Wesley Collins

Plant Ecology
Causes of Spatial Diversity in Terrestrial Plants

Introduction:
Plant communities across the world are primarily defined by climate. There are many
factors that determine what types of plants will grow where, but nothing trumps climate.
Climatic variability/overall climate, habitat heterogeneity, history, energy, competition,
predation, and disturbance are all factors that are hypothesized to influence species richness
(Currie 1991). With advancements in ecological discoveries and other related sciences, there
have been major strides in learning more about what causes plants to grow where they do and
how diversity is maintained, however there is still a poor understanding overall in the
mechanisms that affect species diversity. Ecosystems and Biomes are often characterized by
what plants grow there and the typical climate; but within these broad areas, there is often
extreme spatial diversity among the plants that are present there. Learning how these systems
maintain diversity and discovering why there is such differential diversity in different systems is
important in making sound management decisions in a world that is becoming increasingly
developed.
In early studies of species richness, the rainforest was often referred to as the most
diverse and therefore the most productive. In one early study, the rainforest was considered to be
the most diverse because it provided the most diverse types of habitat (Dobzhansky 1950). This
type of assessment has failed to hold water over time, however. Niche-theory and competition
alone cannot explain diversity because they are often confounded by natural contradictions such
as some deserts or areas like the Texas Hill Country with high diversity and/or endemism (Davis
et al. 1994, Bowles and Thomas 1993). In another study from the 1960s, the author pointed to
primary productivity driving diversity in small areas, but among large areas, diversity was higher

Wesley Collins

Plant Ecology

in the tropics. The reason for higher diversity in the tropics was explained by finer subdivisions
in habitat rather than increased diversity within habitats (MacArthur 1965). While it makes sense
that productivity can drive endemism or diversity in a small area (i.e. an oasis in the desert), it
does not make sense that habitat is further delineated in the tropics, just because. There are
plenty of big trees and lots of productivity in the Pacific North West, however the diversity of
flora and fauna in the area is dwarfed by what can be found in the tropical rainforests. Why have
the species in the Pacific North west not divided their habitat into finer subdivisions?
Washington State does not have a year round growing season like Costa Rica, but that does not
explain why there are such differences in diversity.
It has been hypothesized that energy, or available energy in a given area may account for
some diversity present in an area. In the case of one energy-diversity study, AET and PET
(actual and potential evapotranspiration) were used to measure the energy in an area. Because
evapotranspiration is directly related to moisture availability and temperature which relates
directly to plant production, it could be suitable to use as a quantified form of available energy in
scientific study. The results of this study showed that across North America, there is often
replacement of one species for another across environmental gradients, and while available
energy limits the upper end of species richness, an abundance of energy in an area does not
necessarily mean there will be higher diversity (Curie 1991). Despite some evidence that seems
to show a relation to evapotranspiration and species richness, other studies have presented
contrary evidence. In a study that sought to explore the energy-diversity hypothesis further, it
was discovered that the number of tree species in 26 different large areas of moist forests show
continental differences unrelated to any geographical patterns in evapotranspiration (Latham and
Ricklefs 1993). Though energy has some effects on diversity, research has shown that by and

Wesley Collins

Plant Ecology

large this is largely localized and, diversity of a large area cannot be attributed to available
energy alone.
The most convincing hypothesis on the maintenance of species diversity may be habitat
heterogeneity. Diversity of animals, especially birds, has been shown to be positively correlated
with habitat heterogeneity (Freemark and Merriam 1986). However, studies focusing on habitat
heterogeneity often ignore diversity of plants. Research has shown that tropical tree diversity
often does not seem to follow any recognizable pattern across soil gradients as would be
expected (Wong and Whitmore 1970). Animal diversity may be attributed to habitat
heterogeneity due to a higher abundance of potential niches to be filled, but it is not a valid
explanation for plant diversity.
Consideration should also be given to the scale at which a factor may act upon a system;
regional or local. Many ecological inquiries have assumed that local diversity was the outcome
of local processes within a small homogeneous community (Rickfels 1987). When diversity is
being considered in an area, understanding at what level (local or regional) a process acts on a
community is extremely important to understanding causes of variation in diversity across an
area. Many studies in diversity have failed to consider how the relative effects of local and
regional processes affect diversity. Until these types of questions are answered, attempts at
solving mysteries about the causes and maintenance of species diversity may difficult (Huston
1999).
While there is much contradictory evidence as to how species diversity is maintained, and
why there is such differential diversity across the globe; it is recognized that these processes are
controlled by many factors. Rather than one single factor driving and maintaining diversity, all of
the factors mentioned in the first paragraph from Curries study (and more?) likely act in a

