Exploring Relationships Between Plant

Diversity and Insect Diversity in an

Experimental Grassland

March 18, 2014

Author: Rachelle Bisaillon
Independent Research Paper (ENVS*3410)
University of Guelph
Abstract

The world's flora and fauna are disappearing at rates greater than the mass extinction

events whose collapses punctuate the fossil record (Reid, 1997). Understanding biodiversity and

interaction between trophic levels is necessary to devise conservation strategies that can

encompass whole ecosystems and reduce the amount of extinction taking place. It is widely

believed that the diversity of plants will affect the biodiversity of insects and vice versa (Sherber

et al. 2010). We examined this supposition at a moderate spatial scale in a restored grassland

community in Guelph (Ontario), by comparing the biodiversity patterns of invertebrates living in

the community to those of plants. Using both Shannon`s and Simpson`s diversity index to

calculate plant diversity and only Shannon`s to calculate invertebrate diversity. We found neither

measure of plant diversity has any relationship with invertebrate diversity for either of the

indices, however after running the analysis by order for the invertebrate, a positive effect was

shown between the Coleoptera species diversity and the plant diversity using Simpson`s index.

We concluded that plant diversity does not have an effect on invertebrate diversity, but that both

are probably responding to similar environmental factors (Hawkins and Porter, 2002).

Additionally, the Coleoptera diversity positive effects with plant diversity could possibly be due

to influences from both top down (Predatory Hypothesis) and bottom up controls (Zou et al.

2013). These results support observations from other grassland studies demonstrating that

changes in diversity of a single trophic level can cascade to effects of diversity at other adjacent

trophic levels (Pearson and Dyer, 2006).

Introduction
Grasslands have become one of the most imperiled ecosystems worldwide, facing one of

the largest threats from human activities, with the tall grass prairie type, as the most threatened in

North America (Ceballos et al. 2010; Samson et al. 2004). These ecosystems are very valuable

because they contain a high diversity of unique grass species that provide habitat to many

organisms, such as insects and birds, including many species at risk (Ceballos et al. 2010). Yet,

many grasslands continue to be converted to croplands, in order to support the world's growing

population (Ceballos et al. 2010). We have continued to do this repeatedly though history under

the assumption that ecosystems have inherent properties that give it resilience to such

perturbations (McCann, 2000). However, the diversity loss from these grasslands is causing the

disappearance of uncountable numbers of species (Celballos et al. 2010). It is hypothesized that

we are currently experiencing the 6th mass extinction in the earth's history, with an estimated

decline of an additional 20-50% of biodiversity in the next half century alone (Rahbek and

Colwell, 2011). Along biodiversity decline, changes in essential ecosystem processes will occur,

which may adversely affect primary productivity, nutrient cycling, and trophic interactions in

grassland environments (Sherber et al. 2010). Understanding the unique interactions between the

individual traits of a species with that of a changing ecosystem community is complicated and

not evident. Especially, the plant-insect interaction, who's implication is fundamental to

understand many other biological interfaces in the ecosystem, which may help us better evaluate

the current state of grassland environment and adopt management strategies to restore it.

Species diversity has long been used to characterize healthy ecosystems, as well as to

predict how populations will respond to disturbances (Dejong, 1975). Diversity consists of two

distinct components: species richness, and species abundance. Species richness is the number of
the species per unit area or per sample, whereas species abundance (or evenness) is the relative

abundance of species (Dejong, 1975). Species diversity increase when the number of species

increases and as the abundance of species become more even (DeJong, 1975). Different indices

have been developed to express diversity (DeJong, 1975; Newman et al. 2011). The Shannon-

Weiner Index is the most commonly used in ecology because it is easy to calculate, and is

sensitive to site difference (DeJong, 1975). This index measures the probability of an individual

picked at random from an infinite population will be of a particular species (DeJong, 1975). The

Simpson's Index measures concentration of species by yielding a probability (from 0 to 1),

representing the chance that any two individuals sampled randomly from the population will

belong to a different species (DeJong, 1975). This index is not sensitive to species richness, and

is strongly affected by the abundance of a few common species in the sample (DeJong, 1975).

