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A. Malcolm Campbell
Ecological Interactions
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.
All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any otherexcept for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.
First published in 2016 by

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Abstract
Food webs, energy flow, indirect effects, and nutrient cycling are described
as properties that emerge in ecological systems. Several of these proper-
ties are shown in this book to result from indirect effects and interac-
tions between species and abiotic components of ecological systems. For
instance, top predators affect organisms with which they do not directly
interact, including plants and non-prey animals. In some other interac-
tions, including competition, the nonliving components of ecological
systems (the abiota) can alter the outcome of a biotic interaction. A lim-
iting resource often results in competition, but varying environmental
conditions allow for species coexistence. Finally, this book illustrates how
energy flows in ecological systems, why it is rather inefficient, and how
species interactions relate to homeostasis and emergent properties. In the
course of that discussion, primary production, secondary production, and
trophic levels are defined. Energy flow in ecological systems is tied to the
carbon cycle.
Keywords
carnivores, food webs, energy flows, energy, nutrients, resources, popu-
lation, competition, cooperation, trade-off, consumption, resource use
overlap, competitive exclusion principle, environmental gradients, com-
petitive ability, limiting resource, ecological system, homeostasis, primary
production, trophic level, trophic pyramids, primary consumer, herbi-
vore, secondary consumers, primary consumers, respiration, assimilation
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Vegetation Diversity Increased When Wolves Were
Reintroduced into Yellowstone National Park....................1
Ethical, Legal, Social Implications: There are
Arguments For and Against Species Reintroductions....13
Chapter 2 The Outcome of Competition for a Resource Often
Depends Upon Environmental Conditions......................19
Chapter 3 Energy Flows Through FoodWebs...................................31
Chapter 4 Ecological Systems Are Not Very Efficient at
Transferring Energy From the Sun and Carbon
Dioxide From the Air to Predators...................................41
Conclusion............................................................................................47
Glossary................................................................................................49
Index....................................................................................................53
Preface
This book about ecological interactions is part of a thirty book series that
collectively surveys all of the major themes in biology. Rather than just
present information as a collection of facts, the reader is treated more like
a scientist, which means the data behind the major themes are presented.
Reading any of the thirty books by Paradise and Campbell provides read-
ers with biological context and comprehensive perspective so that readers
can learn important information from a single book with the potential to
see how the major themes span all size scales: molecular, cellular, organ-
ismal, population and ecologic systems. The major themes of biology en-
capsulate the entire discipline: information, evolution, cells, homeostasis
and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about ecological interactions and some
of the supporting evidence behind our understanding. The historic and
more recent experiments and data will be explored. Instead of believing
or simply accepting information, readers of this book will learn about
the science behind ecological interactions the way professional scientists
dowith experimentation and data analysis. In short, data are put back
into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turning
them into powerful and elegant Bio-Math Explorations. I learned a lot
from both of them. While the math is largely absent from this book, our
collaboration with her made this a better book. Nancy Stamp at Bing-
hamton University, and Bill Dunson and Richard Cyr at The Pennsyl-
vania State University influenced me greatly in how I think as a scientist
and approach my teaching. Finally, I thank my students in Integrated
Concepts in Biology II, who enthusiastically participated in our experi-
ment to redesign introductory biology, starting with the text and ending
with a new approach to teaching biology.
Introduction
A hawk swoops down and catches a bird eating seeds at a birdfeeder. This
interaction between two individuals, a predator and its prey, affects both
populations and can cause properties to emerge in ecological systems. The
predator-prey interaction leads to energy flow in the ecosystem. Popula-
tion growth and energy flow are emergent properties that arise because of
the actions of individuals and the coexistence of many species in ecological
systems, which often compete for limited resources. Interactions between
individuals often depend upon the location of the individuals, which may
vary randomly. It is difficult, if not impossible to predict exactly where an
animal will be within an ecosystem, although one can make a reasonable
guess based on its habits. The hawk may search near feeders for potential
prey. The location of an organism determines its interactions with other
organisms, populations, or ecological systems. Location leads to flexibility
in the response of ecological systems, because if all of the individuals in
a population were in the same place at the same time, they might all get
wiped out by some natural disaster or predator. In contrast, if individu-
als of a species are widely distributed, it is less likely that the entire spe-
cies will die. If individual birds do not group together, predators such as
hawks are unlikely to detect every bird, and some individuals will survive.
In this book, emergent properties of ecological systems will be examined.
CHAPTER 1
Vegetation Diversity
Increased When Wolves
Were Reintroduced into
Yellowstone National Park
Emergent properties of ecological systems can be examined by looking

closely at the impact of wolves in the Yellowstone National Park eco-
system. In ecological systems, there are predators and prey. A predator
is any organism that consumes another living organism in such a way
that the other organism, the prey, is killed. Predators can be classified by
the type of other organism they eat, and wolves are carnivores (animals
that eat other animals). Wolves hunt in packs and eat a variety of other
animals. Wolves feed mostly on large mammals, such as deer, elk, and
moose, which are all plant-eating herbivores. If more trees grow in the
presence of wolves as this section heading suggests, the wolves actions
go beyond its direct effect as a predator. Ecologists often construct food
webs (Figure 1) to depict how energy flows in an ecological system. Food
webs are diagrams that show who eats whom and how energy flows in
an ecological system. Energy flow refers to the movement of energy-
containing chemicals from one organism to another. Arrows are drawn
between prey and predator with the arrow pointed in the direction of en-
ergy flow. Energy and nutrients, in the form of carbohydrates, proteins,
and fats, flow upward toward the predators. Organisms that are prey con-
tain energy and nutrient resources for predators.
To understand why more trees grew when wolves were introduced,
the Yellowstone National Park ecological system must first be exam-
ined. To do that, consider the population of wolves, the populations of
2 ECOLOGICAL INTERACTIONS
Figure 1 A food web. The arrows point in the direction of the energy
flow. Here plants are eaten by grasshoppers and rodents, which are
eaten by an assortment of predators, ending in the owl and fox, which
have no predators.
Source: Copyright C. Paradise.
trees, and the entire ecological system. Most of the park is more than
7,500 feet above sea level with forests filled with conifers (trees with
cones). There are also areas of mixed deciduous forests, with trees that lose
their leaves each year, and grasslands. Grasslands and shrubby plant com-
munities predominate at lower elevations, which grade into coniferous
WOLF REINTRODUCTION INCREASED VEGETATION DIVERSITY 3
forests at higher elevations. Deciduous trees in this part of the world

typically grow along floodplains near rivers, but they also grow in higher
elevations. Winters are very cold, and summers are mild. In addition,
wildfires are a natural part of the Yellowstone National Park ecosystems.
As European settlers expanded across most of the United States in the
late eighteenth and early nineteenth centuries, they hunted large preda-
tors to extinction or near extinction. Within Yellowstone National Park,
the gray wolf was hunted out of existence before the National Park Ser-
vice was established to protect the natural resources. In the early days of
the park, poachers routinely hunted and logged within the boundaries of
the forest. From the late 1800s through the mid-1920s, hunters elimi-
nated the gray wolf from all of the lower 48 states, with the exception of
Minnesota. Minnesota, Alaska, and much of Canada became the only
refuge for the remaining gray wolves in North America.
After wolves were eliminated, their prey faced less predation pressure.
In Yellowstone National Park, the wolves primarily eat elk, large deer-like
herbivores. Although an individual elk is much larger than a wolf, an
emergent property of wolves is their social behavior of hunting in packs,
which allows them to work as a coordinated unit to kill animals much
larger than themselves. Predation is an example of our theme that biologi-
cal systems require resources, which results in competition or coopera-
tion. In this case, wolves require food resources, and they have evolved to
work cooperatively in hunting packs to acquire those resources. The social
behavior of wolves is an emergent property because their social behavior
cannot be predicted by studying solitary wolves.
An emergent property of interactions between individual predators
and prey is that the populations of these species are affected. The gain in
resources by a predator may provide the energy needed for reproduction,
leading to population growth. The loss of an individual in a population
of prey could affect evolution or growth rate of the population. After
removal of predators such as wolves from an area, prey populations often
increase in abundance very quickly. This population boom may require
the need for humans to control the rate of growth of the prey population.
One of the tasks of the National Park Service is to monitor the popula-
tions of many species within the parks, and this is especially important
for populations (such as, elk) that lack major predators (Figure 2). Elk
20,000 200
180
16,000 160
140
winter elk count
wolf abundance
12,000 120
= elk 100
= wolves
8,000 80
60
4,000 40
20
0 0
1990 1995 2000 2005 2010
year
Figure 2 Elk and wolf population counts in Yellowstone National

Park. Park rangers count elk from small planes. Years of missing data
for elk counts are either due to counts attempted during times of bad
visibility, or from counts not being made at all in 1996 and 1997.
Source: Data from US Fish & Wildlife Service and National Park Service.
populations had been monitored for decades within Yellowstone National

Park, and until the late 1960s, their populations had been managed by
human hunters inside of the park. Without the wolves, humans had to
limit the elk population to prevent starvation and disease. After 1968,
the National Park Service adopted a natural regulation strategy and no
longer allowed hunting within the park.
Natural regulation meant that park managers would let natural fac-
tors (such as, predation, disease, or lack of food) be the primary causes of
elk death. Note that other predators besides wolves (such as, grizzly bears
and mountain lions) are still in the park and they will eat elk. Hunting
was allowed outside of the park, and because elk migrate, some of the
Yellowstone National Park population encountered hunters when they
left the park. During the time when there was no hunting inside of the
park and no wolves, the elk population increased steadily to a maximum
of about 19,000 individuals.
The gray wolf was reintroduced to Yellowstone National Park in two
separate releases during 1995 and 1996, which was done to help the wolf
population of North America recover but could have an added benefit of
naturally controlling elk populations. As of 2012, about 80 to 90 wolves

in several packs lived within the park (see Figure 2), and over 200 were
in the Greater Yellowstone Ecosystem, which is an area that includes
the park and the surrounding National Forests and other federal lands.
Figure2 compares changes in the wolf population with changes in the elk
herd. The amount of human hunting varied from year to year, but dur-
ing the 20-year period shown in the graph, hunting pressure was fairly
consistent; humans killed an average of 1,148 elk from 1987 to 1995 and
an average of 1,297 elk from 1996 to 2004. Elk have steadily declined,
suggesting a possible relationship with the number of wolves.
So far, two key species have been considered, but the trees have not
been examined yet. When wolves were absent from Yellowstone National
Park, several species of trees (aspen, two species of willows, and two spe-
cies of cottonwood trees) failed to produce significant numbers of off-
spring trees. The adult trees produced seeds, the seeds germinated and
grew, but the seedlings did not reach maturity.
Narrowleaf cottonwood and black cottonwood trees produce many
thousands of seeds every year. Regardless of the number of wolves, many of
their seeds never become seedlings, and only a few seedlings become mature
trees. Every species evolves a reproductive strategy, which may include
such traits as age at maturity, number and survival probability of offspring,
and lifespan. A reproductive strategy is a suite of evolved life cycle-related
traits that taken together lead to successful existence of a species in the
context of that species environment. Natural selection influences each of
these traits. A common reproductive strategy in plants is to produce more
seeds than the environment could support as mature individuals. It might
seem as if producing more seeds than a plant needs to ensure survival of one
replacement individual is a waste of resources, but there are many reasons
that seeds fail to germinate and seedlings die. Long-lived plants such as
cottonwood trees that produce many seeds each year continue to live even
when a very small percentage of offspring in any one year live to maturity.
Although cottonwoods produced many more seeds than could sur-
vive, ecologists had noted that even fewer cottonwood seedlings and
trees were surviving than expected in Yellowstone National Park com-
pared to historical trends. One researcher, Robert Beschta, examined
the cottonwood trees in the Lamar Valley, in the northeastern quadrant
of Yellowstone National Park in the years following wolf reintroduc-

