MASTER THESIS The limiting similarity of seed size and resource- based suppression: A glasshouse experiment Phillip Hughes Matriculation Number: 03637295 Study Programme: (MSc) Master of Science Sustainable Resource Management Supervisors: Prof. Dr. Johannes Kollmann & Florencia Yannelli Chair of Restoration Ecology Technische Universität München
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MASTER THESIS The limiting similarity of seed size and resource-
based suppression: A glasshouse experiment
Phillip Hughes Matriculation Number: 03637295
Study Programme: (MSc) Master of Science Sustainable Resource
Management
Supervisors: Prof. Dr. Johannes Kollmann & Florencia Yannelli Chair of Restoration Ecology Technische Universität München
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Table of Contents
Chapter 1: Introduction and Background ....................................................................................... 2
Introduction to the driver for the research ........................................................................................ 2
Background knowledge pertinent to the research ............................................................................. 3
Restoration and succession ............................................................................................................ 3
2012; de Bello 2012; Chalmandrier et al. 2013; Laliberté et al. 2013).
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Figure 1. The range of scales proposed ecological filters that can act to lead to an actual observed community (Adapted from Götzenberger et al., 2012.).
The theory is such, that from the global species pool, a regional species pool is then defined
by historical and biogeographical processes (‘phylogeographic assembly’; (Kembel 2009). This
can be viewed as the gradients of species richness associated with latitude and elevation, in
that plants species richness often increases near the equator and decreases with elevation,
the historical mechanisms for which are numerous (time from colonisation, rates of
extinction/diversification, etc.). From the regional species pool dispersal and the abiotic
environment (e.g. climate) act to filter adapted species into the local species pool. From the
local species pool, habitat filtering and biotic assembly rules will define the actual community
of plant species (Figure 1). Articulately put, assembly rules are a community level analogue of
natural selection (Keddy 1992). At the actual community level, which is the purpose of this
paper to explore, species interactions (e.g. competition) are driving assembly (Mouchet et al.
2010).
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Functional Traits
The use of functional traits when understanding assembly rules is persuasively surmised with
the following comparison between species based assembly rules and those that are trait
based:
“A species form of ‘rule’ might be: if a community has species A, then it usually will not also have species B or C unless species D or E and either species F or G are present, while if a
community has species D, then it will not also have species E unless species F or G and either species A, B, or C are also present, etc. A more trait based rule is clearer: the proportion of
species from each functional group will tend to remain constant for each observation.” (Weiher & Keddy 1995)
From this quotation it is easy to understand why there is growing recognition that
classification of plant species based on their function, as opposed to their taxonomic identity,
is a more pragmatic approach to addressing ecological questions such as community structure
and vegetation response to environmental change (Cornelissen et al. 2003).
Violle et al. (2007) have concisely formulated a definition of functional trait with respect to
plants, suggesting it is any trait which impacts the fitness indirectly via its effects on growth,
reproduction and survival. The paper also makes the further distinction of a so called
‘performance trait’, that being a direct measure of fitness, and in plants only three such traits
and plant survival (Violle et al. 2007). In this respect, seed mass is an interesting trait for the
establishment phase of plant reproduction. Larger seeds store more resources, which are
therefore able to help young seedlings to survive and establish when there is low resources
(drought, deep shade) and high competition. Smaller seeds can be produced in larger numbers
with the same reproductive effort, and therefore raises the likelihood of establishment of a
seedling where competition is low and resource availably is high. (Cornelissen et al. 2003).
Limiting Similarity
The term limiting similarity was introduced by MacArthur & Levins (1967). This was work that
was based on Lotka-Volterra equations and MacArthur & Levins provided a mathematical
explanation in which they posited that “there is a limit to the similarity of competing species
which can coexist. The total number of species is proportional to the total range of the
environment divided by the niche breadth of the species” (MacArthur & Levins 1967). This
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early position was then more generalised away from single resource models, to be a term that
is applied to the relationship between some measure of the difference in the competitive
abilities of two competitors and the similarity in resource utilisation (Abrams 1983).
The use of the word limiting similarity in the context of assembly rules suggests that species
assemble within a community based on resource use functional traits, and that theys can more
readily coexist if they differ in their traits, such that competition is reduced (Götzenberger et
al. 2012) (Figure 2).
Figure 2: A graphical representation of the limiting similarity hypothesis depicting two invaders establishment success based on functional divergences with a native community (source:
Another important aspect to consider in terms of resource based suppression is the increase
in resource availability commonly found in roadsides. The availability of soil nutrients, along
with water and light are the three basic requirements for plant growth (Raven et al. 2013). An
increase in soil nutrient levels will lead to increased biomass of the plants growing in it (Poorter
& Nagel 2000). On the other hand, for a native plant community, nutrient addition will
increase its vulnerability to invasion by alien species by increasing the amount unused
resources in the system (Davis et al. 2000).
We are interested to see the effects that soil nutrient addition will have during the
simultaneous establishment of a native community and Invasive species and how it affects the
competition parameters between the two.
We designed a greenhouse experiment to reduce the knowledge gap that exists with respect
to applying the limiting similarity hypothesis to guide optimal seed mix choices. Additionally,
we are interested to test if a limiting similarity exists in seed mass of establishing individuals
of IAS compared to the average seed mass of the native community it is competing with. It is
hoped the results from this experiment could be applied to future grassland restoration
projects. Hence, the aim of the study is to investigate the contribution plant seed size and soil
nutrient content have in affecting competitive success of invasive alien species in a
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greenhouse environment. Or conversely and more importantly for restoration ecology,
whether seed size is an adequate functional trait to be used to infer wether a native
community will show greater resistance to invasion from invasive alien species.
