi ng attention for several reasons. Threatened or endangered species prot ected under the Endangered Species Act are at risk due to interference from, and predation by, invasive species (Pimentel et al. 2000). In addition, the cost of invasive sp ecies control has nega tively affected the budgets of many state and federal agencies, and private landholders. In the state of Florida, more than $300 million has been spent to control invasive plant species between 1980 and 2006 (Ferriter et al. 2006), and the state of Florida is currently spending $30 milli on annually to control invasive species (D. Schmitz, DEP, pers. comm.). There are approximately 17,000 na tive plant species in the Un ited States and about 2,500 species native to Florida (Pimentel et al. 2000; Pimentel et al. 2005). In contrast, about 25,000 non-native plant species have b een introduced for cultivation in Florida alone (Frank and McCoy 1995). Of these species introduced to Florida, more than 925 have escaped and become established in surrounding natural areas (G ordon 1998). Driving the importation of new nonnative species is a $15.2 billon environmental horticultural industry in Florida (Hodges and

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14 Haydu 2005). And finally, about 85% of all plants imported into the United States arrive thorough the Miami International Airport (Frank and McCoy 1995). Ruellia tweediana is a non-native species that was in troduced to Florida sometime before 1940 from Mexico. Its native distri bution in Mexico is San Luis Potosi, Tamaulipas, Hidalgo, Puebla, and Veracruz (GRIN 2004). It also extends into South America with a distribution in western Bolivia, Paraguay, Uruguay, and northeast ern Argentina, limited areas of southern Brazil, and in open areas of eastern Chaco, locate d in wet or periodically flooded, sunny places, such as ditches, streams, riverside lawns, or temporarily inundate d areas (Ezcurra 1993). Since its introduction it has naturalized in Alab ama, Florida, Georgia, Hawaii, Louisiana, Mississippi, South Carolina, Texa s, the Virgin Islands, and Puerto Rico (USDA 2007), in USDA hardiness zones 8 through 11 (Gillman 1999). The popularity of R. tweediana in the horticultural trade appears to have increased in recent years and it is availa ble for purchase in many nurseries across Florida. This species holds great attractio n to the homeowner because of its ability to grow in many environmental conditions, from a water plant to growing in well-drained, almost xeric conditions. The genus Ruellia is in the family Acanthaceae whic h contains approximately 256 genera (Zomlefer 1994). Ruellia is a large genus containing about 250 species found mostly in the tropics and subtropics (Long and La kela 1976). Five native species ( R. caroliniensis R. ciliosa R. noctiflora R. pedunculata subsp. pinetorum and R. succulenta) and three naturalized nonnatives ( R. malacosperma R. tweediana and R. ciliatiflora ) occur in Florida (Wunderlin and Hansen 2007). There are seven synonyms for Ruellia tweediana : Arrhostoxylon microphyllum Cryphiacanthus angustifolius Ruellia brittoniana R. coerulea (R. caerulea) R. ignorantiae R. microphylla, and R. spectabilis (Wunderlin and Hansen 2007) a nd eight cultivars: Chi Chi,

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15 Katie Pink, Katie Purple, Katie Variegated, Katie White, Morado Chi, Purple Showers, and Snow White (Wilson and Mecca 2003). A new cu ltivar Oh What a Feeling, is presumed to be a hybrid of R. tweediana x R. caroliniensis (S. Wilson, UF, pers. comm.) and is being sold by http://www.plantdelights.com. Ruellia tweediana is an erect herbaceous perennial with one to many stems which are either green or purple depending on the light conditions in which it grew (pers. obs.). Stems have prominent nodal swelling and are documented to grow up to 1 m tall (Tobe et al. 1998), but plants have been observed that are over 1.5 m tall (pers. obs.). Th e leaves are opposite, linear to lanceolate and up to 30 cm long and less than 2 cm wide. The fl owers are solitary or in fewflowered cymes on axillary peduncles with a lavende r or blue tubular corolla 2 to 4 cm long and five-lobed. Cleistogamy (self-fer tilization without flow er opening) is known to occur and is characterized by small, tubular greenish brown corollas (Long and Lakela 1976). Plants bloom throughout the year in Florida (Tob e et al. 1998). Capsules are 2 to 2.5 cm long, cylindrical, with suborbicular seeds about 2 mm wide (Long and Lakela 1976). Ruellia tweediana produces an average of 20.6 seeds per capsule, each weighi ng an average of 1.78mg (Wilson et al. 2004). Explosive dehiscence of the seed capsules results in seed dispersal distances from the parent plant of 2.5-3 m (Witztum and Sc hulgasser 1995). A common family trait, the seed coat of R. tweediana becomes mucilaginous when moistened (Z omlefer 1994). Some seed predation has been noted in Florida by the Melanagromyza ruelliae fly (Huey et al. 2007). Populations of R. tweediana have appeared in natural area s across the state of Florida in recent years. Wunderlin and Hansen (2007) reported R. tweediana in twenty-eight Florida counties (Table 1-1). The Florida Exotic Pest Plant Council Distribu tion Database (FLEPPC 2007a), reported R. tweediana in 19 different natural areas (T able 1-2) in five different

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16 community types: 1) pine flat woods, prairies, 2) hardwood (ha mmocks, tree islands, etc.), 3) freshwater marshes, 4) rivers, springs and 5) salt marsh. In 2001, FLEPPC upgraded R. tweediana from a Category II (potential problem) to Category I due to altering native plant communities by displacing native species, chan ging community structures or ecological functions, or hybridizing with natives and it s status has not changed since (FLEPPC 2007b). The Institute for Regional Conservations data base, The Floristic Inventory of South Florida Database Online contains distribution records of more than 2,400 species of plants in South Florida conservation areas. Ruellia tweediana is listed in 18 conservation areas in South Florida (Table 1-3) (Gann et al. 2007). Ruellia tweediana has been observed in other Florida natural areas which have not been repo rted to these databases (pers. obs.). According to the Institute for Food and Ag ricultural Science (IF AS) Assessment of the Status of Non-Native Plants in Floridas Natural Areas, R. tweediana is concluded to be invasive and not recommended by University of Florida-IFAS (UF-IFAS) faculty, in the north and central parts of Florida. In southern Fl orida this species may be recommended by UF-IFAS with caution but should be managed to prevent its escape (Fox et al. 2005). Experts questioned for the IFAS Assessment process indi cated that in several locations R. tweediana coverage constitutes 50% of the infested stratum and th at it is changing commun ity structure by adding a new stratum or increasing plant dens ity in the stratum by 5-fold. It is also likely that it is altering the hydrology within a community (C. Gantz, UF, pers. comm.). At Blackwater Creek in Hills borough County the population of R. tweediana is a very dense monoculture in the swamp next to the creek. While it is not growing in the permanent waterways, it is growing on expos ed sandbars in the flowing wate r (A. Fox, UF, pers. comm.). It is commonly found growing in narrow zones along waterways, for example at Frog Creek in

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17 Manatee County (L. Huey, UF, pers. comm.), Hogt own Creek and Paynes Prairie Preserve State Park in Alachua County (pers. obs.), Ocklawah a River in Marion County (L. Huey, UF, pers. comm.), and The North Fork Buffer Preserve in St. Lucie County (A. Fox, UF, pers. comm.). At Fakahatchee Strand State Park in Collier County, R. tweediana is growing down to the swamp edge along the side of well-traveled, dry lime-ro ck roads. Where the swamp has flooded into the road, R. tweediana can be found growing in the middle of the road (pers. obs.). At Tradewinds Park in Broward County, a population of R. tweediana is growing along the side of a horse trail in an old dry stream bed (pers. obs.). At L ong Key Natural Area in Broward County, it is persisting under the canopy of trees of an abandoned homestead (pers. obs.). In Seminole County at Lake Jessup, R. tweediana has formed a large monoculture under the cabbage palm hammock even with grazing and trampling by cows (per s. obs.). These observations suggest most R tweediana populations in natural area s are associated with wate r, and upland populations are likely to have persisted after deliberate planting and cultivation, or from populations that were established under wetter conditions. The native species R. caroliniensis grows in all but seventeen of Floridas counties (Wunderlin and Hansen 2007). Despite having this widespread distribution, it is typically found as only a few scattered plants in any one place, not in dense monocultures. Ruellia caroliniensis does not exhibit the narrow zonal wa ter-related distribution exhibited by R. tweediana although under cultivation the two speci es exhibit similar habitat tolerances (pers. obs.). Study Site and Zones Field research on R. tweediana was conducted near Gainesville Florida (Alachua County) in small tributaries of Sweetwater Branch Cr eek on approximately 1.33 hectares in Paynes Prairie Preserve State Park (F igure 1-1). Study sites were se lected so the survival of R. tweediana within the zone of R. tweediana monocultures could be compar ed with that in adjoining

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18 elevation zones in which R. tweediana is not found. The adjoining zones were either more upland with a diversity of native plants, or at a lower elevation and frequently/constantly submerged. These three zones will be called Rue llia, Upland, and Submerged respectively (Figures 1-2, 1-3). The Ruellia and Submerged zones were typically of 2-4 m wide while the Upland zone varied in size but was generally wide r than 10 m. On average (n = 60 per zone), the number of species in a 1 m2 plot of the Upland zone was 15.5 while in the Ruellia zone it was 6.7. The aboveground dry biomass for the Upland zone for R. tweediana was 0.0 g and other species was 62.8 g and for the Ruellia zone for R. tweediana was 321.2 g and other species was 12.4 g. The average number of R. tweediana stems per plot in the Upland zone was 0.3 and the 250.8 in the Ruellia zone (Table 1-4, Appendix Table A-1 and Table B-1). Research Questions Considering that R. tweediana is found in a range of horticul tural conditions, from aquatic to almost xeric, the principle goal of this re search was to determine why the natural area distribution of R. tweediana was limited to a narrow zone alo ng the banks of waterways rather than in a broader distribution that includes both wetter and drier adjacent habitats. This research investigated whether 1): th e observed distribution of R. tweediana could be explained by limitations due to environmental conditions: soil mo isture, light, and soil temperature, and/or by the biotic interactions of co mpetition with native and non-nativ e plants; and 2) at which lifestage, seeds or seedlings, these limitations were most influential. There were six experimental components: 1) a growth chamber experiment Seed Viability in Submerged Conditi ons and 2) a greenhouse experiment Soil Effects on Seed Germination are both described in Chapter 2. Four of the experiments were conducted in the field, 3) a Seed Burial experiment to assess the longevity of seed s in the soil, is described in

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19 Chapter 3; 4) a Seed Germination experiment, 5) a Survival experiment and 6) a Seedling Transplant experiment, are descri bed in Chapter 4, to address the primary question in the field.

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20 Table 1-1. Florida counties in which Ruellia tweediana occurs, (Wunderlin and Hansens online database, Atlas of Florid a Vascular Plants, 2007). Counties Alachua Levy Brevard Manatee Broward Marion Charlotte Miami-Dade Collier Monroe Mainland Escambia Orange Franklin Palm Beach Hendry Pinellas Highlands Putnam Hillsborough Sarasota Indian River Seminole Lake St. Lucie Lee Sumter Leon Volusia

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21 Table 1-2. List of reported natural areas in which Ruellia tweediana occurs Florida, (The Florida Exotic Pest Plant Council Distribution Database, 2007). Natural Areas County Tradewinds Park Broward Forman Wilderness Preserve Broward Myrtle Slough @ CR 74 Charlotte Big Cypress National Park Collier Fakahatchee Strand State Park Collier Blackwater Creek Hillsborough Marsh Branch Hillsborough East Side Canal Hillsborough Anclote River Hillsborough Ocklawaha River Marion Long Key State Park Monroe Lake Park Scrub Natural Area Palm Beach No. Jupiter Flatwoods Natural Area Palm Beach Anclote River Pasco Alligator Creek Conservation Area Pinellas Cameron Property (SJRWMD) Seminole Fort Mose State Park St. Johns No. Fork Buffer Preserve St. Lucie Center Hill Sumter

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22 Table 1-3. List of conservation areas in which Ruellia tweediana occurs in South Florida, (The Floristic Inventory of South Florida Database Online, Gann et al. 2007). Conservation Area Counties Arch Creek Park Miami-Dade Big Cypress National Preserve Collier, Miami-Dade, Monroe Caloosahatchee Regional Park Lee Camp Owaissa Bauer Miami-Dade Castellow Hammock Park Miami-Dade Collier-Seminole State Park Collier Deering Estate at Cutler Miami-Dade Dry Tortugas National Park Monroe Dupuis Reserve Martin, Palm Beach Enchanted Forest Park Miami-Dade Everglades National Park Collier, Miami-Dade, Monroe Fred C. Babcock-Cecil M. Webb W ildlife Management Area Charlotte Greynolds Park Miami-Dade Hugh Taylor Birch State Park Broward Little Hamaca Park Monroe Long Key/Flamingo Road Natural Area Broward Secret Woods Buffer and Nature Center Broward Simpson Park Miami-Dade

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23 Figure 1-1. Map showing Gainesville creeks. Dark circle in the enlarged insert is the study site in Pa ynes Prairie Preserve State Park, (City of Gainesville 2006, http://www.cityofgainesville.or g/comdev/plan/gis/gis_lib.shtm1 ).

