PLANT DIVERSITY: A STUDY OF ECOLOGICAL DISTURBANCE

Diversity and Richness: a study of ecological disturbance along a trail

Melisa Robinson
Department of Ecology, Oklahoma State University,
Stillwater, OK 74078

1
melisa.robinson@okstate.edu

Melisa Robinson
Abstract
Environmental disturbance is a cause of great concern for ecologists. Discovering what
effect disturbances will have on an ecosystem has been the focus of a multitude of ecological
studies over the years. This experiment puts the intermediate disturbance hypothesis to the test.
By laying out belt transects along a trail in a nature preserve, this experiments tries to prove that
species diversity is highest at an intermediately disturbed quadrat. Our results did not support
that. There was not a statistically significant link between species diversity and intermediately
disturbed quadrats. However, species richness was found to be statistically significant. If we
were to use species richness to calculate the state of the ecosystem instead of species diversity,
then the intermediate disturbance hypothesis is still statistically supported. As it stands, our
results are likely due to a number of unknown variables and a relatively small sample size.
Keywords: ANOVA, intermediate disturbance hypothesis, Shannon-Weiner Diversity index
(H), species diversity, species dominance, species evenness, species richness, trail disturbance

Melisa Robinson
Introduction
Nature is constantly undergoing a series of conflicts. The ecosystem is in a constant fight
for resources, for space, sunlight, and nutrients. If one species falters, another can rise to take its
place in an ecosystem. In this conflict, ecosystems can find an unstable form of equilibrium.
Disturbances within an ecosystem are an important part preventing a few dominant species from
tipping that equilibrium perpetually in their favor. When disturbances occur naturally, such as a
fire or flood, species diversity can actually increase as the dominant competitive species are
destroyed (Ikeda 2003), increasing the availability of resources for other less competitive plants.
Natural disturbances are a part of nature, but there is an ever growing concern among
ecologists about the effects of anthropogenic disturbances (Pickering & Growcock 2009). When
humans explore nature, we leave an imprint. These disturbances are not naturally occurring, but
instead are the results of human interference on the natural world. For instance, the increase in
recreational use of natural parks has pushed scientists to try to find a healthy balance between
preserving and protecting the natural ecosystem while still encouraging recreational use
(Pickering & Growcock 2009). There have been numerous studies carried out that have tried to
discover at what point anthropogenic disturbances can permanently alter an ecosystem (Connell
1978, Pickering & Growcock 2009, Kondoh 2001, Ikeda 2003).
A certain level of disturbance can increase diversity. When the area is undisturbed or
infrequently disturbed, fewer types of plants grow, as certain plants are more efficient at utilizing
available resources and crowd out competitors. On the other hand, frequent, intense disturbances
reduce or even eliminate the presence of local plants, allowing for a wider variety of other plants
to have access to resources (Connell 1978). This means that the most resources are available for