tion (Beschta, 2003). Cottonwoods grow primarily near water, so most
of the trees were found along the Lamar River that runs through the
Yellowstone valley. Beschta measured the diameter of 700 cottonwood
trees from two species that were 5 cm or more in diameter.
Diameters of over 98% of these mature cottonwood trees were be-
tween 30 and 110 cm (685 of the 700 trees). Less than 2% (15 trees) had
diameters between 5 and 29 cm. In addition to sizing all 700 mature trees,
Beschta estimated the density and heights of seedlings. He found most
seedlings to be between 0.1 and 0.6 meters high with some rare individu-
als 1 to 2 meters tall. Based on the sizes and calculations of density, Beschta
estimated that these seedlings were between 1 and 5 years old with densi-
ties from 4,000 to 70,000 per hectare (1 hectare = 10,000 square meters).
Of the thousands of seedlings produced every year, none were maturing
past this 5 year old seedling stage.
Beschta used tree ring data on narrowleaf cottonwoods to determine
the relationship between diameter of trees and tree age. Hollow cores
extracted from the center of a subset of the trees allowed determination
of tree age. Beschta used the relationship between diameter and age for
the subset to determine age of all trees. He could also determine when a
particular tree had germinated from its seed. The scientist developed an
expected distribution of trees of different ages if seedlings were maturing
into adult trees at a rate normal for cottonwoods. He then compared the
actual and expected distributions.
Before 1919, the number of cottonwood trees in each age category
was within the expected distribution because seedling maturation oc-
curred normally. This means that new trees were surviving as expected
up until about 1919. However, the number of new trees decreased dra-
matically by 1920, and the trees that remained were mostly large and
old. Thus there appeared a large gap between the observed distribution of
trees of different sizes and the expected distribution as the high expected
density of middle-aged trees was missing from the forest.
What led to the missing cottonwood trees in the Lamar Valley? From
his analysis of the history of the area, Beschta concluded that 1) fire was
unlikely to have affected the particular groves of trees he studied, 2) long-
term changes in climate had been insufficient to affect woody tree species,
and 3) it was unlikely that insect attack, frost, or disease would have had
the same effect on both species in each and every grove in the Lamar
Valley. Beschta concluded that, despite changes in management strategies
and sizes of elk populations in Yellowstone National Park, the effect of the
elk browsing on cottonwood seedlings was great enough to account for
the observed gap in seed production and seedling maturation.
Other studies have shown heavy plant feeding by elk on aspen and
willow trees with a similar pattern of gaps in the frequency distribution
of ages. Beschta also studied stands of cottonwoods where seedlings did
grow to maturity and he found many trees in the size ranges that were
missing from the Lamar Valley study (Beschta, 2005).
The patterns of frequency distributions at the different sites illustrate
the randomness and variation of emergent properties, which is one of the
themes of the idea of emergent properties. Randomness and variation
within biological systems allows populations to be flexible in their re-
sponse to changing environmental conditions. Some locations of trees led
to exclusion of elk, whereas others did not. The topography of La Duke
Spring and Devils Slide effectively excluded elk. Both sites are bordered
by a river and a road with fairly steep slopes. Although there were not
many very large trees because the grove of trees was established after the
new road was built, there were high proportions of smaller trees present,
which would not be expected if the site were frequented by feeding elk.
The exclusion of elk allowed the narrowleaf cottonwoods to have a
flexible response to the elk-cottonwood interaction. Cottonwood popula-
tions survived in some locations where elk cannot eat saplings, allowing
long-term survival of cottonwood within the entire region, even if some
other populations may not survive. The concept of a flexible growth re-
sponse by a cottonwood population was used by Beschta to illustrate the
pattern of tree growth in areas where elk were excluded, which parallels
the presence of wolves. Also, by demonstrating the growth of young trees
in elk-free areas, he was able to conclude that climate changes to the entire
ecosystem were not responsible for lack of seedling growth.
So far, elk-wolf interactions and elk-tree interactions have been ex-
amined, but the connection between wolves and the resurgence of trees
within the park has not been made. All of the data should be exam-
ined to see how wolves can influence tree growth. From Figure 2, it was
determined that as wolf numbers increased, the elk herd declined, al-
though the correlation was far from perfect.
However, tree maturation was low even when humans restricted the
size of elk herds through hunting and relocation programs that occurred
from the 1920s through the 1960s. It was not until after the natural
regulation of 1968 began that elk populations reached their highest lev-
els. Yet narrowleaf cottonwood sapling growth is absent throughout much
of the Lamar Valley from 1920 to the mid-1990s, which indicates the
number of elk is not the only determining factor for tree survival. The
wolf was the only component of the ecological system missing during that
time period. What is the difference between when humans controlled elk
population (1926 to 1968) and when wolves controlled elk populations
(mid-1990s and on)?
When a group of scientists tried to determine the difference between
human and wolf regulation of elks, they initially focused on the direct
effects of wolf predation. However, predation did not sufficiently answer
the question about the reemergence of aspen, willow, and cottonwood
trees. If the number of elk killed was the only factor that determined the
success of trees, then human control would have yielded similar results
to the natural control. They reasoned that indirect effects of predation
might be the key to understanding the emergent property of tree survival
in the presence of wolves. Indirect effects are another emergent property
of ecological systems. Indirect effects in ecological systems are effects of
one species on other species mediated through shared interactions with a
third species or group of species.
Once wolves were reintroduced to Yellowstone National Park, elk had
to face a predator that they had not encountered in many decades, and
certainly no elk living in the park in 1995 had ever confronted a pack of
hungry wolves. The elk that survived this new predation threat would be
the individuals that were more vigilant for these predators. Are there any
costs to the surviving elk with increased vigilance?
Scott Creel and his colleagues used radiotracking to study the locations
of 14 elk that were part of herds living in Gallatin Canyon, during 2002 and
2003 (Creel et al., 2005). Radiotracking is a method in which scientists fit
individual members of a population with collars that transmit a radio signal.
The scientists pick up the radio signal with receivers, which they used to
60
= forest
50 = edge
= grass
percent of observations
40
30
20
10
0
kills wolves wolves not
detected detected
Figure 3 Radiotracked elk locations and elk kills in Gallatin Canyon

within the greater Yellowstone ecosystem (outside of Yellowstone
National Park). Locations were in conifer forests, the edge between
forest and open grass, and open grass habitat. Wolves were either
detected nearby or not when each elk observation was made.
Source: Data from Creel et al., 2005, Figures 1a and b.
determine their locations. Creel and his colleagues also tracked wolves in two
out of three packs that used the same geographic area. The scientists assessed
the habitat use of elk during times when wolves were present and when
wolves were not detected in the area of the herd (Figure 3). When these data
(which are from just one area the scientists studied) are put into a computer
model with all of their other data to predict habitat use when wolves are
present or not detected, they were able to estimate the probability of grassy
areas or forest occurring where elk are located (Figure 4).
Creel and his colleagues concluded that the elk alter their habitat use
when they detect the presence of wolves. From Figures 3 and 4, it can be
seen that the elk behavior is not perfectly correlated with the presence
of wolves; there is variation in elk behavior. Elk are not perfect in their
detection of wolves, nor are they perfect in selecting habitats based on
their assessment of the risk of predation. Furthermore, the scientists had
not placed a radio transmitter on every wolf, and so some could be in the
area without detection. Elk cannot know where every wolf is, nor do they
know the best place to avoid detection by their predators. If they did,
0.8 = wolves not detected

= wolves detected
probability of occurence
0.6
0.4
0.2
0
elk presence in grass elk presence in forest
Figure 4 Effects of wolf presence on habitat use by elk. The

probabilities shown are the probabilities that grassy areas or
coniferous areas were prevalent where elk were found. Bars show
means and 95% confidence intervals.
Source: From Creel et al., 2005, Figure 3.
wolves would starve for lack of prey. In addition, prey animals often assess
their risk not only on the presence of predators but also on their need for
food. For instance, a hungry elk might take more chances on feeding in
the grassy areas (where their preferred food is) than an elk with a full belly
that may remain in the relative safety of the conifer forest.
The scientists concluded that elk move in response to the presence of
predators within 1 kilometer of their location. Again, variability in elk
movements results from variability in wolf detection by elk. Elk are more
likely to occupy sites, such as grassy areas, where they can forage with
confidence that wolves are absent (that is, they havent detected them).
Conversely, elk are more likely to seek protective cover in coniferous for-
ests when they detect wolves nearby. Altering habitat preference based on
the presence of wolves is likely to have several effects, including a decrease
in the energetic resources obtained by elk (not eating grass all the time), an
increase in energetic costs by elk (more vigilance when eating grass), and a
possible release of herbivore pressure on tree seedlings (fewer elk feeding in
the grass where new seedlings could sprout). Recall that the grassy areas are
often at lower elevations, in floodplains or near rivers, and this is exactly
where the groves of cottonwood, aspen, and willows are often located.
Now that a connection between trees, elk and wolves has been made,
it would be wise to determine if any other factors influence where the
elk eat and indirectly what they eat. To determine whether the changes
in elk behavior that occur in the presence of wolves have an indirect ef-
fect on tree growth, one would need to know whether more trees like
cottonwood, aspen, and willow are growing in areas where elk are spend-
ing less time. In another study, Beschta and a colleague, William Ripple,
measured stands of aspen trees in elevated and floodplain sites (Ripple
and Beschta, 2007). Like the cottonwoods, there are gaps in the growth
of aspen forests that correspond to the time when wolves were absent
from the park. The scientists determined the recent history of browsing
and measured the height of the five tallest young aspen in each of several
stands. They estimated the annual heights of these trees for the 9 previous
years based on the pattern of browsing damage of the highest branches.
At each tree they also measured the number of downed large logs that
were within 3 meters of the tree, because logs might impede elk escaping
through the forest. The scientists hypothesized that aspen trees would be
taller nearer streams than in upland areas if wolves are frequenting upland
areas. Aspens also were predicted to be taller at sites with downed logs than
sites with fewer or no downed logs due to behavior modification of the elk
caused by the reintroduction of wolves. Of four habitat types studied, they
predicted that stream-side sites with logs would have the highest predation
risk, whereas upland sites without logs would have the lowest predation
risk. Elk might be able to assess the risk of predation in different habitats,
and predation risk on elk should then correlate with aspen growth.
The scientists documented a consistent pattern. The percent brows-
ing declined the most in stream-side sites with logs, indicating that elk
are avoiding those areas after wolves were reintroduced. This decrease in
browsing led to significantly taller aspen trees by the end of the study.
Upland sites had the most browsing, although slightly more browsing
occurred at the site without logs. Elk changed their foraging behavior in
response to the presence of the predator, spending more time eating aspen
at sites that were not where wolves frequent and that had fewer downed
logs, making it easier to escape if a wolf were to appear. This change in
feeding pattern then led to regrowth of aspen (and other trees) where
wolves were and elk werent.
It is uncommon for a policy change to be measured so carefully to de-

termine whether the policy was a success or not. Many scientists studying
Yellowstone National Park ecological system concluded that the regrowth
of the vegetation was likely due to a combination of altered elk behavior
and reduced elk numbers in response to the reintroduction of wolves.
Much of the regrowth occurred near rivers, which wolves frequently use,
and would thus present a high predation risk to the elk. Aspen stands
with downed logs would present a further risk of predation because areas
with lots of downed logs would be difficult to run through while escaping
wolves.
The change in elk behavior leads to indirect responses throughout the
system due to emergent properties. As aspen, willow, and cottonwood re-
generate and begin reproducing again, other aspects of ecological systems
are predicted to change. Beschta and Ripple analyzed the communities
of berry-producing shrubs that often grow under aspen trees (Beschta
and Ripple, 2012). Among other things, they examined the relationship
between shrub height and aspen height and the number of different spe-
cies of shrubs under aspen stands with different heights. Some stands
contained many small aspens, and others had aspens that were taller than
what elk could usually reach and graze upon, as discussed above. As aspen
height increased, so did the height of shrubs underneath. In addition, the
number of shrub species increased with increased aspen height.
Other emergent properties appeared after wolves were reintroduced
to Yellowstone National Park. For instance, coyotes had enjoyed being
the top predator in Yellowstone National Park during the wolves long ab-
sence. Wolves interact with coyotes, and they may compete for resources
because they are both predators. Coyotes scavenge for food, feeding on
carcasses left behind by other predators. Wolves are larger and hunt in
packs, whereas coyotes are often solitary with a smaller body size. When
the wolves came back, the coyote numbers plummeted, and the average
size of surviving coyotes also increased. In addition, beavers have been
sighted in Yellowstone National Park where they have not been seen in a
long time. This recolonization of beavers is a consequence of regrowth of
a prime beaver food source, willow trees, along rivers.
Biological systems, from wolves to elk to beaver, require resources.
In order to obtain their resources, wolves hunt cooperatively in packs,
although different packs must compete for territory. Elk have to maintain
vigilance in order to avoid predation while foraging for the food they
need; as they alter their behavior to reduce the risk of predation, they may
exhibit a trade-off. A trade-off is a compromise that occurs as organisms
exchange one behavior or resource for another. Here, elk exchange abun-
dant food consumption for increased survival.
The entire ecological system responded to the presence of wolves in
ways that no one predicted when wolves were reintroduced to the park.
Beaver were not present in the park for many years in part because their
preferred trees (young willow trees) were not present in high abundance;
the number of beaver colonies went from one to twelve between 1999
and 2010. The ecological system contained within Yellowstone National
Park exceeds the sum of the individual species living in the park.
The randomness of this large ecological system provides flexibility of
response. For instance, the locations of downed logs, the meandering of a
river, or the movements of a herd of elk and a pack of wolves may all play
a role in whether a cottonwood or aspen can grow to maturity. Depend-
ing on the abiotic conditions at a particular location (such as, an upland
area or a floodplain), trees may or may not grow regardless of the elk.
The carcasses left behind by wolves provide a bounty to other scavengers,
including coyotes, which may explain the larger sizes of individual surviv-
ing coyotes that were not killed due to resource competition with wolves.
There may be more variability of response among scavengers depending
on abundance and location of the resource, possibly leading to more com-
petitive interactions as scavengers attempt to gain these resources. The
interactions between predators and their prey can lead to emergent prop-
erties. In the next chapter, how emergent properties arise from competi-
tive interactions will be examined.
Ethical, Legal, Social Implications: There are

Arguments For and Against Species Reintroductions
Over the last several decades, many species have disappeared through-
out all or part of their original ranges. The most common reasons for
loss of species are habitat loss, pollution, competition with invasive spe-
cies, and global climate change, which are all primarily caused by human
overpopulation and overconsumption of resources. Many people argue

that to restore ecological systems humans must reintroduce species to
their natural ranges, assuming they still exist somewhere. Perhaps the
most successful case of a species that was restored to its habitat is the
California condor with the largest wing span of any bird (9 feet). All 22
of the remaining condors were captured in 1987 and placed into a cap-
tive breeding program with the intent to rerelease them into the wild.
Since 1991, over 150 young adults have been released, and 150 are still in
the breeding program. Other species, such as gray wolves, may require
some assistance from humans to expand back into their original ranges.
Reintroducing the gray wolf to Yellowstone National Park was the
result of a long debate over the value of reintroducing a predator. Wolves
have been reintroduced in other places, and similar debates have occurred
in these places too, including Scandinavia and the Adirondack Park in
New York State. In addition to the biological data required to make a suc-
cessful reintroduction, social scientists conducted many surveys of public
attitudes and perceptions toward wolf reintroductions. Although 60% of
the general population supports reintroduction, only 35% of ranchers do.
As might be imagined, ranchers were nervous and opposed to the idea,
fearing for their livestock. Although a majority of the general popula-
tion has a positive attitude toward wolves, more of those people live in
urban areas and have little or no experience with wolves. People who
have a negative attitude toward wolves live near the wolves. Others argue
against reintroduction for other reasons, saying that money spent on cap-
tive breeding and reintroduction would be better spent solving societal
problems, like homelessness and poverty.
As these arguments are considered, one can appreciate the complexi-
ties of conservation. Because humans eliminated top predators, do we
now have an ethical obligation to restore those species to their former
ranges? When predators were eliminated, not much was known of the
ecological roles of those species. The direct and indirect roles of wolves
have been demonstrated in their return to Yellowstone National Park.
Gray wolves were the only native vertebrate species that was not still pres-
ent in Yellowstone prior to its reintroduction. However, local residents
are concerned about loss of life or economic damage to crops or livestock
that may result from wolf reintroduction. Humans eradicated gray wolves
from the park and the surrounding area because they perceived the wolves
as a threat to livestock and family and that perception persists. Should
federal tax dollars be used to conduct additional research to determine
whether these perceived threats were justified or not?
Ethicists have argued that if we honor our ethical duty to a past eco-
system by reintroducing a missing species, then we create a new conflict
with our ethical duties to present and future ecosystems, because a miss-
ing species may cause harm to other species. They ask if our duties to ex-
isting ecosystems outweigh our duty to past ecosystems. If we reintroduce
a species to an area where it is no longer present, species that currently
thrive there may be harmed. In the case of wolves, prey populations were
harmed, but some species increased in abundance after reintroduction. It
is well understood that indirect effects and emergent properties are dif-
ficult to predict. How can someone weigh these possible outcomes when
these properties cant even be predicted?
Finally, there is a concern among ecologists that a plan by humans
to restore an ecosystem leads the public to believe that ecosystems are
naturally static. Ecological systems are naturally dynamic and constantly
undergo changes. Extinction of a species can be caused by non-human
processes, and if humans are perceived as a natural part of an ecosystem,
then humans are just another natural cause of extinction, as natural as an
ice age or a volcano. Thus, both reintroductions and eliminations may be
viewed as equally harmful or beneficial to ecological systems. People with
different vested interests will fall on different sides of this ethical issue.
These arguments and the emotions they generate have led to changes
in laws and policy. The concern of the late 1960s and early 1970s was
caused in part by the publication of Silent Spring by Rachel Carson, sev-
eral environmental disasters, and the first Earth Day in 1970. Important
environmental laws were enacted during those times, including the En-
dangered Species Act (ESA) in 1973. The ESA is a very powerful law
aimed at preserving endangered species. The federal government defines
an endangered species as one at risk of extinction through all or a sig-
nificant portion of its natural habitat. A threatened species is at less risk,
but is likely to become endangered in the foreseeable future. The ESA re-
quires the US government to have a recovery plan for any species listed as
endangered or threatened. Biologists from the Fish and Wildlife Service
within the US Department of Interior are charged with developing recov-

ery plans to protect species and increase their populations.
Gray wolves were listed as endangered in the lower 48 states but not
in Alaska or Canada. The reintroduction of wolves into Yellowstone Na-
tional Park was part of the federal recovery plan. Part of the reintroduc-
tion debate centered on the question of whether humans should restore a
species that is abundant in Alaska to an area where they have become lo-
cally extinct. In addition to questions based on emotions and value judg-
ments, the recovery plan presented many legal questions too.
The cost of the reintroduction program to Yellowstone National Park
was a few hundred thousand dollars with no clear economic benefit at the
time. Now that the recovery is in full swing, the benefits are estimated to
be millions of dollars due to increased tourism to Yellowstone National
Park. The financial benefit takes into consideration lost income from
reduced hunting and compensation for ranchers who lost livestock. In-
terestingly, tourism is also an emergent property that has an economic im-
pact that had not been predicted when recovery plan was implemented.
In early 2008, the gray wolf was removed from the ESAs threatened
list for the lower 48 states, even though wolves are only in a few states.
Gray wolves were removed because about 1,500 wolves are thriving in
the greater Yellowstone ecosystem and several other areas in the Northern
Rockies. Ironically, the success of the wolves has produced unexpected
controversy, because their removal from the endangered species list meant
that wolves can be shot and killed once they step out of Yellowstone Na-
tional Park. An emergent property argument could be made that the only
way to protect the wolves is to keep their population low. Even while
acknowledging the success of wolf conservation, environmental groups
feared that losing their protected status could be a step backward, espe-
cially if hunters and ranchers begin killing wolves with impunity. Even
while ranchers complain that wolves eat their livestock and that they
should be allowed to kill wolves to defend their stock, environmental
groups have sued the federal government over the decision to remove
protection of the gray wolf. The judge reversed the Bush Administrations
decision to remove protection for the wolves. Rather than appeal the deci-
sion, the Bush Administration removed its support for the delisting, ef-
fectively placing gray wolves back on the endangered species list. During
the time that protection was removed, more than 100 wolves were killed
in Idaho, Wyoming, Montana, and parts of Oregon. This series of events
is an ongoing drama and debate about the ethical, legal, and social impli-
cations of humans trying to be good stewards of an endangered species
and the entire ecosystem.
Bibliography
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coyotes in northwestern Montana, J Mammal 83(3):754766, 2002.
Bangs EE, Fritts SH: Reintroducing the gray wolf to central Idaho and
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Beschta RL: Cottonwoods, elk, and wolves in the Lamar Valley of Yellow-
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Beschta RL: Reduced cottonwood recruitment following extirpation of
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CHAPTER 2
The Outcome of
Competition for a Resource
Often Depends Upon
Environmental Conditions
Within ecological systems, different species compete for resources that are
in short supply. Over evolutionary history, competition for resources has
led to evolution of specific adaptations for efficient gathering of those
resources. For instance, in the examination of the effects of wolves on
Yellowstone National Park in the previous chapter, it was mentioned that
they evolved social behavior, which led to cooperation within a pack.
When social behavior of wolves first evolved, the wolf ancestors that
exhibited cooperative behavior were more successful than those that
hunted alone. Cooperative behavior is behavior that involves several
individuals and is mutually beneficial, and just like any other trait has
a genetic component to it. Wolves that hunted cooperatively produced
more offspring, who in turn inherited the alleles associated with coopera-
tive hunting. Cooperation spread throughout the wolf population due
to increased resource acquisition, which increased the efficiency of food
gathering for all subsequent wolves. It might be predicted that the organ-
ism most efficient in gathering a specific resource will outcompete other
organisms and drive less efficient populations toward extinction. If wolves
are better hunters than coyotes, why hasnt the wolf driven the coyote to
extinction?
Two species such as wolves and coyotes that consume similar resources
have high resource use overlap. Resource use overlap is a measure of the
resources shared between any pair of species. Coexistence is possible, but
how do they strike a balance? According to the competitive exclusion

principle, one competitor excludes the other from a commonly used
resource when resource use overlap is high. The competitive exclusion prin-
ciple states that no two species that consume the exact same set of resources
can coexist. In other words, the two species need to evolve a mechanism to
eat slightly different foods (or otherwise divide up the resources), or one
of them will become extinct. Resource overlap leads to either extinction of
the losing competitor species or an evolutionary adaptation to use a differ-
ent resource, thus reducing overlap. The competitive exclusion principle
assumes that other ecological factors (such as, resource abundance and abi-
otic conditions) are constant. Competitive exclusion has many exceptions
that are almost always based on the fact that ecological factors frequently
change over time because biological systems are dynamic.
A scenario of high resource use overlap is not likely to lead to long
term sustainability of the predator populations. One of them will likely
go extinct, at least locally, and environmental factors other than resources
may be important in determining which one survives and which one goes
extinct. Measuring resource use overlap helps scientists assess the stability
of an ecosystem like Yellowstone National Park, predict future resource
usage, and plan conservation efforts. For example, using data on resource
use, ecologists could predict ways in which wolf and coyote behaviors
might evolve to reduce the degree of resource use overlap in Yellowstone
National Park. Similarly, conservation biologists could assess the resources
and predict the possibility of extinction of wolf competitors or the success
of wolves when wolves were reintroduced to the Park.
In addition, species that have high resource use overlap coexist by
altering their resource use when the competing species is present. Coyotes
may include a higher proportion of elk in their diet when wolves are
absent, but then switch to consuming more squirrels, rabbits, and mice
when wolves are present. Resource use overlap can be relatively high in
two coexisting species but cannot be 100%. As overlap increases, species
involved in competition may switch to unused, less desirable resources,
especially if they are abundant in the environment. This switching behav-
ior is a mechanism for species coexistence.
The outcome of competition depends on the species involved, but
it may also depend on variation in abiotic factors in the environment.
CONDITIONS CAN DETERMINE THE OUTCOME OF COMPETITION 21
Ecological systems change across space, whereas conditions related to an

abiotic factor change as location changes (for example, average tempera-
tures become cooler as one moves from the equator toward the poles, or
the moisture in soil may decrease higher up a slope from a body of water).
Another type of change is across time, where the condition changes as
time passes. A common example is the variation in the amount of rain as
seasons change from dry to rainy season. Some changes take place over
time and space. Consider an estuary, the region where rivers meet oceans.
The salt content of estuaries may change not only across space, moving
from a river delta toward the ocean, but also across time, tides push in
salty water or yield to fresh water in the river. Changes in environmental
properties across time or space are called environmental gradients, and
biologists often study how organisms respond to these gradients. An envi-
ronmental gradient is a change in an abiotic variable across time or space.
Some conditions are harsher than others; for most plants, drier soils
with fewer nutrients are more stressful than wet, fertile soils. Distur-
bances (such as, fires or strong storms) may also affect the ability of an
organism to survive. Some scientists have hypothesized that organisms
adapted to living in harsh conditions, either because of high levels of
disturbance or low levels of essential nutrients, are poor competitors in
less harsh conditions. If this hypothesis is correct, species adapted to con-
ditions of high nutrients or high resources would be better competitors
when living in conditions with high nutrients or resources.
One study that investigates this hypothesis about adaptation to harsh
conditions will be examined in more detail, but before that, the three
main ideas about emergent properties should be reviewed and applied to
this hypothesis. One main idea about emergent properties is that biologi-
cal systems require resources, which results in competition or coopera-
tion for resources. Another main idea about emergent properties is that
randomness and variation within an ecological system allows coexistence
of species. Different species that consume the same resource (e.g., plants
competing for sunlight) may be adapted to live in different environmen-
tal conditions. For example, plants compete for access to sunlight, but
some plants are adapted to direct sunlight and others prefer indirect light,
which minimizes resource use overlap. Tall oak and pine trees are adapted
to more direct light conditions, whereas shade-loving mosses and ferns
specialize in low light photosynthesis and yet all four plants consume the
same resource. These plants dont coexist at the exact same location, but
they can live near each other within in a wider ecological system, such as
a forest. The third main idea about emergent properties is that biologi-
cal systems exceed the sum of their parts. These ecological systems do so
by increasing biodiversity in the forest, causing indirect interactions, and
changing the abiotic conditions of the forest.
Ecologists Scott Wilson and Paul Keddy tested the hypothesis that
there is a trade-off between competitive ability and adaptations to harsh
conditions (Wilson and Keddy, 1986). They studied the distributions of
seven plant species along the shores of Axe Lake in Ontario, Canada,
where there exists an environmental gradient. This environmental gradi-
ent included both changes in disturbance in the force of waves hitting the
shore and the amount of organic matter and nutrients in the soil. The sci-
entists quantified the organic matter content in many plots as a measure
of the gradient because they had found in previous research that organic
matter negatively correlated with wave action intensityshores exposed
to high wave action have lower levels of nutrients. Their study sites ranged
from exposed beaches with high wave action and low nutrient concentra-
tions to sheltered shores in small inlets with nutrient-rich soils.
Wilson and Keddy divided the plots into seven organic matter cat-
egories. Within each category, the researchers determined the percentage
of plots of that contained each species, and represented those values as
a frequency distribution. Because plots often contained more than one
species, the percentages in any one category did not add up to 100%.
Regardless, they were able to compare frequency distributions of the dif-
ferent species against the organic matter content.
Once Wilson and Keddy plotted their data, they observed clear dif-
ferences in the distributions of the plants. Some plants had higher fre-
quencies of occurrence at lower organic matter levels, whereas others had
higher frequencies at the high end of the organic matter scale. Pipewort
(Eriocaulon septangulare) had a higher frequency of occurrence at low or-
ganic matter content, and generally breaksedge (Rhynchospora.fusca) and
brownfruit rush (Juncus pelocarpus) did, too. Loosestrife (Lysimachia ter-
restris) had higher frequency of occurrence in sites with higher organic
matter content, as did three-way sedge (Dulichium arundinaceum).
St. Johns wort (Hypericum ellipticum) had overall low frequencies of oc-
currence throughout, although slightly higher in medium organic matter
content sites. Finally, sundew (Drosera intermedia) had high frequencies
of occurrence at all medium and high organic matter content sites.
Wilson and Keddy designed an experiment to determine the competi-
tive abilities of each plant species compared to the other six species. The
scientists placed two plants, one from each of two different species or
two from the same species, in a small bucket filled with sand and organic
shoreline sediment, simulating a sheltered shore with high organic matter.
They had ten buckets of each pair-wise combination of species for a total
of 490 buckets.
The scientists measured relative growth, growth in the presence of an-
other species compared to growth in the presence of another member of the
same species, as an indication of competitive ability. They quantified their
comparative growth using a value they called relative increase per plant (RIP).
Growth was determined by measuring biomass at the time of planting and
again at the time of harvest about 3 months later. Biomass is the total mass
of a living organism after removal of water, which is often expressed or
referred to as dry mass. Because the biomass measurement kills the plants,
the initial measurements were made on a random sample of plants of the
same size as those planted, which were sacrificed for that purpose. To cal-
culate RIP at the end of the experiment, they determined the average final
biomass of plants of species i grown in the presence of plants of species j
minus the average initial biomass of plants of species i. They then divided
that quantity by the average final biomass of a plant of species i grown in
the presence of a plant of species i minus initial biomass of species i.
(final biomass of i with j) (initial biomass of i)

RIPij = (final biomass of i with i) (initial biomass of i)
RIPii, which assigns a value for plants grown in the presence of an-
other plant of the same species, will always equal 1. If RIPij is less than 1,
the plant of species i accumulated less biomass when grown with a plant
of species j than when grown with a plant of its own species. For instance,
brownfruit rush accumulated less biomass in the presence of three-way
sedge than when grown with another brownfruit rush. Conversely if RIPij
is more than 1, species i accumulated more biomass when grown with a
plant of species j than when grown with another plant of species i. Three-
way sedge accumulated more biomass when grown with brownfruit rush
than it did when grown with another three-way sedge.
In order to determine the performance of each species compared to
all others, Wilson and Keddy calculated the target score for each species.
They did this by averaging the RIPij values for all other species j in com-
parison to the target species. A target value above 1 indicates a species is a
good competitor and outcompetes all or most of the other six. Three-way
sedge did well in the presence of other species because the sedge accumu-
lated more biomass on average in the presence of other species than it did
in the presence of other sedge (average target score = 1.21). Brownfruit
rush, loosestrife and St. Johns wort all also had average target scores above
1, breaksedge and sundew had averages close to 1 and pipewort had an
average target score below 1 (average target score = 0.93).
Similarly, Wilson and Keddy calculated a neighbor score. To find this
value they averaged the RIPij values of all six other species i grown in the
presence of a particular species j. A neighbor score above 1 indicates that
other species i fared well when paired with species j. If a neighbor is a
good competitor on the other hand, other species will do poorly, and the
mean neighbor score will be less than one. Three-way sedge is not a good
neighbor, because the average neighbor did worse in its presence than it
did in the presence of a plant of its own species (average neighbor score =
0.90). Brownfruit rush is also not a good neighbor (average neighbor
score = 0.93). Loosestrife had an average neighbor score right at 1, mean-
ing that neighbors did equally well, on average, as if loosestrife was grow-
ing alongside another loosestrife. The other four plants, St. Johns wort,
breaksedge, sundew, and pipewort, all had average neighbor scores above
one, ranging from 1.15 to 1.35, suggesting that these species are good
neighbors and other species did well when growing next to them.
Finally, the scientists compared each target score and each neighbor
score to their average position on the sediment organic matter scale, and
they calculated the correlation coefficient for each set of scores versus the
percent sediment organic matter content. Average target scores were posi-
tively correlated with average percent organic matter content (r = 0.77,
P = 0.03). Average neighbor scores were negatively correlated with average
percent organic matter content (r = 0.68, P = 0.07). For these analyses,
the sundew was excluded because it was an outlier, having a target score
well below and a neighbor score well above where it would be predicted
based on its average position on the sediment organic matter scale.
Wilson and Keddy concluded that the sundew is not a very good
competitor in high organic matter soils, having the second lowest target
score, and only does better in head-to-head competition with pipewort.
And sundew is also a very good neighbor, with the other six species out-
competing sundew in the experiment. In addition, sundew is insectivo-
rous. Because of sundews ability to acquire extra nutrients from insects,
the researchers did not include this species in the correlation analysis.
Under the conditions of the experiment, sundew did not perform well,
but along the shores of the lake, sundew actually does compete well and
tends to be found in more favorable conditions. The nutrients from in-
sects give it a competitive edge.
This chapter began by asking whether there were emergent properties
associated with competition for limited resources. The plants investigated
here with high competitive ability tend to occupy sheltered, nutrient-rich
shores, whereas species with low competitive ability occupy disturbed
shorelines with low nutrient content soils. This make sense because those
who compete well get the best growing conditions, whereas those that
compete poorly must adapt to worse soil conditions or face extinction.
Adaptations to particular environmental conditions (such as, distur-
bance and soil nutrient concentrations) determine where species can live,
which directly or indirectly determine the outcome of competition. The
property that emerges from all this is a diverse community of plant species
that coexist at a location (such as, the shore of a lake) and yet use a variety of
microhabitats. Plants provide resources to many microbial and animal spe-
cies that benefit from these different plants. The beneficiaries of the plants
have, in turn, different competitive abilities and adaptations that they need
to feed on particular plants. All of these different interactions lead to in-
creased complexity and the potential for indirect interactions in ecological
systems, which is a good example of an emergent property at the ecological
system scale.
Although the seven plants exhibit a significant amount of overlap in
distributions at Axe Lake, Ontario, the sundew has adapted very well to
living in conditions in which the other species cannot exist. It would not
be expected that plants would evolve to be less competitive and adapt to

