Leaf Size and Its Relation To The Environment of Terrestrial Plants

Wesley Collins
Leaf Size Distribution Project
Plant Ecology
Leaf Size and its Relation to the Environment of Terrestrial Plants

Introduction:
When the health of a plant is considered, the first things that are looked at are
often the leaves. During times of drought or mineral deficiency, leaves can indicate the
problem. Leaves are also extremely diverse and much of the diversity is dependent on the
environment that the leaves are found in. In plant ecology, plants from mesic and xeric
environments are often compared to illustrate how different the plant physiologies can be
from one another. Because of global changes in climate, comparing leaf size on a local
and global level may be able to demonstrate patterns that are occurring across large areas
of vegetation. Leaf size and morphology can differ greatly within a small area and within
the same species just based on water availability. So, by looking at leaf size and
morphology over time in an area, the effects of drought upon forest structure and
distribution can be analyzed with quantifiable data (Damesin et al 26).
Methods:
Here in central Texas, the habitat is extremely diverse and plants from both mesic
and xeric ecosystems are often found nearby each other. The karst topography and unique
artesian hydrology make central Texas a great place to study the effects of drought.
Because the ground water in the Edwards Plateau region is so dependent on being
replenished by rain and the roots of trees often reach down into the aquifers, changes in
water availability for trees can be seen easily (Jackson et al 11,387). In this experiment,
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Plant Ecology
leaves were compared from two very different sites. The first site was adjacent to Spring
Lake, in San Marcos, Texas. Spring lake is a small lake that is impounded by a man made
dam. The lake is fed by several massive artesian springs coming out of the Edwards
Aquifer, which is recharged by runoff from the surrounding hill country to the west. The
springs come out of the base of the Balcones escarpment, and create a very unique
ecosystem with several species endemic to the springs and river. The first area where
leaves were collected, located at the bottom of the escarpment (BES), is riparian with
high water availability. Some of the species present here included Populus deltoids
(Eastern Cottonwood), Platanus occidentalis (American sycamore), many species of vine
(Virginia creeper and mustang grapes), and green briar. Because the lowland area around
the lake accepts runoff from upland areas to the west, the soil is likely nutrient rich
allowing some of the trees to reach exceptional heights with unlimited water availability.
The canopy amongst these lowland trees reached 50 to 60 feet in many of the trees in
sight.
The second site was chosen up the hill and is a much drier area. This site is located
in the Sink Creek Greenbelt in San Marcos. Sink Creek is a large creek bed that is usually
dry, but during massive rain events west of San Marcos, becomes a raging river that
floods into Spring Lake. The study site was labeled TES for being at the top of the
Balcones Escarpment. The vegetation here was very different than at the BES site, with
many brushy species present. Also, evergreen trees and conifers were present here. These
types of trees are often associated with drier areas with poor soils. Many of the trees and
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Plant Ecology
woody plants do not shed leaves in order to maintain energy and to not waste water.
Species present included live oak, juniper, cedar elm, prickly pear and tasajillo, Texas
persimmon, and agarita. The association of these species in the density that they were
found at the TES was a dead giveaway that the area was over grazed at some point in
recent history. A common sight in the Texas Hill country is the encroachment of woody
species into what were formerly large open grasslands where woody species were the
exception, not the rule. Historically, fires were a large part of the ecology in the Edwards
Plateau region. Fires swept through the area and kept woody species to a minimum. Only
the most well established trees could remain through fire, so the brushy species were kept
at bay. When the grass was all eaten, there was no fuel for fires to spread and so woody
species took a firm grasp on the region (Fuhlendorf 819). Wildfires are of major concern
now due to massive fuel loads associated with dense brush and prolonged drought killing
many trees throughout the region. Even the different species of oak trees present, beloved
by many, are a major contributor to the encroachment of woody plants in the grasslands
of the Hill Country (Van Auken 214). Overabundance of deer has also led to selection for
woody plants in the hill country. Because deer eat forbs and broad leaf weeds for the
most part, those types of plants are over browsed while many unpalatable woody species
are left unchecked (Augustine et al 995).
Samples at each site were taken using a standard 22.9 cm. by 22.9 cm. square
collection frame with a wire handle. A spot with a layer of leaf litter on top of the soil
was chosen and the collection frame was placed down on the leaves. Leaves within the
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Plant Ecology
square frame were then collected and bagged. Leaves whose area were less than half in
the square were not collected nor were leaves that were seriously decayed. Once the
leaves were collected, they were stored in water and then sorted by species. The length
and width of all leaves that were suitable for use (less than 25% decomposed) was
recorded, and 10 leaves from each species were selected. The 10 leaves were then traced
on grid paper, and the area of each leaf was counted. Using linear regression, an equation
setting length and width as the independent variable and area as the dependent variable
was surmised. At this point, the theoretical area for every leaf based on their length and
widths was calculated so that leaf area and species location could be compared and
analyzed.
At each site, differences in leaf morphology could immediately be seen. The
leaves from the BES site were large, with massive surface area to volume ratios. The
leaves at this site were not tough either. By comparison, many of the leaves from the TES
site were small, thick, and waxy, all signifiers of leaves that conserve water more
efficiently. There is a cost associated with smaller leaves and a cost associated with larger
leaves, and both are associated with water availability (Kubiske et al 1405). Smaller
leaves sacrifice surface area in order to prevent water loss. In doing so they sacrifice
photosynthetic ability by diminished surface area to collect light. On the other hand,
leaves that are extremely large have much higher photosynthetic rates, but with large
leaves come more stomata and a higher respiration rate.
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Plant Ecology
Results:
In order to see if there were significant differences in leaf size and area dependent