Melisa Robinson
areas that are only disturbed infrequently. This phenomenon is known as the intermediate
disturbance hypothesis.
Put forth by Connell (1978) this hypothesis supports the idea that species diversity should
be highest when nature finds a balance between stagnation and destruction. Scientists have been
trying to discover at what point this balance occurs. In this experiment, we put the intermediate
disturbance hypothesis to the test. By placing belt transects perpendicular to a trail, we were able
to measure the degree of species diversity at varying levels of disturbance. We hypothesized that
species diversity would highest among the intermediately disturbed quadrats along the trail.
Methods
We performed our experiment on September 23rd at the James K. McPherson Botanical
Reserve in Stillwater, OK along a 100 meter stretch of trail that our instructor had previously
marked with a measuring tape. Our instructor assigned us three points along the trail at various
distances from the trailhead, and established our belt transects at these three points. Our transects
consisted of 10 continuous 10cm x 50cm quadrats that extended out perpendicular from the
center of the trail, as marked by the previously laid tape. The direction that the belt transects
were laid out was chosen at random by flipping a coin. This allowed us to measure the level of
species diversity along varying levels of disturbance as the closer to the middle of the trail, the
higher the disturbance of the habitat.
After laying out the belt transect at the selected point, we then recorded all plant species
with green material that were under a meter tall in each quadrat. We excluded trees and shrubs
from our data unless they were under a meter tall. We also excluded mosses, liverworts and algae
if they were present. We recorded how many different species were present and also how many
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times each species occurred. This was done in all ten quadrats of each belt transect. Several
teams over the course of the week followed this process to collect all the data used in this
experiment.
To analyze the data, we used ANOVA testing to determine four different metrics of
species diversity. Our instructors entered our classs collective data into the SPSS program which
we were then able to analyze. The metrics that we analyzed were the Shannon-Weiner Diversity
index (H), species richness, species evenness, and species dominance. The Shannon-Weiner
Diversity index is commonly used to measure the relationship between species richness and
evenness, or simply species diversity. The ANOVA test was attempting to determine if there is a
statistically significant relationship between any of the four metrics between quadrats and
therefore whether or not our data supported our hypothesis. For this test, we were able to
formulate our null and alternative hypothesis for each metric we analyzed. Either there is no
significant difference between species diversity, richness, evenness, or dominance between
quadrats - our null hypotheses- or there is a significant difference between species diversity,
richness, evenness, or dominance between quadrats our alternative hypotheses.
Results
Our results did not support our biological hypothesis regarding species diversity.
Although species diversity was higher at an intermediate level of disturbance (Figure 1), the p
value for H was 0.071, which is not statistically significant. We failed to reject our null
hypothesis, there is no significant difference between species diversity between quadrats. We
also failed to reject the null hypothesis for species evenness and species dominance. The p value
for each was 0.195 and 0.648, respectively. Species evenness was higher at an intermediate level

Melisa Robinson
(Figure 3), but was still not statistically significant. The figure for species dominance (Figure 4)
did not indicate any level of species dominance in any of the quadrats, which supports our
statistical data. Single species dominance did not occur in any of our quadrats.
The only result that was statistically significant was species richness. As seen in Figure 2,
species richness is higher at an intermediate level of disturbance. This supports our p value,
which was 0.022. We rejected our null hypothesis and determined that there is a significant
difference between species richness between quadrats. We then ran a Post HOC test for species
richness and determined that, in regards to species richness, terminal quadrats (with major or
minor disturbance) are significantly different than central, intermediately disturbed quadrats
(Table 1).
Discussion
Our results did not support the intermediate disturbance hypothesis. While species
diversity was shown to be higher within the intermediate quadrats, the results were not
statistically significant. The only metric that had statistical significance was species richness,
demonstrating that there was an increased variety of plant species in intermediately disturbed
quadrats, but not that there was an increase in plant diversity in the same quadrats. According to
this data, we must reject our initial hypothesis. There is not a statistical link between
intermediate disturbance and diversity.
Species diversity accounts for both species richness and species evenness, and how they
relate to each other. In this experiment, richness was the only metric considered statistically
significant. There have been several studies that focus on using species richness at the
predominant method of determining ecological productivity (Kondoh 2001) rather than species
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Melisa Robinson
diversity. Species richness simply studies the number of species within a studied area. However,
diversity accounts for not only the number of different species, but also the number of
individuals present within a species. Most argue that species diversity is a more accurate reading
of the state of the disturbed area, because species richness does not record how many of each
species is present which could form a skewed idea of the studied area. But an increase in species
richness does still indicate a higher level of available resources in the intermediately disturbed
area. Resources once hoarded by more dominant species are made available for other individuals
due to the initial destructive disturbance and as we move away from that disturbance, the more
dominant types of species make themselves known. Essentially the same intermediate
disturbance phenomenon holds true for both diversity and richness. If this holds true, then our
experiment still somewhat supports the intermediate disturbance hypothesis; it just uses richness
instead of diversity as the metric to determine that.
There have been many studies before this one have proven that there is a link between
diversity and intermediate disturbance. Therefore, we must examine whether or not there is a
flaw in our data or experiments methods. There are several ways that our data may have been
skewed. Firstly, our data was rather limited in many ways. Unlike other experiments where
scientist deliberately trampled areas at a certain recorded frequency (Kondoh 2001), we had no
real way of knowing other than visual inspection- how or where disturbance occurred along this
trail. We also cannot account for how long ago this area was disturbed. We simply do not know
what the actual level of disturbance along the trail was. For instance, if the disturbance frequency
was wider in some areas, then transects in those areas may never have left an area of frequent
disturbance, skewing our results.