low-nutrient growth conditions. But growth in nutrient poor conditions
is better than going extinct after competing and losing out to a more
competitive species. This reveals an evolutionary mechanism involving
competing species: A species may adapt to exploit a resource more ef-
ficiently and become a superior competitor (target score > 1). Alterna-
tively, a species with low target scores may adapt to live in an environment
from which superior competitors are excluded, or it may adapt to exploit
a different resource altogether. Adapting to an alternative means to ac-
quire a limited resource is an explanation for sundew, the carnivorous
plant. In the next case study, competition for a different type of resource,
but one that also involves the introduction of new species to an area and
habitat alterations by humans, will be examined.
To examine the impacts of an introduced species and habitat altera-
tion on the outcome of competition, researchers at The Australian Na-
tional University studied birds that set up nests in tree cavities. These
cavities are often in short supply because many other animals (such as,
mammals and bees) also use these cavities as habitat, so competition may
be strong. If forests are cleared by humans that may well reduce the num-
ber of available cavities. If a tree-cavity nesting bird species is introduced
to an area where competition is already fierce, the outcome of compe-
tition may be altered. Researchers studied the invasive common myna
bird (Acridotheres tristis) and two native parrot species, the crimson rosella
(Platycercus elegans) and the eastern rosella (Platycercus eximius).
The goal of the research was to determine the impact of com-
mon myna use of nest boxes, which are artificial cavities set up by the
researchers, on the abundance of the two parrots in forests with differ-
ent tree densitiesfrom highly modified open grassy woodlands with
low tree densities to native forests with high tree densities. The research-
ers studied the abundance, percent occupancy of nest boxes, and egg
success (percentage of laid eggs that hatched) of each species across the
gradient of tree density (Figure 5). They determined that abundances
of common myna and crimson rosella were correlated with the per-
centages of nest boxes occupied across all forest types (compare Figures
5A and B). This is important to know to determine how competition
for nest boxes affects abundance of species. If a superior competitor or
140
bird density (#/km2 1SE)

120 = myna
= crimson rosella
100 = eastern rosella
80
60
40
20
0
A low medium high
60
nest box occupancy (% 1SE)
50
40
30
20
10
0
B low medium high
100
egg success (% 1SE)
80
60
40
20
0
C low medium high
tree density
Figure 5 Responses of three cavity-nesting bird species to forests of

different density. A, Mean bird densities. Change in abundance across
forest density types was statistically significant for all three species.
B, Mean percent occupancy of nest boxes. Change in occupancy
across forest density types was statistically significant for all three
species. C, Mean percent egg success. Change in egg success across
forest density types was statistically significant for common myna and
crimson rosella. SE, standard error.
Source: From Grarock et al., 2013, Tables 2, 3, and 4.
poor habitat is shown to reduce nest box occupancy, then that reduction
leads to a decrease in the population.
They then compared abundance of the two parrot species to nest box
occupancy of myna birds across forests of different densities (Figure 6).
Overall, the researchers hypothesized that tree density would influence
the three species differently and that increased occupancy of nest boxes
by common myna would reduce the abundance of the parrots, especially
at low tree densities.
Similar species often overlap in their geographic ranges and their diet.
This leads to potential competition between the two species. As would be
the case with the wolf and coyote, different combinations of prey items
in the diet lead to more or less overlap in diet and, thus, more or less
competition. When wolves were reintroduced to Yellowstone National
Park, there was suddenly competition in the park between wolves and
coyotes. This led to a change in the coyote population that was present
in the park. Competition can lead to adaptations to environmental con-
ditions: A species may adapt to some environmental condition, such as
high temperature or high tree density, in order to avoid competition with
another species.
With the myna and two parrots, the overlap in nest cavity use was
100%, or very close to it. All three species utilized the nest boxes, but
they may be more selective in natural cavities, reducing competition. The
introduction of a new species, the common myna, which overlapped in
nesting requirements so much with native species could, and did, led to
competition.
However, the strength of competition varied with tree density and
the variability in tree density at different sites allowed all three species
to coexist, despite the dramatic disturbances associated with introduc-
tion of a strong competitor and forest clearing by humans. These two
disturbances appear to work together, because clearing of forests leads
to habitats with lower densities of trees and thus fewer nesting cavi-
ties. The common myna preferred low density forests and had densi-
ties about ten times higher in those forests. High densities of common
myna led to high percent occupancy of nest boxes and greater success
of eggs. Something in the low density forests allowed myna birds to be
more successful.
160
crimson rosella abundance (#/km2)

140 = high tree density
= medium tree density
120 = low tree density
100
80
60
40
20
0
0 20 40 60 80 100
A common myna nest box occupancy (%)
120
eastern rosella abundance (#/km2)
100
80
60
40
20
0
0 20 40 60 80 100
B common myna nest box occupancy (%)
Figure 6 Relationships between the percentage of nest boxes occupied

by myna birds at a site and abundance of rosellas. A, Crimson
rosella. B, Eastern rosella. The best fit regressions were statistically
significant for both rosella species.
Source: From Grarock et al., 2013, Figures 6 and 7.
Low tree density forests might naturally have low densities of rosellas.
However, the researchers were able to show that in those habitats high tree
densities of myna birds led to high occupancy of nest boxes and that re-
duced the abundance of crimson and eastern rosellas. When mynas were
less abundant as in the medium and high tree density forests, abundance,
nest box occupancy, and egg success all went up to different degrees in
the two rosella species.
Changes in an environmental condition (in this case, forest tree den-

sity) strongly influenced the outcome of competition for a limiting re-
source (in this case, nest boxes). A limiting resource is a resource (such as,
food, light, nutrients, or space), which is in short supply and restricts the
growth of an organism or population. The outcome of competition was
determined partly by the differential use of habitat by the birds determined
by their different genetically encoded responses to variable forest types.
This chapter considered how competition for resources was affected
by variability in environmental conditions. The results of the studies dem-
onstrated all three of the main ideas of emergent properties. Adaptations
to different habitats led to the evolution of diverse species that utilize
similar resources within an ecosystem in such a way that they can coexist
in a broad geographical region. Species diversity is an emergent property
of ecological systems; the ecosystem, with its individual species all living
in the same region, is more than the sum of its parts. Competition for
resources led species to evolve adaptations to exist in different microhabi-
tats. The three bird species have adapted to slightly different conditions,
which often helps avoid direct competition and possible extinction. The
three bird species were able to coexist together, even though densities of
rosellas were reduced in the presence of high myna density, possibly be-
cause the birds competed less for other resources, such as food.
Bibliography
Grarock K, Lindenmayer DB, Wood JT, et al.: Does human-induced
habitat modification influence the impact of introduced species?
A case study on cavity-nesting by the introduced common myna
(Acridotheres tristis) and two Australian native parrots, Environ Man-
age 52:958970, 2013.
Pianka ER: The structure of lizard communities, Ann Rev Ecol Syst 4:
5374, 1973.
Pianka ER: Niche overlap and diffuse competition, Proc Nat Acad Sci
71(5):21412145, 1974.
Wilson SD, Keddy PA: Species competitive ability and position along a
natural stress disturbance gradient, Ecology 67:12361242, 1986.
CHAPTER 3
Energy Flows Through

FoodWebs
In Chapter 1, the effects of reintroduction of a top predator, the gray

wolf, on its prey and the plants on which the prey feed in Yellowstone
National Park were examined. The energy pathways in that ecological
system changed when a new species was added or an old one was reintro-
duced, and this can cause a disruption to ecological system homeostasis.
Food webs were considered in the context of trophic levels. In this chapter
and the next, cycling of carbon and the efficiency of energy transfer in
ecological systems will be considered in more detail. In this chapter, the
pathways of energy in a food web will be examined to better understand
species interactions and how food webs are an emergent property in eco-
logical systems.
Robert Paine performed experimental manipulations of ecological
systems (Paine, 1966). His studies focused on rocky intertidal communi-
ties. Specifically, Paine collected information on feeding relationships in
rocky intertidal zone food webs that exist at two geographic latitudes.
The food webs Paine studied are only a portion of the entire food web
of the coastal ecological system and primarily comprise the macroscopic
species that are normally intertidal and live on the rocks. He did not in-
clude microscopic species or species that actively swim through the water.
Paine studied these food webs and made observations on the quantity
and composition of each species diet. Specifically, the biologist examined
individual carnivores exposed by low tide, and he recorded their identity,
their prey, and the length of both predator and prey and also made gen-
eral observations of the interaction.
Paines most northern study area was a rocky shore off the coast of
Washington State, where the community is dominated by mussels,
barnacles, snails, and one starfish. Paine observed communities in this

area over an entire year, focusing on predation events by the two carnivo-
rous species in the web, Pisaster ochraceus, a starfish, and Nucella emar-
ginata, a small snail.
Paine determined the caloric content of each average sized prey type
and determined the percentage of calories, and consequently the percent-
age of energy, acquired by each predator when consuming each prey type.
A calorie is the amount of energy, or heat, it takes to raise the temperature
of 1 gram of water 1o C, or 4.184 joules. For some species for which Paine
did not measure caloric content, he estimated it based on similar species
that he did measure. In addition, the scientist did not distinguish between
species within certain prey categories. In these cases, Paine assumed that
the carnivores exercised no preference in their choice of species within
that prey category.
Based on over 1,300 observations of predation events, Paine found
that Pisaster consumed chitons (2 species grouped together), limpets
(2 species), bivalves, acorn barnacles (3 species), and Mitella, another spe-
cies of barnacle. Chitons constituted 3% of the prey items eaten (41% of
calories), limpets 5% (5% of calories), bivalves 27% (37% of calories),
acorn barnacles 63% (12% of calories), and Mitella 1% (3% of calories).
Pisaster also consumed Nucella occasionally (less than 0.01% of prey items
eaten, constituting 2% of calories consumed). These data were compiled
from 1049 observations of Pisaster feeding.
Pain made another 287 observations of Nucella feeding and found
that 5% of the prey items they consumed were bivalves (constituting 10%
of their caloric intake) and 95% were acorn barnacles (constituting the
rest of their caloric intake). Thus Nucella and Pisaster both consumed bi-
valves and acorn barnacles. For Pisaster these prey made up 90% of their
prey, by abundance, but only 49% of their calories. Nucella ate only these
types of prey, but focused more heavily on acorn barnacles, which also
made up a large proportion of the prey items consumed by Pisaster (63%).
Paine constructed an analogous web for a subtropical coastal habitat
from the Northern Gulf of California. The top carnivore in this ecologi-
cal system is also a starfish (Heliaster kubiniji), the next two trophic levels
contain carnivorous snails, and the main prey are herbivorous snails, bi-
valves (such as, clams), and barnacles. The percentage data on numbers
Energy Flows Through FoodWebs 33
Table 1 Percentages of prey consumed (before the slash) and