We hypothesize that: 1) Large-seeded native communities are more effective at supressing
the invasion success of small- and large-seeded invasive alien species; 2) Small-seeded native
communities are only effective at supressing the invasion of small-seeded invasive alien
species; 3) Nutrient enriched soils will increase the invasion success of all invasive alien species
within the small- and large-seeded native communities.
.
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Materials and Methods
Species Selection
Invasive Alien Species
Eight invasive alien plant species were selected in order to be seeded in native seed
communities. The small group included seeds under 0.5 mg and the large group those over 2
mg (Table 1). The experimental seed bank of these species was designed using average
values published for Ambrosia artemisiifolia and extrapolated for the other species for
better comparison (Rothrock et al. 1993; Fumanal et al. 2008)
Table 1: The chosen target invasive alien species were collected from the field (see Appendix 1 for locations)
Genus species Family Seed size group
Solidago gigantea Asteraceae Small
Solidago canadensis Asteraceae Small
Conyza canadensis Asteraceae Small
Erigeron annuus Asteraceae Small
Ambrosia artemisiifolia Asteraceae Large
Fallopia japonica Polygonaceae Large
Impatiens glandulifera Balsaminaceae Large
Heracleum mantegazzianum Apiaceae Large
Native Species
A pool of species with their corresponding traits was prepared, based on the roadside mixes
of the Rieger-Hofmann catalogue for Germany. The community composition was randomly
selected from this pool according to seed mass and life span for each replica, consisting of five
combinations of 5 native grassland species. The species were selected according to the mean
seed mass, including two species of grasses in both target community mixtures (Table 2). Two
levels were chosen for seed mass (i.e. small, large) and these were defined according to the
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0.25–0.75 percentile from the average seed mass of all species of the roadside seed mixture
(0.3 mg and 1.3 mg; Rieger-Hofmann GmbH)
Table 2: The native species chosen to make up the native community purchased from the commercial source
Genus species Family Seed Size Group
Achillea millefolium Asteraceae Small
Agrostis capillaris Poaceae Small
Origanum vulgare Lamiaceae Small
Papaver rhoeas Papaveraceae Small
Plantago media Plantanginaceae Small
Poa compressa Poaceae Small
Bromus erectus Poaceae Large
Centaurea cyanus Asteraceae Large
Centaurea scabiosa Asteraceae Large
Helictotrichon pubescens Poaceae Large
Knautia arvensis Dipsacaceae Large
Salvia pratensis Lamiaceae Large
Sanguisorba minor Rosaceae Large
Seed Preparation and Germination Trials
Invasive Species
Previous to the main experiment, all invasive species seeds were collected (see Appendix 1),
stratified when needed and tested for potential germination at ideal conditions. The amount
of 50 seeds were placed in Petri dishes, evenly distributed on filter papers (9 cm diameter)
moistened with de-ionized water and replicated five times. The stratification was carried at
4.5 °C and the period varied among the species (Table 3). After stratification, they were
placed into a climatic cabinet under day conditions for 16 hours (20 °C with artificial light)
and night conditions for 8 hours (12 °C; no light).
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Table 3: Time period required for stratification according to the species. This was defined as the period of time after more than 10% of the seeds started germination.
Species Stratification (4°C) Heracleum mantegazzianum 4 weeks
Fallopia japonica 3 weeks
Ambrosia artemisiifolia 3 weeks
Impatiens grandulifera 2 weeks
Erigeron annuus 4 weeks
Solidago gigantea No stratification needed
Solidago canadensis No stratification needed
Conyza canadensis No stratification needed
Seed were prepared for the main experiment by placing them in little biodegradable
transplanting pots for being stratified. The information on stratification requirements was
used later for the preparing the seeds for the main experiment
Emergence trials In order to avoid a density effect due to differences in emergence of the
native species, emergence trials were performed. These trials aimed to estimate the final
number of seed that had to be sown according to how each species performed. For this, 30
seeds from each native species were place on top of a 36 cm2 pot filled with low nutrient [pH
times and placed into a climatic chamber for germination. The trials were carried out initially
under day conditions for 12 hours (20 °C with artificial light) and night conditions for 8 hours
(10 °C; no light), but three weeks later was changed to day conditions for 16 hours (22 °C with
artificial light) and night conditions for 8 hours (10 °C; no light).
Experimental Design
The greenhouse experiment was established in the Centre of Greenhouses and Laboratories
Dürnast, part of the Life Science Centre Weihenstephan, Technische Universität München in
Munich, Germany. Following from recommendations by Snaydon (1991) an additive
experimental design was chosen in order to investigate the competitive ability of the IAS
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target species. This way, the density of the target species in the pure stands (controls) was the
same as the density of the target species in the mixtures (‘treatments’). The experiment ran
for 8 weeks (from mid-May until early July) in an unheated greenhouse which had a
temperature of 22 ± 3 °C (mean ± SD).
The experiment had a randomised factorial block design. Eight IAS were planted within
different native plant communities, this being the first factor, made up of two levels, large
seeded and small seeded native community (small with average mass <0.5 mg and large-
seeded mix > 1.9 mg). The next factor was nutrient addition and was made up of two levels,
nutrient addition or none. The nutrient addition consisted of the addition of 10 ml at a 1.5‰
concentration of fertilizer Ferty 3 (15% N, 10% P2O5, 15% K2O) to each pot (3 l). The treatments
were then replicated five times to make a total of 160 pots. Controls were also implemented
whereby the eight Invasive alien species were subjected to only nutrient additions without
the native community influence and the native communities experienced nutrient addition
treatment without the presence of invasive species (Figure 3).
Figure 3: The experimental design of the study. Eight invasive plant species are subjected to influences from two types of native community based on seed size. The impact of nutrient addition is also tested. Controls are used to compare performance of species subject to the treatments with respect to no treatment. (Illustration: M. Brockard and H. Vogt)
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This design allows for the observation of the competitive effect that large and small seeded
native communities impose on establishment by large and small seeded IAS and whether
nutrient levels also exert an effect on the establishment success.