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24 Figure 1-2. Photo on left is aligne d with diagram on right to indi cate the labeling of the three zones, Upland, Ruellia, and Submerged, in th e field taken from the Submerged zone. Upland Zone Ruellia Zone Submerged Zone

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25 Ruellia Zone Upland Zone Figure 1-3. The edge between the Upland zone and Ruellia zone, from the perspective of the Upland zone, at Paynes Prairie Pres erve State Park, (January 2006).

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26 Table 1-4. Dry above ground biomass data for 1 m2 plot, Ruellia weight is R. tweediana and Others weight = all other species in plot. Ruellia count = number of R. tweediana stems found in 1 m2 plot (Raw data in Appendix Table B-1). Factor Treatment n Ruellia weight Others weight Units Ruellia count Zone Ruellia 60 321.212.4g 250.8 Upland 60 062.8g 0.4 Size Large seedlings 40 160.445.2g 118 Small seedlings 40 154.436g 124.8 Seed experiment 40 167.231.6g 130.8 Competition Cleared (Aug. 06) 60 179.243.6g 123.6 Not-Cleared (Nov. 06) 60 14231.6g 125.6

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27 CHAPTER 2 SEED VIABILITY IN SUBMERGED CONDI TIONS AND SOIL EFFECTS ON SEED GERMINATION Introduction In horticultural situations, Ruellia tweediana can grow in a range of s ites from an aquatic state to well-drained, almost xeric conditions. Mature plants have also been shown to survive for several months growing in submerged conditions in laboratory trials (A. Fox, UF, unpublished data). But observations from sites across Florida have shown that this sp ecies is not found in the waterways but alongside them. A factor that migh t be causing this is a family trait of the Acanthaceae, a mucilaginous gel that forms around some species seeds when moistened. Gutterman et al. (1973) indicate that the gel ai ds long-distance disper sal of the seeds and adhesion of seeds to the soil surface. A previous germination trial on R. tweediana seeds, with and without the mucilaginous gel present, indi cated that there was no difference in percent germination (L. Huey, UF, unpublished). But Gutterma n et al. (1973) also i ndicated that this gel can inhibit germination when seeds are in exces s water certain species within this family. Ruella tweediana seeds produce a mucilaginous gel (pers. obs ), so it is possible that continuous submersion of seeds covered with this gel would inhibit germination in this species. There are no reports in the literature that this possibi lity has been examined for this species. Seeds can be in different physiological states after leaving the parent plant (Booth et al. 2003). Ruellia tweediana seeds do not exhibit primary dormancy but are instead non-dormant after leaving the parent plant (Wilson and M ecca 2003). After dispersal, seeds that become buried can have a forced quiescence due to e nvironmental conditions in the soil that are unfavorable for germinati on. It is not known whether R. tweediana seeds buried in unfavorable environmental conditions exhibit secondary dormanc y, where they still do not germinate even if placed in optimal germination conditions (Booth et al. 2003).

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28 For successful reproduction, germination of qui escent, non-dormant seed s has to occur at the appropriate season and soil de pth. Other factors that can a ffect germination include soil structure, which determines the distribution and av ailability of water, solutes and gases, and the chemical effects of minerals in the soil, although this is a poorly understood relationship. Although most inorganic ions in soils do not have any specific effect on seed germination, nitrate ions have been shown to stimulate germination of some seeds (Hilhorst and Karssen 2000). A submersion experiment was conducted to determine if submerged R. tweediana seeds were capable of germination. If not, th is could be limiting the distribution of R. tweediana in the constantly submersed sites. A second experiment tested the effects of se veral soil types including those from the three field site zones on R. tweediana seed germination. Chemical and physical characteristics of the field site soils were al so measured to provide information on the site conditions and for comparison if th ere were significant differences in seed germination from soil samples from the different zones. Hypotheses Seed Viability in Submerged Conditions 1. Null Hypothesis: The percentage of submerged seeds that germinate or survive during 50 days in deionized water will not be different then zero. Alternative Hypothesis: The per centage of submerged seeds that germinate or survive during 50 days in deionized water will be significantly different then zero. Soil 2. Null Hypothesis: Seeds will have the same per centage germination in each tested soil type. Alternative Hypothesis: S eeds will not have the same percentage germination in each tested soil type. 3. Null Hypothesis: If there are any significant diffe rences in seed germination in the soils from different sources, this will co rrelate significantly with differe nces in soil characteristics. Alternative Hypothesis: If there are any significant differences in seed germination in the soils from different sources, this will not correlate with differences in soil characteristics.

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29 Materials and Methods Seed Viability in Submerged Conditions Ruellia tweediana seeds were collected November 4, 2005 from Paynes Prairie Preserve State Park. All seeds were coll ected on the same day, mixed thor oughly, and stored together in a covered plastic container unde r refrigerated conditions. At beginning of the experiment (3/10/06), these seeds had a viab ility of 95% when tested with tetrazolium dye (Peters 2000 as described below). Ten seeds were submerged in oxygenated dei onized water in each of 10 replicate petri dishes, which were sealed with parafilm. Petri di shes were incubated at 20/30 C (12h/12h cycle) and a 12 hour day/night cycle. Ev ery ten days (3/10/06 4/20/06) all germinated seeds were removed from the Petri dishes and placed on Farfa rd soil (Conrad Fafard, Inc.) in a greenhouse. Two of the replicate Petri dishes were then removed from the experiment and all remaining ungerminated seeds were tested with tetrazolium to see if they were still viable but dormant. For this viability test, seeds were moved to a new pe tri dish lined with filter paper, they were cut laterally and stained with 1.0% concentration of tetrazolium and placed in a dark, 30 C incubator for 12 hours, following the Peters (2000) Tetrazolium Tes ting Handbook. An entirely stained embryo indicated that th e seed was viable but dormant. Soil Soil effects on seed germination Five replicates of soil slabs of approximately 30 cm x 30 cm x 6 cm depth were collected from Paynes Prairie Preserve St ate Park from all three zones. Soil slabs and Fafard soil (Conrad Fafard, Inc.) (used as a control so il) were placed in tr ays and air dried for ten days. Fifty seeds (from the same batch of seeds used in th e Seed Viability in Submerged Conditions experiment), were placed on t op of the soil in each of four replications per zone. The 5th soil

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30 replicate had no R. tweediana seeds added to test for any germination from the field seed bank. Trays were moistened, covered w ith plastic and re-watered as needed. During the 30 day period, trays were monitored at least weekly, and seed lings were counted and removed as they were observed. This experiment was repeated using the pr eviously described methods, but was expanded to include a soil mixing treatment. For the se ed mixing treatment, the soil was mixed to redistribute seeds to depths ranging from 1 cm to approximately 3 cm. The factor effects tested in this experiment were soil s ource and seed position in soil. Soil characteristics Ten replicate sites were chosen at Paynes Prairie Preserve State Park based on the presence of the Upland, Ruellia, and Submerged zones. A 1-liter volume of soil was collected from each zone for each of the ten sites and was analyzed for organic matter (OM), cation exchange capacity (CEC), so il pH, estimated nitrogen releas e, NO3-N (nitrates), readily available phosphorus, total inorganic phosphor us, water soluble phosphorus, exchangeable calcium, exchangeable hydrogen, exchangeable magnesium, exchangeable potassium, and exchangeable sodium. A second set of soil samples was collected from each zone in a random selection of 5 of the 10 replicat e sites and was analyzed for text ure. All analyses were conducted by A & L Southern Agricultural Laboratories, Inc, Pompano Beach, FL (http://www.al-labsplains.com). Statistical Analyses Analyses of variance (ANOVA) and Tukeys multiple comparison tests were performed to investigate differences in seed germination be tween the different soil sources and seed position in the soil, and to compare soil characteristic s from the different zones. Differences between

PAGE 31

31 means will be reported with an alpha ( ) value and ANOVA results will be reported with a pvalue. All statistical analyses were done with SA S Version 9.1 software (2007). Results Seed Viability in Submerged Conditions Out of the 100 seeds that were submerged in deionized water, 42 seeds were tested for viability of which 41 were viable. The other 58 seeds had germinated in the chamber during the 50 day study and had been placed on soil (Figur e 2-1). These seedlings were followed for 30 days after the last seed germinate d, and only 4 of these seedlings died. Soil Soil effects on seed germination In the first soil slab experiment, a total of 7 s eeds from the field seed bank germinated, 2 of which were in soil to which no seeds had been ad ded and 5 germinated from a visible capsule in one replication of the Ruellia zone soil to wh ich 50 seeds were added (this was known because the seedling count was greater than 50). In th e Upland, Ruellia, Submerged, and Control soil on average 91.0%, 94.5%, 95.5%, 88.6% of added seed s (assumes no resident seed germination) germinated respectively. These values are not sign ificantly different from one another (p = 0.2) (Figure 2-2). In the second experiment, 77%, 73.5%, 78% 88% of seeds germinated (assumes no resident seed germination) respectively in th e Upland, Ruellia, Submerged and Control soil and these values were not significantly different from one another at an of 0.05. Although the soil source in this experiment was not a significant factor, the soil mixing treatment was significant (p = 0.0001), with seeds on top of the soil havi ng a higher average germination (88.8 %) than mixed seeds (69.2 %) (Figure 2-2).

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32 Soil characteristics Soil pH, estimated nitrogen release, total inorganic phosphorus, water soluble phosphorus, exchangeable hydrogen, exchangeable magnesium and exchangeable potassium were not significantly different between zones (Table 21). While OM, CEC, NO3N (nitrates), readily available phosphorus, exchangeable calcium, an d exchangeable sodium were different among zones (Table 2-1, Figures 2-3). Specifically, cati on exchange capacity was highest in the Ruellia zone with lower, and similar values in the Up land and Submerged zone (Figure 2-3). Compared to optimal concentrations for crop growth, NO3 -N (nitrates) rates were Medium, Medium, and Low in the Upland, Ruellia, Submerged zones re spectively (Table 2-1 and Figure 2-3). Readily available phosphorus and exchangeable calcium we re above the optimum rate for crops yield in all zones (Table 2-1). Total inorganic phosphorus levels were all higher than expected in crop soils while water available phospho rus was very low for all zones (A & L Southern Agricultural Laboratories). High levels of salin ity can be a limiting factor on plant growth (Warrence et al. 2002). The sodium gradient on these transects had higher concentrations in the submerged zone and decreases to the Upland zone (Figure 2-3). Soil texture was not significan tly different among zones ( of 0.05). The soil classification is sand with an average composition of 91.5% sand, 5.6% silt, and 2.9% clay. Discussion Seed Viability in Submerged Conditions The maximum time that seed submersion was test ed was only 41 days instead of the expected 50 days, because all seeds had germinated by day 41. Therefore the null hypothesis, the percentage of submerged seeds that germinate or survive during 50 days in deionized water will not be different then zer o, has to be rejected.

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33 Soil Soil effects on seed germination It was excepted that there would be different germination rates among the zonal soil samples and the control soil, but that was not the case. Flooding and soil type effects on seed germination do not appear to be responsible for limiting the distribution of R. tweediana Soil characteristics Given that there were no differe nces in seed germination on the soil from different zones, it can be concluded that differences in soil charac teristics are not significantly influencing shortterm seed survival and seed germination. The NRCS (2007) soil series for the region within Paynes Prairie Preserve State Park where the field research was conducted is Pomona sand, depressional (Figure 2-4). This soil type has na tive vegetation composed primarily of cypress ( Taxodium distichum (L.) L.C. Rich.), swamp maple ( Acer rubrum L.), tupelo (Nyssa sylvatica Marshall var. biflora (Walter) Sar g.), bay, and some scattered pond pine ( Pinus serotina Michx.), and typical soil parameters as id entified in Table 2-2 (NRCS 2007). Aerial photographs of the study site from 1982 showed that it was completely surrounded by farmlands. This compares to the 2005 aerial photograph in Figure 2-4. A particularly interesting result in the soil analyses was that the Submerged zone had the lowest value for percent organic matter and that all zones were above the NRCS expected range. The soil pH was also higher than expected for this soil type (NRCS 2007) but horticulturists have reported that R. tweediana can grow within the found range (Daves Garden 2007). While there were significant differences between zones fo r cation exchange capacity, all values were considered within normal range fo r crop growth but were higher th an recorded for this soil type by NRCS (Table 2-2). Even with some of the soil parameters not being in the normal crop or

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34 NRCS soil survey ranges or having significant differences in the soil characteristics among zones, it appears that this did not affect seed survival and germination in the two experiments.

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35 0 10 20 30 40 50 60 70 80 90 100 10203040 Days Submerged# Germinated or Viable Figure 2-1. Percent germination and percent viability of seeds over time in the Seed Viability in Submerged Conditions study. The percent germin ated of seeds remaining at each time is represented in black and the percent viable of seeds tested each time is represented in grey.