percentages of calories in diet (after the slash) by each predator.
Prey are grouped by type with the result of lumping of multiple prey
species, as denoted by spp. Sample sizes were 2,245 (Heliaster),
113 (Muricanthus), 62 (Hexaplex), 14 (A. tuberculata), 432
(A. angelica), 39 (Morula), and 8 (Cantharus). A, Higher trophic
level carnivores that consume at least some carnivores. B, Carnivores
that consume only herbivores/detritivores.
A. prey Heliaster Muricanthus A. tuberculata Hexaplex
Muricanthus <0.01 / 4
Acanthina tuberculata <0.01 / 1 1/6
Hexaplex <0.01 / 4
Morula 1/2 <0.01 / 2 2/2
Cantharus 3/9
Acanthina angelica 1/6 57 / 74
Columbellidae 3/6 3/4 6/7
bivalves 31 / 35 36 / 21 14 / 4 77 / 53
herbivorous snails 8 / 30 24 / 48 28 / 22 16 / 37
barnacles 51 / 9 31 / 3
chitons 1/3
brachiopod 5/4
B. prey Morula Cantharus A. angelica
barnacles 100 / 100 100 / 100 100 / 100
Source: From Paine, 1966, Figure 2.
and calories is presented in Table 1. If all the numbers are added up to

the left or right of the slash in each column (that is, for each predator),
it adds up to close to 100%. In addition, one could reconstruct the food
web from the data in this table.
The predators in the Washington State rocky intertidal food web rely
to a large extent on barnacles and bivalves, but Pisaster also relies heavily
on chitons. The numbers in the food webs show the percentage of either
individuals or total caloric content of a particular prey type. The numbers
are not exactly related to rates of energy flow, but because calories mea-
sure the heat released by a food item when it is entirely combusted, the
percentages of calories in a pathway from predator to prey should indicate
which pathways are major and which minor for energy flow. For instance,
63% of all prey individuals eaten by Pisaster are acorn barnacles, but that
represents only 12% of the calories that flow into Pisaster. Bivalves and
chitons combined provide 88% of Pisaster calories, even though they
make up only 30% of the prey consumed.
If all prey items had caloric content percentages equal to their num-
bers consumed, then one could conclude that they are approximately
equal in terms of individual caloric content. But chitons, making up
only a small percentage of individuals consumed in the Washington State
rocky intertidal food web account for 41% of calories in the Pisaster diet.
Conversely 63% of prey eaten is barnacles, which accounts for only 12%
of Pisaster calories. This indicates that individual chitons have many more
calories than an individual barnacle. In fact, chitons are larger animals
than barnacles.
In the Gulf of California rocky intertidal food web, the top predator
starfish (Heliaster) relies heavily on bivalves and herbivorous snails for
two-thirds of calorie intake. Muricanthus and Hexaplex, two predatory
snail species also rely heavily on these prey. Acanthina tuberculata, another
predatory snail, relies on herbivorous snails but 74% of its calorie intake
comes from Acanthina angelica, a snail that preys on barnacles. Three
other snails observed by Paine feed only on barnacles. Although barnacles
provided 100% of the calories to these three species, other carnivores that
consumed barnacles derived much less of their total energy from these
prey, even if they made up a large part of the diet.
Comparing the two food webs, there are three trophic levels in the
Washington State rocky intertidal food web and five in the Gulf of
California rocky intertidal food web. The Washington State food web
contained 11 species, 18.2% of which were carnivores, and the Gulf of
California food web contained 45 species, 15.6% of which were carni-
vores. The analysis presented for these food webs is a bit simplistic because
it turns out that some species change their feeding habits as they mature;
some species are prey to carnivores only in younger, smaller stages; and,
of course, there are species that are not shown, as mentioned earlier. Yet,
it has been found for other ecological systems that diversity tends to be
higher closer to the equator.
Paine next performed a manipulation in the Washington State rocky
intertidal food web by excluding Pisaster from an area about 8 2 meters.
An adjacent area was monitored, but Pisaster was allowed to hunt nor-
mally. The scientist sampled both plots at irregular intervals and counted
the number and density of resident large attached invertebrate and algal
species (Table 2).
Table 2 Species present in the presence or absence of Pisaster.

species present in Pisaster Pisaster
notes on exclusion plot
plots (# possible) present excluded
bivalve (1) 1 1 dominant species
acorn barnacles (3) 3 3 barnacles being crowded out
limpets (2) 2 0
Mitella (1) 1 1 exist in scattered clumps
chitons (2) 2 0
algae (4) 4 1
anemone (1) 1 1 much reduced in density
sponge (1) 1 1 much reduced in density
Source: From Paine, 1966, text.
In the control area, as Paine had found in other areas, there existed the
nine attached species in the food web (not counting the motile Nucella)
plus four species of algae, one anemone species, and a sponge species.
Immediately following removal of Pisaster, one species of acorn barnacle
established a large population and occupied between 60% and 80% of
the available space on the rocks. The following summer, the barnacles
were being crowded out by rapidly growing bivalves and Mitella, a dif-
ferent kind of barnacle. The process of continued replacement of existing
individuals by the bivalve continued up to the end of the experiment.
The species that became dominant filled up much of the space available
to them on the rocks.
The removal of the starfish Pisaster resulted in a decrease in the num-
ber of species in the food web, including species that did not interact
with Pisaster at all. The number of species went from fifteen to eight.
The density of those fewer species was much higher than in the control
rock, measured simply by the amount of space they took up on the rocks.
Predation by Pisaster increases diversity. Paine predicted that several of
the eight remaining species would eventually be eliminated, although his
study did not go on long enough to observe those effects. In fact, all spe-
cies with the exception of the dominant bivalve, a mussel, might eventu-
ally be eliminated from the Pisaster removal plot. Paine concluded that
Pisaster, the top predator, functioned to reduce the density of the compet-
itive dominant species, the bivalve mussel. Much like the gray wolf when
it was eliminated from Yellowstone Park, the starfish has indirect effects
on diversity. The indirect effects differed, as the gray wolf elimination led
to increased herbivory by elk and a reduction in growth of trees, whereas
the elimination of the starfish led to increased resource use by mussels,
and a reduction or elimination in other species.
The removal of Pisaster led to an accumulation of energy within the
bivalve population. That energy did not flow upward to the top trophic
level as it normally would with Pisaster present. In addition to accumula-
tion of energy in a lower trophic level, one might also have predicted that
another predator might move into an area devoid of its top predator, or
that energy would flow more into the decomposer food web as individual
bivalves or other prey died from overcrowding or space limitation.
In an ecological system with many predator-prey interactions, the ef-
fect of the loss of a predator may depend on the strength of the inter-
action between predator and prey, or the amount of energy that flows
through that link in the food web. Jordi Bascompte and his colleagues
studied the homeostasis of a Caribbean marine ecological system through
analysis of food webs (Bascompte et al., 2005). In addition, the biologists
attempted to predict the effects of overfishing of top predators on ecologi-
cal system homeostasis. Bascompte and his colleagues analyzed patterns
in interaction strengths among all predator-prey interactions. Interac-
tion strengths are not exactly the same as energy flow, but the two mea-
sures are related. Interaction strength is the impact of one species on the
biomass or abundance of another species, usually in a consumerresource
relationship. The scientists analyzed a large food web by quantifying the
per capita interaction strengths between all predatorprey links.
For their marine food web, Bascompte and colleagues compiled data
on 249 species and 3,313 predatorprey interactions. For the most part,
their data of feeding relationships, size and biomass estimates, and in-
teraction strength came from or were calculated from data from other
published studies. The scientists included all bottom and open water com-
munities from the surface to 100 meters in depth in a 1,000 km2 area.
Bascompte and his colleagues calculated a standardized measure of
interaction strength of predators on their prey, which was the estimated
proportion of prey biomass consumed per unit of predator biomass per
day for each predatorprey interaction. Their goal was to use estimates of
consumption to biomass ratios to obtain measures of energy transfer that
the scientists then standardized to per capita effects of predators on prey

populations. The equation used was [(Q/B)j DCij]/Bi. In this formula,
(Q/B)j is the number of times a population of predator j consumes its
own weight per day, accounting for individuals of different age and size.
Bj is biomass of predator j. Multiply (Q/B)j Bj to obtain the biomass
that the predator population requires per day.
DCij is the proportion of prey i in the diet of predator j, and these
data were obtained from studies examining predator stomach contents.
By multiplying (Q/B)j Bj DCij, they obtained the total biomass
of prey i consumed daily by populations of predator j. To calculate the
per capita, or per individual, biomass of prey i consumed, they divided
(Q/B)j Bj DCij by Bj. This then results in the numerator of the equa-
tion used by Bascompte and his colleagues.
The scientists defined Bi as the biomass of prey i, determined, as for
predators, by the average number of individuals per square meter multi-
plied by average body mass. By dividing the per capita biomass of prey i
consumed [(Q/B)j DCij] by the total biomass of prey i, they obtained
the standardized measure of per capita interaction strength for a predator-
prey interaction. Rather than amount of prey consumed by one indi-
vidual predator, this standardized measure of interaction strength is the
proportion of a prey population consumed daily by one unit of predator
biomass. Once the measure of interaction strength was made for each
predatorprey pair, the scientists generated a frequency distribution of
interaction strengths (Figure 7).
In addition, the scientists studied how interaction strengths are com-
bined in known two link feeding chains embedded in their food web. A
two-link chain has three trophic levels and three speciesfor instance, a
plant, its herbivore, and a carnivore. In Paines Washington State rocky in-
tertidal food web, there are two two-link chains, for instance. Bascompte
and his colleagues also examined food chains that exhibited omnivory.
The two two-link food chains from Paines study include omnivory be-
cause Pisaster consumes Nucella but also Nucellas prey (bivalves and acorn
barnacles). The scientists considered all two-link food chains within their
complex web.
The scientists classified the strength of the two interactions within
each food chain into one of four classes. The 25% of interactions that
1000
800
600
frequency
400
200
0
7 6 5 4 3 2 1 0 1
log of interaction strength
Figure 7 Frequency distribution of non-zero predator-prey interaction