The exact mixtures were randomised (Appendix 2), with the ratio of seeds present in each
kept constant. The seeding density was kept constant at 15 g m-2. This density was chosen to
keep results comparable between a preliminary experiment (F. Yanelli, unpubl. work) and
this experiment. The seed producer recommends a seeding density of 3 g m-2 (Kiehl et al.
2010), as we had five times more soil volume than the previous experiment but overall
smaller surface, we have in fact created high density communities. In order to achieve this
constant density, the average seed mass was calculated for the native seeds (2.18 mg). It
was calculated that approximately 213 seeds were required per pot. As we were using five
species per pot, this equated to 43 seeds per species per pot. Using the germination data
obtained before experiment begin, we were able to adjust the seed count required, relative
to the germination rate per species as shown in Table 4.
Table 4: Depicting the germination rates that were obtained from germination tests prior to
experiment begin. These rates were then used to adjust the amount of seeds used per species to
keep the density of the native species in the community’s constant at 15 g m-2.
Genus species Seed size Germination rate (%) Total seeds
Bromus erectus Large 80 52
Centaurea cyanus Large 83 51
Centaurea scabiosa Large 78 53
Helictotrichon pubescens Large 41 69
Knautia arvensis Large 88 48
Salvia pratensis Large 46 66
Sanguisorba minor Large 80 52
Achilliea millefolium Small 90 48
Agrostis capillaris Small 68 57
Origanum vulgare Small 65 58
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Papaver rhoeas Small 56 62
Plantago media Small 58 61
Poa compressa Small 65 58
Potting up of the experiment involved filling 20 cm diameter pots with low nutrient potting
soil (‘Einheits Erde Special’). Initially, an empty 200 ml plastic cup was left empty in the
centre of the pots to create a void in the soil. The same 200 ml plastic cups were filled
separately. The filled pots and cups were then brought into the greenhouse and randomly
distributed onto one of the tables. These were left for 1 week to be watered in and allow the
soil to settle. Extra soil was added to individual pots and cups that required more soil so that
soil amount was kept constant. Further on, each of the 200 ml plastic cups was sown with
one invasive species seeds and left to germinate. At the same time, native species mixes
were evenly sown into the larger pots. After 4 weeks, the invasive species that were alive
and developed first true leaves were then translocated into the respective pots (Appendix
3). In effect within each 20 cm diameter pot, one seedling of the Invasive species was
surrounded by a native community made up of either small or large seeded members (Figure
4).
Figure 4: The distribution of the native communities in each pot. Placed in the middle of the community in treatment pots was the invasive alien species. In control pots, either the native community or the invasive alien species was not present. (Image source: Florencia Yannelli)
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After set-up, the pots were separated across two tables within the greenhouse. One table
would receive the nutrient treatments and the other table would not receive nutrient
treatments, for preventing the non-nutrient addition treatments to be affected when
watering. In order to control for edge effects and micro climatic conditions, the pots were
redistributed within their respective tables once per month. The pots were initially watered
every day from above until germination and establishment of the plants, then changed to
watering every second day from beneath allowing the pots to soak for 1 hour.
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Variables and Measurement
The response variables that were measured are summarised in the following table:
Variable Description
TOTAL BIOMASS Total above ground dried mass
IAS LEAVES MASS Combined leaves dried mass
IAS STEMS MASS Combined stems dried mass
IAS SPECIFIC LEAF AREA Ratio of Leaf area to dry mass
IAS HEIGHT Height to apical meristem
The chosen plant characteristics are considered acceptable ways of measuring plant
responses to soil resources, response to disturbance, plant defence/protection and
competitive strength (Cornelissen et al. 2003). Specific leaf area was measured using the
approach as per the Cornelissen paper. Directly removing three healthy leaves (including
petiole) and scanning them into the computer software ImageJ for precise area
measurements (courtesy; Wayne Rasband, Research Services Branch, National Institute of
Mental Health, Bethesda, Maryland, USA.) Moreover, the drying procedure was to take all
collected material and place it into an oven at 75 oC for a minimum of 48 hours.
Data Analysis
The relative competition index (RCI) was used to estimate the competitive effect of native
species on the invasive species, with the following formula (Weigelt & Jolliffe 2003):
RCIγ = γControl − γTreatment
γControl
Using this formula, a particular variable is measured in the controls, i.e. biomass, and it is
compared to the value an individual gains under treatment conditions, making direct
comparison of competitive performance achievable
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The RCI was calculated for all response variables measured i.e. native biomass, invasive total
biomass, leaves biomass, steams biomass, SLA and height. Subsequently, RCI and row values
were tested for normality using the modified Shapiro-Wilk test (Rahman & Govindarajulu
1997). Only Ambrosia artemisiifolia provided normally distributed data across all but one
variable (RCI-SLA). As the assumption for normal distribution failed across the majority of
species, it was decided that in order to make comparisons using all species, only non-
parametric testing would be applied. Therefore, the effects of the treatments were verified
using the non-parametric Kruskal-Wallis test (Kruskal & Wallis 1952) to explore multi-factor
interactions and Wilcoxon test for pairwise comparison across the two factors, seed size and
nutrient addition. Each invasive species was analysed separately, but then all small and large
seeded IAS were grouped together to seek for general results on the effect of limiting
similarity in terms of the seed size.
All statistical analysis were performed using Infostat software (Di-Rienzo et al. 2013).
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Results
From the initially decided upon eight species of IAS, there was no data collected for Conyza
canadensis, Fallopia japonica, Heracleum mantegazzianum and Impatiens glandulifera.