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36 0 10 20 30 40 50 ControlUplandRuelliaSubmerged Soil TypeNumber Germinated Figure 2-2. Number of germinated seeds in the Soil effect on s eed germination experiments. Experiment 1 seed germination is presented in the hash marked bars. Experiment 2 is presented with black for seeds on top of the soil and dark grey for seeds mixed in the soil.

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37 Table 2-1. Soil analyses results for 10 replicat es from Upland, Ruellia, and Submerged zones. Analysis conducted by A & L Southern Agri cultural Laboratories, Inc. on May 15, 2006. Zone Soil Parameters Upland Ruellia Submerged Units Optimum crop nutrient rangeb A&L ratingsb Organic matter 8.22 8.69 6.9 % Sig.a Cation exchange capacity 18.56 24.53 15.31 meq/100g Sig. 5 to 35 Soil pH 7.6 7.17 6.76 NS 6.0-7.0 H, VH, H Estimated nitrogen release 208.3 217.7 197 Lbs./A NS VH, VH, VH NO3-N (nitrates) 13 7.55 5.35 ppm Sig. M, M, L Readily available phosphorus 76.8 39.3 58.1 ppm Sig. 20-30 VH, H, VH Total inorganic phosphorus 147.45 145.7 135.15 ppm NS 40-60 VH, VH, VH Water soluble phosphorus 0.44 0.45 0.27 ppm NS VL, VL, VL Exchangeable Calcium 3269.5 4322 2513.5 ppm Sig. VH, VH, H Hydrogen 0.29 0.44 0.63 meq/100g NS Magnesium 169.9 185.2 228.4 ppm NS 30-70 M, M, M Potassium 40.3 40.85 27.25 ppm NS 90-125 VL, VL, VL Sodium 35.25 100.4 111.5 ppm Sig. L, H, H aSig. = Significant (p=0.05), NS = Not Significant between zones (Figure 2-1) bMost soil test readings on the report are given a rating of very low (VL), low (L), medium (M), high (H), or very high (VH). The purpose of these readings is to provide a guideline for determining optimum nutrient levels for crop growth.

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38 0 20 40 60 80 100 120 % Organic matterCation exchange capacity meq/100g NO3-N ppmReadily Available Phosphorus ppm Calcium ppbSodium ppmFor units see 'x' axis Upland Ruellia Submerged ab a b a ab b a b ab b a b b a b b a a Figure 2-3. Soil parameters with different letter s indicated significant differences among zones within parameters ( of 0.05), n=10.

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39 Figure 2-4. Natural Resources Conservation Se rvice and United States Department of Agriculture map of soil series superimposed over an aerial photograph of a section of Paynes Prairie Preserve St ate Park. Soil type 25 indica tes Pomona sand, depressional the area in which this fi eld research was conducted.

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40 Table 2-2. Soil survey results of Ponoma Sand, depressional from Natural Resources Conservation Service and United States Department of Agriculture October 2006 (http://websoilsurvey.nrcs.usda.gov) Soil Parameters Units Effective cation-exchange capacity .3-3.5Meq/100g Soil reaction pH 3.5-5.5pH Salinity 0-2mmhos/cm Sodium adsorption ratio 0-4-USDA texture sand-Sand -a% Silt 0-15% Clay 1-6% Moist bulk density 1.2-1.5g/cc Saturated hydraulic conductivity 42.34-141.14Micro m/sec Available water capacity .05-.10In/In Organic mater 1-3% aValue not provided

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41 CHAPTER 3 SEED BURIAL Introduction A soil seed bank is the reserve of viable seeds present in the soil and on its surface (Roberts 1981). A transient seed bank is characteri zed by a lack of viable seeds after a year under field conditions; whereas a persistent seed ba nk has viable seeds afte r a year (Thompson and Grime 1979). Once the mature plant population is controlled, the management of an invasive species will be much easier if it has a transient seed bank. Re-inspe ction and re-treatment is only needed for two years to ensure that none of the germinating s eeds in the soil develop to produce seeds and replenish the seed bank (Magda et al 2004). Conversely, the management of a species with a persistent seed bank will be more long-te rm and depend directly on the longevity of the seed bank (Gardener et al. 2003; Swanton and Booth 2004). There is limited knowledge of the importance seed banks should have in invasive species management in natural areas (Swanton and B ooth 2004). This is because there are not many studies on seed banks in natural areas, but there is a large body of literature dealing with seed banks in agricultural systems due to the economic importance of agricultural weeds and the fact that many of them reproduce only by seeds (Robe rts 1981). These studies can be reviewed to help understand what might be occurring in natural systems. In agriculture it is common to apply the s hort-term management of removing the above ground biomass of weeds that are competing with crops and limiting the overall yield (Swanton and Booth 2004). In the management of the above ground vegetation of ag ricultural weeds, the crop is protected and all other plan ts are removed. This is achieved by selective herbicides that do not kill the crop and by altering of the genes of the crop to make them resistant to, and hence permit the use of, certain herbicides (e.g. R oundup Ready crops). This differs from natural

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42 ecosystems where only the invasive species need to be removed and all other plants need to be protected. This difference makes the use of herbicid es in natural areas diffi cult and the altering of genes not an appropriate concept. But Jordan et al. (1995) showed that s eed mortality had greater effect on weed populations in crops than plant mort ality or decrease in seed production per plant. The management of invasive species in natura l ecosystems should include the management of the species seed bank (Swanton and Booth 2004). For invasive species management it is importa nt to have an understanding of the seed viability, dormancy, and longevity of each invasive species (Gardener et al. 2003; Lutman et al. 2002; Mennan and Zandstra 2006). Understanding seed longevity help s to predict the rate of loss of seeds from the seed bank (Baskin and Baskin 2006). Dormancy is important because it is the reason that seeds can remain viable in the seed bank for extended lengths of time (Booth et al. 2003). As summarized by Baskin and Baskin (2006) one manner of determining longevity is based on inferring the seeds age based on site in formation. This can be done in archeological sites where viable seeds can be dated based on the layer of ear th in which the seeds are found. Age can also be inferred in agricultural fields where the field has been sown in one crop for an extended period of time and the seed s that lie beneath can be assume d to be older than the sown crop. Another method to determine longevity of a seed bank is to bury se eds in the soil, wait various periods of time, exhume the samples, a nd check the seeds for viability. Experiments of this nature have been conducted si nce the late nineteenth century. In a previous study it was determined that fewer than 2% of Ruellia tweediana seeds were viable after 13 months of burial in field condi tions (Barnett, unpublished data). This suggested that R. tweediana essentially has a transient seed bank and that long-term management needs

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43 would be reduced. The first compon ent of the present study was to replicate this previous work, with seeds of R. tweediana buried in the field in areas that contained mature populations of R. tweediana to determine their survival for at least fi fteen months. The second component of this study was to expand the area of interest to incl ude adjoining zones in the field that did not contain mature R. tweediana plants, to determine if seeds have different rates of germination or mortality in these zones and if these zones could maintain transient or persistent seed banks. The overall objective of this study was to de termine if the field distribution of R. tweediana was limited by seed bank dynamics. Hypotheses 1. Null Hypothesis: Plots will have the same mean physical characteristics such as: Gap Fraction, Light, Soil Moisture, and/or Soil Temper ature in the Upland, Ruellia, and Submerged zones. Alternative Hypothesis: Plots wi ll NOT have the same mean phys ical characteristics such as: Gap Fraction, Light, Soil Moisture, and/or So il Temperature in the Upland, Ruellia, and Submerged zones. 2. Null Hypothesis: Buried R. tweediana seeds will NOT survive for more than a year. Alternative Hypothesis: Buried R. tweediana seeds will survive for more than a year. 3. Null Hypothesis: Buried R. tweediana seeds will have the same number in these categories: Missing, Dead, Total Germinated, Dormant, a nd Total Viable in the Upland, Ruellia, and Submerged zones. Alternative Hypothesis: Buried R. tweediana seeds will NOT have the same number in these categories: Missing, Dead, Total Germinated, Dorm ant, and Total Viable in the Upland, Ruellia, and Submerged zones. Materials and Methods Study Site and Zone Characterization Ten replicate sites were located at Paynes Prairie Preserve State Park on December 15, 2005 based on the presence of adjacent Upland, Ru ellia, and Submerged zones. A transect was run from the center of the Submerged zone, th rough the Ruellia zone, and 5m into the Upland zone at each of the ten replicate sites. Three 0.25 m2 plots along these transects were created, one

PAGE 44

44 in the center of each zone. The experimental desi gn was Zone as the factor with three treatment levels Upland, Ruellia, and Submerged. Canopy photos and light measurements. Digital canopy photographs were taken above every plot with a Nikon Coolpix 450 0 digital camera on a tripod with a fish-eye lens attachment. The camera was leveled with the top faci ng north for each photograph on a non-sunny day. Photographs were analyzed with the Hemi View 2.1 (Delta-T Devices, Cambridge, UK) computer program to classify each pixel in each photograph as either sky or non-sky to calculate canopy closure, also know n as Gap Fraction. Gap Fracti on is a value that ranges from 0 (a closed canopy) to 1 (an open sky). HemiViews settings were calibrated to site conditions at Paynes Prairie Preserve State Park based on th e latitude, longitude, declination, and altitude. Photos were imported into the program, sized to include the whole image, aligned to have north in the photo match north in the program, and the threshold (brightne ss of photograph) was specified. Light measurements were taken with a LI-C OR LI-250 (LI-COR, Lincoln Nebraska) light meter. Readings were taken at ground/water leve l in the center of each plot by averaging the light readings over a 30 second period of time. Soil moisture and temperature measurements. Soil moisture measurements 6 cm deep were taken periodically duri ng the study with a Dynamax Th eta Probe type ML2x (Delta-T Devices, Cambridge, UK) attached to a Delta-T Mo isture Meter. The probe was calibrated to the site conditions following the manufacturers instru ctions. The measurements were recorded as a dimensionless parameter that is expressed as percent volume. Thus 0 % corresponds to a completely dry soil, and pure water gives a r eading of 100 %. Soil temperature measurements

PAGE 45

45 were taken at 8 cm depth with a VersaTuff Plus 396 Atkins (techniCAL Systems, Hamilton, Ontario) temptec probe (C). Seed Burial Study Ruellia tweediana seeds were collected November 4, 2005 from Paynes Prairie Preserve State Park. All seeds were coll ected on the same day, mixed thor oughly, and stored together in a covered plastic container under re frigerated conditions, and at the beginning of the experiment (March 2006) had a viability of 95% when tested with tetrazolium dye (Peters 2000 as described below). On March 9-10, 2006, fifty seeds were mixe d with ~4 g of sand (Quikrete Premium Play Sand which was sterilized at 100 C for five days ), then rolled tightly in a 15.2 cm by 15.2 cm section of nylon mesh and attached to a surveyor s flag with a zip tie (Figure 3-1). Sand was mixed with the seeds to decrease the spread of disease and fungus from on e seed to another (Van Mourik et al. 2005). On March 13, 2006, one seed bag was buried 10 cm deep in the ground at the center of each plot and the other nine bags were buried in a circle surrounding this bag. The area that the bags were buried in was approximately 0.25 m2 (Figure 3-2). Based on high rates of seed death in the Submerged zone in a preliminary study (not re ported), the number of sa mpling bags placed in the Submerged treatment was limited to five. One bag was removed from each plot after 1, 3, 6, 9, 12, and 15 months (March 2006 to June 2007). Bags were brought to th e lab, zip ties were cut off and the sand was washed from the seed bags. The number of seeds recovered was re corded, as was the number of seeds that had germinated in the ground. Recovered, ungerminated seeds were placed in a petri dish lined with filter paper, moistened with deionized water, and sealed with parafilm. Petri dishes were incubated at 20/30 C (12h/12h cycle) and a 12 hour day/night cycle for 30 days. These conditions have been shown to promote successful germination (Wilson and Mecca 2003).