strengths from a marine food web (n 5 3,313) spanning seven orders
of magnitude.
Source: From supplemental data file from Bascompte et al., 2005.
were weakest were designated Class 1, the next 25% Class 2, the next
25% Class 3, and the 25% of interactions that were strongest were desig-
nated Class 4 (also called strong interactions). The species of fish involved
in most strongly interacting two-link food chains included sharks and
groupers as top predators preying on a variety of fishes.
Bascompte and his colleagues examined the two link food chains with
and without omnivory where the interaction strengths were all strong, rel-
ative to all other two-link food chains. To determine whether two strong
interactions occur in the same two-link food chain (leading to a major en-
ergy pathway) more often than expected by chance within two-link food
chains, the scientists randomly shuffled all the interaction strengths from
the original food web. They repeated this process for 50,000 simulated
food webs, each time determining the number of two-link food chains
with two strong interactions.
From the distribution of these values from the simulations, Bascompte
and colleagues determined that two strong interaction strengths in the
same two-link food chain occurs less frequently than expected by chance;
99.8% of the simulated food webs had greater numbers of two-link food
chains with two strong interactions. And within two-link food chains
with two strong interactions, there was a high probability that the food
chain included omnivory (the top predator preyed on both other species).
Only 0.01% of the simulated food webs had higher frequencies of two-
link food chains with omnivory where the omnivory interaction strengths
were strong.
There are 25 two-link food chains in Paines Gulf of California food
web, many of which include omnivory because the starfish Heliaster eats
10 of the 12 groups of prey shown. Certain pathways are more impor-
tant in terms of energy flow, but it is possible that if one pathway was
eliminated the starfish would adjust its diet and another pathway might
come to dominate. Although Paine did not measure interaction strengths,
pathways through which more energy flowed would likely be classified as
strong interactions, and one could predict them based on the number and
percentage of calories consumed in each predator-prey interaction.
In the Caribbean marine food web analyzed by Bascompte and his
colleagues, most species did not interact with one another; 95% of pair-
wise interaction strengths were zero. Only 3,313 out of a possible 61,752
interactions were more than zero. Extremely weak non-zero interactions
indicate that a predator rarely consumes a particular prey, either because
it is rare in the community, hard to catch, distasteful in some way, or not
preferred as prey.
The 25% of interaction strengths classified as strong (Class 4) were less
likely to be found together in the same two-link chain than by chance.
This might help maintain energy flow homeostasis in ecological systems.
Much of the energy of an ecological system flows through pathways of
predators and prey that interact strongly, but because strong interactions
are dispersed throughout the food web in the Caribbean marine food
web, eliminating one predator or another high on the food chain may not
result in a drastic effect on energy flow in the system.
Bascompte and his colleagues were interested in this question, but
they were also interested in what would happen to the biomass at the
base of a food chain if a top predator were overfished in a strongly in-
teracting two-link system. Removal of a lone top predator can lead to
increased biomass of its prey, as Paine showed in his study of the removal
of Pisaster. Overfishing tends to eliminate species higher in food chains,
so the scientists simulated the fishing of top predators and explored the
subsequent change in biomass at the base of food chain. They explored
two-link food chains with and without omnivory. If removal of a predator
affects the amount of biomass of a plant, then a trophic cascade has oc-
curred. A trophic cascade is when a change in the size of one population
has an effect on populations in other trophic levels.
The scientists concluded that when a two-link food chain has two
strong interactions a trophic cascade would be more likely. When a preda-
tor is removed, biomass of prey should increase, as Paine showed, and
that increased biomass should cause a reduction in biomass of the preys
food. This has been shown for some ecological systems. However, Bas-
compte and colleagues showed that the magnitude of the trophic cascade
is reduced in the presence of strong omnivory; and because most of their
two-link food chains contained only one or no strong interactions, the
likelihood of trophic cascades due to the loss of a top predator is reduced
in this marine system.
This suggests homeostasis of ecological systems might be more easily
maintained, but Bascompte pointed out that fishing selectively targets a
set of species in upper trophic levels that tend to be part of strongly in-
teracting two-link food chains. The scientists found that ten fish species
currently under heavy fishing pressure accounted for 48% of the strong
interactions in two-link food chains. Of these food chains, 31% had a
buffering effect from strong omnivory, and 69% had potential for tro-
phic cascades. Thus the impacts on ecological system homeostasis may
be stronger than expected; because fishing practices preferentially target
species whose removal can lead to trophic cascades, and this can upset
homeostasis of an ecological system. In the next chapter, more about the
efficiency of energy flow in ecological systems will be examined.
Bibliography
Bascompte J, Melin CJ, Sala E: Interaction strength combinations and
the overfishing of a marine food web, Proc Nat Acad Sci 102(15):5443
5447, 2005.
Paine RT: Food web complexity and species diversity, Am Nat 100(910):
6575, 1966.
CHAPTER 4
Ecological Systems Are

Not Very Efficient at
Transferring Energy From
the Sun and Carbon
Dioxide From the Air to
Predators
In the first two chapters of this book, the focus was on emergent properties
that arise in ecological systems as a result of predatorprey or competitive
interactions. Each of these types of interactions affects the flow of energy
and nutrients within an ecological system. Energy flows are emergent
properties, and these will be examined in more detail here. Ecological sys-
tems are characterized by interactions and interdependent relationships.
Emergent properties at the ecological systems level arise when species live
together in the same habitat, obtaining their resources in diverse ways.
Energy, in the form of sunlight, carbohydrates or fats, as well as nu-
trients (such as, carbon dioxide, nitrates, and phosphates) are the re-
sources that all organisms must obtain to survive and maintain stable
or constant internal conditions (homeostasis). Ultimately, energy from
the sun powers the growth of animals at the top of most food webs (see
Figure1 in Chapter 1). All of the interactions discussed in this chapter
revolve around obtaining resources, from wolves eating elk or plants com-
peting for soil nutrients.
Each individual plant, bacteria, fungus, and animal goes about its
business, obtaining its own resources, confronting its own predators, prey,
competitors, parasites, and so on. When an elk chews on a cottonwood

sapling, there is a transfer of energy and material from the plant to the
herbivore. Plants convert light energy from the sun and carbon dioxide
from the atmosphere into the chemical energy of carbohydrates, and this
conversion of energy is called primary production. Primary production
is the production of organic compounds from carbon dioxide, principally
through the process of photosynthesis. Adding up all the light energy in
primary production will equal the amount of energy available in the first
trophic level. A trophic level is a feeding position in a food web, which
describes what an organism eats and what eats it. Light energy and car-
bon dioxide flow from the abiotic part of the environment to the living
primary producers (Figure 8).
Summing up all individuals and species, resources of energy and mat-
ter move from the bottom of trophic pyramids to the top. A trophic
pyramid is a representation of all the energy or biomass at each trophic
carbon cycle
atmosphere
light
energy CO2
respiration
photosynthesis
fossil fuels &

cement production
vegetation animals
decomposition
soils rivers
surface ocean
marine biota
deep
dissolved organic ocean
carbon
sediments
Figure 8 The carbon cycle, showing how light energy and carbon
dioxide (CO2) are taken up by plants as the first step of producing
chemical energy that can later flow into higher trophic levels (the
consumers).
Source: Modified from http://earthobservatory.nasa.gov/Library/ CarbonCycle/carbon_cycle4
.html, this file is in the public domain (created by NASA).
Ecological Systems Are Not Very Efficient 43
level. When considering matter, scientists tend to measure the amount of

biomass, or the amount of living matter mass minus the mass of water.
For energy, scientists measure the amount in different food sources, often
in units of calories. When all of the energy is added up, there is a consis-
tent relationship observed across different trophic levels (Figure 9).
Spiders, dragonflies, and raccoons are carnivores, and they could be
secondary, tertiary, or quaternary consumers. Secondary, tertiary, and
quaternary consumers are organisms that get their energy from other
consumers. These are often called carnivores. Secondary consumers eat
primary consumers, tertiary consumers eat secondary consumers, and so
on. Primary consumers, or herbivores, eat plants or other primary pro-
ducers, secondary consumers eat primary consumers, and tertiary con-
sumers eat secondary consumers. Quaternary consumers are usually the
top predators that consume tertiary (and often secondary and primary)
consumers. Going back to our original question, how does light energy
from the sun get incorporated into a dragonflys body? Lets consider a
classic study in ecology by John Teal who studied salt marshes of Georgia
along the southeastern coast of the United States. A salt marsh is part of
an estuary, the waters between oceans and rivers.
Salt marshes are bounded on one side by sea islands and on the other by
the mainland. The salt concentrations of the marsh fluctuate with the amount
of rainfall and the height of the tides. Some species have adapted to these
harsh and dynamic conditions. Salt-tolerant smooth cordgrass makes up most
of grass in the salt marshes. However, there is a gradient of habitat types.
quaternary consumers
carnivores tertiary consumers
secondary consumers
herbivores primary consumers
plants = primary producers
Figure 9 Distribution of biomass or energy at different trophic levels

in an ecological system. Primary consumers are those animals that eat
plants, also known as herbivores. A secondary consumer eats primary
consumers, and a tertiary consumer eats secondary consumers. Secondary
consumers and above are also known as carnivores or predators.
Source: Copyright C. Paradise.
Cordgrass dominates in tidal creek, levee, and low marsh zones and,
thus, is the dominant primary producer. Algae species floating in the
water are also important primary producers, whereas other plants domi-
nate the drier zones.
Teal listed the obvious species occurring in each zone, although he
focused his research on the low saltmarsh. From knowledge of these ani-
mals, plants, and microbes, he constructed a food web.
Teal and his colleagues estimated 34,580 kilocalories/m2/yr of primary
production in the salt marsh, which is how much energy was converted
from carbon dioxide and sunlight into carbohydrates. The actual amount of
plant biomass growth, however, was estimated to be only 6,580 kcal/m2/yr.
Dividing the biomass energy (6,580) by total chemical energy (34,580)
tells us that only 19% of the total chemical energy is retained in the
plant biomass. Teal measured algal production and biomass too. The ef-
ficiency of conversion of primary production to biomass was higher for
algae, although the total amount of production was lower (1,620 (bio-
mass growth) 1,800 (total primary production) kcal/m2/yr = 90%).
The total biomass growth for all primary producers (6,580 + 1,620 =
8,200 kcal/m2/yr) is what is available to primary consumers (herbivores).
Along just one food chain with Teals food web, Teal documented
that smooth cordgrass was consumed by planthoppers and katydids, pri-
marily. The latter two organisms are types of insects. Spiders, wrens and
dragonflies all consumed planthoppers and katydids. Focusing on the
small jumping insects called planthoppers and katydids, a type of grass-
hopper, Teal and his colleagues estimated the annual amount of energy
in the insects consumption, production, respiration, assimilation, and
waste (Figure 10).
After eating, the food is either assimilated into the body via the diges-
tive system, or eliminated as feces. Of the assimilated food, some goes to
growth (production) and some goes to respiration (adenine triphosphate
[ATP] production and CO2 waste). Respiration is sort of the opposite of
photosynthesis; carbon dioxide is a waste product of cellular respiration,
which puts carbon dioxide back into the environment for plants to ab-
sorb again during photosynthesis. Thus, carbon cycles in the environment
(i.e., CO2 biomass CO2). Nutrient cycling is an emergent property
of ecological systems and occurs as organisms grow, consume, and respire
(Figure 11).
Ecological Systems Are Not Very Efficient 45
350
323.5
= planthoppers
300 = katydids
275.0
energy (kcal / m2 / year)
250
205.0
200
150
99.4
100
70.0 70.0
48.5
50
29.4
10.8 18.6
0
consumption assimilation production respiration feces
Figure 10 Energy relationships in saltmarsh planthoppers and

katydids. Estimates are from Teal (1962). He estimated 85%
assimilation efficiency for planthoppers because they feed on liquid
sugar solutions from plants (which are easy to digest), not cellulose
(which is a plant structural chemical that is not easy to digest).
The assimilation efficiency for grasshoppers, which consume mostly
cellulose, was estimated to be 30%.
Source: From Teal, 1962, data in text.
The amount of production of the two herbivorous insect species is

available for consumption by spiders, wrens, and dragonflies. Spiders are
the most abundant predator of these two insects, but Teal lumped all
of the predators together for his analysis. Teal estimated that together
predators consumed 28 out of 80.8 kcal/m2/yr available to them in the
form of planthopper and katydid biomass. The rest of the insect herbivore
biomass (52.8 kcal/m2/yr) ended up as food for bacteria or was washed
out of the saltmarsh when the insects died. The predators converted those
28 kcal into only 5 kcal/m2/yr of biomass production.
As Teal no doubt realized, ecological systems are not very efficient. Most
of the energy captured by primary producers is lost through respiration. No
organisms can use all of the energy they consume to grow bigger; each organ-
ism must use energy to maintain basic metabolic functionsto move or to
reproduce. Much of the chemical energy used for metabolism or reproduc-
tion is lost to as heat, which is waste energy. Because of this waste, the amount
of energy or biomass in predator trophic levels will always be smaller than the
amount of energy or biomass of the producer trophic level. After using the
CO2 in respiration
CO2 in atmosphere respiration heat energy
smooth
cordgrass
production = planthoppers spiders