There are two reasons for this omission: First, Heracleum mantegazzianum, Impatiens
glandulifera, and Conyza canadensis did not germinate adequate to fulfil the requirements
for all replicates required, this means that these species were removed before beginning and
transfer of these species into the native species communities did not occur. Second,
although Fallopia japonica did show adequate germination to fulfil the replicates
requirement; this species experienced during the course of the experiment for unknown
reasons an excessive rate of mortality and so there were not enough individuals left to be
statistically valid. Hence, I show results only for three small-seeded species (Solidago
Canadensis, Solidago gigantea and Erigeron annuus) and one large-seeded species
(Ambrosia artemisiifolia).
Species-specific results
There was no significant effect of either seed size of the native community or nutrient
addition on Solidago canadensis (p > 0.05). On the other hand, Solidago gigantea produced
observable differences in RCI-Native Biomass (H = 7.9, p = 0.048) for both factors seed mass
and nutrient addition. Wilcoxon tests showed that the statistical difference for the relative
competition index in native biomass was a result of the nutrient addition treatment. RCI-
native biomass was lower in no nutrient additions (mean = -0.13) when compared to
nutrient addition treatments (mean = 0.15; W = 121, p = 0.011).
In Erigeron annus differences were observed in the RCI-leaf area (H = 9.63, p = 0.022).
Further, pairwise testing with the nutrient factor using Wilcoxon tests did not produce any
significant difference. There was however a statistical difference observed in the RCI-leaf
area (LSM mean 0.93, SSM mean 0.76, [W = 94, p = 0.0025]), RCI-leaf mass (LSM mean 0.95,
SSM mean 0.77, [W= 90, p=0.0084]) and RCI-height (LSM mean 0.71, SSM mean 0.51, [W=
86.5, p=0.0217]).
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In terms of the only large-seeded invasive species Ambrosisa artemisiifolia, multifactor
comparisons testing did not indicate any observable differences between the treatments.
Wilcoxon pairwise comparisons on nutrient addition indicated a significant difference in
relative competition index of SLA (W = 71, p = 0.010). No nutrient addition treatments
produced lower competition indexes (mean = -0.12) than the nutrient additions (mean =
0.13). Seed mass of the native community was not found to be significant in terms of each
of the five measured variables.
Biomass of all invasive species was lower in all treatments when compared to the controls,
this is reflected in the relative competition index value for all treatments across all species
being above one [suppression] (Figures 5–8). There was no statistical difference between the
factors nutrient addition and native community seed size (both, p > 0.05).
Figure 5: Relative competition index for Solidago canadensis biomass showing similar levels of suppression across all treatments. (Lsm= Large seeded community; Ssm= Small seeded community; No-nut is no nutrient additions; Nut= Nutrient additions)
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Figure 6: Relative competition index for Solidago gigantea biomass showing similar levels of suppression across all treatments. (Lsm= Large seeded community; Ssm= Small seeded community; No-nut is no nutrient additions; Nut= Nutrient additions)
Figure 7: Relative competition index for Erigeron annuus biomass showing similar levels of suppression across all treatments. (Lsm= Large seeded community; Ssm= Small seeded community; No-nut is no nutrient additions; Nut= Nutrient additions)
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Figure 8: Relative competition index for Ambrosia artemisiifolia biomass showing similar levels of suppression across all treatments. (Lsm= Large seeded community; Ssm= Small seeded community; No-nut is no nutrient additions; Nut= Nutrient additions)
Combined Results
Nutrient additions did not significantly affect any of the measured response variables for
either the large or small IAS (W = 3415.5; p > 0.05). The total biomass of all invasive species
tested was significantly lower in both the large- and small-seeded communities when both
are compared to the control treatments (p < 0.0001) (Figure 9).
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Figure 9: Large and small seed communities were indistinguishable in their ability to reduce the total biomass of both large and small seeded invasive alien species when compared to the control treatments (p < 0.0001).
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Discussion
Specifically the results indicate that regardless of seed size, with no nutrient addition the
native communities produced more biomass in the treatments where Solidago gigantea was
present. The opposite was true for the nutrient addition treatments, in this case the native
communities produced more biomass in the treatments where Solidago gigantea was not
present. Leaf area of Erigeron annus was least supressed in the small seeded communities
with nutrient addition.
The results of the experiment indicate that none of the three postulated hypotheses could be
accepted. Native communities made up of large seed mixes were not more effective at
supressing the invasion success of either small or large seeded invasive alien species when
compared to the supressing effect exerted by small seeded native communities. Furthermore,
nutrient enrichment in this experiment did not indicate that the invasion success of alien
species was increased in either large or small seeded communities. A limiting similarity
between native community seed size and IAS seed size was not observed. An important point
to note however, is that a true comparison between effects on large- and small-seeded
invasive alien species could not be determined with the experiment as it only was left with
one large-seeded species (Ambrosia artemisiifolia) for the comparison.
A result which was a little less anticipated was seen where the small-seeded mixes supressed
the biomass of the invasive alien species equally as well as the large-seeded mixes. This result,
although disproving our hypothesis on limiting similarity of seed size, does however indicate
the role interspecific competition plays in community dynamics. In all cases, the invasive
species produced significantly more biomass in the control pots where no native community
was present. This can only be explained by the absence of the community competing for
resources with the IAS (Aerts 1999). An interesting addition to explore this result next time
may be to include measurement of below ground characteristics as they have been shown to
be stronger (Aerts 1999)(Casper & Jackson 1997). Also following this, the rather symmetrical
effects observed between the two treatments for seed size in this experiment also correlate
with results from other studies that have shown that the effects of belowground competition
act more symmetrically when compared to above ground competition effects (Casper &
Jackson 1997).
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Another explanation for the suppression affected by both large- and small-seeded
communities could also be linked in to the seeding density that was used for the experiment.