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46 Germination was then recorded and all remain ing ungerminated seeds were tested with the vitality stain tetrazolium to see if they were sti ll viable but dormant. Seeds were moved to a new petri dish lined with filter paper, they were cu t laterally and stained with 1.0 % concentration of tetrazolium and placed in a dark, 30 C incu bator for 12 hours, following the Peters (2000) Tetrazolium Testing Handbook. The seeds were evalua ted for an entirely stained embryo which indicated that the seed was viable but dormant. Statistical Analyses Dependent variables related to the seeds were the percent missing, percent germination, and percent dead. Total viable seeds was the su m of the numbers of seeds germinating in soil, capable of germinating, and dormant. Analyses of variance (ANOVA) and Tukeys multiple comparison tests were performed to investigate changes in the dependent seed variables over time and the differences between zones in: ca nopy photos, light measurements, soil moisture, soil temperature, and dependent seed variables. Differences between means will be reported with an alpha value and ANOVA results will be reported w ith a p-value. All statistical analyses were done with SAS Version 9.1 software (SAS 2007). Results Zone Characterization Canopy photos and light measurements. Canopy photographs revealed mean (+ standard deviation) Gap Fractions of 0.1147 (+ 0.007), 0.115 (+ 0.009), 0.1153 (+ 0.0010), for the Upland, Ruellia, and Submerged treatments respectively. Th ese indicated an almost closed canopy with no significant difference between zones (p= 0.99). This was supported by the mean (+ standard deviation) measurements of light in the understory, 67.9 mols s-1 m-2 (+ 50.5), 60.3 mols s-1 m-2 (+ 28.3), and 50.2 mols s-1 m-2 (+ 15.1) for the Upland, Ruellia, and Submerged treatments respectively, which also showed no difference be tween the zones (p=.52). Full sunlight at solar

PAGE 47

47 noon on a clear summer day is 2224 mols s-1 m-2 (+ 90) in Miami, Florida (Lee and Downum 1991). Soil moisture and temperature measurements. Percent soil moisture was significantly different between each zone at each sampling inte rval and when averaged across the study period ( = 0.05) Figure 3-3. Soil moistures, averaged ove r all sampling intervals, were 32 %, 52 %, and 91 %, in the Upland, Ruellia, and Submerged treatments, respectively. Average soil temperature in the Upland trea tment (21.4 C) was significantly different from the Submerged (20.9 C) treatment ( of 0.05), while the Rue llia treatment (21.1 C) was not different from the Upland or Submerged treatments. Seed Burial Study Samples were exhumed from the field April 14, 2006 (Month 1), June 16, 2006 (Month 3), September 13, 2006 (Month 6), December 16, 2006 (Month 9), March 15, 2007 (Month 12) and June 19, 2007 (Month 15), (the last sampli ng time only included the Upland and Ruellia treatments because of the reduced number of sa mple bags in the Submerged zone). Overall, a low percentage of seeds were missing within e ach zone and there was no significant difference between zones for this variable. However, there was an increase in the number missing over the 15 month period of burial (F igure 3-4, Table 3-1). The percentage of seeds that were recovered bu t were dead varied significantly with burial time and were significantly high er each month in the Submerged treatment compared to the Upland and Ruellia treatments (Figure 3-5). There was a two-way interaction between treatment and burial time with a p-value of 0.058 (Table 32) because Upland and Ruellia zones were only different during some months. The same twoway interaction (p = 0.088) was found in the percentage of seeds that germinated (Table 3-3 and Figure 3-6) because Upland and Ruellia zones were similar during some months. Seeds that were viable but presumed dormant consisted

PAGE 48

48 of 3.2 % of the total seed. There was a two-way interaction betwee n treatment and burial time for dormant seeds (p = 0.003) (Figure 3-7, Table 34), again because differences between Upland and Ruellia zones only occurred at some sampling times. The percentage of total viable seeds (total viable seeds includ ed seeds germinating in soil, capable of germinating, and dormant) averaged over burial times were 74.9 % for the Upland, 66.3 % for the Ruellia, 8.9 % for the Submerge d (Figures 3-7 and 3-8). During the first 12 months the percentages of vi able seeds for the Upland and Ruellia treatments were not significantly different ( of 0.05), while the Submerged treatm ent was different from them both. There was a significant difference between the Upland and Ruellia treatments during month 15 (the Submerged treatment was not sampled), with higher viability in the Upland treatment. In the Upland and Ruellia treatments, by nine months afte r burial, percent total viable was significantly reduced from its highest percenta ge (three months after burial in the Upland and one month after burial in the Ruellia treatment Figure 3-8). Discussion Zone Characterization Canopy photos and light measurements were take n for site and zone characterization and to determine if differences in light could explain the fi eld distribution of R. tweediana. All habitats, however are under the same cypre ss canopy and no differences in Gap Fraction and light measurements were found. Thus, the first null hypothesis has to be accepted for these two variables. Soil moisture decreased and soil temperature increased moving from the Submerged zone to the Upland zone. This was expected due to th e slight elevational incr ease along this transect, and the thermal stability of water and the higher specific heat of water compared to land. These

PAGE 49

49 data lead to the rejection of th e null hypotheses that soil moistu re and soil temperature in the Upland, Ruellia, and Submerged zones would be the same. Seed Burial Study A previous study (E. Barnett, unpublished) indicated that R. tweediana had a transient seed bank within the Ruellia z one but results here suggest ot herwise. The high (34.4 % (SD + 9.1 %)) total seed viability in the Ruellia zone after 15 months of burial indicated that the seed bank is persistent. This difference in results between those two studies could be due to many factors. Ruellia tweediana viability was sensitivity to storage c onditions. Seeds had a rapid decline in viability (~95% down to 0% 8%) when stored under ambient laboratory conditions when containers are completely sealed or comple tely open for approximately 4 months. Higher viability rates (95%) were found wh en seeds were stored in refr igerated conditions and covered but not completely sealed. The previous study did not test the viability of the seeds at time of burial. Another factor that was different was the addition of sand to the seed burial bags. This buffer between buried seeds could have reduced disease spread and fungal growth during the experiment. Soil moisture, previously thought to be an indicator of where R. tweediana could survive, was lower in the previous study than the current one (soil moisture of th e Ruellia zone in the previous study was similar to that in the Upland zone the current study). Thus, this variable is unlikely to explain differences in seed viabi lity between the two studies. The results of the current study falsified the hypothesis that R. tweediana would not survive over a year, and this species has a persistent seed bank. The expansion of the experiment to include the seed survival in the adjoining zones resulted in unanticipated outcomes. It was expected that there would be st atistical differences in viability between the Upland and Ruellia treatments because of the absence of R. tweediana

PAGE 50

50 plants and the decreased level of moisture in th e Upland zone. Instead, th e percent viability was not different between zones during the first 12 mont hs and seed survival was actually greater in the Upland zone after 15 months of burial. This suggests that under the conditions of this study, seed survival is unlikely to be the factor limiting R. tweediana in the Upland zone. Because of the results of the two experime nts Soil effects on Seed Greenhouse and Seed Viability in Submergence Conditions (Cha pter 2), the high percentages of dead seeds recovered from the Submerged zone were also unexpected. Reasons why this occurred could include the anaerobic conditions in the field comp ared to the oxygenated deionized water used in the Seed Viability in Submerge nce Conditions experiment. It al so could have been due to some toxicant or adverse biochemical conditions in the water that caused the seeds to die. The tributaries of Sweetwater Branch Creek in wh ich the research was conducted are directly downstream of a sewage plant and a large homele ss community that uses the stream. Additional studies would be needed to determine what condi tions in the Submerged zone caused the death of the R. tweediana seeds. For example an analysis of water quality, including redox potential might be instructive. In this burial experiment, seed s were collected from may different plants, and were almost certainly in different physiologi cal states. It was not believed that the seeds had a primary dormancy (Wilson and Mecca 2003) and once seeds were collected from parent plants many seeds would be non-dormant and consequently could germinate in the bags in the soil. Some of the buried seeds did not germinate in the soil but germinated once placed in the incubation chamber. It is likely that these seeds had forced quiescent due to envir onmental conditions in the soil unsuitable for germination. Other seeds di d not germinate even after placement in the

PAGE 51

51 germination chamber and those seeds exhibite d induced secondary dormancy (called dormant but viable in this text). This experiment indicates that soil moisture and light are not the primary factors that exclude R. tweediana from the Upland zone. It also demons trated that there is potential for a persistent seed bank to exist in the Upland a nd Ruellia zones. The viability of buried seeds decreased over time but not as quickly as had be en expected, and seeds did survive burial for a year in each of the zones, although at very low percentages in the Submerged zone.

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52 Figure 3-1. Nylon mesh seeds bags attached to the bottom of surveyors flags with zip ties.

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53 Figure 3-2. Seed burial plot, clos est flags in Submerged zone, center flags in Ruellia zone, background flags in Upland zone.

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54 0 10 20 30 40 50 60 70 80 90 100 FMAMJJASONDJFMPercent Soil Moisture Upland Ruellia Submerged Figure 3-3. Average percent soil moisture for each zone February 2006 to March 2007. There were significant differences between zones at each sampling interval and averaged across time.

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55 0 5 10 15 20 25 30 35 0246810121416 Months After Burial% Missing of Seed Total U R S bc a ab b a bc Figure 3-4. Percent missing of seed total in each zone. U= Upland zone, R= Ruellia zone, and S= Submerged zone. Different letters indicat e significant difference between months after burial ( of 0.05) for % missing averaged over all zones. Line fitted to averages with an R2 = 0.93. (Submerged zone not sampled at 15 months.) Table 3-1 ANOVA table for percent missing of seed total. Source of Variation Degree of Freedom F Value P Value Zone 2 0.290.7497 Month 5 25.990.0001 Zone*Month 9 0.940.4952

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56 0 10 20 30 40 50 60 70 80 90 100 0246810121416 Months After Burial% Dead of Seed Total U R S Figure 3-5. Percent dead of seed total in each zone. U= Upland zone, R= Ruellia zone, and S= Submerged zone. Percent dead were significan tly higher at each month after burial in the Submerged zone compared to the Upla nd and Ruellia zones. (Submerged zone not sampled at 15 months.) Table 3-2 ANOVA table for percent dead of seed total. Source of Variation Degree of Freedom F Value P Value Zone 2 212.70.0001 Month 5 3.710.0034 Zone*Month 9 1.880.0588

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57 0.0 10.0 20.0 30.0 40.0 50.0 60.0 70.0 80.0 90.0 100.0 0246810121416 Months After Burial% Germinated of Seed Total U R S Figure 3-6. Percent germinated of seed total in each zone. U= Upland zone, R= Ruellia zone, and S= Submerged zone. (Submerged zone not sampled at 15 months.) Table 3-3 ANOVA table for percent germinated of seed total. Source of Variation Degree of Freedom F Value P Value Zone 2 126.850.0001 Month 5 10.660.0001 Zone*Month 9 1.730.0875

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58 Table 3-4 ANOVA table for per cent dormant of seed total. Source of Variation Degree of Freedom F Value P Value Zone 2 4.910.0086 Month 5 8.040.0001 Zone*Month 9 2.970.0028

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59 0% 20% 40% 60% 80% 100% U-1U-3U-6U-9U-12U-15R-1R-3R-6R-9R-12R-15S-1S-3S-6S-9S-12 Zone-MonthPercent of Seed Total # Missing # Dead Total # Germinated # Dormant Figure 3-7. Percentages of total se ed (50) that were recovered from the field that were 1) missing, 2) dead, 3) germinated, or 4) viable but presumed dormant. The Zones are represented by U= Upland R= Ruellia, S= Submerged. The numbers represent the time buried, e.g. 9=nine months buried.

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60 0 10 20 30 40 50 60 70 80 90 100 UplandRuelliaSubmergedPercent Viable of Seed Total Month 1 Month 3 Month 6 Month 9 Month 12 Month 15 ab a b b ab ab bc c a ab b b b a bc ab ab Figure 3-8. Mean percent viable seed for 10 repl icate samples per zone. (Percent Viable included seeds that germinated in bag/chamber and dormant seeds.) Different letters indicate significant differences at of 0.05, within a zone over time.

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61 CHAPTER 4 FIELD STUDY Introduction Persistence and spread of most plant species is commonly limited by a combination of seed and microhabitat availability (E riksson and Ehrlen 1992; Scherff et al. 1994; Kollmann et al. 2007). Scherff et al. (1994) proposed that all it would take to overcome these limitations and expand a species distribution is successful seed dispersal coupl ed with opportunistic seedling growth in a new habitat. Understa nding what limits pers istence and spread is of great importance for understanding distribution of invasive non-nati ve species. Currently, however, the specific seed and microhabitat limitations are unknown fo r most non-native plants (Turnbull et al. 2000; Kollmann et al. 2007). Initial establishment processe s are thought to be of great importance in determining the distribution of plants. Several st udies (cited by Foster 1999) indicate that it is in the early establishment stages of life history when a plant may be most sensitive to competition. Studies of colonization potential beyond the boundaries of an existing plant population are typically achieved by comparing the establishment of seed lings from seeds transplanted into foreign microsites to that of controls planted in to the home habitat (Scherff et al. 1994). The overall objective of the field study was to investigate whethe r the observed field distribution of R. tweediana could be explained 1) by lim itations due to environmental conditions: soil moisture, light, and soil temperatur e, and/or by the bio tic interactions of competition with native plants and 2) at which life-stage, seeds or seedlings, these limitations were most influential. The first component of the present study was to determine if seeds of R. tweediana could germinate in the Upland or Ruelli a zone. If they germinated, the next component was to determine if the seedlings coul d survive over a three month period of time.