Melisa Robinson
Another variable that we cannot account for is potential productivity levels in the soil. If
certain areas have poor nutritional content, it could affect species diversity. We also have to
consider that our sample size may have been too small to record an accurate average. We only
recorded transects along a single stretch of trail; if that trail existed as an outlier, our results
could be skewed. If we were to return to the same area and increase the number of belt transects
that we recorded, our data would likely follow the trends recorded in numerous experiments
before ours.
In the end, our data was not nearly conclusive enough to challenge the intermediate
disturbance hypothesis. And if we use species richness as a measure to determine ecological
health along a disturbance, our data supports the hypothesis. Scientists still have much to
discover about maintaining proper ecological balance against anthropogenic disturbances, but
hopefully with more research that is slightly more comprehensive than this experiment, we will
find a healthy balance between humanity and nature.
Acknowledgements
I would like to thank the professors of Ecology 1604, Dr. Henry Adams and Dr. Arpad Nyari.
Without them, this paper would never have been written. I would also like to thank my Teaching
Assistant, Alissa Freeman, for remembering to bring the duct tape that got most of the ticks off
of me!

Melisa Robinson
Literature Cited
Cole, J.N. 1995. Experimental trampling of vegetation. I. Relationship between trampling
intensity and vegetation response*. Journal of Applied Ecology 32 : 203-214
Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199(4335) : 130210.
Ikeda, H. 2003. Testing the intermediate disturbance hypothesis on species diversity in
herbaceous plant communities along a human trampling gradient using a 4-year
experiment in an old-field. Ecological Research 18 : 185197
Kondoh, M. 2001. Unifying the relationships of species richness to productivity and disturbance.
Proc Biol Sci 268(1464) : 269271.
Pickering, C.M., Growcock A.J. 2009. Impacts of experimental trampling on tall alpine
herbfields and subalpine grasslands in the Australian Alps. Journal of Environmental
Management 91 : 53254.

Melisa Robinson
Table 1: Post HOC table analysis using Fishers Least Significant Difference (LSD),
determining which quadrats differ from each other.

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Multiple Comparisons
Dependent Variable: richness
LSD
Mean Difference
(I) distance

(J) distance

one

two

-.18421

.16430

.263

-.5068

.1383

three

-.23684

.16430

.150

-.5594

.0857

-.40789

.16430

.013

-.7304

-.0853

five

-.43421

.16430

.008

-.7568

-.1117

six

-.30263

.16430

.066

-.6252

.0199

seven

-.18421

.16430

.263

-.5068

.1383

eight

-.11842

.16430

.471

-.4410

.2041

nine

-.03947

.16430

.810

-.3620

.2831

ten

.09211

.16430

.575

-.2304

.4147

one

.18421

.16430

.263

-.1383

.5068

three

-.05263

.16430

.749

-.3752

.2699

four

-.22368

.16430

.174

-.5462

.0989

five

-.25000

.16430

.129

-.5725

.0725

six

-.11842

.16430

.471

-.4410

.2041

seven

.00000

.16430

1.000

-.3225

.3225

eight

.06579

.16430

.689

-.2568

.3883

nine

.14474

.16430

.379

-.1778

.4673

ten

.27632

.16430

.093

-.0462

.5989

one

.23684

.16430

.150

-.0857

.5594

two

.05263

.16430

.749

-.2699

.3752

four

-.17105

.16430

.298

-.4936

.1515

five

-.19737

.16430

.230

-.5199

.1252

six

-.06579

.16430

.689

-.3883

.2568

seven

.05263

.16430

.749

-.2699

.3752

eight

.11842

.16430

.471

-.2041

.4410

nine

.19737

.16430

.230

-.1252

.5199

.32895

.16430

.046

.0064

.6515

one

.40789

.16430

.013

.0853

.7304

two

.22368

.16430

.174

-.0989

.5462

three

.17105

.16430

.298

-.1515

.4936

five

-.02632

.16430

.873

-.3489

.2962

six

.10526

.16430

.522

-.2173

.4278

seven

.22368

.16430

.174

-.0989

.5462

four

two

three

ten
four

(I-J)

95% Confidence Interval

Std. Error

Sig.