We had chosen 15 g m-2 which was approximately five times the recommended density from
the seed supplier. Higher density, which ties in with the idea of reducing the spatial resources
available to invading species may be the key driver for the suppression ability of newly
established plant communities in this experiment. This was the finding of an experiment in
which a doubling of the recommended seeding rate of wheat plants lead to a halving of weedy
species present in the crop (Lemerle et al. 2004)
Interestingly, the nutrient additions in this experiment did not induce a biomass increase in
either the IAS or the native communities present. This is contrary to previous work in this area
suggesting that nutrient increases should lead to an increase in biomass (Poorter & Nagel
2000). A possible explanation for this is that the soil used for the experiment was already over
abundant in nutrients relative to what the plants required. In essence, even the treatments
that were considered no nutrient additions actually were abundant with unused resources,
creating a situation where nutrient availability was not a limiting resource, even in the no
nutrient treatments. This would tie in with a secondary point, in that an increase in limiting
resource’s should increase that invasibility of a native community (Davis et al. 2000) and this
was not observed with our data.
Conclusion
Our results although not allowing us to accept our original hypotheses, do provide important
results that can be adopted by land managers tasked with restoring grassland communities.
Owing to the effect we observed that small seeded mixes supressed invasive species equally
as well as large seeded communities. This suggested that density has a very important role.
Land managers who need to purchase seed do so by weight. It would be the logical choice for
them, based on our results to purchase a mix that includes smaller seeded species, as for the
same weight of seed they will end up with more individuals and therefore, a higher plant
density for a lower cost.
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Chapter 3: Final Discussion and Conclusion
The damage caused by invasive species are one of the most significant challenges that land
managers and restoration ecologists will have to deal with leading into the future (Hobbs &
Harris 2001). The invasion of exotic species poses a two folded problem, not only do they
contribute to environmental degradation, they also substantially impede efforts to restore
ecosystems (Funk & Cleland 2008).
Regardless of the causes for degradation, effective restoration and management will require
increased levels of understanding of the processes and properties of ecosystems that convey
resistance and resilience from invasive alien species (Falk et al. 2006). In essence, in the case
of de novo establishment of a grassland communities in roadsides it is imperative to recreate
communities with these processes and properties (Saunders & Hobbs 1991). The task for
ecologists is to integrate and obtain relevant ecological information and test these approaches
through continued experimentation.
Integrated weed management is the deliberate selection, integration and implementation of
effective weed control measures with due consideration of economic, ecological and
sociological consequences (Hobbs & Humphries 1995). In using an integrated weed
management approach the first step in actually preventing the infestation is the designing of
roadside communities based on properties that convey resilience. This is where sound
ecological understanding in areas such as succession, assembly rules, functional traits and
limiting similarity theory play a vital role (Walker et al. 2007).
A central principle in integrated weed management is the idea of an economic threshold
(Figure 10) (Swanton & Weise 1991). Prior to reaching this threshold, natural weed controlling
methods should be manipulated to their fullest. Once this threshold has been reached action
in the form of chemical control must be taken in order to prevent an outbreak of a pest species
before it can cause an economic injury (Swanton & Weise 1991). The term economic injury
implies that after breaking a certain threshold, the cost to eradicate the invading population
becomes exponentially more than the preventative actions (Harker & O’Donovan 2013).
Integrated weed management sustainable management practices such cultural controls that
maintain pests at low levels. This keeps pesticide use to a minimum by using them only in an
outbreak situation (Harker & O’Donovan 2013).
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Figure 10: The figure illustrates the principle of the economic threshold and is based on a logistical growth curve. (Image source: http://www.weedcenter.org/textbook/8_jacobs_options.html)
Roadsides are a hotspot for the development of IAS infestations as a result of high seed rain
and favourable environmental conditions for their growth (Haan et al. 2012). The dispersal
vector of passing vehicles cannot be controlled. It is therefore inevitable that there will be
some level of infestation of invasive alien species present even in the most well planned
restored roadside sites. Coupled with sound ecology theory, integrated weed management is
a sustainable management choice for invasive alien species in roadside grassland
communities.
Ecological theory, and the significant research that supports it is a solid foundation for devising
management and restoration methods that will create or maintain sustainable ecosystems
(Clewell et al. 2004). Adding to the theoretical suite of options available to restoration
ecologists is the theory of limiting similarity (Funk & Cleland 2008). Our experimentation did
indicate that seed mass is an inappropriate functional trait for use in a limiting similarity
approach. Therefore the experimentation is still a step forward for scientific understanding
and the integration of limiting similarity theory into sustainable management strategies and
practical restoration techniques.
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Acknowledgements
I wish to extend my utmost gratitude to Florencia Yanelli for her limitless knowledge, support
and guidance throughout the experimental and writing stage of this thesis work and not giving
up on me. I wish to thank the staff at Dürnast Research Laboratory and Greenhouse for their
technical guidance and support. Thank you to Martina Brockard and Hannah Vogt for assisting
in establishing the experiment. Lastly I wish to thank Prof. Dr. Johannes Kollmann for accepting
my thesis proposal and providing me with his expert knowledge, technical guidance and the
chairs resources to fulfil the thesis requirements.
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References
Abrams, P., 1983. The theory of limiting similarity. Annual review of ecology and systematics, (34). Available at: http://www.jstor.org/stable/2096978 [Accessed March 29, 2014].
Aerts, R., 1999. Interspecific competition in natural plant communities: mechanisms, trade-offs and plant-soil feedbacks. Journal of Experimental Botany, (January 1999). Available at: http://jxb.oxfordjournals.org/content/50/330/29.short [Accessed December 15, 2014].
De Bello, F., 2012. The quest for trait convergence and divergence in community assembly: are null-models the magic wand? Global Ecology and Biogeography, 21(3), pp.312–317. Available at: http://doi.wiley.com/10.1111/j.1466-8238.2011.00682.x [Accessed July 12, 2014].