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62 The third component was to determ ine if transplanted seedlings could survive in the Upland or Ruellia zone. Hypotheses Seed Germination and Survival Experiments 1. Null Hypothesis: Plots will have the same mean Gap Fraction at both levels of Zone and/or Competition. Alternative Hypothesis: Plots wi ll NOT have the same mean Ga p Fraction at both levels of Zone and/or Competition. 2. Null Hypothesis: R. tweediana seeds will have the same perc entage germination at both levels of Zone and/or Competition. Alternative Hypothesis: R. tweediana seeds will NOT have the same percentage germination at both levels of Zone and/or Competition. If Alternative Hypothesi s is accepted then: o Null Hypothesis: Values of Light, Soil mois ture, and/or Soil temperature will be the same for at both levels of Zone and/or Competition Alternative Hypothesis: Values of Light Soil moisture, and/or Soil temperature will NOT be the same at both levels of Zone and/or Competition. 3. Null Hypothesis: Germinated R. tweediana seedlings will have the same percentage survival at both levels of at both levels of Zone and/or Competition. Alternative Hypothesis: Germinated R. tweediana seedlings will NOT have the same percentage survival at both levels of Zone and/or Competition. If Alternative Hypothesi s is accepted then: o Null Hypothesis: Values of Light, Soil mois ture, and/or Soil temperature will be the same for at both levels of Zone and/or Competition. Alternative Hypothesis: Values of Light Soil moisture, and/or Soil temperature will NOT be the same at both levels of Zone and/or Competition. Seedling Transplant Experiment 4. Null Hypothesis: Plots will have the same mean Gap Fraction at both levels of Zone, Size, and/or Competition. Alternative Hypothesis: Plots will NOT have the same mean Gap Fraction at both levels of Zone, Size and/or Competition. 5. Null Hypothesis: Transplanted R. tweediana seedlings will have the same percentage Survival, Growth, Root biomass, and/or Shoot biomass at both levels of Zone, Size and/or Competition. Alternative Hypothesis: R. tweediana seedlings will NOT have the same percentage Survival, Growth, Root biomass, and/or Shoot biom ass at both levels of Zone, Size and/or Competition. If Alternative Hypothesi s is accepted then:

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63 o Null Hypothesis: Values of Light, Soil mois ture, and/or Soil temperature will be the same for at both levels of Zone, Size and/or Competition Alternative Hypothesis: Values of Light Soil moisture, and/or Soil temperature will NOT be the same at both levels of Zone, Size and/or Competition. Material and Methods Study Site and General Plot Establishment Research was conducted on small tributaries of Sweetwater Branch Creek located in Paynes Prairie Preserve State Park, Alachua County, Florida. Te n replicate sites were located during May 2006 based on the presence of the Upland, Ruellia, and Submerged zones. A transect was established from the center of the Submerge d zone, through the Ruellia zone, and 5 m into the Upland zone at each of the te n replicate sites. In the experimental design, Zone was the mainplot factor with two treatments: Upland and Ruellia (Submerged zone was excluded from these experiments). Within each replicate site, six 1 m2 plots were laid out on a transect parallel to the zone orientation with a 0.5 m wide buffer surr ounding each plot as seen in Figure 4-1. These plots were paired (2 x 1 m2 plots) for the sub-plot factor which was Competition, with two treatments: Cleared and Not-Cleared (Figure 4-2) In the Cleared treatment all vegetation was removed with a gas-powered weed trimmer to ground level. One week la ter all regrowth and remaining vegetation was removed by hand, resulting in a vegetation-free pl ot, and this condition was maintained throughout the stud y. Vegetation was not disturbed in the Not-Cleared treatment. Prior to vegetation removal within the Cleared treatment, a 0.25 m2 area was sampled where species were recorded and colle cted for above ground biomass. At the end of the study it was repeated in the Not-Cleared treatment (Table 1-4, Appendix Table A-1 and Table B-1). This created a set of three paired plots, with the members of the each pair immediately next to one another while the three se ts of pairs were not always ad joining one another (Figure 4-1). Treatments were located randomly (Figures 4-1 and 4-3). Light intensity at the soil surface, soil

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64 moisture, and soil temperature were recorded at each plot every two weeks during the study, for use as covariables. Canopy photos were taken in the plots at the beginning of the Seed Germination and Survival Experiments and Seedling Transplant Experiment for site characterization. For details about the methods and instruments used see Chapter 3. Seed Germination and Survival Experiments Seed germination A seedling germination study was conducted using the five of the 10 re plicate sites on the eastern side of the Prairie study area. This was a split -plot experimental design with Zone (main plot) and Competition (sub-plot) as factors. A 3 x 2 grid was created using string on a 1 m2 PVC frame and was placed over one member of each pa ired plot. Six empty 10 cm diameter plastic flower-pots, with bottoms removed, were inserted into the ground with one pot in each section of the grid (Figure 4-4). Ten seeds were placed on th e soil surface in five of the pots, leaving one for a check to see if there was a seed bank alread y in the soil or if new seeds fell into the pots. This study was conducted the first time from August 13 through October 4, 2006 with each plot watered with 7.5 liters of creek water applied with a watering can on rain-free days for the first two weeks. The study was conducted a second time October 4 through November 20, 2006 but the plots were not watered. Seedlings were counted as they emerged and were removed from the pots. Seed germination and seedling survival An experiment identical to the first (Augus t 13-October 4) Seed Germination study was conducted in the remaining five replicate sites on the western side of the Prairie study site from August 13 through November 16, 2006. Pots were wa tered with 7.5 liters on rain-free days for the first two weeks. Seedlings were counted bi-weekly but not removed.

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65 Seedling Transplant Experiment Soil from the Upland and Ruellia zones was collected May 23, 2006 and June 2, 2006. The soil was air dried and sifted. Four hundred R. tweediana seeds were placed in seedling trays of each soil type on each of June 1 and June 30, 2006 to create two age groups for the Size factor, Large (15 cm tall) and Small (3 cm tall) seedlings. On June 30, Large seedlings (June 1 group) were repotted into 10 cm wide, square pots and seedlings were transplanted to the field in August. A split-split plot experimental design was used with Zone as the main-plot factor with Upland and Ruellia as treatments. The sub-plot factor was Competition with Cleared and NotCleared treatments, and the subsub plot factor was Size with Large and Small seedlings. The two sets of paired plots in each zone that were not used for the germination studies were used for this study. A 4 x 3 grid was created using string on a 1 m2 PVC frame which was placed over each plot (Figure 4-5). In one set of paired plot s, 10 Large seedlings were randomly transplanted into the grid squares and in the other set of pa ired plots the 10 Small seedlings were randomly transplanted into the grid squa res. Thus a total of 80 seedlings, 40 Large and 40 Small, were planted in each replicate site (F igure 4-6 and 4-7). The five east ern replicate sites were planted the week of August 1 and the western five repl icate sites were planted the week of August 7, 2006. Seedling plots were watered with 7.5 liters of creek water on rain-free days for the first two weeks to minimize transplant shock. This experiment was conducted over 3.5 months ending on November 16. During this period, survival of seedlings was noted and biweekly measurements of seedling height were reco rded. On November 16, 2006 all surviving plants were collected, separated into shoots and root s and dried before weighing for above-ground and below-ground biomass.

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66 Statistical Analyses ANOVA of canopy photo measurements was used to investigate differences in Gap Fraction between zones. A mixe d-model ANOVA was used for hypot hesis testing in the Seed Germination and Survival Experiments and the Seedling Transplant Experiment. This was done with SAS Mixed procedure for all experiments except the Seedling Survival Data, which was analyzed as binary data, 1=lived 0=dead, w ith the SAS Glimmix macr o. Light intensity, soil moisture, and soil temperature measurements were averaged over time and analyzed as covariables with Proc Mixed. All statistical anal yses were done with SAS Version 9.1 software (SAS 2007). The Zone factor was analyzed as a one-sided test because of the preexisting knowledge that the Ruellia treatment (the zo ne in which vast populations of mature R. tweediana occur) would have high rate s of seed germination and seedling survival. Results Seed Germination and Survival Experiments Canopy photos. Canopy photographs revealed mean (+ SD) Gap Fractions of 0.113 (+ 0.013) and 0.112 (+ 0.005), respectively for the Zone treatments Upland and Ruellia, and 0.113 (+ 0.009) and 0.112 (+ 0.01), respectively for the Competition treatment Not-Cleared and Cleared. This indicated an almost closed canopy with no significant difference between zone treatments (p= 0.7) or competition treatments (p=0.9). Checks. The purpose of the checks wa s to determine if there was germination from a seed bank or if seeds were being deposited and germinating duri ng the experiments. Only one seedling was removed from one check pot during th e first (August) germination trial. Given this extremely rare occurrence, the checks were remove d from all datasets before the analyses were conducted.

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67 Seed germination During the first germination experiment, 1 of the 100 pots had germination over 100 % (11 seeds instead of 10 indicating the presence of an outside seed). Du ring the course of the experiments, a few pots were disturbed by armadillos, 3 in the first experiment and 1 in the second. Data from these pots were removed from the analyses because there was a high number of replicate pots and no pattern wa s found in the disturbance. In the ANOVA for the first (watered) experiment Zone (Upland, Ruellia ) had a p-value of 0.06, with germination rates of 67.8 % (Ruellia) and 57.9 % (Upland) (Figure 4-8). In the second experiment, Zone had a p-value of 0.04, with th e Ruellia treatment having a higher percentage germination rate (35.4 %) than the Upland treat ment (21.2 %), Figure 4-9. The p-value for the Competition factor in the first experiment wa s 0.37 and 0.56 for the second experiment and the Zone x Competition interacti on was 0.8 and 0.12, respectively for either the first or second experiment. In the Proc Mixed analysis all three covari ables, light intensity, soil moisture, and soil temperature, had an influence on the Zone factor in the first experiment, while only soil moisture had an influence on the second experiment. If these covariables could be set to an equal level in all treatments, the average germination would be the same between the zones in the first experiment. The means for all the covariables light, soil moisture, and soil temperature are presented in Table 4-1. The overall lower percen t germination for the second experiment (note different scales on y axis in Figures 4-8 and 4-9) was possibly because the seeds were not watered, but could have been due to other factors. The analysis of the covariables supports the suggestion that soil moisture was influential.

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68 Seed germination and seedling survival During the course of this experiment, 6 pots were disturbed by armadillos. For these pots, no further data were collected and the data co llected until that point were removed from the analyses. During the germination part of this experiment, the Competition factor had a p-value of 0.08, with the Cleared treatment having a higher percentage germination (65.4 %) than the NotCleared treatment (55.0 %) (Figure 4-10). The fa ctor of Zone, and the Zone x Competition interaction had p-values of 0.17 and 0.2, respectively. These result s were unexpected since this experiment was very similar to the first seed ge rmination experiment except that it lasted one month longer. The covariables that were analyzed using Proc Mixed did no t have an influence on this experiment. At the end of the three months, 49 pots ( out of the 100 pot total) had no surviving seedlings. These 49 pots were excluded from the st atistical analysis so the relationship between factor level and survival could be assessed, (i nclusion of the 49 pots would have created a zeroinflated dataset, which is very complex to analyze). Zone for percent survival had a p-value of 0.36, while Competition was a highly significant factor (p= 0.0003), the Cleared having a larger percen t survival (85 %) than NotCleared treatment (40 %) (Figure 4-11). The Z one x Competition interaction had a p-value of 0.35. The covariables analyzed for this experime nt did not show any influence on the study. Seedling Transplant Experiment Canopy photos. Canopy photographs revealed mean Gap Fractions that did not vary for any factors of Zone (p= 0.8), Competition (p= 0.85), or Size (p=0.59). Th e average Gap Fraction of 0.114 (+ 0.011 SD) indicated an almost closed canopy.

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69 Survival and Height. Of the 800 seedlings transplanted, 662 seedlings survived (83% survival). Zone and Competition had p-values of 0.055 and <.0001, respectively (Table 4-2). The mean survival was higher in the Ruellia treatments (0.85 + 0.36 SD) than it was in the Upland treatment (0.81+ 0.39 SD). The mean survival was highe r in the Cleared treatments (0.93 + 0.26 SD) than it was in the Not-Cleared treatment (0.73 + 0.45 SD). The interaction between the Zone and Competition factors had a p-value of 0.002 (Table 4-2). In the Cleared treatment the survival was higher in the Ruellia zone and in the Not-cl eared treatment the survival was higher in the Upland zone (Figure 4-12:A). The final inte raction was between the Size and Competition factors with a p-value of 0.047 (Tab le 4-2). In the Cleared treatmen t the survival was nearly the same for with Large and Small plants while in the Not-Cleared treatment the survival of Large was greater than Small plants (Figure 4-12:B). In the ANOVA of the change in seedling hei ght over the 3.5 month st udy, Zone, Size, and Competition were all significant (p<.0001) (Table 4-3). The mean change in height was greater in the Ruellia treatments (1.63 + 1.8 SD) than it was in the Upland treatment (0.59 + 0.65 SD). The mean change in height was gr eater in the Large treatments (1.49 + 1.78 SD) than it was in the Small treatment (0.73 + 0.95 SD). The mean change in he ight was greater in the Cleared treatments (1.53 + 1.76 SD) than it was in the Not-Cleared treatment (0.69 + 0.95 SD). The interaction between the Zone and Competition f actors had a p-value of 0.0003 (Table 4-3). The difference in change of height between the Ruel lia and Upland zones was greater in the Cleared than the Not-Cleared treatment (Figure 4-13:A). The final interaction wa s between the Zone and Size factors with a p-value of 0.0095 (Table 4-3). The difference ch ange in height between the Ruellia and Upland zones was greater in the Large than the Small seedling size (Figure 4-13:B).