on species, one-way ANOVA test was used on JMP. Using species as the treatment, leaf
length, measured leaf area, and estimated leaf area were all compared. In all three results,
a p value of less than .001 was observed, showing that there are significant differences
among species based on leaf size and area and these differences are not due to chance.
Additional analysis was done in order to see if there were differences in the
measurements between groups. Group and species were used as the treatments and
measured area was the dependent variable. These results showed a p value smaller than
.001 once again, showing that there are significant differences between groups not due to
chance. Some of these differences between groups could be a source of error, but overall
the differences in leaf size are still significant. With the mean expected leaf area at the
BES being 15.22 cm2 and 5.14 cm2 at the TES, an ANOVA was run and p value of less
than .001 was calculated. This shows that there are significant differences between leaf
areas between sites. All graphs and tables can be found in the Appendix.
Discussion:
As the results of analysis showed, there are significant differences in leaf size
between species, sites. These types of results can be expected as long as there is
environmental variation among locations where plants are sampled. The same
expectation should hold true for species within genera who are spread across large areas
with varying environments. One of the species present at the TES site was Rhus
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Plant Ecology
lanceolata, the Prairie Flame-leaf sumac. There are about 13 extant species of sumac in
North America, with some species being naturally rare and covering very small areas in
their distribution. Members of Rhus grow in tree and shrub form. The largest species can
reach up to 10 meters in height. At least one species, Rhus michauxii, is on the federal
endangered species list. Species in this genus are diverse in their habitat, with many
found across the Eastern United States, from tropical areas in Florida to high latitudes in
Canada. Additionally, several species are found west of the Rockies and throughout the
deserts of southwest North America (information gathered from USDA Plant Database).
Many of these desert species are found in extremely small populations naturally, as is the
case with many types of desert flora and fauna (Bevill 496).
The Prairie Flame-leaf sumac found at the TES site is perfectly adapted to being
where it is. The species is highly drought and heat tolerant, can grow on or near rocky
outcrops (top of escarpment), and can grow in slightly alkaline soils, something present
throughout the Hill Country. The center of the distribution of the prairie flame-leaf sumac
is in west Texas in the transition zone between the Edwards Plateau and Trans-Pecos ecoregions. This places the Prairie-leaf sumac nearly in the middle of the E-W distribution of
all sumacs in North America, ranging from Maine to California (Digital Representations
of Tree Species Range Maps from "Atlas of United States Trees). When leaf size and
geographic location is compared in sumacs in North America, some patterns appear to
reveal themselves (see table 1). Sumacs found throughout the eastern part of the United
States have considerably larger leaves than those found in the desert regions of the
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Plant Ecology
southwest. One of the largest differences is between Rhus aromatica, with leaf area as
high as 28 cm2 and Rhus microphylla, having leaves with an area anywhere from .5 to 4
cm2. The contrast between these two species is stark, however, a subtler pattern appears
across all species. Overall, as the species change from east to west, the leaves get smaller.
This change can be explained by the issues discussed in the methods section concerning
water availability. Moving west from the East Coast, average rainfall totals get lower and
lower, reaching their lowest points in the desert Southwest region (see fig. 1). Since
water loss through stomata is somewhat negated by abundant water resources, the species
of sumacs that have adapted to the eastern most areas in the country have large leaves to
maximize photosynthesis. Conversely, sumacs found in the desert areas have extremely
small leaves in order to minimize stomatal water loss. These exact same patterns can be
observed by comparing different leaves from different genera of plants collected from the
two sites in this study. The leaves from the trees at the BES had very large leaves relative
to the leaves present at the TES site.
Another study area of the various Rhus species was to find data on seed size. All
species in this genus produce a drupe containing one seed. When comparing seed sizes
across the different species, it was difficult to find data for some of the uncommon
members of the genus; however, the species for which that data was available showed
seeds to be around 3.5 mm2 in area. Additionally, while researching the seed sizes, drupe
size was often shown rather than seed size. For all species, the length of the drupe ranged
from 3 mm to 10 mm (USDA PLANTS, Aggie Horticulture, USGS Tree Survey).
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Plant Ecology
Because the seed takes up such a large volume of the drupe, the seed size closely
resembles drupe size and should not be overlooked as sufficient data for determining
relative seed size. It should also be noted that the smallest drupe size belonged to Rhus
microphylla, the species with the smallest leaf size and one of the species found in desert
climates. Though the data is not complete, it is likely that there is some association
between the climate and seed size in this genus, or that there are certain adaptations for
some of the desert dwelling species. Additionally, seeds are not necessarily the primary
mode of reproduction as all sumacs in North America readily reproduce via root suckers
(Doust 741). This mode of reproduction could explain some of the lack of diversity in the
investment of nutrients that this genus puts into its seeds based on where it is located
geographically, since seeds are not necessary for reproduction. Because this is a common
mode of reproduction for members of Rhus, some of the distribution patterns of the
extremely isolated and rare species make sense (Sherman-Broyles 556). These plants rely
on animals in order to reproduce sexually. Animals eat the drupe, and if an animal
dispersed a seed into these isolated areas, then these populations could propagate and
expand if the resources are available to do so.
Leaf and seed morphology appears to be closely related to water availability. By
examining leaves across two different sites with two very different microclimates,
differences in tree and leaf morphology were visually clear and easily explained. Upon
examination of the species of Rhus found throughout North America, water availability
clearly plays a large role in determining leaf size across vast areas. Even subtle
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Plant Ecology
differences in water availability can dramatically change the composition of vegetation in