Casper, B.B. & Jackson, R.B., 1997. Plant Competition Underground. Annual Review of Ecology and Systematics, 28(1), pp.545–570. Available at: http://arjournals.annualreviews.org/doi/abs/10.1146%2Fannurev.ecolsys.28.1.545.
Chalmandrier, L. et al., 2013. A family of null models to distinguish between environmental filtering and biotic interactions in functional diversity patterns. Journal of vegetation science : official organ of the International Association for Vegetation Science, 24(5), pp.853–864. Available at: http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=4003529&tool=pmcentrez&rendertype=abstract [Accessed August 1, 2014].
Cleland, E.E., Larios, L. & Suding, K.N., 2013. Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap. Restoration Ecology, 21(3), pp.390–398. Available at: http://doi.wiley.com/10.1111/j.1526-100X.2012.00896.x [Accessed March 28, 2014].
Clewell, A., Aronson, J. & Winterhalder, K., 2004. The SER International primer on ecological restoration. , 2(2), pp.206–207. Available at: http://scholar.google.com/scholar?hl=en&btnG=Search&q=intitle:The+SER+International+Primer+o+n+Ecological+Restoration+O+verview#1 [Accessed October 22, 2014].
Cody, M.L. & Diamond, J.M., 1975. Ecology and Evolution of Communities, Belknap Press of Harvard University Press. Available at: http://books.google.de/books?id=j_idbVxwzpQC.
Connor, E. & Simberloff, D., 1979. The assembly of species communities: chance or competition? Ecology, 60(6), pp.1132–1140. Available at: http://www.jstor.org/stable/1936961 [Accessed November 7, 2014].
Cornelissen, J.H.C. et al., 2003. A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Australian Journal of Botany, 51(4), p.335. Available at: http://www.publish.csiro.au/?paper=BT02124.
Costanza, R. et al., 2014. Changes in the global value of ecosystem services. Global Environmental Change, 26(0), pp.152–158. Available at: http://www.sciencedirect.com/science/article/pii/S0959378014000685.
Davis, M., Grime, J. & Thompson, K., 2000. Fluctuating resources in plant communities: a general theory of invasibility. Journal of Ecology, pp.528–534. Available at:
Master Thesis
33
http://onlinelibrary.wiley.com/doi/10.1046/j.1365-2745.2000.00473.x/full [Accessed November 15, 2014].
Falk, D.A. et al., 2006. Foundations of Restoration Ecology, Society for Ecological Restoration International.
Fumanal, B., Gaudot, I. & Bretagnolle, F., 2008. Seed-bank dynamics in the invasive plant, Ambrosia artemisiifolia L. Seed Science Research, 18(02), pp.101–114. Available at: http://journals.cambridge.org/article_S0960258508974316.
Funk, J. & Cleland, E., 2008. Restoration through reassembly: plant traits and invasion resistance. Trends in Ecology & …, 23(12), pp.695–703. Available at: http://www.ncbi.nlm.nih.gov/pubmed/18951652 [Accessed March 29, 2014].
Gerhold, P. et al., 2013. Functional and phylogenetic community assembly linked to changes in species diversity in a long-term resource manipulation experiment F. de Bello, ed. Journal of Vegetation Science, 24(5), pp.843–852. Available at: http://doi.wiley.com/10.1111/jvs.12052 [Accessed March 25, 2014].
Götzenberger, L. et al., 2012. Ecological assembly rules in plant communities--approaches, patterns and prospects. Biological reviews of the Cambridge Philosophical Society, 87(1), pp.111–27. Available at: http://www.ncbi.nlm.nih.gov/pubmed/21692965 [Accessed March 24, 2014].
Grube, A. et al., 2011. Pesticides Industry Sales and Usage 2006 and 2007 Market Estimates, Washington DC.
Haan, N.L., Hunter, M.R. & Hunter, M.D., 2012. Investigating Predictors of Plant Establishment During Roadside Restoration. Restoration Ecology, 20(3), pp.315–321. Available at: http://dx.doi.org/10.1111/j.1526-100X.2011.00802.x.
Herben, T. & Goldberg, D.E., 2014. Community assembly by limiting similarity vs. competitive hierarchies: testing the consequences of dispersion of individual traits A. MacDougall, ed. Journal of Ecology, 102(1), pp.156–166. Available at: http://doi.wiley.com/10.1111/1365-2745.12181 [Accessed March 25, 2014].
Hobbs, R.J. & Harris, J. a., 2001. Restoration Ecology: Repairing the Earth’s Ecosystems in the New Millennium. Restoration Ecology, 9(2), pp.239–246. Available at: http://doi.wiley.com/10.1046/j.1526-100x.2001.009002239.x.
Hobbs, R.J. & Humphries, S.E., 1995. An integrated approach to the ecology and management of plant invasions. Conservation Biology, 9(4), pp.761–770.
Keddy, P.A., 1992. Assembly and response rules: two goals for predictive community ecology. Journal of Vegetation Science, 3(2), pp.157–164. Available at: http://dx.doi.org/10.2307/3235676.
Kembel, S.W., 2009. Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests. Ecology letters, 12(9), pp.949–60. Available at: http://www.ncbi.nlm.nih.gov/pubmed/19702749 [Accessed March 19, 2014].
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Kiehl, K. et al., 2010. Species introduction in restoration projects – Evaluation of different techniques for the establishment of semi-natural grasslands in Central and Northwestern Europe. Basic and Applied Ecology, 11(4), pp.285–299. Available at: http://www.sciencedirect.com/science/article/pii/S1439179109001455.
Kruskal, W.H. & Wallis, W.A., 1952. Use of Ranks in One-Criterion Variance Analysis. Journal of the American Statistical Association, 47(260), pp.583–621. Available at: http://www.tandfonline.com/doi/abs/10.1080/01621459.1952.10483441.