Wesley Collins

Plant Ecology

synergistic fashion to affect diversity (Kreft and Jetz 2007). The weight of each factor has some
variation spatially. Because of this, there is differential resource availability and habitat
heterogeneity along with various gradients (sunlight, rainfall) across the globe, which has caused
differential diversity maintained by local and/or regional factors such as disturbance frequency or
predation/herbivory.
Methods:
In order to study variation in plant species across topographic gradients in a landscape, a
site at near the Supple Science Building at the Texas State University campus was selected. The
site was chosen because it is a storm water retention basin which has an elevation gradient that
could affect plant growth. There are many anthropogenic influences upon the site that could
affect what plants will grow and establish there, including adjacent parking lots (impervious
cover), grass seed blankets, and occasional mowing in some parts. The 2014 Plant Ecology class
at Texas State used standard sampling methods to determine many characteristics of the species
present at the retention basin. Sampling was done using parallel transects 6 m apart, and plots
along the transects were spaced 4 m apart. Quadrats of 0.25 m^2 were used at the plots in
sampling. Percent cover of each species present in the quadrat was estimated visually as the
percentage of the quadrat that is covered by each species. Bare ground percentage recorded along
with the measurements of the maximum heights of each species present.
Once the data was collected, it was entered onto an excel spreadsheet by quadrat and
transferred to JMP for statistical analysis. Additionally, data from the same data for the same site
from the 2013 plant ecology class was used in order to compare data from this year to last year.

Wesley Collins

Plant Ecology

Results:
Upon visiting the research site, there were several obvious gradients that could influence
the plants growing there. One of the most obvious changes in the setting was the elevation. There
was a steep slope going from around 500 cm. in elevation at the top of the basin, to 0 cm. at the
bottom. This change could have profound effects on water availability at various times
throughout the year. Several hypothesis could be formed based on the elevation gradient. It
would be expected that due to increased water availability at a lower elevation, a higher total
percentage of plant cover would be seen at the bottom of the basin than at the top (see fig. 1).
There appears to be a negative relationship between elevation and the total percent of plant
cover, however the correlation is very weak.
Fig. 1 Bivariate Fit of TotPlantCov By Elevation (cm)

Another interesting question posed during this study was how elevation may affect
species richness in this anthropogenic setting. The expectation was that species richness would
decrease at the bottom of the basin, however when regression analysis was run on a scatterplot of

Wesley Collins

Plant Ecology

elevation vs. number of species, there was a bi-modal distribution of species richness with one
mode at the bottom of the basin and the other at the top (fig. 2).
Fig. 2 Bivariate Fit of 2014NumSp By Elevation (cm)

In order to search for more correlations between elevation and the plant community,
weighted max height of plants in each quadrat were graphed against elevation. The expected
outcome of this analysis was that there would be taller plants at the bottom of the basin than at
the top. While the results from the 2014 plant ecology seemed to support that hypothesis (fig. 3),
the data from the 2013 collection did not (fig. 4).
Fig. 3

Fig. 4

Wesley Collins

Plant Ecology

In addition to analysis on the entire basin, several comparisons were made between
environmental gradients and single species. Cynodon dactylon, Bermuda grass, was a common
grass at the basin, so it was used in analysis to compare changes over the course of two years
worth of data. A bubble map for each year (2013-fig.5, 2014-fig. 6) was constructed using the
plot coordinates as the x and y axis, and the size of the bubbles reflected the percent cover of
Cynodon. Over the course of the two years, there were differences in the distribution of Bermuda
grass, with the highest percent cover moving east in the plot over time.
Fig. 5 Location vs. Percent Cover of Cynodon 2013

Fig. 6 Location vs. Percent Cover Cynodon 2014

Wesley Collins

Plant Ecology

Another common grass at the basin, Bromus carthaticus, was examined in relation to
elevation over the course of the two year study. Percent cover of bromus seemed to increase
substantially at the bottom of the basin over the course of the year (fig 7-8).
Fig 7-8

Wesley Collins

Plant Ecology

Discussion:
While there were several noticeable patterns in the data, there was not major differences
across the gradients. Although the results may not be statistically significant, they are important
in that they show true environmental gradients on a small scale and the responses of plants as a
result. This type of study can help glean knowledge on the workings of true natural systems,
because experimental settings such as the drainage basin offer an easy site to monitor over the
course of a number of years. Additionally, experimental modification can be performed in order
to compare responses do various inputs (fertilization, mowing, differed mowing, etc.).
The focus on this experiment was the elevation gradient, and its effects on the number,
type, and density on plant species. It was hypothesized that the total percent cover of plants
would be higher at the bottom of the basin than the top, however the data did not seem to support
this. Overall, percent cover was relatively uniform across the basin, with only a slight negative
regression line. The primary reason that it was hypothesized that percent cover would be higher
at the bottom was because there would presumably be more water available at the low elevation
of the basin. Water tends to carry nutrients and minerals in sediment that it washes, as well as
pollutants, so water in addition to nutrients, minerals, and pollutants would in theory have a
greater impact (good or bad) on the plants at the bottom of the basin than the top. The data does
not support this to the point that it can be said it is significant, but the subtle pattern is there
regardless.
A pattern from the results that seemed to refute one of the null hypothesis of the
experiment was the bi-modal distribution of species richness. For several reasons, it was
hypothesized that there would be higher species richness at the top, rather than the bottom. The
primary reason that this was hypothesized was again due to water. In many natural ecosystems