Lower Bound

Upper Bound

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Melisa Robinson

eight

.28947

.16430

.079

-.0331

.6120

.36842

.16430

.025

.0459

.6910

.50000

.16430

.002

.1775

.8225

one

.43421

.16430

.008

.1117

.7568

two

.25000

.16430

.129

-.0725

.5725

three

.19737

.16430

.230

-.1252

.5199

four

.02632

.16430

.873

-.2962

.3489

six

.13158

.16430

.423

-.1910

.4541

seven

.25000

.16430

.129

-.0725

.5725

eight

.31579

.16430

.055

-.0068

.6383

.39474

.16430

.017

.0722

.7173

ten

.52632

.16430

.001

.2038

.8489

one

.30263

.16430

.066

-.0199

.6252

two

.11842

.16430

.471

-.2041

.4410

three

.06579

.16430

.689

-.2568

.3883

four

-.10526

.16430

.522

-.4278

.2173

five

-.13158

.16430

.423

-.4541

.1910

seven

.11842

.16430

.471

-.2041

.4410

eight

.18421

.16430

.263

-.1383

.5068

nine

.26316

.16430

.110

-.0594

.5857

ten

.39474

.16430

.017

.0722

.7173

one

.18421

.16430

.263

-.1383

.5068

two

.00000

.16430

1.000

-.3225

.3225

three

-.05263

.16430

.749

-.3752

.2699

four

-.22368

.16430

.174

-.5462

.0989

five

-.25000

.16430

.129

-.5725

.0725

six

-.11842

.16430

.471

-.4410

.2041

eight

.06579

.16430

.689

-.2568

.3883

nine

.14474

.16430

.379

-.1778

.4673

ten

.27632

.16430

.093

-.0462

.5989

one

.11842

.16430

.471

-.2041

.4410

two

-.06579

.16430

.689

-.3883

.2568

three

-.11842

.16430

.471

-.4410

.2041

four

-.28947

.16430

.079

-.6120

.0331

five

-.31579

.16430

.055

-.6383

.0068

six

-.18421

.16430

.263

-.5068

.1383

seven

-.06579

.16430

.689

-.3883

.2568

.07895

.16430

.631

-.2436

.4015

nine
ten
five

nine

six

seven

eight

nine

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Melisa Robinson

nine

ten

.21053

.16430

.200

-.1120

.5331

one

.03947

.16430

.810

-.2831

.3620

two

-.14474

.16430

.379

-.4673

.1778

three

-.19737

.16430

.230

-.5199

.1252

-.36842

.16430

.025

-.6910

-.0459

five

-.39474

.16430

.017

-.7173

-.0722

six

-.26316

.16430

.110

-.5857

.0594

seven

-.14474

.16430

.379

-.4673

.1778

eight

-.07895

.16430

.631

-.4015

.2436

ten

.13158

.16430

.423

-.1910

.4541

one

-.09211

.16430

.575

-.4147

.2304

two

-.27632

.16430

.093

-.5989

.0462

-.32895

.16430

.046

-.6515

-.0064

-.50000

.16430

.002

-.8225

-.1775

-.52632

.16430

.001

-.8489

-.2038

six

-.39474

.16430

.017

-.7173

-.0722

seven

-.27632

.16430

.093

-.5989

.0462

eight

-.21053

.16430

.200

-.5331

.1120

nine

-.13158

.16430

.423

-.4541

.1910

four

ten

three
four
five

*. The mean difference is significant at the 0.05 level.

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Melisa Robinson
Figure 1: Error plot for Shannon-Weiner Diversity index; analyzing levels of species diversity
between quadrats using ANOVA testing.
Figure 2: Error plot for species richness; analyzing levels of species richness between quadrats
using ANOVA testing.
Figure 3: Error plot for species evenness; analyzing levels of species evenness between quadrats
using ANOVA testing.
Figure 4: Error plot for species dominance; analyzing levels of species dominance between
quadrats using ANOVA testing.

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Melisa Robinson
Figure 1

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Figure 2

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Figure 3

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Figure 4

18