an area. By examining leaf and seed morphologies across large areas, trends in changing
vegetation structure due to climate change may be uncovered. Additionally, comparing
current data of leaf and seed morphology and distribution to data from historical
specimens, models of long-term vegetation change can be created. These models could
then be used to compare vegetation responses from year to year to climate data over a
large area within a genus. This type of study would be beneficial to those located in areas
where drought or desertification is a concern, including areas of many areas of Texas.
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Works Cited:
Damesin, C., S. Rambal, and R. Joffre. "Between-tree Variations in Leaf s13 C of
Quercus Pubescens and Quercus Ilex among Mediterranean Habitats with
Different Water Availability." Oecologia 111.1 (1997): 26-35. Print.
Jackson, R. B. "Ecosystem Rooting Depth Determined with Caves and DNA."
Proceedings of the National Academy of Sciences 96.20 (1999): 11387-1392.
Print.
Fuhlendorf, Samuel D., and Fred E. Smeins. "Long-term Vegetation Dynamics Mediated
by Herbivores, Weather and Fire in Asavanna." Journal of Vegetation Science 8.6
(1997): 819-28. Print.
Auken, O. W. Van. "Shrub Invasions Of North American Semiarid Grasslands." Annual
Review of Ecology and Systematics 31.1 (2000): 197-215. Print.
Augustine, David J., and Lee E. Frelich. "Effects of White-Tailed Deer on Populations of
an Understory Forb in Fragmented Deciduous Forests." Conservation Biology 12.5
(1998): 995-1004. Print.
Kubiske, Mark E., and Marc D. Abrams. "Photosynthesis, Water Relations, and Leaf
Morphology of Xeric versus Mesic ecotypes in Central Pennsylvania in Relation
to Moisture Stress." Canadian Journal of Forest Research 22.9 (1992): 1402-407.
Print.
"Welcome to the PLANTS Database | USDA PLANTS." Welcome to the PLANTS
Database | USDA PLANTS. N.p., n.d. Web. 09 Feb. 2014.
Bevill, R. L., & Louda, S. M. (1999). Comparisons of related rare and common species in
the study of plant rarity. Conservation Biology, 13(3), 493-498.
"Digital Representations of Tree Species Range Maps from "Atlas of United States
Trees" by Elbert L. Little, Jr. (and Other Publications)." GECSC: Tree Species
Distribution Maps for North America. N.p., n.d. Web. 10 Feb. 2014.
Climate Prediction Center - Global Monsoons: North American Precipitation." Climate
Prediction Center - Global Monsoons: North American Precipitation. N.p., n.d.
Web. 10 Feb. 2014.
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Doust, John Lovett, and Lesley Lovett Doust. "Modules of Production and Reproduction
in a Dioecious Clonal Shrub, Rhus Typhina." Ecology 69.3 (1988): 741. Print.
Sherman-Broyles, S. L., Gibson, J. P., Hamrick, J. L., Bucher, M. A., & Gibson, M. J.
(1992). Comparisons of allozyme diversity among rare and widespread Rhus
species. Systematic Botany, 551-559.
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Appendix:
Table 1: List of Rhus Species found in North America, with approximate distribution edges in
latitude and longitude
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Fig. 1 Showing Typical Rainfall Gradients Across North America
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Analysis and Graphs
Oneway Analysis of Length By Species (Variable name)
Oneway Anova
Summary of Fit
Rsquare 0.405492
Adj Rsquare 0.392548
Root Mean Square Error 2.832042
Mean of Response 5.570233
Observations (or Sum Wgts) 1972
Analysis of Variance
Source DF Sum of Squares Mean Square F Ratio Prob > F
Species (Variable name) 42 10552.515 251.250 31.3262 <.0001*
Error 1929 15471.468 8.020
C. Total 1971 26023.983
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Oneway Analysis of ExpectedrArea By Species (Variable name)
Oneway Anova
Summary of Fit
Rsquare 0.316419
Error 1929 293054.89 151.92
C. Total 1971 428705.29
Oneway Analysis of Measured Area By Species (Variable name)
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Oneway Anova
Summary of Fit
Rsquare 0.609766
Error 893 63909.05 71.57
C. Total 935 163770.98
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Oneway Analysis of Measured Area By Species (Variable name)
Oneway Anova
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Summary of Fit
Rsquare 0.609766
Oneway Analysis of ExpectedrArea By Site (BES/TES)
Oneway Anova
Summary of Fit
Rsquare
Adj Rsquare
Root Mean Square Error
Mean of Response
Observations (or Sum Wgts)
0.079507
0.07904
14.15326
13.03168
1972
t Test
TES-BES
Assuming equal variances
Difference
Std Err Dif
Upper CL Dif
-10.077 t Ratio
0.772 DF
-8.562 Prob > |t|
-13.0444
1970
<.0001*
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Lower CL Dif
Confidence
-11.592 Prob > t