Laliberté, E., Norton, D. a. & Scott, D., 2013. Contrasting effects of productivity and disturbance on plant functional diversity at local and metacommunity scales N. Mason, ed. Journal of Vegetation Science, 24(5), pp.834–842. Available at: http://doi.wiley.com/10.1111/jvs.12044 [Accessed March 28, 2014].
Laughlin, D.C., 2014. Applying trait-based models to achieve functional targets for theory-driven ecological restoration. Ecology letters, 17(7), pp.771–84. Available at: http://www.ncbi.nlm.nih.gov/pubmed/24766299 [Accessed July 9, 2014].
Lemerle, D. et al., 2004. Reliability of higher seeding rates of wheat for increased competitiveness with weeds in low rainfall environments. The Journal of Agricultural Science, 142(4), pp.395–409. Available at: http://www.journals.cambridge.org/abstract_S002185960400454X [Accessed December 15, 2014].
Lindenmayer, D. & Burgman, M., 2005. Vegetation loss and degradation. In Practical Conservation Biology. CSIRO Publishing.
Lonsdale, W., 1999. Global patterns of plant invasions and the concept of invasibility. Ecology, 80(5), pp.1522–1536. Available at: http://www.esajournals.org/doi/abs/10.1890/0012-9658(1999)080%5B1522:GPOPIA%5D2.0.CO%3B2 [Accessed November 15, 2014].
MacArthur, R. & Levins, R., 1967. The limiting similarity, convergence, and divergence of coexisting species. American naturalist, 101(921), pp.377–385. Available at: http://www.jstor.org/stable/2459090 [Accessed October 22, 2014].
May, F. et al., 2013. Plant functional traits and community assembly along interacting gradients of productivity and fragmentation. Perspectives in Plant Ecology, Evolution and Systematics, 15(6), pp.304–318. Available at: http://linkinghub.elsevier.com/retrieve/pii/S1433831913000590 [Accessed March 28, 2014].
Mouchet, M. a. et al., 2010. Functional diversity measures: an overview of their redundancy and their ability to discriminate community assembly rules. Functional Ecology, 24(4), pp.867–876. Available at: http://doi.wiley.com/10.1111/j.1365-2435.2010.01695.x [Accessed March 20, 2014].
Philipp Schledorn, M.K., 2014. Detection of Glyphosate Residues in Animals and Humans. Journal of Environmental & Analytical Toxicology, 04(02). Available at: http://www.omicsonline.org/open-access/detection-of-glyphosate-residues-in-animals-and-humans-2161-0525.1000210.php?aid=23853 [Accessed October 31, 2014].
Poggio, S.L., 2005. Structure of weed communities occurring in monoculture and intercropping of field pea and barley. Agriculture, Ecosystems & Environment, 109(1-2), pp.48–58. Available at:
Master Thesis
35
http://linkinghub.elsevier.com/retrieve/pii/S0167880905001180 [Accessed November 15, 2014].
Poorter, H. & Nagel, O., 2000. The role of biomass allocation in the growth response of plants to different levels of light, CO2, nutrients and water: a quantitative review. Functional Plant Biology, 27(12), p.1191. Available at: http://www.publish.csiro.au/paper/PP99173_CO.
Price, J. & Pärtel, M., 2013. Can limiting similarity increase invasion resistance? A meta‐analysis of experimental studies. Oikos, 122(5), pp.649–656. Available at: http://doi.wiley.com/10.1111/j.1600-0706.2012.00121.x [Accessed March 29, 2014].
Quarles, W., 2003. Native Plants and Integrated Roadside Vegetaion Management. IPM Practitioner, XXV(3).
Rahman, M.M. & Govindarajulu, Z., 1997. A modification of the test of Shapiro and Wilk for normality. Journal of Applied Statistics, 24(2), pp.219–236. Available at: http://dx.doi.org/10.1080/02664769723828.
Raven, P.H., Evert, R.F. & Eichhorn, S.E., 2013. Biology of Plants, W.H. Freeman Publishers. Available at: http://books.google.de/books?id=6MjduQAACAAJ.
Rothrock, P.E., Squiers, E.R. & Sheeley, S., 1993. Heterogeneity and Size of a Persistent Seedbank of Ambrosia artemisiifolia L. and Setaria faberi Herrm. Bulletin of the Torrey Botanical Club, 120(4), pp.417–422. Available at: http://www.jstor.org/stable/2996745.
Saunders, D.A. & Hobbs, R.J., 1991. Roads, Roadsides and Wildlife Conservation: A Review. In NATURE CONSERVATION 2: THE ROLE OF CORRIDORS. Chipping Norton, N.S.W.: Surrey Beatty & Sons, pp. 99–117.
Secretariat of the Convention on Biological Diversity, 2010. Global Biodiversity Outlook 3, Montreal. Available at: www.cbd.int/GBO3.
Smith, C.C. & Fretwell, S.D., 1974. The Optimal Balance Between Size and Number of Offspring. AMERICAN NATURALIST, 108(962), pp.499–506.
Snaydon, R.W., 1991. Replacement or Additive Designs for Competition Studies? The Journal of Applied Ecology, 28(3), p.930. Available at: <Go to ISI>://A1991HA12300013\nhttp://www.jstor.org/stable/2404218?origin=crossref.
Stubbs, W.J. & Bastow Wilson, J., 2004. Evidence for limiting similarity in a sand dune community. Journal of Ecology, 92(4), pp.557–567. Available at: http://doi.wiley.com/10.1111/j.0022-0477.2004.00898.x.
Swanton, C.J. & Weise, S.F., 1991. Integrated Weed Management: The Rationale and Approach. Weed Technology, 5(3), pp.657–663. Available at: http://www.jstor.org/stable/3987055\nhttp://www.jstor.org.ezproxy.sussex.ac.uk/stable/pdfplus/3987055.pdf?acceptTC=true.