Additionally, species diversity will also depend on choosing a clearly defined taxonomic group

(family, order, class), since classifying the insects by order may result in a different diversity

interpretation than classifying them by a more specific taxa, such as family (Cousins, 1991). The

definition of biodiversity is linked to the spatial scale, since small local responses at the

community level will be different than responses examined at a more global scale (Newman et

al. 2011). The choice of index, taxon or spatial scale used will yield different ways to interpret

diversity, therefore it is important to make the right choice of aspect, and measurement according

to guidelines from previous studies for better comparison.

A decrease in biodiversity in any environment is generally known to be detrimental

regardless of the variation that its definition implies (Newman et al. 2011). Biodiversity is

necessary to increase ecosystem functioning through different mechanisms, such as

complementarity, facilitation and sampling effect (Newman et al. 2011). First, complentarity
states that a more diverse community should be able to use resources more completely; therefore

there is greater provision of services within that environment, leading to increased productivity

(Newman et al. 2011). The facilitation mechanism occurs when one species modifies the

environment in a way that makes it favorable for co-occurring species (Bruno, 2003). Finally, the

sampling effect of diversity is the magnitude of influence each species has on ecosystem

functioning (Newman et al. 2011). With higher biodiversity, there is a greater chance of

sampling a species that is well adapted to a particular set of conditions (Newman et al. 2011;

Wardle, 1999).

Furthermore, the insurance effect, or the biodiversity-stability hypothesis, hypothesizes

that greater biodiversity not only provides greater productivity but might also provide greater

ecosystem stability, because the more species present, the greater the likelihood for the

ecosystem to adapt (Newman et al. 2011; Macdougall et al. 2013). Thus, high plant species

diversity provides a buffering effect to anthropogenic influences by stabilizing ecological

systems though functional complementarity (Newman et al. 2011). This means that if the

stability hypothesis is true, diverse plant ecosystems have increased protection in potential

herbivore outbreaks (Macdougall et al. 2013; Haddad et al. 2011). This was first considered in

1958, when frequent pest outbreaks were observed in monoculture agricultural systems (Haddad

et al. 2011). High local biodiversity also has the capacity to resist invasion, because there are less

available niches in diverse ecosystems, giving the community stability (Knops et al. 1999).

Biodiversity loss, specifically at lower trophic levels, will affect nutrient and autotrophic

production, which will essentially have profound effects on the integrity, and functioning of that

whole ecosystem (Knops et al. 1999). It is hypothesized that change in diversity of a low trophic

level organism like plants should influence the level of diversity of higher members of the food
chain such as insects (Siemann, 1998). Compelling evidence have shown plant diversity having a

strong bottom up effect on biotic interactions, and species situated at higher trophic levels having

indirectly mediated effects by the bottom up trophic cascade (Sherber et al. 2010) . Sherber et

al.'s 2010 study observed that when there is a decline in plant species richness and abundance,

species richness of insects and biotic interactions were also affected. These results strengthen the

bottom up model of plant species richness, where organisms are stimulated by the trophic level

beneath them, and plant diversity cause effects that propagate up the food chain (Haddad et al.

2011; Scherber et al. 2010). Plants and insects largely influence one another, since herbivores are

situated low in the food web, and plant diversity will have a dampened trophic interaction with

increasing distance up the food chain (Haddad et al. 2011; Scherber et al. 2010). Many studies

emphasize the relationship between plant and insect richness and many models agree that a

diverse resource base will naturally support a higher biodiversity of consumers (Knops et al.

1999; Colwel and Rahbek, 2011). Increases in plant diversity will increase the abundance of its

predators, which in turns will increase the energy transferred between the trophic levels. This

increases connectivity within the food web, and increase food web complexity, which then

continues to increase herbivore species richness (Haddad et al. 2011). Higher plant diversity

stabilizes insect communities by providing food and habitat resources, allowing many different

herbivorous insects specialised in specific plant species to persist locally, and preventing other

insects to establish competitive dominance in the community (Haddad et al. 2011; Siemann,

1998). With each additional type of resource introducing new specialized consumers, and with

many insects exhibiting a high degree of host specialization, higher levels of these herbivores

may live in the same area (Colwel and Rahbek, 2011). However, this also means that if a species

of plant is lost, the specialized insect will most likely have to relocate in order to survive, and
diversity of the community will be driven down. This highlights plants as one of the fundamental

organism groups to have the potential to impact many different organisms in an ecosystem

(Sherber et al. 2010).