Wesley Collins

Plant Ecology

that are highly productive, natural monocultures form (Huston, plant ecology lecture). This
occurs because in areas of high resources, there is often one or a few species that will be more
efficient than all other species, so they will have the advantage and choke the others out. In
settings where there are less resources or more frequent disturbance, no single species is
operating optimally, and so competition prevents one species from taking over.
In order to compare data over two years, the weighted max height of plants in each
quadrat was graphed against elevation for both years of data. It was hypothesized that the tallest
plants would be at the bottom, for the same reasons that the last two hypothesis were formed:
water and the particulates carried in runoff. The data from this year seemed to wholly support
that hypothesis, however the data from last year did not. While it is hard to ignore the regression
line on the graph from this year (fig 3), the graph from the 2013 data (fig 4) contradicts the 2014
data. Obviously, there are many outside influences that could be operating on a more regional
scale that would affect these changes (winter/spring rainfall, seed production last year due to
environmental conditions, days since last mowing could be different over the years). When
examining data that shows temporal changes such as these, it is important to consider that local
facors-such as location in the basin, and regional factors-such as seasonal rainfall, have shared
control over the diversity in a location.
In addition to whole-basin analysis, several individual species were selected for analysis
in order to compare changes over time in distribution. The first individual selected was Bermuda
grass. The percent cover of Bermuda grass seemed to change spatially over the course of the
year, with more plots reporting higher percent covers of the grass in 2014 compared to 2013.
There could be several reasons for this, however because there were sod blankets placed at the
top of the basin after some construction, I would hypothesize that Bermuda grass seed washed

Wesley Collins

Plant Ecology

from the blankets, down into the basin. Though this is a completely anthropogenic situation, it
has value ecologically because it shows a natural phenomenon occurring which changes the
distribution of plants. Using these types of situations as models allows ecologists to hypothesize
on natural occurrences, predict changes in the spatial arrangement in plants before they happen,
and design management strategies if necessary.
The final pattern examined in an individual species was done using percent cover data for
the grass Bromus. Data for percent cover of Bromus was graphed against elevation for both
years. There was a dramatic increase in percent cover of Bromus in the bottom of the basin, but
not much change in the top. This type of change once again shows how there can be large
changes in species composition in relatively short periods of time in herbaceous plants. There
may have been a seed source of Bromus in the seed blankets, or there could have been a timely
rainfall even that promoted seed germination of bromus early this year. The potential answers to
questions regarding changes such as these are many.
Conclusion:
While there are many questions that this type of experiment fails to account for in terms
of what is controlling and maintaining diversity in this anthropogenic setting, there are definitely
some useful correlations that have ties to broad concepts in ecology. Changes and patterns across
the topographical gradient are likely due to multiple gradients created by differences in elevation.
These include solar radiation hitting the leaves and water availability, but it is likely there are
other gradients such as soil moisture and nutrient availability across the site. This small-scale
approach shows how major shifts can occur from year to year in plant communities and
consideration must be given to both local and regional factors in order to disseminate temporal

Wesley Collins

Plant Ecology

flux in diversity so that conservation questions can be answered and management decisions
enacted.
Literature Cited:
Bowles, David E., and Thomas L. Arsuffi. "Karst aquatic ecosystems of the Edwards Plateau
region of central Texas, USA: a consideration of their importance, threats to their
existence, and efforts for their conservation." Aquatic Conservation: Marine and
Freshwater Ecosystems 3.4 (1993): 317-329.
Currie, D. J. (1991). Energy and large-scale patterns of animal-and plant-species richness.
American Naturalist, 27-49.
Davis, S. D., Heywood, V. H., & Hamilton, A. C. (1994). Centres of plant diversity. NATURAL
HISTORY, 111(1), 01-1.
Dobzhansky, T. (1950). Evolution in the tropics. American Scientist, 38(2), 209-21.
Freemark, K. E., & Merriam, H. G. (1986). Importance of area and habitat heterogeneity to bird
assemblages in temperate forest fragments. Biological Conservation, 36(2), 115-141.
Huston, M. (1979). A general hypothesis of species diversity. American naturalist, 81-101.
Huston, M. A. (1999). Local processes and regional patterns: appropriate scales for
understanding variation in the diversity of plants and animals. Oikos, 393-401.
Kreft, H., & Jetz, W. (2007). Global patterns and determinants of vascular plant diversity.
Proceedings of the National Academy of Sciences, 104(14), 5925-5930.

Wesley Collins

Plant Ecology

Latham, R. E., & Ricklefs, R. E. (1993). Global patterns of tree species richness in moist forests:
energy-diversity theory does not account for variation in species richness. OIKOSCOPENHAGEN-, 67, 325-325.
MacArthur, R. H. (1965). Patterns of species diversity. Biological Reviews, 40(4), 510-533.
Ricklefs, R. E. (1987). Community Diversity: Relative Roles of Local and Regional Processes.
Wong, Y. K., & Whitmore, T. C. (1970). On the influence of soil properties on species
distribution in a Malayan lowland dipterocarp rain forest. Malayan Forester, 33(1), 4254.