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70 Biomass. For shoot biomass, Zone had a p-value of 0.051 (Table 4-4). The mean dry shoot biomass was higher in the Ruellia treatments (0.108 g + 0.114 SD) than it was in the Upland treatment (0.079 g + 0.84 SD). Size and Competition both ha d a p-value of <.0001 (Table 4-4). The mean dry shoot biomass was highe r in the Large treatments (0.144 g + 0.113 SD) than it was in the Small treatment (0.042 g + 0.019 SD). The mean dry shoot biomass was higher in the Cleared treatments (0.129 g + 0.115 SD) than it was in the Not-Cleared treatment (0.058 g + 0.068 SD). There was an interaction between the Zone, Size, and Compe tition factors with a pvalue of 0.007 (Table 4-4, Figure 4-14). In the Cleared treatment the dry shoot biomass was higher in the Ruellia zone and in the Not-cleare d treatment the dry shoot biomass was higher in the Upland zone (Figure 4-14). For root biomass, Size and Competition bot h had a p-value of <.0001 (Table 4-5). The mean dry root biomass was higher in the Large treatments (0. 072 g + 0.050 SD) than it was in the Small treatment (0.019 g + 0.019 SD). The mean dry root bi omass was higher in the Cleared treatments (0.059 g + 0.051 SD) than it was in the Not-Cleared treatment (0.032 g + 0.037 SD). There a interaction between the Zone, Size, and Competition factors with a p-value of 0.008 (Table 4-5). In the Cleared treatment the dry shoo t biomass was higher in the Ruellia zone and in the Not-cleared treatment the dry shoot biomass was higher in the Upland zone (Figure 4-15). Covariables. In the analysis of th e covariables light, temperature, and moisture, only moisture had a significant effect on the S eedling Transplant Study. For seedling growth, moisture influenced the zone treatment. For sh oot biomass, moisture influenced one, two-way interaction, Ruellia, Notcleared Upland, clea red; and one three-way interaction, Ruellia, Small, Cleared Upland, Large, Cleared. For root biomass, moisture influenced one two-way

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71 interaction, Ruellia, Not Cleared Upland, Not-Clear ed. The extent to which the covariable of moisture influenced the experiment warrants further investigation. Discussion In the Seed Germination and Seedling Survival Experiments, the absence of seedlings in all but one check pot suggests that natural seed rain and seed bank of R. tweediana was low during the study period, indicating either little seed production during the 3-month study period, dormancy, or possibly a general limit ation in seed availability. When seeds are sown in the field, germination occurs and juvenile seedlings are f ound. However, it must be noted that seeds that were mixed with the soil in th e greenhouse study (Chapter 2) had decreased rates of germination compared to when they were on the soil surface. If there are strong seed limitation factors such as seed production, seed dispersal, and/or s eed predation, they are possibly important for controlling spread of R. tweediana If seeds were present in th e soil, dormancy was not broken under the conditions of these studies. A seed bank study in the Upland zone is needed to determine if R. tweediana has a seed bank there. A seed prod uction and dispersal study needs to be conducted in the existi ng mature populations of R. tweediana Both of these experiments could further explain the seed limitation that is occurring in the Upland zone. Another factor that may be involve d is that seed predation by the Melanagromyza ruelliae fly causes a 25 % reduction in seed viability and al so inhibits the seed capsules from explosively dehiscing (Huey et al. 2007). How widely this is occurring and its eff ect on seed production and dispersal or seedbanks warra nts further investigation. The Competition factor did not have an effect in the firs t or second Seed Germination experiment but it did in the Seed Germination and Seedling Survival experiment. This positive effect of the disturbance of clearing away competing vegeta tion in both Ruellia and Upland zones indicates a microhabitat limitation and sugg ests competitive suppression by the resident

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72 vegetation. This is also seen in the Seedling Tran splant data, where the survival of transplanted seedlings, change in height, and root and shoot bi omass were always higher in cleared plots and were more self-limited by R. tweediana than by native vegetation. Survival rates of the seedlings indicated that the seedling size or zone in which seedlings were tr ansplanted had little influence if plots were cleared (Fig ure 4-12). While there was mi crohabitat limitation, it was not completely limiting the seedlings and they were st ill able to survive and grow in the Not-Cleared plots. Environmental conditions that were measured during the experiment to determine if they explained the zonation of R. tweediana did not reveal conditions that prevented seed germination or seedling emergence beyond the current population edge. Moisture did influence some of the results of the experiments, for example in the second Seed Germination experiment where germination was probably reduced by the abse nce of watering at the beginning of the experiment, but the germination pe rcentages were still above 20%. Conclusions If there is an area that has not been invaded by R. tweediana but is otherwise similar to the study site at the Prairie and seed reaches that site, it would likely become invaded in a similar manner as the Prairie, and the invasion woul d start around the waterways. The seeds would germinate even if there was some native plan t competition, and would survive and grow. There are environmental conditions around the waterways that differ from the Upland zone, that could not be specifically characterize d, resulting in higher germination, survival, growth, and biomass. But even with these high values in the Ruellia zo ne and the seed and microhabitat limitations that were identified in the Upland zone, there was stil l some germination and survival in the Upland zone when seeds or plants were sown there with competition from native vegetation. It seems that the full extent of the invasion is not yet realized at this site and R. tweediana plants will

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73 move into the Upland if seed re aches it, especially if there is limited competition. During the experiment the beginnings of this expansion in to the Upland by seedlings (10-15cm tall) were observed, a few of which appeared in Upland plots (Appendix Table 1-A). Also R. tweediana extensively reproduces vegetative ly with rhizomes and layering and rooting at the nodes (pers. obs.). There was exte nsive expansion, by layering, of populations into the waterways when the water was low during 2006 a nd this changed the course of the water, as a once moving waterbody became two separate pools. Germinated seeds were also monitored in one of the creek beds that had little water fo r a few months. The seedlings survived for over a month until they were completely submerged (and 2 seedlings survived for 2 months submerged), but in one of the plots there was a ~25 cm long R. tweediana stem that layered into the plot and never died. The existing monocultures of mature R. tweediana plants with the abrupt line between zones can suggest the population is no longer expanding. However, during the course of these studies, vegetative expansion was observed into the Upland zone. A future study to experimentally monitor the rate at which the pop ulation is expanding at this local scale would allow better predictions of the extent and rate of spread into the Upland zone once the habitat closest to the waterways has been colonized. Determining the mechanism by which the habitat closest to the waterway (at this stage of invasion in this site, the habitat that became the Ruellia zone) was initially colonized, was not an objective of this study. However, the presence of R. tweediana within the study site and creeks across Gainesville, Florida, is likely due to the widespread use of this popular ornamental plant (pers. obs.), which creates a source of propagules and seeds in residential areas upstream of the invaded sites. The mucilaginous ge l produced by these seeds when wetted, promotes dispersal of

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74 this nature by allowing the seeds to stay buoyant in the water. Once the seed leaves the water, the gel dries and glues the seed to th e soil (Gutterman et al. 1973). While R. tweediana plants grown from seed are easy to kill with herbicides in greenhouse experiments (pers. obs.) this is not necessarily true once it be comes vegetatively established in natural communities (R. Stocker, UF, pers. comm.). This understanding, along with the knowledge of R. tweedianas ability to invade and expand on ce it has arrived in a natural community, means that early detection and rapi d removal of new populations in natural areas should be a priority. A large scale study to look at common characteristics of invaded sites across Florida would be needed to get a complete understanding of the eco logical and propagule pressure factors that influence the invasion of new sites. The results presented herein can contribute to improved predictions of where R. tweediana is likely to invade on a state-wide basis in the future. Importantly, these results pr ovide more definitive information to guide local level predications about how R. tweediana invasions will proceed once the species becomes established within a site.

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75 Figure 4-1. Plot layout map for Seed Germina tion and Survival Experiments and Seedling Transplant Experiment at Paynes Prairie Preserve State Park.

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76 Figure 4-2. Example of one set of paired plots in the Seedling Transplant Experiment with competition factor applied. Le ft side of photograph = NotCleared treatment, right side = Cleared treatment. This is within the Ruellia treatment, planted with a Small seedling treatment.

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77 Figure 4-3. Example of the plot layout for one replicate site.

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78 Figure 4-4. Example of a Seed Germination Experime nt plot within a Ruellia treatment and with the Cleared treatment applied.

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79 Water Cleared Not Cleared Not Clear Cleared x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Ruellia Zone x x x x x x x x x x Large Large Small Small Not Cleared Cleared Cleared Not Cleared x x x x x x x x x x x x x x x x x x x x x x x x x x x x x x Upland Zone x x x x x x x x x x Small Small Large Large Figure 4-5. Example of one replicat e site plot layout for the Seed ling Transplant Experiment. In this split-split plot experime ntal design. Zone is the main -plot factor, Competition is the sub-plot factor, and Size is the sub-sub-plot factor.

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80 Figure 4-6. Example of a plot from the Seedling Transplant Experiment with Upland, Cleared, and Large seedling treatments.

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81 Figure 4-7. Example of a plot from the Seedli ng Transplant Experiment with Upland, Notcleared, and Small seedling treatments.

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82 0 10 20 30 40 50 60 70 80 RuelliaUplandClearedNot ClearedPercent Germination Figure 4-8. Percent germination rates for the firs t experiment of the Seed Germination Study. A) The dark grey bars represent the data av eraged over the two treatments (Upland and Ruellia) of the Zone factor. B) The light grey bars represent the data averaged over the two treatments (Cleared and Not-Clear ed) of the Competition factor. Bars represent standard erro r and within-factor p-va lues are at the top. p = 0.06 p = 0.37 A B

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83 0 10 20 30 40 50 RuelliaUplandClearedNot ClearedPercent Germination Figure 4-9. Percent germination rates for the sec ond experiment of the Seed Germination Study. A) The dark grey bars represent the data averaged over the tw o treatments (Upland and Ruellia) of the Zone factor. B) The lig ht grey bars represent the data averaged over the two treatments (Cleared and NotCleared) of the Competition factor. Bars represent standard erro r and within-factor p-va lues are at the top. Table 4-1. Means of covariates for Seed germination in field experiments. RuelliaRuelliaUplandUpland Units ClearedNot-ClearedClearedNot-Cleared Soil Temp 19.2719.3819.3019.32 C Light 24.3424.2924.5324.39 mols s-1 m-2 First experiment Soil Moisture 24242324 % Soil Temp 23.7023.7723.6023.61 C Light 23.4814.2812.9512.32 mols s-1 m-2 Second experiment Soil Moisture 60543228 % p = 0.04 p = 0.56 A B

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84 0 10 20 30 40 50 60 70 80 RuelliaUplandClearedNot ClearedPercent Germination Figure 4-10. Percent seed germination rates over three months in the Seed Germination and Seedling Survival experiment. A) The dark gr ey bars represent the data averaged over the two treatments (Upland and Ruellia) of the Zone factor. B) The light grey bars represent the data averaged over the two trea tments (Cleared and Not-Cleared) of the Competition factor. Bars represent standard error and within-factor p-values are at the top. p = 0.17 p = 0.08 A B

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85 0 10 20 30 40 50 60 70 80 90 100 RuelliaUplandClearedNot ClearedAverage Percent Survival Figure 4-11. Average percent surv ival over three months in the Seed Germination and Seedling Survival experiment. A) The dark grey bars represent the data averaged over the two treatments (Upland and Ruellia) of the Zone factor. B) The light grey bars represent the data averaged over the two treatmen ts (Cleared and Not-Cleared) of the Competition factor. Bars represent standard error and within-factor p-values are indicated. p = 0.36 p = 0.0003 A B

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86 0.6 0.65 0.7 0.75 0.8 0.85 0.9 0.95 1 1.05 ClearedNot-ClearedSurvival Ruellia Upland 0.6 0.65 0.7 0.75 0.8 0.85 0.9 0.95 1 1.05 ClearedNot-ClearedSurvival Large Small Figure 4-12. Seedling Transplant Experiment twofactor interactions of seedling Survival. A) Zone x Competition interact ion with p=0.002. B) Size x Comp etition interaction with p=0.05. Table 4-2. ANOVA table for seedling Survival ov er 3.5 months in Seedling Transplant Study Source of Variation Degree of Freedom F ValuePr > F Zone 9 4.840.0553 Size 18 1.30.2698 Competition 37 20.52<.0001 Zone*Size 18 1.80.1964 Zone*Competition 37 10.790.0022 Size*Competition 37 4.230.0468 Note: Not capable of calculating three-way interaction A B p =0.002 p =0.05