Top down effects can also be seen if the herbivores are not food limited, though this

interaction does not yield as strong of an interaction as a bottom up effect (Scherber et al. 2010).

The Janzen -Connell hypothesis suggests that highly specialized enemies of plants, such as insect

herbivores, help maintain high diversity of the plant community by elevating mortality of plant

species occurring in high density (Bagchi et al. 2014). This indicates that herbivores have the

potential to control populations of plants just like plants have the capability of controlling insect

populations (Macdonald and Kotanan, 2010).

According to theory, a combination of both environmental change and diversity loss

induced primarily by human influences dramatically increased the risk of abrupt and irreversible

ecosystem collapse (MacDougall et al. 2013). Rapid change in the environment will influence

species diversity and may lead to homogenization of our planet. While, there is also evidence

that it is particular species that are important to ecosystem functioning and stability, and not

biodiversity, we do not know which species are critical, because the relationship between trophic

interaction is still not fully understood (Newman et al. 2011). Species interaction is a subject for

much needed research, and that of plant-insect interaction should be the focus, since it is the

bottom of the food web, relatively easy to measure and can help us understand other biological

relationships within an ecosystem. Many observational studies show positive correlation between

plant and insect diversity (Murdoch et al. 1972; Nagel, 1979; Southwood et al. 1979; Knops et

al. 1999), but this should be tested in a restored grassland ecosystem under field conditions. My

research question will be looking at the relationship patterns existing between plant and insect
diversity in an experimental and restored grassland community. I hypothesize to see a direct

interaction between both plant and insects. We would expect to see if plant diversity is high,

insect diversity will be high as well, and if the experimental plots contain low levels of plant

diversity, it will be natural for that community to support a low level of insect consumers.

Materials and Methods

Study Site

Twenty experimental plots of 4x4m in a grassland ecosystem at the Guelph Turfgrass

Institute in Guelph, Ontario were used for the purpose of the study. Before planting in May 2010,

Roundup (glyphosate) was sprayed twice. Cultivation and leveling was done using a Blecavator.

Plant Species Selection

The species selected were 15 tallgrass prairie species native to the area, and one non-

native grass species, which were carefully selected based on availability for restoration purposes

and the common functional diversity in tallgrass prairies. The seeds were dry stored for seven

weeks at 17C.

Study design

Plots were planted in a randomized complete block design and each had the same initial

seed density. This was a correlation based study, thus no variable were directly manipulated

other than the initial planting of the grass species.

Plant Data Collection

At the end of the growing season in September 2010, the above ground biomass was

clipped at 5cm above the soil surface. The biomass was dried at 60C and weighed.

Invertebrate Data Collection

The invertebrate sampling was completed in June, 2011 at 1pm. This collection was done

using a vortis sampler (inverted leaf blower) at 5 different points in each plot for 60sec each (see

Figure1 and Figure 2). These points were 1m diagonally from each corner of the plot, and

directly in the middle. All the insects collected from the leaf blower from each plot was stored in

a plastic bag and labelled. In order to preserve the specimens, they were stored in a freezer at a

constant temperature of 0 C.

Invertebrates were identified down to the family level for each of the plot using a

microscope. After identification they were stored by order. Results were recorded by family for

each of the plots.

Plant and Insect Biodiversity Index Calculation

Shannons Index of diversity

The Shannons index of diversity was calculated for both the invertebrates and the plants. In

this equation (1), C is a constant, and pi represents a proportion of the number of individuals

belonging to a species (ni) divided by the total number of individuals.

(1) Shannons Index of diversity

Simpsons index of diversity

The Simpsons index of diversity was calculated only for the plant diversity because it is

hypothesized as being a better index of biodiversity for plants (Wilsey et al. 2005). Simpson`s

index of diversity is not a good indicator for insect diversity because this index is strongly

affected by the abundance of a few common species in the sample, and our invertebrate samples

had many common species (DeJong, 1975). In this equation (2), S represents the number of

species, ni represents the number of individuals in the sample belonging to the ith species, and N

represents the total number of individuals in the sample.