6,580 kcal/m2/yr katydids wrens
dragonflies
feces
death feces
death
respiration =
28,080 kcal/m2/yr
bacteria & other

decomposers
other consumers
that eat bacteria
Figure 11 One pathway of energy and carbon in the Georgia

saltmarsh. Carbon cycles (atmosphere plants herbivores
carnivores atmosphere), and each component sends carbon dioxide
back to the atmosphere. Decomposers also respire or provide biomass
to predators. The pathway of carbon is combined with energy flow
(sunlight plants herbivores carnivores) with respiration
energy used for metabolism and lost as heat. The size of the boxes for
cordgrass, insects, herbivores, and predators is proportional to the
energy content in each trophic level.
Source: From Teal, 1962, Figure 4 and data in text.
energy in carbohydrates, the carbon is released as carbon dioxide into the

atmosphere, where it is available for plants to consume again in a continu-
ous cycle. The heat energy is no longer available for organisms to use as a
source of energy for creating biomass, and so this energy is biologically out of
circulation. This chapter has revealed how energy flows and nutrients cycle
through ecological systems and how their movement are emergent properties
that arise from the species that interact within an ecological system.
Bibliography
Teal JM: Energy flow in the salt marsh ecosystem of Georgia, Ecology 43(4):
614624, 1962.
Conclusion
Emergent properties at any level of biological hierarchy cannot be studied
in isolation. The other fundamental ideas of biology (such as, evolution,
cells as functional units, information, and homeostasis) all contribute
to emergent properties. For instance, any interaction between two spe-
cies may result in selective pressures and possible extinction, which is an
emergent property. Information is used by individuals to assess the envi-
ronment while foraging or maintaining homeostasis. Indirect effects arise
from biotic interactions; the wolfelk predatorprey interaction led to
effects on trees and several other animal species in Yellowstone National
Park. Cooperation is an emergent property exhibited by packs of wolves
because they compete in an ecological system. Species adapt to local envi-
ronmental conditions, which allows species to coexist even when they are
competing for similar resources. This leads to increased species diversity
within an ecological system. Finally, nutrients cycle and energy flows in
ecological systems, and those movements of matter and energy arise from
interactions between species because herbivores eat plants and carnivores
eat herbivores.
Glossary
adaptations. A change or the process of change by which a species becomes better
suited to its environment via the mechanism of natural selection.
assimilation. The process of absorbing nutrients and incorporating them into the
body.
biomass. The total amount of organic matter in an organism, population, or
other ecological system after water is removed.
calories. A calorie is the amount of energy, or heat, it takes to raise the tempera-
ture of 1 gram of water 1C, or 4.184 joules.
carbon dioxide. A colorless, odorless gas composed of one carbon atom bonded
to two oxygen atoms and produced by burning carbon and organic compounds
and by respiration.
carnivores. Carnivores are animals that eat animals.
competition. The interaction that results from the demand by two or more or-
ganisms for limited environmental resources.
competitive ability. Possession of the qualities required to compete well for
resources.
competitive exclusion principle. The competitive exclusion principle states that
no two species that consume the exact same set of resources can coexist.
consumption. The act of eating or drinking resources.
cooperation. The act of working together for mutual benefit.
cooperative behavior. Cooperative behavior is behavior that involves several indi-
viduals and is mutually beneficial.
disturbances. Temporary changes in environmental conditions that cause pro-
nounced changes in an ecosystem.
ecological system. An ecological community together with the abiotic environ-
ment, usually considered to function as a unit.
energy. A fundamental aspect of nature that is transferred between parts of a
system in the production of physical or chemical change within the system and
usually regarded as the capacity for doing work.
energy flows. Energy flow refers to the movement of energy-containing chemi-
cals from one organism to another.
environmental gradients. The gradual changes in environmental factors from
one place to another.
evolved. To be produced by natural evolutionary processes or mechanisms, such
as natural selection, genetic drift, or gene flow.
extinction. The permanent loss of a species from the planet when no individuals
of that species are left living.
50 GLOSSARY
food chain. A series of feeding links showing who eats whom in a direct line from
one species that is eaten by a second that is eaten by a third.
food webs. Food webs are diagrams that show who eats whom and how energy
flows in an ecological system.
homeostasis. Maintain internal conditions within a range of acceptable extremes.
indirect effects. Indirect effects in ecological systems are effects of one species on
other species mediated through shared interactions with a third species or group
of species.
interaction strengths. The impact of one species on the biomass or abundance of
another species, usually in a consumer-resource relationship.
intertidal. The area of the shore between the low point and the high point of the
tide.
limiting resource. A limiting resource is a resource (such as, food, light, nutrients,
or space), which is in short supply and restricts the growth of an organism or
population.
nitrates. Ions of NO32 that have a negative one charge.
nutrients. Food or chemicals that organisms obtain from their environment and
need to live and grow.
phosphates. Ions of PO432 derived from phosphoric acid H3PO4.
population. A population is a group of individuals of the same species living in
the same place at the same time.
predators. Organisms that kill and feed on other organisms.
prey. Prey are organisms that are consumed by predators, either in whole or part.
primary consumers. Primary consumers are organisms that get their energy from
primary producers (plants), also known as herbivores.
primary production. The production of organic compounds from carbon dioxide.
quaternary consumers. Top predators that consume tertiary consumers.
reproductive strategy. A reproductive strategy is a suite of evolved life cycle-
related traits that taken together lead to successful existence of a species in the
context of that species environment.
resource use overlap. A measure of the resources shared between any pair of
species.
resources. Natural features or organisms used as a supply of energy or nutrients
to other organisms.
respiration. Cellular respiration is a process in organisms involving the trans-
duction of energy, typically with the intake of oxygen and the release of carbon
dioxide from the oxidation of complex organic substances.
secondary consumers. Predators that eat primary consumers or herbivores.
tertiary consumers. Predators that consume secondary consumers, which are also
predators.
trade-off. A trade-off is a compromise that occurs as organisms exchange one
behavior or resource for another.
GLOSSARY
51
trophic cascade. It occurs when a change in the size of one population has an
effect on populations in other trophic levels.
trophic level. A trophic level is a feeding position in a food web, which describes
what an organism eats and what eats it.
trophic pyramids. Representations of all the energy or biomass at each trophic
level.
Index
Aspen trees, growth, 1113 Environmental gradients, 21, 22
Environmental laws, 15
Beschta, Robert, 57, 11, 12 Extinction of species, 3, 15
Biomass
distribution, 43 Food webs, 1, 2
of species, 23
Global climate change, 13
Calories, definition of, 32
Carbon cycle, 42, 44 Heat energy, 46
Carnivores. See Secondary consumers Herbivores. See Primary consumers
Common myna, use of nest box, Homeostasis, 31, 36, 39, 40
2629
Competitive ability, 22 Indirect effects, of
Competitive exclusion principle, 20 predation, 8
Cooperative behavior, 19 Insects
Cottonwood assimilation, 44
growth, 7, 8, 11 respiration, 44, 45
seedlings, 57, 10 Interaction strengths, 3639
seed production, 5
Coyotes scavenge, 12 Keddy, Paul, 2225
Creel, Scott, 89
Limiting resource, 30
Ecological systems, 2, 8, 1215, 31,
32, 34, 36, 39, 40 Natural habitat, 15
energy flows in, 1, 2 Natural regulation, 4, 8
predators in, 1 Nest box occupancy, 2629
prey in, 1 Nucella feeding, 32
social behavior, 3 Nutrients, 1
Elk growth of, 26
behavior, 9, 1113
cottonwood interaction, 78 Paine, Robert, 3140
food consumption, 13 Pisaster
habitat use, 10 feeding, 32
population, 3, 4, 7, 8 food web, 3239
wolf interactions, 78 Plant biomass growth, 44
Endangered Species Act (ESA), 1516 Predator-prey interaction
Energy flows through food strengths,3638
webs,3140 Prey consumption in diet, 33
Energy relationships, in saltmarsh Prey population, 3, 13, 15
planthoppers and Primary consumers, 43
katydids,45 Primary production, defined, 42
54 INDEX
Radiotracking, 8 Tree densities, 2629

elk locations, 9 Trophic cascade, 40
Relative increase per plant Trophic level, 42
(RIP),2324 Trophic pyramids, 42
Reproductive strategy, of species, 5
Resource use overlap, 19 US Department of Interior, 16
Ripple, William, 11, 12
Rocky intertidal food web, 31, 33, Vegetation diversity, in Yellowstone
34, 37 National Park, 117
Secondary consumers, 43 Wilson, Scott, 2225

Species present of Pisaster, 35 Wolf reintroductions, 1416
Strong interactions, definition of, 38 Wolves
hunting, 1, 35, 8, 12, 16
Teal, John, 4345 population, 4, 5
Trade-off, 13 predation, 3, 8, 9, 11
Transferring energy, from sun
and carbon dioxide to
predators,4146
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Food webs, energy flow, indirect effects, and nutrient cycling
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are described as properties that emerge in ecological systems.
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THE CONTENT this book illustrates how energy flows in ecological systems,
Energy Physics why it is rather inefficient, and how species interactions relate
Engineering to homeostasis and emergent properties. In the course of that
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Christopher J. Paradise, PhD

All rights reserved. No part of this publication may be reproduced, stored

First published in 2016 by

ISBN-13: 978-1-60650-955-5 (print)

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Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

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Emergent properties of ecological systems can be examined by looking

forests at higher elevations. Deciduous trees in this part of the world

Figure 2 Elk and wolf population counts in Yellowstone National

populations had been monitored for decades within Yellowstone National

naturally controlling elk populations. As of 2012, about 80 to 90 wolves

of Yellowstone National Park in the years following wolf reintroduc-

Figure 3 Radiotracked elk locations and elk kills in Gallatin Canyon

0.8 = wolves not detected

Figure 4 Effects of wolf presence on habitat use by elk. The

It is uncommon for a policy change to be measured so carefully to de-

Ethical, Legal, Social Implications: There are

overpopulation and overconsumption of resources. Many people argue

within the US Department of Interior are charged with developing recov-

how do they strike a balance? According to the competitive exclusion

Ecological systems change across space, whereas conditions related to an

(final biomass of i with j) (initial biomass of i)

be expected that plants would evolve to be less competitive and adapt to

bird density (#/km2 1SE)

Figure 5 Responses of three cavity-nesting bird species to forests of

crimson rosella abundance (#/km2)

Figure 6 Relationships between the percentage of nest boxes occupied

Changes in an environmental condition (in this case, forest tree den-

Energy Flows Through

In Chapter 1, the effects of reintroduction of a top predator, the gray

barnacles, snails, and one starfish. Paine observed communities in this

Table 1 Percentages of prey consumed (before the slash) and

and calories is presented in Table 1. If all the numbers are added up to

Table 2 Species present in the presence or absence of Pisaster.

the scientists then standardized to per capita effects of predators on prey

Figure 7 Frequency distribution of non-zero predator-prey interaction

Ecological Systems Are

competitors, parasites, and so on. When an elk chews on a cottonwood

fossil fuels &

level. When considering matter, scientists tend to measure the amount of

carnivores tertiary consumers

herbivores primary consumers

plants = primary producers

Figure 9 Distribution of biomass or energy at different trophic levels

Figure 10 Energy relationships in saltmarsh planthoppers and

The amount of production of the two herbivorous insect species is

production = planthoppers spiders

bacteria & other

Figure 11 One pathway of energy and carbon in the Georgia

energy in carbohydrates, the carbon is released as carbon dioxide into the

Radiotracking, 8 Tree densities, 2629

Secondary consumers, 43 Wilson, Scott, 2225

Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

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