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87 0 0.5 1 1.5 2 2.5 3 ClearedNot-Clearedchange in height (cm) Ruellia Upland 0 0.5 1 1.5 2 2.5 3 LargeSmallchange in height (cm) Ruellia Upland Figure 4-13. Transplant study twofactor interactions of seed ling change in height over 3.5 months. A) Zone x Competition interacti on with p=0.003 B) Zone x Size interaction with p=0.0095. Table 4-3. ANOVA table for Increase in Height over 3.5 months in Seedling Transplant Study. Source of Variation Degree of Freedom F Value Pr > F Zone 18 26.05<.0001 Size 54 19.23<.0001 Competition 54 23.48<.0001 Zone*Size 54 7.240.0095 Zone*Competition 54 14.680.0003 Size*Competition 54 1.280.2621 Zone*Size*Competition 54 2.750.1032 p =0.003 p =0.0095 A B

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88 Figure 4-14. Seedling Transplant Experiment aver age shoot biomass data. Three-way interaction: Aseparated by Zone factor; Bseparate d by Competition factor; Cseparated by Size factor. Large0.000 0.040 0.080 0.120 0.160 0.200 0.240 RuelliaUplandshoot biomass ( g Clear Not-Clear Ruellia0 0.04 0.08 0.12 0.16 0.2 0.24 ClearedNot-Clearedshoot biomass ( g Large Small Upland0 0.04 0.08 0.12 0.16 0.2 0.24 ClearedNot-Clearedshoot biomass ( g Large Small Cleared0.000 0.040 0.080 0.120 0.160 0.200 0.240 LargeSmallshoot biomass ( g Ruellia Upland Not-Cleared0.000 0.040 0.080 0.120 0.160 0.200 0.240 LargeSmallshoot biomass ( g Ruellia Upland Small0.000 0.040 0.080 0.120 0.160 0.200 0.240 RuelliaUplandshoot biomass ( g Clear Not-Clear A B C

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89 Figure 4-15. Seedling Transplant Experiment aver age root biomass data. Three-way interaction: Aseparated by Zone factor; Bseparate d by Competition factor; Cseparated by Size factor. Ruellia 0 0.02 0.04 0.06 0.08 0.1 ClearedNot-Clearedroot biomass Large Small Uplan d 0 0.02 0.04 0.06 0.08 0.1 ClearedNot-Clearedroot biomass ( Large Small Cleared0 0.02 0.04 0.06 0.08 0.1 LargeSmallroot biomass Ruella Upland Not-Cleared0 0.02 0.04 0.06 0.08 0.1 LargeSmallroot biomass Ruella Upland Large 0 0.02 0.04 0.06 0.08 0.1 RuellaUplandroot biomass Clear Not-Clear Small 0 0.02 0.04 0.06 0.08 0.1 RuellaUplandroot biomass Clear Not-Clear C B A

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90 Table 4-4. ANOVA table for Shoot Bi omass in Seedling Transplant Study Source of Variation Degree of FreedomF ValuePr > F Zone 95.020.0518 Size 1887.82<.0001 Competition 3652.79<.0001 Zone*Size 180.470.4999 Zone*Competition 3632.47<.0001 Size*Competition 366.000.0193 Zone*Size*Competition 368.110.0072 Table 4-5. ANOVA table for Root Bi omass in Seedling Transplant Study Source of Variation Degree of FreedomF ValuePr > F Zone 90.000.9907 Size 18158.30<.0001 Competition 3646.38<.0001 Zone*Size 187.990.0112 Zone*Competition 3624.70<.0001 Size*Competition 365.770.0216 Zone*Size*Competition 367.870.0081

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91 APPENDIX A SPECIES LIST Table A-1. Species recorded from a 0.25 m2 plot from the Cleared (C) (Aug 2006) and NotCleared (NC) (Nov 2006) treatments of the paired plots of the Seed Germination and Survival Experiments and the Seedling Transplant Experime nt. L = Large seedling treatment, SM = Small seedling treatment, SE = Seed germin ation and survival experiments. Plot 1 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott x Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x Phytolaccaceae Petiveria alliacea L. x x x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 Poaceae Grass #4 Poaceae Grass #5 x Poaceae Oplismenus hirtellus (L.) Beauv. Ranunculaceae Ranunculus hispidus Michx. var. nitidus (Chapman) T. Duncan x x x x x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. x Ulmaceae Celtis occidentalis L. x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 92

92 Table A-1. Continued. Plot 2 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. x x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen x x x x Asteraceae Verbesina virginica L. x x x x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. x Oxalidaceae Oxalis L. x x x Phytolaccaceae Petiveria alliacea L. x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) x Poaceae Grass #1 Poaceae Grass #2 x Poaceae Grass #3 Poaceae Grass #4 x Poaceae Grass #5 x x Poaceae Oplismenus hirtellus (L.) Beauv. x x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x Rosaceae Prunus serotina Ehrh. Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. x Ulmaceae Celtis occidentalis L. x x x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 93

93 Table A-1. Continued. Plot 3 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. x Anacardianceae Toxicodendron radicans (L.) Kuntze x Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. x Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x Phytolaccaceae Petiveria alliacea L. x x x x x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 x Poaceae Grass #4 Poaceae Grass #5 x Poaceae Oplismenus hirtellus (L.) Beauv. Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x x Rosaceae Prunus serotina Ehrh. x x x x x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. Ulmaceae Celtis occidentalis L. x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 94

94 Table A-1. Continued. Plot 4 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. x x x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x x x Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli x Commelinaceae Commelina sp. L. x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. x Lamiaceae Salvia L. x Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x Phytolaccaceae Petiveria alliacea L. Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) x Poaceae Grass #1 Poaceae Grass #2 x Poaceae Grass #3 Poaceae Grass #4 x x x x Poaceae Grass #5 x Poaceae Oplismenus hirtellus (L.) Beauv. x x x x x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x x x Rosaceae Prunus serotina Ehrh. x x Rosaceae Rubus argutus Link x x x x Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. Ulmaceae Celtis occidentalis L. x x x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x x Vitaceae Vitis rotundifolia Michx. ? species 1 x x ? species 2

PAGE 95

95 Table A-1. Continued. Plot 5 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x x Aceraceae Acer negundo L. x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x x Asteraceae Youngia japonica (L.) DC. x Asteraceae Vernonia Schreb. Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x Blechnaceae Woodwardia Sm. x x Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x x x Phytolaccaceae Petiveria alliacea L. x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. Ulmaceae Celtis occidentalis L. x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2 x

PAGE 96

96 Table A-1. Continued. Plot 6 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x x x Aceraceae Acer negundo L. x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. x Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen x x Asteraceae Verbesina virginica L. x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x Asteraceae Elephantopus elatus Bertol. x x Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. x Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. x x x Lamiaceae Salvia L. x Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x x x Phytolaccaceae Petiveria alliacea L. x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. x x Ulmaceae Celtis occidentalis L. x x x x x x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x Vitaceae Vitis rotundifolia Michx. x ? species 1 ? species 2

PAGE 97

97 Table A-1. Continued. Plot 7 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. x x x x x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. x x Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen x x Asteraceae Verbesina virginica L. x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x x Asteraceae Elephantopus elatus Bertol. x Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau x x x Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x Blechnaceae Woodwardia Sm. x Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x x x Phytolaccaceae Petiveria alliacea L. x x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 x x Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. x x x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x x x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. x x x x x Ulmaceae Celtis occidentalis L. x x x x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 98

98 Table A-1. Continued. Plot 8 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. Anacardianceae Toxicodendron radicans (L.) Kuntze x x x Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. x x x x Asteraceae Elephantopus elatus Bertol. x Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x Cucurbitaceae Melothria pendula L. x Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. x x x Fagaceae Querus nigra L. x Hamamelidaceae Liquidambar styraciflua L. x x Lamiaceae Stachys floridana Shuttlw. ex Benth. x x x Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x x Phytolaccaceae Petiveria alliacea L. x x Pinaceae Pinus L. x Poaceae Dichanthelium (A.S. Hitchc. & Chase) x x x x x Poaceae Grass #1 Poaceae Grass #2 x x x x x Poaceae Grass #3 x Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. x x x x x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x Rosaceae Prunus serotina Ehrh. x x x Rosaceae Rubus argutus Link x Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. x x x Ulmaceae Celtis occidentalis L. x x x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x Vitaceae Vitis rotundifolia Michx. x ? species 1 x x x x x x ? species 2

PAGE 99

99 Table A-1. Continued. Plot 9 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x Aceraceae Acer negundo L. Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x x x Blechnaceae Woodwardia Sm. x Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. x x Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x Phytolaccaceae Petiveria alliacea L. x x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 Poaceae Grass #3 x x Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. Smilacaceae Smilax sp. L. Ulmaceae Celtis occidentalis L. x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 100

100 Table A-1. Continued. Plot 10 Zone Upland Ruellia Size L SM SE L SM SE Competition C NC C NC C NC C NC C NC C NC Family Genus species Acanthaceae Ruellia tweediana Griseb. x x x x x x x Aceraceae Acer negundo L. x Anacardianceae Toxicodendron radicans (L.) Kuntze Apiaceae Hydrocotyle L. Araceae Colocasia sp. Schott Araliaceae Hedera helix. L. Asteraceae Acmella oppositifolia (Lam.) R.K. Jansen var. repens (Walt.) R.K. Jansen Asteraceae Verbesina virginica L. x x x x x x Asteraceae Youngia japonica (L.) DC. Asteraceae Vernonia Schreb. Asteraceae Elephantopus elatus Bertol. Bignoniaceae Campsis radicans (L.) Seem. Ex Bureau Bignoniaceae Macfadyena unguis-cati (L.) A.H. Gentry x x x x x x x Blechnaceae Woodwardia Sm. Caprifoliaceae Sambucus nigra L. ssp. Canadensis (L.) R. Colli Commelinaceae Commelina sp. L. x x x x x x x x x x Cucurbitaceae Melothria pendula L. Dioscoreaceae Dioscorea bulbifera L. Fagaceae Quercus virginiana P. Mill. Fagaceae Querus nigra L. x Hamamelidaceae Liquidambar styraciflua L. Lamiaceae Stachys floridana Shuttlw. ex Benth. Lamiaceae Salvia L. Oleaceae Ligustrum sinense Lour. Oxalidaceae Oxalis L. x x x Phytolaccaceae Petiveria alliacea L. x x x x Pinaceae Pinus L. Poaceae Dichanthelium (A.S. Hitchc. & Chase) Poaceae Grass #1 Poaceae Grass #2 x Poaceae Grass #3 Poaceae Grass #4 Poaceae Grass #5 Poaceae Oplismenus hirtellus (L.) Beauv. x x Ranunculaceae Ranunculus hispidus Michx. Var. nitidus (Chapman) T. Duncan x x x Rosaceae Prunus serotina Ehrh. x Rosaceae Rubus argutus Link Saururaceae Saururus cernuus L. x Smilacaceae Smilax sp. L. x Ulmaceae Celtis occidentalis L. x x x Vitaceae Parthenocissus quinquefolia (L.) Planch. x Vitaceae Vitis rotundifolia Michx. ? species 1 ? species 2

PAGE 101

101 APPENDIX B SPECIES BIOMASS Table B-1. Biomass and the number of Ruellia tweediana stems in a 0.25 m2 plot cleared at setup of experiment in Cleared treatment (Aug. 2006) and in the Not-Cleared at the end of the experiment (Nov. 2006). Plot Zone Size Competition Ruellia we ightOthers weightRuellia count 1 U L C 0.018.00 1 U L NC 0.04.60 1 U SM C 0.014.00 1 U SM NC 0.016.40 1 U SE C 0.016.00 1 U SE NC 0.020.30 1 R L C 102.014.068 1 R L NC 51.43.047 1 R SM C 74.08.062 1 R SM NC 8.40.119 1 R SE C 58.02.049 1 R SE NC 47.30.143 2 U L C 0.010.00 2 U L NC 0.08.80 2 U SM C 0.08.00 2 U SM NC 0.03.50 2 U SE C 0.022.00 2 U SE NC 0.09.00 2 R L C 58.08.049 2 R L NC 46.36.434 2 R SM C 84.04.056 2 R SM NC 38.60.044 2 R SE C 58.00.057 2 R SE NC 26.35.538 3 U L C 0.028.00 3 U L NC 0.038.30 3 U SM C 0.026.00 3 U SM NC 0.033.60 3 U SE C 0.026.00 3 U SE NC 0.013.80 3 R L C 44.02.035 3 R L NC 80.70.989 3 R SM C 112.02.064 3 R SM NC 70.83.164 3 R SE C 92.02.057 3 R SE NC 56.50.464 4 U L C 0.046.00 4 U L NC 0.037.80 4 U SM C 0.030.00 4 U SM NC 0.017.10