(2) Simpsons index of diversity

Statistical Analysis

The analysis was comprised of a two step process. A visual inspection of the histograms,

box plots and normal Q-Q plots yielded results that were approximately normally distributed

(p>0.05), thus they do not differ significantly from normality for the Shannons invertebrate

index data, Simpsons plant diversity index data, and for the Shannons plant diversity index

data. Since the data is normally distributed, a linear regression test was used to test for

significance. The analysis was again performed between orders. With the assumption that all the

data is normally distributed, a linear regression was used to test for significance.
Results

Across all invertebrate species, neither measure of plant biodiversity (Simpson and

Shannon) was a significant predictor of relationship between insects and plants diversity

(p>0.05) (see Figure 3 and Figure 4). Performing transformation to the data did not improve the

significances between the two groups either.

The analysis was again performed after calculating the diversity between orders.

Although most did not yield any significant relationship with both plant diversity indexes

(p>0.05), the diversity of the Coleoptera order was significant with the Simpson index of plant

diversity (p= 0.0062) (see Figure 5), however was not significant with the Shannon index of

plant diversity (p>0.05).

Discussion

These results lead us to conclude that total plant diversity has almost no effect on the

diversity of invertebrates (Hawkins and Porter, 2002). Our results are consistent with previous

studies reporting that insect biodiversity are often limited by factors other than the distribution of

their food plants (Hawkins and Porter, 2002). Large scale associations between invertebrates and

plant diversity responding to the same driving variables have been previously examined

(Hawkins and Porter, 2002). Dixon et al.`s 1987 study observed the ratio of aphid biodiversity to

plant biodiversity across 22 countries decreased with increasing plant diversity, indicating that

plant and animal diversity are not necessarily positively associated at large spatial extents. The

main conclusion is that resource diversity has no correlation on the contribution it provides to

consumer diversity (Hawkins and Porter, 2002). They are mainly driven by the underlying
changes in environmental factors (Zou et al. 2013). Additionally, specifically herbivore diversity

shows no evidence to be constrained by plant diversity.

Interestingly, Coleoptera had a positive correlation with plant diversity (only with

Simpsons index). Niemela and Halmes study in 1992 determine that one of the most important

factors affecting diversity of carabid (coleoptera) is vegetation. A positive relationship between

plant diversity and the diversity of predatory invertebrates is hypothesized by the Enemy

Hypothesis, a top down control effect where members situated high in the food chain have an

impact on species in the lower levels of the food chain (Zou et al. 2013). Coleopteras are also

known to be affected through bottom up effects and can be indirectly mediated through a bottom

up trophic cascade (Zou et al. 2013). An increase in plant diversity will lead to increased

amounts of leaf litter, enhancing net primary productivity, and providing greater quantities of

food to the Coleoptera (Magura et al. 2005). Greater availability of resources can lead to an

overall increase in the biodiversity of Coleoptera consumers (Zou et al. 2013). Highly diverse

plant communities can provide more refuges, and more stable prey availability, for the

Coleoptera (Zou et al. 2013). This results in the Coleopteras catching their prey more effectively,

hence higher biodiversity of predatory species (Zou et al. 2013). This hypothesis has been

supported by a large number of experimental studies (Scherber et al. 2010; Wenninger and

Inouye, 2008; Haddad et al. 2011; Moreira et al.2012; Jonsson et al. 2009; Varchola and Dunn,

1999).

Coleopteras are known for being both herbivorous and carnivorous (Niemela et al. 1992).

Increases in plant diversity, along with an assumed increase of herbivore species will provide an

increase in food resources which will initially increase the carnivorous Coleoptera population.

However, competing predatory spiders and ants can increase the overall competition for prey in
the long run, and drive down the overall diversity of carnivorous Coleoptera (Zou et al. 2013).

Our experiment was based in a restored grassland community, a very young community that

might not have had sufficient time for predatory species to fill all the available niches (Zou et al.