0.95 Prob < t
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1.0000
<.0001*
Source
Site (BES/TES)
Error
C. Total
DF Sum of Squares
1
34085.09
1970
394620.20
1971
428705.29
Mean Square
34085.1
200.3
F Ratio
170.1576
Means for Oneway Anova

Level
BES
TES
Number
1543
429
Mean
15.2238
5.1470
Std Error
0.36031
0.68333
Lower 95%
14.517
3.807
Std Error uses a pooled estimate of error variance
Oneway Analysis of ExpectedrArea By Site (BES/TES)
Oneway Anova
Summary of Fit
Rsquare
Adj Rsquare
Root Mean Square Error
0.079507
0.07904
14.15326
Upper 95%
15.930
6.487
Prob > F
<.0001*
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Mean of Response
Observations (or Sum Wgts)
Plant Ecology
13.03168
1972
t Test
TES-BES
Assuming equal variances
Difference
Std Err Dif
Upper CL Dif
Lower CL Dif
Confidence
-10.077
0.772
-8.562
-11.592
0.95
t Ratio
DF
Prob > |t|
Prob > t
Prob < t
-13.0444
1970
<.0001*
1.0000
<.0001*
Source
Site (BES/TES)
Error
C. Total
DF Sum of Squares
1
34085.09
1970
394620.20
1971
428705.29
Mean Square
34085.1
200.3
F Ratio
170.1576
Means for Oneway Anova

Level
BES
TES
Number
1543
429
Mean
15.2238
5.1470
Std Error
0.36031
0.68333
Std Error uses a pooled estimate of error variance
Lower 95%
14.517
3.807
Upper 95%
15.930
6.487
Prob > F
<.0001*

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