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102 Table B-1. Continued. Plot Zone Size Competition Ruellia weightOthers weightRuellia count 4 U SE C 0.026.00 4 U SE NC 0.020.90 4 R L C 170.04.0115 4 R L NC 143.70.0125 4 R SM C 126.00.0106 4 R SM NC 124.00.0132 4 R SE C 224.02.0169 4 R SE NC 132.20.1116 5 U L C 0.012.00 5 U L NC 0.02.71 5 U SM C 0.04.00 5 U SM NC 0.013.90 5 U SE C 0.016.00 5 U SE NC 0.02.60 5 R L C 108.02.062 5 R L NC 63.20.060 5 R SM C 58.00.059 5 R SM NC 76.90.049 5 R SE C 76.02.056 5 R SE NC 53.60.150 6 U L C 0.00.00 6 U L NC 0.022.00 6 U SM C 0.02.00 6 U SM NC 0.011.80 6 U SE C 0.022.03 6 U SE NC 0.01.91 6 R L C 46.034.025 6 R L NC 73.30.156 6 R SM C 234.04.096 6 R SM NC 119.80.168 6 R SE C 102.00.0missing 6 R SE NC 104.10.0595 7 U L C 0.014.00 7 U L NC 0.09.60 7 U SM C 0.012.00 7 U SM NC 0.014.70 7 U SE C 0.011.00 7 U SE NC 0.013.20 7 R L C 76.04.049 7 R L NC 54.52.850

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103 Table B-1. Continued. Plot Zone Size Competition Ruellia weightOthers weightRuellia count 7 R SM C 18.012.015 7 R SM NC 12.45.326 7 R SE C 44.04.044 7 R SE NC 52.30.873 8 U L C 0.010.00 8 U L NC 0.014.10 8 U SM C 0.022.00 8 U SM NC 0.00.10 8 U SE C 0.020.00 8 U SE NC 0.015.30 8 R L C 32.02.019 8 R L NC 57.80.152 8 R SM C 60.04.096 8 R SM NC 43.01.559 8 R SE C 54.012.028 8 R SE NC 89.60.449 9 U L C 0.06.00 9 U L NC 0.014.20 9 U SM C 0.028.00 9 U SM NC 0.011.20 9 U SE C 0.02.00 9 U SE NC 0.00.50 9 R L C 86.02.060 9 R L NC 87.72.681 9 R SM C 74.06.064 9 R SM NC 69.10.377 9 R SE C 62.00.062 9 R SE NC 71.80.957 10 U L C 0.028.00 10 U L NC 0.035.50 10 U SM C 0.010.00 10 U SM NC 0.022.20 10 U SE C 0.016.00 10 U SE NC 0.057.51 10 R L C 130.04.055 10 R L NC 91.82.448 10 R SM C 84.06.045 10 R SM NC 56.51.647 10 R SE C 142.02.096 10 R SE NC 128.10.0567

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104 LIST OF REFERENCES Baskin C. C. and J. M. Baskin. 2006. Symposium: The natural history of soil seed banks of arable land. Weed Science 54:549-557. Booth, B. D., S. D. Murphy, and C. J. Swant on. 2003. Weed ecology: in natural and agricultural systems. Cambridge, MA: CABI Publishing. 303 pp. City of Gainesville. 2006. GPS map library. http://www.cityofgainesville .org/comdev/plan/gis/gis_lib.s html. Accessed: June 2007. DAntonio, C. M. and P. M. Vitousek. 1992. Bi ological invasions by exotic grasses, the grass/fire cycle, and global change. Annua l Review of Ecology and Systematics 23: 63-87. Daves Garden. 2007. Daves garden guides and in formation. http://davesgarden.com. Accessed: October 2007. Eriksson, O. and J. Ehrlen. 1992. Seed and micr osite limitation of recruitment in plant populations. Oecologia 91: 360-364. Ezcurra, C. 1993. Systematics of Ruellia (Acanth aceae) in Southern South America. Annual of the Missouri Botanical Garden 80: 787-845. Ferriter, A., B. Doren, C. Goodyear, D. Thayer, J. Burch, L. Toth, M. Bodle, J. Lane, D. Schmitz, P. Pratt, S. Snow and K. Langela nd. 2006. The status of nonindigenous species in the South Florida environment. Pp. 9-1 9-101. In South Florida Environmental Report. West Palm Beach, FL: South Florida Water Management District. http://www.sfwmd.gov. Accessed: July 2007. Florida Exotic Pest Plant Council. 2007a. Ea rly detection and dist ribution mapping system. http://www.fleppc.org/list/05Lis t.htm. Accessed: June 2007. Florida Exotic Pest Plant Council. 2007b. Flor ida Exotic Pest Plan t Council's 2005 list of invasive species. http:// www.fleppc.org/EDDMapS/florida. cfm. Accessed: August 2007. Foster, B. L. 1999. Establishment, competition and the distribution of native grasses among Michigan old-fields. Journal of Ecology 87: 476-489. Fox, A.M., D.R. Gordon, J.A. Dusky, L. Tyson, and R.K. Stocker. 2005. IFAS Assessment of the Status of Non-Native Plants in Floridas Natural Areas. http://plants.ifas.uf l.edu/assessment. Accessed: June 2007. Frank, J. H. and E. D. McCoy. 1995. Introduction to insect behavioral ecology: the good, the bad, and the beautiful: non-indi genous species in Florida. In vasive adventive insects and other organisms in Florida. Th e Florida Entomologist 78: 21-35.

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105 Gann, G.D., K.A. Bradley, and S.W. Woodmansee. 2007. The Flor istic inventory of south Florida database online. http://regionalconservation.org/ir cs/index.asp. Accessed: June 2007. Gardener, M. R., R. D. B. Whalle y, and B. M. Sindei. 2003. Ecology of Nassella neesiana Chilean needle grass, in pastures on north ern Tablelands of New South Wales. II. Seedbank dynamics, seed germination, and seed ling recruitment. Au stralian Journal of Agricultural Research. 54: 621-626. Gillman, E. F. 1999. Ruellia brittoniana, Fact Sh eet FPS-513. http://edis.ifas.ufl.edu. Accessed: August 2007. Gordon, D. R. 1998. Effects of invasive, non-indi genous plant species on ecosystem processes: lessons from Florida. Ecol ogical Applications 8: 975-989. [GRIN] Germplasm Resources Information Network. 2004. National Germplasm Resources Laboratory, Beltsville, Maryland. http://www.ars-grin.gov/cgibin/npgs/html/taxon.pl?419324. Accessed: November 2007 Gutterman, Y., A. Witztum, and W. Heydecker. 1973. Studies of the surfaces of desert plant seeds. II. Ecological adaptations of the seeds of Blepharis persica Annuals of Botany. 37: 1051-1055. Hillhorst H. W. M. and C. M. Karssen 2000. Effect of chemical environment on seed germination. Pp. 293-310. In M. Fenner, ed. Seeds: The ecology of regeneration in plant communities 2nd edition. Wallingf ord, Oxon: CABI Publishing. HemiView. 1999. HemiView user manual. Versi on 2.1. Cambridge, UK: Delta-T Devices Ltd. Hodges, A. W., and J. J. Haydu. 2005. Economic impacts of the Florida environmental horticultural industry in 2005. IFAS, UF L Economic Information Report FE675. Huey, L. A., G. J. Steck, and A. M. Fox. 2007. Biological notes on Melanagromyza ruelliae (Diptera: Agromyzidae), A seed feeder on the invasive Mexican Petunia, Ruellia tweediana (Acanthaceae). The Florida Entomologist 90: 763-765. Kollmann, J., L. Frederiksen, P. Vestergaard, H. H. Bruun. 2007. Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system. Biological Invasions 9: 31-42. Jordan, N., D. A. Mortensen, D. M. Prenzlow, and K. C. Cox. 1995. Simulation analysis of crop rotation effects on weed seedbanks. Am erican Journal of Botany 82: 390-398. Lee, D. W. and K. R. Dowmun. 1991. The spect ral distribution of biologically active solar radiation at Miami, Florida, USA. Internat ional Journal of Biom eteorology 35: 45-54. Long, R. W. and O. Lakela. 1976. A flora of tropi cal Florida: a manual of the seed plants and ferns of Southern Peninsular Florida. Miami, FL: Banyan Books. 962 pp.

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106 Lutman, P. J. W., G. W. Cussans, K. J. Wri ght, B. J. Wilson, G. McN Wright, and H. M. Lawson. 2002. The persistence of se eds of 16 weed species over six years in two arable fields. Weed Research 42: 231-241. Mack, R. N., D. Simberloff, W. M. Lonsdale H. Evans, M. Clout, and F. A. Bazzaz. 2000. Biotic invasions: causes, epidemiology, globa l consequences, and control. Ecological Applications 10: 689-710. Magda D., M. Duru, and J. P. Theau. 2004. Defi ning management rules for grasslands using weed demographic characterist ics. Weed Science 52: 339-345. Mennan, H. and B. H. Zandstra. 2006. The eff ects of depth and duration of seed burial on viability, dormancy, germination, a nd emergence of ivyleaf speedwell ( Veronica hederifolia ). Weed Technology 20: 438-444. [NRCS] Natural Resources Cons ervation Service and United States Department of Agriculture 2007. Web soil survey. http://websoilsurvey.nrcs.usda.gov. Accessed: September 2007. Peters, J. 2000. Tetrazolium testing handbook: contribution No. 29 to the handbook on seed testing. Association of Offi cial Seed Analysts. 21 pp. Pimental, D., L. Lach, R. Zumiga, and D. Mo rrison. 2000. Environmental and economic costs of nonindigenous species in the United States. BioScience 50(9): 53-65. Pimental, D., R. Zumiga, and D. Morrison. 2005. Update on environmental and economic costs associated with alien-invasive species in the United States. Ecological Economics 52: 273288. Roberts, H. A. 1981. Seed banks in so il. Advances Applied Biology 6: 1-55. [SAS] Statistical Analysis Syst ems Institute, Inc. 2007. SAS/STAT Users Guide. Version 9.1.3. Cary, NC: SAS Institute. 375 pp. http://support.sas.com/documentation/ onlinedoc/91pdf/sasdoc _913/whatsnew_10304.pdf Accessed: September 2007. Scherff, E. J., C. Galen, and M. L. Stanton. 1994. Seed dispersal, seedling survival and habitat affinity in a snowbed plant: limits to the distribution of the snow buttercup, Ranunculus adoneus Oikos 63: 405-413. Swanton, C. J. and B. D. Booth. 2004. Manageme nt of weed seedbanks in the context of populations and communities. Weed Technology 18: 1496-1502. Thompson, K. and J. P. Grime. 1979. Seasonal vari ation in seed banks of herbaceous species in ten contrasting habitats. J ournal of Ecology 67: 893-921.

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107 Tobe, J. D., K. C. Burks, R. W. Cantrell, M. A. Garland, M. E. Sweeley, D. W. Hall, P. Wallace, G. Anglin, G. Nelson, J. R. Cooper, D. Bi ckner, K. Gillbert, N. Aymond, and K. Greenwood, N. Raymond. 1998. Florida wetlan d plants: An identification manual. Tallahassee, FL: Florida Department of Environmental Protection. 598 pp. Turnbull, L. A., M. J. Crawly, and M. Rees. 2000. Are plant populations seed-limited? A review of seed sowing experiments. Oikos 88: 225-238. [USDA] United States Department of Agricu ltural and National Re sources Conservation Services. 2007. The PLANTS database. http://plants.usda.gov. Accessed: May 2007. Van Mourik, T. A., T. J. Stomph, and A. J. Murdoch. 2005. Why high seed densities within buried mesh bags may overestimate depletion rates of soil seed banks Journal of Applied Ecology. 42: 299-305. Warrence, N. J., J. W. Bauder, and K. E. Pear son. 2002. Basics of Salinity and Sodicity Effects on Soil Physical Properties. Department of Land Resources and Environmental Sciences, Montanta State University-Bozeman. 30pp. Wilson, S. B., P. C. Wilson, and J. A. Alba no. 2004. Growth and development of the native Ruellia caroliniensis and invasive Ruellia tweediana HortScience 39: 1015-1019. Wilson, S. B. and L. K. Mecca. 2003. Seed production and germination of eight cultivars and the wild type of Ruellia tweediana : A potentially invasive ornamental. Journal of Environmental Horticulture 21: 137-143. Witztum, A. and K. Schulgasser. 1995. The me chanics of seed expulsion in Acanthaceae. Journal of Theoretical Biology 176: 531-542. Wunderlin, R. P. and B. F. Hansen. 2007. Atlas of Florida Vascular Plants. http://www.plantatlas.usf.edu. Accessed: June 2007. Zomlefer, W. B. 1994. Guide to Flowering Plant Fa milies. Chapel Hill, NC: University of North Carolina Chapel Hill Press. 430 pp.

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108 BIOGRAPHICAL SKETCH Karen V. S. Hupp, the daughter of Stephen and Kathryn Shepherd was born on 1981, in Chicago, Illinois. The third of four children w ho mostly grew up in Spartanburg, South Carolina, she graduated from Paul M. Dorman High School in 1999. She received a BS degree in Environmental Science from Catawba College in North Carolina in 2003. During that program she worked on invasive plant removal at the Cowpens National Battle Field and conducted research on the restoration of granite outcr ops at Dunns Mountain, NC. Since September 2003, Karen has worked as an OPS assistant with Alison Foxs research t eam, providing excellent support to many projects related to invasive plants. In Fall 2005, she started her MS program at the University of Florida with the objective of furthe r increasing her knowledge of invasive plant ecology and management. Also that fall she married Jason R. Hupp on September 18.