2013). Therefore, if this experiment would run for multiple years following the restoration, we

will most likely see a change in biodiversity levels of Coleoptera. Beetles are also believed to

generally benefit from high plant diversity through improved ecosystem functioning (Zou et al.

2013). High plant diversity can enhance shade and soil moisture, which can create favourable

microhabitats for Coleopteras and their larvae (Niemela et al. 1992).

Environmental stresses are speculated to produced less change in the biodiversity of the

Coleoptera community (Bulan and Barrett, 1971). In a study by Bulan and Barrett in 1971,they

observed that herbivorous Coleoptera species diversity along with other herbivorous insects was

significantly reduced in a plot subject to environmental stress such as mowing, however the

carnivorous Coleoptera population was not measurably affected. This indicates that secondary

consumer Coleopteras (carnivores) could be a more disturbance-tolerant group (Bulan and

Barrett, 1971). Since our samples comprised both herbivore and carnivore Coleopteras, and we

observed their increase in diversity in relationship with the increase in diversity of plants, this

effect could be due to using the carnivorous Coleopteras in our analysis. Vegetation may have a

more significant effect on the carnivorous Coleoptera's biodiversity than environmental factors

(Bulan and Barrett, 1971). Additionally, situations in the trophic level, can be a factor in the

resilience to such environmental perturbation in invertebrates (Bulan and Barrett, 1971).

Determination of the Coleoptera's food preference is not as simple as it seems, because many

would require us to identify down to the species level and some Coleopteras may be omnivorous.

However, we could follow the guidelines and scheme by Borror and Delong (1964) to try to
separate the Coleoptera into two distinct categories (herbivore and carnivore) and rerun the

analysis to see if the herbivorous Coleoptera diversity and carnivorous Coleoptera diversity

would still show the same significance level with plant diversity.

There are limitations in the experimental design that may lead to the lack of evidence

presented in this experiment. The most apparent one was that this was a short run experiment, in

order to get meaningful results it is best to have long-running experiments. Without having

multiple years in the experiment, there is higher chance that observations are due to climatic

stressor.

Many studies show that invertebrates are good indicators of ecosystem integrity and not

necessarily biodiversity of the resource base (Carignan and Villard, 2001). Invertebrates easily

respond to stressors, and have a varied lifespan that is useful when looking into chronic and

acute disturbances (Adamus, 1995).Their presence is more associated with environmental factors

than biological ones; such as competition, predation and parasitism (Carignan and Villard, 2001).

Both plants and invertebrates mainly react to disturbances at fine spatial scales and not larger

scale disturbances (Carignan and Villard, 2001).

All in all, I will have to reject my hypothesis that plant diversity will have a positive

effect on total insect diversity because only one order showed significance with one index of

diversity. This does not mean that plants do not matter to the insects because they clearly do.

Since, removing all plants from the area would cause the insect population in that area to

disappear. This means that the simple idea that there are so many species of animals because

there are so many species of plants may not be so simple after all (Hawkins and Porter, 2002).
 Figures and Tables

Figure 1: place in the plots were the invertebrates

Figure 2: Vortis sampler used in this experiment
were collected (1 meter diagonally from each corner
and in the middle)

2.6 2.6
2.4 2.4
2.2 2.2
Insect Diversity

Insect Diversity

2 2
1.8 1.8
1.6 1.6
1.4 1.4
1.2 1.2
1 1
2 4 6 8 10 1 1.2 1.4 1.6 1.8 2 2.2
Simpson's Plant Diversity Index Shannon's Plant Diversity Index

Figure 3: Insect diversity (Shannon`s Index) vs Plant Figure 4: Insect diversity (Shannon`s Index) vs Plant
diversity (Simpson`s Index), showing no significant diversity (Shannon`s Index), showing no significant
relationship (p>0.05). relationship (p>0.05).
 1.8
1.6
1.4
Coleoptera Diversity
1.2
1
0.8
0.6
0.4
0.2
0
0 1 2 3 4 5 6 7 8 9
Simpson's Plant Diversity Index

Figure 5: Coleoptera diversity (Shannon`s Index) vs Plant diversity (Simpson`s Index), showing significant relationship
(p= 0.0062).
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