Temperton, V.M. et al., 2013. Assembly Rules and Restoration Ecology: Bridging the Gap Between Theory and Practice, Island Press.
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Thompson, K. et al., 2010. Little evidence for limiting similarity in a long-term study of a roadside plant community. Journal of Ecology, 98(2), pp.480–487. Available at: http://doi.wiley.com/10.1111/j.1365-2745.2009.01610.x [Accessed March 25, 2014].
United Nations Environmental Programme, 2005. Millenium Ecosystem Assesment, Washington DC.
Violle, C. et al., 2007. Let the concept of trait be functional! Oikos, 116(5), pp.882–892. Available at: http://dx.doi.org/10.1111/j.0030-1299.2007.15559.x.
Violle, C. et al., 2011. Phylogenetic limiting similarity and competitive exclusion. Ecology letters, 14(8), pp.782–7. Available at: http://www.ncbi.nlm.nih.gov/pubmed/21672121 [Accessed March 19, 2014].
Walker, L.R., Walker, J. & Hobbs, R.J., 2007. Linking Restoration and Ecological Succession, Springer. Available at: http://books.google.de/books?id=RYZUe7I7BZcC.
Weigelt, A. & Jolliffe, P., 2003. Indices of plant competition. Journal of Ecology, 91(5), pp.707–720. Available at: http://doi.wiley.com/10.1046/j.1365-2745.2003.00805.x.
Weiher, E., Clarke, G. & Keddy, P., 1998. Community assembly rules, morphological dispersion, and the coexistence of plant species. Oikos, 81, pp.309–322. Available at: http://www.jstor.org/stable/3547051 [Accessed November 1, 2014].
Weiher, E. & Keddy, P., 1995. Assembly rules, null models, and trait dispersion: new questions from old patterns. Oikos, 74(1), pp.159–164. Available at: http://www.jstor.org/stable/3545686 [Accessed November 7, 2014].
Wilson, J., 2007. Trait-divergence assembly rules have been demonstrated: Limiting similarity lives! A reply to Grime. Journal of Vegetation Science, (1974), pp.451–452. Available at: http://onlinelibrary.wiley.com/doi/10.1111/j.1654-1103.2007.tb02557.x/abstract [Accessed April 21, 2014].
Wilson, J.B. & Stubbs, W.J., 2012. Evidence for assembly rules: limiting similarity within a saltmarsh. Journal of Ecology, 100(1), pp.210–221. Available at: http://doi.wiley.com/10.1111/j.1365-2745.2011.01891.x [Accessed March 27, 2014].
World Wildlife Fund, 2014. Living Planet Report 2014 R. McLellan, ed.,
Young, T., Chase, J. & Huddleston, R., 2001. Community succession and assembly comparing, contrasting and combining paradigms in the context of ecological restoration. Ecological restoration, pp.5–18. Available at: http://er.uwpress.org/content/19/1/5.full.pdf [Accessed November 7, 2014].
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Appendices
1. Seed Collection Locations
SPECIES LOCATION SOLIDAGO GIGANTEA
River Isar at Freising (coordinates: 48.400951, 11.757429) -> 7 whole ramets were collected in 7 patches separated by at least
100m (February 2014).
SOLIDAGO CANADENSIS North of Freising (48.412119, 11.739362) & Marienhof
(48.291204, 11.689709) -> 6-7 ramets were collected from around 4-5 patches from each site (October 2013).
CONYZA CANADENSIS
Seed were acquired from Herbiseed (from Serbia) -> I don't have specific info on this species because we bought the seeds.
ERIGERON ANNUUS
TUM Weihenstephan (48.399546, 11.719602) & Marienhof (48.291204, 11.689709) -> more than 50 flowers collected at
each location (October 2013).
IMPATIENS GLANDULIFERA Isar close to Neufahrn (48.312114, 11.700218), Freising - Votting (48.395274, 11.713096), Talham (48.478938, 11.774057) ->seeds
collected from 3-4 fruits per plant from around 5 plants per patch, moved 50m and collected the same amount and so on
(October 2013).
H. MANTEGAZZIANUM
A place close to Hohenkammer (48.403167, 11.536919). All the available seeds collected from 5 individuals in only one patch
(October 2013).
AMBROSIA ARTEMISIIFOLIA
Seeds were collected in a mud-deposit location at 'Neue Donau' in Austria (48°16'38.47"N 16°22'2.55"E). Date of harvest:
12.10.2012
FALLOPIA JAPONICA
Seeds were collected in Freising from only one patch and several ramets (48°23'52.2"N 11°44'54.1"E). Data harvest: October 2013
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2. Exact Seed Mixtures per Replicate
Large seeded Small seeded
R1 R1
Bromus erectus Poa compressa
Helictotrichon pubescens Agrostis capillaris
Centaurea cyanus Achillea millefolium
Salvia pratensis Origanum vulgare
Sanguisorba minor Papaver rhoeas
R2 R2
Bromus erectus Poa compressa
Helictotrichon pubescens Agrostis capillaris
Centaurea cyanus Papaver rhoeas
Knautia arvensis Plantago media
Salvia pratensis Achillea millefolium
R3 R3
Bromus erectus Poa compressa
Helictotrichon pubescens Agrostis capillaris
Salvia pratensis Achillea millefolium
Sanguisorba minor Papaver rhoeas
Knautia arvensis Plantago media
R4 R4
Bromus erectus Poa compressa
Helictotrichon pubescens Agrostis capillaris
Sanguisorba minor Achillea millefolium
Knautia arvensis Papaver rhoeas
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Centaurea scabiosa Plantago media
R5 R5
Bromus erectus Poa compressa
Helictotrichon pubescens Agrostis capillaris
Salvia pratensis Papaver rhoeas
Centaurea scabiosa Plantago media
Sanguisorba minor Origanum vulgare
3. The exact timing of the various stages of the experimental set-up.