Professional Documents
Culture Documents
Ecological
Bundlethe more
ing plants and non-prey animals. In some other interactions,
books you buy,
including competition, the nonliving components of ecological
the greater your
Interactions
systems (the abiota) can alter the outcome of a biotic interac-
discount!
tion. A limiting resource often results in competition, but varying
environmental conditions allow for species coexistence. Finally,
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
discussion, primary production, secondary production, and tro-
Biotechnology
phic levels are defined. Energy flow in ecological systems is tied
Biology
to the carbon cycle.
Mathematics
Chemistry Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS
cology and renewable natural resources. He also occasionally
Perpetual access
leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
Unlimited
A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
Downloadable PDFs
Genetics Society of America (2013); American Association for the
Free MARC records
Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell
Ecological Interactions
Ecological Interactions
10 9 8 7 6 5 4 3 2 1
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
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
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
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
8 ECOLOGICAL INTERACTIONS
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
WOLF REINTRODUCTION INCREASED VEGETATION DIVERSITY 9
60
= forest
50 = edge
= grass
percent of observations
40
30
20
10
0
kills wolves wolves not
detected detected
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,
10 ECOLOGICAL INTERACTIONS
0.6
0.4
0.2
0
elk presence in grass elk presence in forest
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.
WOLF REINTRODUCTION INCREASED VEGETATION DIVERSITY 11
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.
12 ECOLOGICAL INTERACTIONS
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.
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
16 ECOLOGICAL INTERACTIONS
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
Arjo WM, Pletscher DH, Ream RR: Dietary overlap between wolves and
coyotes in northwestern Montana, J Mammal 83(3):754766, 2002.
Bangs EE, Fritts SH: Reintroducing the gray wolf to central Idaho and
Yellowstone National Park, Wildlife Soc B 24:402413, 1996.
Beschta RL: Cottonwoods, elk, and wolves in the Lamar Valley of Yellow-
stone National Park, Ecol Appl 13(5):12951309, 2003.
Beschta RL: Reduced cottonwood recruitment following extirpation of
wolves in Yellowstones Northern Range, Ecology 86(2):391403, 2005.
Beschta RL, Ripple WJ: Berry-producing shrub characteristics following
wolf reintroduction in Yellowstone National Park, Forest Ecol Manag
276:132138, 2012.
Carroll N, Dunning JB Jr, Freeman A, et al.: Reintroduction of the gray
wolf to Yellowstone National Park: a case study, Presented at the May
1997 Purdue Bioethics Workshop: http://www.bioethics.iastate.edu/
classroom/graywolf.html. Accessed March 30, 2008.
Creel S, Winnie J Jr., Maxwell B, et al.: Elk alter habitat selection as an
antipredator response to wolves, Ecology 86(12):33873397, 2005.
Pearson B: The recall of the wild, TPM (The Philosphers Magazine) (website):
http://www.philosophersnet.com/magazine/article.php?id=992.
Accessed March 30, 2008.
Ripple WJ, Beschta RL: Wolves and the ecology of fear: can predation
risk structure ecosystems? BioScience 54(8):755766, 2004.
Ripple WJ, Beschta RL: Restoring Yellowstones aspen with wolves, Biol
Cons 138:514519, 2007.
Robbins J: Lessons from the wolf, Sci Am June:7681, 2004.
Smith DW, Stahler DR, Guernsey DS: Yellowstone wolf project: Annual
report, 2005. National Park Service, Yellowstone Center for Resources,
Yellowstone National Park, Wyoming, YCR-200604, 2006.
18 ECOLOGICAL INTERACTIONS
White PJ, Garrott RA: Northern Yellowstone elk after wolf restoration,
Wildlife Soc B 33(3):942955, 2005.
Williams CK, Ericsson G, Heberlein TA: A quantitative summary of atti-
tudes toward wolves and their reintroduction (19722000), Wildlife
Soc B 30(2):575584, 2002.
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
20 ECOLOGICAL INTERACTIONS
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).
CONDITIONS CAN DETERMINE THE OUTCOME OF COMPETITION 23
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.
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
24 ECOLOGICAL INTERACTIONS
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,
CONDITIONS CAN DETERMINE THE OUTCOME OF COMPETITION 25
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
26 ECOLOGICAL INTERACTIONS
140
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
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.
CONDITIONS CAN DETERMINE THE OUTCOME OF COMPETITION 29
160
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 (%)
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.
30 ECOLOGICAL INTERACTIONS
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
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).
Energy Flows Through FoodWebs 35
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
36 ECOLOGICAL INTERACTIONS
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
Energy Flows Through FoodWebs 37
1000
800
600
frequency
400
200
0
7 6 5 4 3 2 1 0 1
log of interaction strength
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
Energy Flows Through FoodWebs 39
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,
40 ECOLOGICAL INTERACTIONS
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
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,
42 ECOLOGICAL INTERACTIONS
carbon cycle
atmosphere
light
energy CO2
respiration
photosynthesis
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
quaternary consumers
secondary consumers
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
CO2 in respiration
CO2 in atmosphere respiration heat energy
smooth
cordgrass
feces
death feces
death
respiration =
28,080 kcal/m2/yr
other consumers
that eat bacteria
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
The Momentum Press digital library is very affordable, with no obligation to buy in future
years.
For more information, please visit www.momentumpress.net/library or to set up a trial in the
US, please contact mpsales@globalepress.com.
EBOOKS Ecological Interactions
FOR THE Christopher J. Paradise A. Malcolm Campbell
APPLIED BIOLOGY COLLECTION
Food webs, energy flow, indirect effects, and nutrient cycling
SCIENCES
are described as properties that emerge in ecological systems.
LIBRARY Several of these properties are shown in this book to result from
Create your own indirect effects and interactions between species and abiotic
Customized Content components of ecological systems. For instance, top predators
affect organisms with which they do not directly interact, includ-
Ecological
Bundlethe more
ing plants and non-prey animals. In some other interactions,
books you buy,
including competition, the nonliving components of ecological
the greater your
Interactions
systems (the abiota) can alter the outcome of a biotic interac-
discount!
tion. A limiting resource often results in competition, but varying
environmental conditions allow for species coexistence. Finally,
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
discussion, primary production, secondary production, and tro-
Biotechnology
phic levels are defined. Energy flow in ecological systems is tied
Biology
to the carbon cycle.
Mathematics
Chemistry Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS
cology and renewable natural resources. He also occasionally
Perpetual access
leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
Unlimited
A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
Downloadable PDFs
Genetics Society of America (2013); American Association for the
Free MARC records
Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell