REPUBLIC OF THE PHILIPPINES

BATANGAS STATE UNIVERSITY

COLLEGE OF ENGINEERING, ARCHITECTURE AND FINE ARTS

ELECTRONICS AND COMMUNICATIONS ENGINEERING DEPARTMENT

ENG-413 ENVIRONMENTAL SCIENCE

AND ENGINEERING

GROUP NUMBER 2

EVOLUTION, BIOLOGICAL COMMUNITIES, AND SPECIES INTERACTIONS

SUBMITTED BY:

NUGUIT, RAPHAEL

CAPOYAN, JOHN CARLO

LAJARA, NATHANIEL

RAMOS, LIZETTE ANGELA

ECE-2103

SUBMITTED TO:

MR. RICHARD AMODAL

SEPTEMBER 2021
EVOLUTION

Evolution in its contemporary meaning in biology typically refers to the changes

in the proportions of biological types in a population over time (see the entries on
evolutionary thought before Darwin and Darwin: from Origin of Species to Descent of
Man for earlier meanings). As evolution is too large of a topic to address thoroughly in
one entry, the primary goal of this entry is to serve as a broad overview of contemporary
issues in evolution with links to other entries where more in-depth discussion can be
found. The entry begins with a brief survey of definitions of evolution, followed by a
discussion of the different modes of evolution and related philosophical issues, and
ends with a summary of other topics in the philosophy of evolution focusing particularly
on topics covered in this encyclopedia.

Although the work of Charles Darwin (see the entry on Darwinism) is usually the
starting point for contemporary understandings of evolution, interestingly, he does not
use the term in the first edition of On the Origin of Species, referring instead to “descent
with modification”. In the early-mid 20th century, the “modern synthesis” gave birth to
population genetics, which provided a mathematization of Darwinian evolutionary theory
in light of Mendelian genetics (see also the entry on evolutionary genetics). This yielded
a prevalent—probably the most prevalent—understanding of evolution as “any change
in the frequency of alleles within a population from one generation to the next”. Note,
however, that this definition refers to evolution only in a microevolutionary context and
thus doesn’t reference the emergence of new species (and their new characteristics),
although it is intended to underlie those macroevolutionary changes.

In a popular textbook, Douglas Futuyma gives a more expansive definition:

[biological evolution] is change in the properties of groups of organisms over the course
of generations…it embraces everything from slight changes in the proportions of
different forms of a gene within a population to the alterations that led from the earliest
organism to dinosaurs, bees, oaks, and humans. (2005: 2)
Note also that Futuyma’s definition, unlike the population genetics’ definition, does not
limit itself to changes in alleles; John Endler’s definition is similar in this respect:

Evolution may be defined as any net directional change or any cumulative

change in the characteristics of organisms or populations over many generations—in
other words, descent with modification… It explicitly includes the origin as well as the
spread of alleles, variants, trait values, or character states. (Endler 1986: 5)

Yet even this definition is not expansive enough; molecular evolution focuses on the
molecular changes within macromolecules such as DNA and RNA.

In a very different vein, Leigh van Valen characterized evolution as “the control of
development by ecology” (1973, 488); this anticipates those who emphasize the
importance of development in evolution, including proponents of “evo-devo” (see the
entry on developmental biology). Today, some have called for an “extended evolutionary
synthesis” in light of developmental biology and other recent findings in evolutionary
biology.

Although this entry focuses on biological evolution, philosophers and biologists have
also sought to extend evolutionary ideas to the cultural realm. Figuring out how and
whether to extend the definition of evolution to this realm is part of the study of cultural
evolution.

In spite of this diversity of definitions, there has been very little philosophical analysis of
the term “evolution” itself. This dearth forms a stark contrast to the voluminous literature
in the philosophy of evolution; indeed, for a long time the philosophy of biology was
focused almost entirely on evolution. Thankfully, that is no longer the case, with
philosophers turning their attentions to issues in genetics, molecular biology, ecology,
developmental biology, and more. It may be, as Theodosius Dobzhansky famously said,
that “Nothing in biology makes sense except in the light of evolution” (1973: 125), but
much of biology is not evolutionary biology. Still, though, philosophy of evolution
remains a growing and vibrant area within the philosophy of biology.

2. Modes of Evolution
It is essential to understand that biologists recognize many ways that evolution can
occur, evolution by natural selection being just one of them, although it is often held to
be the most prevalent one. Evolution can also occur through genetic drift, mutation, or
migration. It can also occur through sexual selection, which some consider to be a form
of natural selection and others consider to be distinct from natural selection (the latter
having been Darwin’s 1859, 1874 view). Evolutionary theory, then, can be taken to be
the study (including, but not limited to, mathematical models) of these and other modes
of evolution.

To see why it makes sense to think of multiple modes of evolution, consider again one
of the definitions of evolution presented above, where evolution is understood as “any
change in the frequency of alleles within a population from one generation to the next”.
With natural selection, the frequency of alleles that confer greater fitness would tend to
increase over those which confer lesser fitness. Sexual selection would be the same,
but with fitness understood strictly in terms of mating ability. With genetic drift, a form of
evolution that involves chance (see the entry on genetic drift for explanation), there
could be an increase in the frequency of alleles that confer greater fitness, an increase
in the frequency of alleles that confer lesser fitness, or an increase in the frequency of
alleles whose manifestation (if any) was neutral. If organisms migrate from one
population to another, it is likely that there will be a change in the frequency of alleles in
both populations. And if there is a mutation from one allele to another, then the
frequency of alleles in the population will likewise change, albeit by a small amount.
Distinguishing these different modes of evolution allows biologists to track the various
factors that are relevant to evolutionary changes in a population.
The careful reader may have noted that the previous paragraph invoked probabilistic
language: what tends to happen, what could happen, what is likely to happen. Indeed,
mathematical evolutionary models today (see the entry on population genetics) are
typically statistical models. This fact about evolutionary models has given rise to a
debate in the philosophy of evolution over whether natural selection and genetic drift
should be be understood as causes of evolution, as most biologists conceive them, or
as mere statistical summaries of lower-level causes: births, deaths, etc. (The natural
selection and genetic drift entries give more information about this debate). It is for this
reason that this entry uses the more neutral phrase “modes of evolution” so as not to
beg any questions under dispute between the causalist and the statisticalist.

Although there is widespread agreement that there are multiple modes of evolution,
much contemporary work in biology and philosophy of biology has been focused on
natural selection. Whether this focus is a good thing or not is in part what the debate
over adaptationism is about. That is, do we have reason to think that natural selection is
the most prevalent or most important mode of evolution? Should scientific
methodologies be geared toward testing natural selection hypotheses or toward a
variety of possible evolutionary modes? The focus on natural selection has also led to a
large literature on the concept of fitness, given that population genetics’ definitions and
other definitions of natural selection typically invoke fitness; a natural selection
explanation of why X was more successful than Y might invoke X’s higher fitness. What
fitness means, what entities it applies to (genes, organisms, groups, individuals, types),
what sort of probabilities it invokes, if any, and how it should be calculated, are all under
philosophical dispute. There is also a large literature on conceptually and empirically
distinguishing natural selection from genetic drift. Migration, mutation (as a mode of
evolution), and sexual selection have received less attention from philosophers of
biology.
Adaptation

Evolutionary adaptation, or simply adaptation, is the adjustment of organisms to

their environment in order to improve their chances at survival in that environment. In
evolutionary theory, adaptation is the biological mechanism by which organisms adjust
to new environments or to changes in their current environment. Although scientists
discussed adaptation prior to the 1800s, it was not until then that Charles Darwin and
Alfred Russel Wallace developed the theory of natural selection.

Organisms can adapt to an environment in different ways. They can adapt

biologically, meaning they alter body functions. An example of biological adaptation can
be seen in the bodies of people living at high altitudes, such as Tibet. Tibetans thrive at
altitudes where oxygen levels are up to 40 percent lower than at sea level. Breathing air
that thin would cause most people to get sick, but Tibetans’ bodies have evolved
changes in their body chemistry. Most people can survive at high altitudes for a short
time because their bodies raise their levels of hemoglobin, a protein that transports
oxygen in the blood. However, continuously high levels of hemoglobin are dangerous,
so increased hemoglobin levels are not a good solution to high-altitude survival in the
long term. Tibetans seemed to have evolved genetic mutations that allow them to use
oxygen far more efficently without the need for extra hemoglobin.

Organisms can also exhibit behavioral adaptation. One example of behavioral

adaptation is how emperor penguins in Antarctica crowd together to share their warmth
in the middle of winter.

Scientists who studied adaptation prior to the development of evolutionary theory

included Georges Louis Leclerc Comte de Buffon. He was a French mathematician who
believed that organisms changed over time by adapting to the environments of their
geographical locations. Another French thinker, Jean Baptiste Lamarck, proposed that
animals could adapt, pass on their adaptations to their offspring, and therefore evolve.
The example he gave stated the ancestors of giraffes might have adapted to a shortage
of food from short trees by stretching their necks to reach higher branches. In Lamarck’s
thinking, the offspring of a giraffe that stretched its neck would then inherit a slightly
longer neck. Lamarck theorized that behaviors aquired in a giraffe's lifetime would affect
its offspring. However, it was Darwin’s concept of natural selection, wherein favorable
traits like a long neck in giraffes suvived not because of aquired skills, but because only
giraffes that had long enough necks to feed themselves survived long enough to
reproduce. Natural selection, then, provides a more compelling mechanism for
adaptation and evolution than Lamarck's theories.

Some creatures, such as this leafy sea dragon fish

(Phycodurus eques) have evolved adaptations that allow them
to blend in with their environment (in this case, seaweed) to
avoid the attention of hungry predators.

What Is Acclimation?

One big problem in studying ecology is that environments rarely, if ever, stay the same.
Temperatures, precipitation, and food sources are always changing. One way that living
things can handle these changes is through a process called acclimation.

Acclimation is a slow, reversible change to the body that allows an organism to handle
a different environment. This change can occur over a few days, several weeks, or even
months.

Acclimation and Adaptation

Acclimation may seem like a similar concept to adaptation which is a feature acquired
in a population that helps them live in a new environment. Adaptations can take
generations to develop and happen to the population. Acclimation, on the other hand, is
something that happens to an individual to allow it to survive when the environment
changes.
Examples of Acclimation

Salmon

Salmon move from place to place to live out their life cycle. They begin their lives in
freshwater streams, then move to saltwater oceans. As the salmon enter water that
becomes saltier, they need to find a way to stay hydrated.

They have a very simple solution: they drink. This helps get their kidneys and gills
working to get rid of the salt. Young salmon will stay at the edge between freshwater
and saltwater for a few days or weeks in order to acclimate gradually.

When salmon near the end of their life cycle, they return to the freshwater streams to
spawn, or reproduce. They'll again spend a few days to weeks in less and less salty
water to become acclimated to their new environment.

Most fish need to live in either a saltwater or a freshwater environment. However,

salmon are able to acclimate to the changing salt concentrations in the water as they
migrate. This is reversible, because they can go from freshwater to salt water and back
again. It is also gradual, because the change to the salmon doesn't happen instantly.

Listeria

While it's easy to think of big things, like animals, having to acclimate to their
environments, small organisms need to do this too. Bacteria, for example, have to
acclimate when they enter new environments.

Listeria monocytogenes is a bacterium that causes food poisoning in humans. It grows

happily at our body temperature, infecting cells and making us sick. But one reason
Listeria is able to make us sick is its ability to acclimate to cold temperatures.

When put into a refrigerator, instead of laying dormant like other bacteria species,
Listeria will turn on a number of genes that allow it to continue growing and multiplying
at lower temperatures. Listeria can also change the specific fatty acids in its membrane
to prevent it from solidifying at low temperatures.
Once Listeria leaves the refrigerator and is eaten, it's suddenly in a warm, but acidic,
environment. In order to survive the acid in your stomach, Listeria can acclimate and
turn on its acid tolerance response.

English naturalist Charles Darwin developed the idea of natural selection after a
five-year voyage to study plants, animals, and fossils in South America and on islands
in the Pacific. In 1859, he brought the idea of natural selection to the attention of the
world in his best-selling book, On the Origin of Species.

Natural selection is the process through which populations of living organisms

adapt and change. Individuals in a population are naturally variable, meaning that they
are all different in some ways. This variation means that some individuals have traits
better suited to the environment than others. Individuals with adaptive traits—traits that
give them some advantage—are more likely to survive and reproduce. These
individuals then pass the adaptive traits on to their offspring. Over time, these
advantageous traits become more common in the population. Through this process of
natural selection, favorable traits are transmitted through generations.

Natural selection can lead to speciation, where one species gives rise to a new and
distinctly different species. It is one of the processes that drives evolution and helps to
explain the diversity of life on Earth.

Darwin chose the name natural selection to contrast with “artificial selection,” or
selective breeding that is controlled by humans. He pointed to the pastime of pigeon
breeding, a popular hobby in his day, as an example of artificial selection. By choosing
which pigeons mated with others, hobbyists created distinct pigeon breeds, with fancy
feathers or acrobatic flight, that were different from wild pigeons.

Darwin and other scientists of his day argued that a process much like artificial selection
happened in nature, without any human intervention. He argued that natural selection
explained how a wide variety of life forms developed over time from a single common
ancestor.
Darwin did not know that genes existed, but he could see that many traits are
heritable—passed from parents to offspring.

Mutations are changes in the structure of the molecules that make up genes, called
DNA. The mutation of genes is an important source of genetic variation within a
population. Mutations can be random (for example, when replicating cells make an error
while copying DNA), or happen as a result of exposure to something in the environment,
like harmful chemicals or radiation.

Mutations can be harmful, neutral, or sometimes helpful, resulting in a new,

advantageous trait. When mutations occur in germ cells (eggs and sperm), they can be
passed onto offspring.

If the environment changes rapidly, some species may not be able to adapt fast enough
through natural selection. Through studying the fossil record, we know that many of the
organisms that once lived on Earth are now extinct. Dinosaurs are one example. An
invasive species, a disease organism, a catastrophic environmental change, or a highly
successful predator can all contribute to the extinction of species.

Today, human actions such as overhunting and the destruction of habitats are the main
cause of extinctions. Extinctions seem to be occurring at a much faster rate today than
they did in the past, as shown in the fossil record.
Mutations Are the Raw Materials of Evolution
By: Joel L. Carlin (Department of Biology, Gustavus Adolphus) © 2011 Nature
Education

A mutation is a change in the sequence of an organism's DNA. What causes a

mutation? Mutations can be caused by high-energy sources such as radiation or by
chemicals in the environment. They can also appear spontaneously during the
replication of DNA.

Mutations generally fall into two types: point mutations and chromosomal aberrations. In
point mutations, one base pair is changed. The human genome, for example, contains
over 3.1 billion bases of DNA, and each base must be faithfully replicated for cell
division to occur. Mistakes, although surprisingly rare, do happen. About one in every
1010 (10,000,000,000) base pair is changed. The most common type of mistake is a
point substitution. More uncommon is the failure to copy one of the bases (deletion), the
making of two copies for a single base (point duplication) or the addition of a new base
or even several bases (insertion). Chromosomal aberrations are larger-scale mutations
that can occur during meiosis in unequal crossing over events, slippage during DNA
recombination or due to the activities of transposable events. Genes and even whole
chromosomes can be substituted, duplicated, or deleted due to these errors (Figure 1).

Mutations can have a range of effects. They can often be harmful. Others have little or
no detrimental effect. And sometimes, although very rarely, the change in DNA
sequence may even turn out to be beneficial to the organism.

A mutation that occurs in body cells that are not passed along to subsequent
generations is a somatic mutation. A mutation that occurs in a gamete or in a cell that
gives rise to gametes are special because they impact the next generation and may not
affect the adult at all. Such changes are called germ-line mutations because they occur
in a cell used in reproduction (germ cell), giving the change a chance to become more
numerous over time. If the mutation has a deleterious affect on the phenotype of the
offspring, the mutation is referred to as a genetic disorder. Alternately, if the mutation
has a positive affect on the fitness of the offspring, it is called an adaptation. Thus, all
mutations that affect the fitness of future generations are agents of evolution.

Mutations are essential to evolution. Every genetic feature in every organism was,
initially, the result of a mutation. The new genetic variant (allele) spreads via
reproduction, and differential reproduction is a defining aspect of evolution. It is easy to
understand how a mutation that allows an organism to feed, grow or reproduce more
effectively could cause the mutant allele to become more abundant over time. Soon the
population may be quite ecologically and/or physiologically different from the original
population that lacked the adaptation. Even deleterious mutations can cause
evolutionary change, especially in small populations, by removing individuals that might
be carrying adaptive alleles at other genes.

Most mutations occur at single points in a gene, changing perhaps a single protein, and
thus could appear unimportant. For instance, genes control the structure and
effectiveness of digestive enzymes in your (and all other vertebrate) salivary glands. At
first glance, mutations to salivary enzymes might appear to have little potential for
impacting survival. Yet it is precisely the accumulation of slight mutations to saliva that is
responsible for snake venom and therefore much of snake evolution. Natural selection
in some ancestral snakes has favored enzymes with increasingly more aggressive
properties, but the mutations themselves have been random, creating different venoms
in different groups of snakes. Snake venoms are actually a cocktail of different proteins
with different effects, so genetically related species have a different mixture from other
venomous snake families. The ancestors of sea snakes, coral snakes, and cobras
(family Elapidae) evolved venom that attacks the nervous system while the venom of
vipers (family Viperidae; including rattlesnakes and the bushmaster) acts upon the
cardiovascular system. Both families have many different species that inherited a slight
advantage in venom power from their ancestors, and as mutations accumulate the
diversity of venoms and diversity of species increased over time.

Although the history of many species have been affected by the gradual accumulation
of tiny point mutations, sometimes evolution works much more quickly. Several types of
organisms have an ancestor that failed to undergo meiosis correctly prior to sexual
reproduction, resulting in a total duplication of every chromosome pair. Such a process
created an "instant speciation" event in the gray treefrog of North America

The consequence of doubling the genome size in plants is often abnormally large seeds
or fruits, a trait that can be of distinct advantage if you are a flowering plant! Most
cereals that humans eat have enormous seeds compared to other grasses, and this is
often due to the genomic duplications that occurred in the ancestors of modern rice and
wheat and, because the mistake occurred in reproductive organs, was successfully
passed on to future generations. Humans themselves have mimicked this process by
interbreeding individual plants with the largest fruits and seeds in the process of artificial
selection, creating many of our modern agricultural crop strains. The idea of evolution
by natural selection, first described by Charles Darwin and Alfred Russell Wallace,
requires differential survival due to some individuals having greater evolutionary fitness.
Whether that fitness is affected by genetic disorders, venomous saliva or enlarged
offspring, heritable variation can only arise by mutation. Evolution is simply not possible
without random genetic change for its raw material.

LIMITING FACTORS

A limiting factor is anything that constrains a population's size and slows or stops
it from growing. Some examples of limiting factors are biotic, like food, mates, and
competition with other organisms for resources. Others are abiotic, like space,
temperature, altitude, and amount of sunlight available in an environment. Limiting
factors are usually expressed as a lack of a particular resource. For example, if there
are not enough prey animals in a forest to feed a large population of predators, then
food becomes a limiting factor. Likewise, if there is not enough space in a pond for a
large number of fish, then space becomes a limiting factor. There can be many different
limiting factors at work in a single habitat, and the same limiting factors can affect the
populations of both plant and animal species. Ultimately, limiting factors determine a
habitat's carrying capacity, which is the maximum size of the population it can support.

Liebig’s Law of Minimum

Definition

In the 19th century, the German scientist Justus von Liebig

formulated the “Law of the Minimum,” which states that if
one of the essential plant nutrients is deficient, plant growth
will be poor even when all other essential nutrients are
abundant.

How It Works

It states that growth is controlled not by the total originality applied to plant growth,
where it was found that increasing the amount of plentiful nutrients did not increase
plant growth. Only by increasing the amount of the limiting nutrient (the most scarce) in
relation to “need”, was the growth of the plant improved.

Importance of Micronutrients

The Law of the Minimum takes on added importance when fertilizer prices — especially
of nitrogen (N) and phosphate (P2O5) products — are high. This may tempt some
growers to reduce or even eliminate applications of micronutrient or secondary nutrient
fertilizers that provide balanced potassium (K), magnesium (Mg) and sulfur (S). But von
Liebig’s “Law” tells us clearly that if a soil is deficient in, say, Mg, yields will be
depressed regardless of how much N-P-K product you apply
Shelford's Law of Tolerance

It is a law stating that a certain organism’s survival and existence depend upon
the multifaceted set of conditions wherein each individual has definite minimum,
maximum and optimum ecological factors to establish success.

The absence of an organism can be limited by the qualitative or quantitative

insufficiency from the various environmental factors which may come up to the limits of
tolerance for that organism. Environmental factors involved climatic change,
topographic location and biological necessities of both plants and animals.

This law is possibly the more precise indication of natural complexity. Each individual or
a population is subject to an ecological change that crop up the minimum and maximum
capacity to any complex environmental factors. The range wherein it carried out from
the minimum to maximum signify the limit of tolerance of an organisms, if all known
factors are actually within the particular range of a certain organisms yet it still fails, it is
important to consider extra factors of interrelationships with other organisms

It is been studied that an organisms may have an extensive tolerance for one factor yet
a slight array for another. When an organism has a wide range on all factors it indicates
that a certain organisms are most widely distributed and are contribute to augment
diversity in the community.

Ecological Niche

Jitka Polechová, David Storch, in Encyclopedia of Ecology (Second Edition), 2019

Introduction
Ecological niche is a term for the position of a species within an ecosystem,
describing both the range of conditions necessary for persistence of the species, and its
ecological role in the ecosystem. Ecological niche subsumes all of the interactions
between a species and the biotic and abiotic environment, and thus represents a very
basic and fundamental ecological concept. The tentative definition presented above
indicates that the concept of niche has two sides which are not so tightly related: one
concerns the effects environment has on a species, the other the effects a species has
on the environment. In most of ecological thinking, however, both meanings are
implicitly or explicitly mixed. The reason is that ecology is about interactions between
organisms, and if persistence of a species is determined by the presence of other
species (food sources, competitors, predators, etc.), all species are naturally both
affected by environment, and at the same time affect the environment for other species.

If we want to treat both of these aspects of ecological niche within one framework, we
can define it more formally as the part of ecological space (defined by all combinations
of biotic and abiotic environmental conditions) where the species population can persist
and thus utilize resources and impact its environment. It is useful, however, to
distinguish three main approaches to the niche. The first approach emphasizes
environmental conditions necessary for a species presence and maintenance of its
population, the second approach stresses the functional role of species within
ecosystems, and the third one a dynamic position of species within a local community,
shaped by species’ biotic and abiotic requirements and by coexistence with other
species.

The Ecological Niche in Aquatic Ecosystems

W.M. Lewis, in Reference Module in Earth Systems and Environmental Sciences, 2021

Aquatic ecological niches

Ecological niches are found in all types of ecosystems. At the level of concept or
definition, there is no distinction between aquatic niches and terrestrial niches. Even so,
aquatic environments are distinctive in that some (but not all) of the niche axes most
likely to be important differ from those of terrestrial environments. Important dimensions
of the ecological niche for aquatic organisms include temperature, dissolved oxygen,
habitat structure, predation, and plant nutrients.
The range of temperature for aquatic ecosystems is much narrower than the range for
terrestrial ecosystems because liquid water has a minimum temperature of 0 °C.
Thermal thresholds are weak for phytoplankton and zooplankton, as shown by the
distribution of species across wide ranges of latitude and elevation. Thermal thresholds
are more important for large invertebrates and especially for fishes. For example, the
family Salmonidae (salmon and trout) contains many species that are intolerant of
waters exceeding 15–20 °C. Similarly, perennially cool waters, such as montane or
subarctic lakes, cannot sustain populations of many kinds of warmwater fishes, such as
most species of the sunfish family (Centrarchidae).

Water holds only approximately 10 mg/L of oxygen at low temperatures and 6–7 mg/L at
high temperatures. Thus, respiration can make water anoxic if it is not offset by
photosynthetically produced oxygen or by contact of water with the atmosphere. High
rates of respiration in water that is in contact with sediment, for example, can remove all
oxygen from water in a matter of a few days.

Some aquatic environments are much more subject to oxygen depletion than others.
Mountain streams of high gradient with low amounts of respiration are on one end of the
spectrum, in that they have very little potential for oxygen depletion, whereas fertile
lakes or wetlands, where there is much respiration and less efficient gas exchange with
the atmosphere, show a much higher probability of oxygen depletion.

The distribution of organisms reflects in part their ability to tolerate oxygen depletion.
Fishes, for example, show a wide range of tolerance to oxygen depletion. On one
extreme are fishes that can obtain oxygen from the atmosphere (e.g., the tropical
labyrinth fishes such as the betta, Betta) and fishes that have a small, upturned mouth
capable of drawing oxygen from the top 1–2 mm of water in contact with the
atmosphere (cyprinodont fishes, including the mosquito fish, Gambusia). Intermediate in
tolerance are fishes that have no special means of obtaining oxygen, but have high
physiological resistance to oxygen depletion (some catfishes, such as the bullhead, an
ictalurid, Ameiurus, and the common carp, a cyprinid, Cyprinus). In contrast, other
fishes are moderately or highly sensitive to oxygen depletion: trout and salmon, as well
as largemouth bass (Micropterus), bluegill (Lepomis) and other sunfishes, and many
others. Thus, the fish fauna of a particular waterbody reflects the likelihood of oxygen
depletion. Invertebrates follow similar patterns. Aquatic larvae of midge species, for
example, vary in oxygen tolerance, which explains their contrasting distributions in
aquatic environments.

Habitat structure (cover) is a consistent component of most terrestrial environments; it is

provided to a large extent by vegetation, which appears on all but the driest surfaces or
on ice sheets. In contrast, the pelagic (open-water) environment of lakes provides no
structure. Thus, organisms living in pelagic zones are specialized for living in an
environment that lacks spatial stability and offers no refuge from predation. Variation in
cover, ranging from the open waters of lakes and sandbed streams to richly vegetated
littoral zones, vegetated wetlands, and rocky streams, offer a very wide range of
possibilities for the niche axes that relate to habitat.

Limnologists and aquatic ecologists have shown repeatedly that predation has strong
effects on the species composition of inland waters. Top predators are especially
important in that they often remove or greatly suppress the abundance of their prey.
Visual predators of lakes in particular may be much more efficient in eliminating their
prey than would be the case in most other environments. In fact, some large
invertebrates are unable to inhabit aquatic ecosystems that contain fishes. Small
fishless lakes, for example, contain a rich abundance of large invertebrates that are not
found in similar lakes that are stocked with fish. Thus, the niche dimension related to
predation is critical to the distribution of many invertebrates.

Aquatic environments show an exceptional range of potential for producing autotrophic

biomass. Waters that have very low concentrations of nitrogen and phosphorus have
production potential that falls well below that of any soil-based environment where
moisture is present. In contrast, aquatic ecosystems with high concentrations of
phosphorus and nitrogen have production potential for autotrophs that may be 1000
times higher than the least productive waters. Competition for nutrients is paramount in
unproductive waters, which contain species having very high affinity for phosphorus and
nitrogen. The most productive aquatic environments also support specialized taxa, but
with different kinds of adaptations. Balance of nutrients may also be an important
determinant of niche. For example, the nitrogen-fixing blue-green algae (heterocystous
cyanobacteria) are, unlike other algae, capable of converting gaseous N2 to ammonia
that can be used in making amino acids. When inorganic nitrogen is in short supply,
these organisms are capable of continuing growth when other autotrophs cannot.

SPECIATION

Speciation is how a new kind of plant or animal species is created. Speciation

occurs when a group within a species separates from other members of its species and
develops its own unique characteristics. The demands of a different environment or the
characteristics of the members of the new group will differentiate the new species from
their ancestors.

An example of speciation is the Galápagos finch. Different species of these birds live on
different islands in the Galápagos archipelago, located in the Pacific Ocean off South
America. The finches are isolated from one another by the ocean. Over millions of
years, each species of finch developed a unique beak that is especially adapted to the
kinds of food it eats. Some finches have large, blunt beaks that can crack the hard
shells of nuts and seeds. Other finches have long, thin beaks that can probe into cactus
flowers without the bird being poked by the cactus spines. Still other finches have
medium-size beaks that can catch and grasp insects. Because they are isolated, the
birds don’t breed with one another and have therefore developed into unique species
with unique characteristics. This is called allopatric speciation.
There are five types of speciation: allopatric, peripatric, parapatric, and sympatric and
artificial.

Allopatric speciation (1) occurs when a species separates into two separate groups
which are isolated from one another. A physical barrier, such as a mountain range or a
waterway, makes it impossible for them to breed with one another. Each species
develops differently based on the demands of their unique habitat or the genetic
characteristics of the group that are passed on to offspring.

When Arizona's Grand Canyon formed, squirrels and other small mammals that had
once been part of a single population could no longer contact and reproduce with each
other across this new geographic barrier. They could no longer interbreed. The squirrel
population underwent allopatric speciation. Today, two separate squirrel species inhabit
the north and south rims of the canyon. On the other hand, birds and other species that
could easily cross this barrier continued to interbreed and were not divided into separate
populations.

When small groups of individuals break off from the larger group and form a new
species, this is called peripatric speciation (2). As in allopatric speciation, physical
barriers make it impossible for members of the groups to interbreed with one another.
The main difference between allopatric speciation and peripatric speciation is that in
peripatric speciation, one group is much smaller than the other. Unique characteristics
of the smaller groups are passed on to future generations of the group, making those
traits more common among that group and distinguishing it from the others.

In parapatric speciation (3), a species is spread out over a large geographic area.
Although it is possible for any member of the species to mate with another member,
individuals only mate with those in their own geographic region. Like allopatric and
peripatric speciation, different habitats influence the development of different species in
parapatric speciation. Instead of being separated by a physical barrier, the species are
separated by differences in the same environment.

Parapatric speciation sometimes happens when part of an environment has been

polluted. Mining activities leave waste with high amounts of metals like lead and zinc.
These metals are absorbed into the soil, preventing most plants from growing. Some
grasses, such as buffalo grass, can tolerate the metals. Buffalo grass, also known as
vanilla grass, is native to Europe and Asia, but is now found throughout North and
South America, too. Buffalo grass has become a unique species from the grasses that
grow in areas not polluted by metals. Long distances can make it impractical to travel to
reproduce with other members of the species. Buffalo grass seeds pass on the
characteristics of the members in that region to offspring. Sometimes a species that is
formed by parapatric speciation is especially suited to survive in a different kind of
environment than the original species.

Sympatric speciation (4) is controversial. Some scientists don’t believe it exists.

Sympatric speciation occurs when there are no physical barriers preventing any
members of a species from mating with another, and all members are in close proximity
to one another. A new species, perhaps based on a different food source or
characteristic, seems to develop spontaneously. The theory is that some individuals
become dependent on certain aspects of an environment—such as shelter or food
sources—while others do not.

A possible example of sympatric speciation is the apple maggot, an insect that lays its
eggs inside the fruit of an apple, causing it to rot. As the apple falls from the tree, the
maggots dig in the ground before emerging as flies several months later. The apple
maggot originally laid its eggs in the fruit of a relative of the apple—a fruit called a
hawthorn. After apples were introduced to North America in the 19th century, a type of
maggot developed that only lays its eggs in apples. The original hawthorn species still
only lays its eggs in hawthorns. The two types of maggots are not different species yet,
but many scientists believe they are undergoing the process of sympatric speciations
SPECIES INTERACTIONS

The species interaction describes the relationships among organisms of different

species living in the same location. Examples include herbivores eating plants,
carnivores eating other animals, and organisms competing for food, space, or mates.
The states of all the abiotic and biotic components of the environment.

COMPETITION
- A type of antagonistic relationship within a biological community.
- Organisms compete for resources that are in limited supply such as energy and matter
in usable forms, living space, and specific sites to carry out life’s activities.
- Competition shapes a species population and biological community by causing
individuals and species to shift their focus from one segment of a resource type to
another.

Competition is most typically considered the interaction of individuals that vie for a
common resource that is in limited supply, but more generally can be defined as the
direct or indirect interaction of organisms that leads to a change in fitness when the
organisms share the same resource. The outcome usually has negative effects on the
weaker competitors. There are three major forms of competition. Two of them,
interference competition and exploitation competition, are categorized as real
competition. A third form, apparent competition, is not. Interference competition
occurs directly between individuals, while exploitation competition and apparent
competition occur indirectly between individuals

When an individual directly alters the resource-attaining behavior of other individuals,

the interaction is considered interference competition. For example, when a male
gorilla prohibits other males from accessing a mate by using physical aggression or
displays of aggression, the dominant male is directly altering the mating behavior of
other males. This is also an example of an intra-specific interaction. Exploitation
competition occurs when individuals interact indirectly as they compete for common
resources, like territory, prey or food. Simply put, the use of the resource by one
individual will decrease the amount available for other individuals. Whether by
interference or exploitation, over time a superior competitor can eliminate an inferior one
from the area, resulting in competitive exclusion (Hardin 1960). The outcomes of
competition between two species can be predicted using equations, and one of the
most well known is the Lotka-Volterra model (Volterra 1926, Lotka 1932). This model
relates the population density and carrying capacity of two species to each other and
includes their overall effect on each other. The four outcomes of this model are: 1)
species A competitively excludes species B; 2) species B competitively excludes
species A; 3) either species wins based on population densities; or 4) coexistence
occurs. Species can survive together if intra-specific is stronger than inter-specific
competition. This means that each species will inhibit their own population growth
before they inhibit that of the competitor, leading to coexistence.

Another mechanism for avoiding competitive exclusion is to adopt alternative life history
and dispersal strategies, which are usually reinforced through natural selection. This
mechanism reduces competitive interactions and increases opportunities for new
colonization and nutrient acquisition. The success of this is often dependent upon
events (such as tide, flood, or fire disturbances) that create opportunities for dispersal
and nutrient acquisition. Consider that Plant Species A is more efficient than Plant
Species B at nutrient uptake, but Plant B is a better disperser. In this example, the
resource under competition is nutrients, but nutrient acquisition is related to availability.
If a disturbance opens up new space for colonization, Plant B is expected to arrive first
and maintain its presence in the community until Plant A arrives and begins competing
with Plant B. Eventually Plant A will outcompete Plant B, perhaps by growing faster
because Plant A is more efficient at nutrient acquisition. With an increasing Plant A
population, the Plant B population will decline, and given enough time, can be excluded
from that area.

The exclusion of Plant B can be avoided if a local disturbance (for example, prairie fires)
consistently opens new opportunities (space) for colonization. This often happens in
nature, and thus disturbance can balance competitive interactions and prevent
competitive exclusion by creating patches that will be readily colonized by species with
better dispersal strategies (Roxburgh et al. 2004) (Figure 2). The success of the
dispersal versus nutrient acquisition trade-off depends, however, on the frequency and
spatial proximity (or how close they are) of disturbance events relative to the dispersal
rates of individuals of the competing species. Coexistence can be achieved when
disturbances occur at a frequency or distance that allows the weaker, but often better
dispersing, competitor to be maintained in a habitat. If the disturbance is too frequent
the inferior competitor (better disperser) wins, but if the disturbance is rare then the
superior competitor slowly outcompetes the inferior competitor, resulting in competitive
exclusion. This is known as the intermediate disturbance hypothesis

So in general, Competition is an interaction between organisms or species in which

both the organisms are harmed. Limited supply of at least one resource (such as
food, water, and territory) used by both can be a factor. Competition both within and
between species is an important topic in ecology, especially community ecology.

Mutualism is defined as an interaction between populations that is favorable to both.

Two main types of competition are identified: intraspecific competition and
interspecific competition.
Intraspecific competition
– competition among members of the same species.

For example, two male birds of the same species might compete for mates in the
same area. This type of competition is a basic factor in natural selection. It leads to the
evolution of better adaptations within a species

Interspecific competition
– competition between members of different species.
For example, sharks, dolphins, and seabirds often eat the same type of fish in
ocean ecosystems. Competition can be direct or indirect.

Intraspecific Competition
Intraspecific competition is a common and important interaction for many aquatic
species. A classic laboratory study by L. B. Slobodkin showed reduced growth, survival,
and reproduction of Daphnia when population size was high, as a result of exploitative
competition, and served as the basis of subsequent studies on competition in
zooplankton.

One outcome of intraspecific competition is logistic population growth (called sigmoidal

or S-shaped growth); population growth is nearly exponential when numbers are low,
but then growth rate is reduced progressively as the population expands, and eventually
the population approaches its carrying capacity. Logistic growth of aquatic populations
has been demonstrated repeatedly in laboratory studies of aquatic algae, bacteria,
protozoans, and metazoans. It has been demonstrated less often in the field, probably
because it is difficult to observe colonization events that usually precede logistic growth.

Interference competition can also be an important mechanism of intraspecific

competition. Many zooplankton taxa make autotoxins, which are chemicals that inhibit
feeding or increase mortality in conspecifics. For example, individuals of the rotifer
Synchaeta pectinata produce an autotoxin that reduces growth rate and increases
mortality of other individuals of the same species. Autotoxin effects have also been
demonstrated in a marine phytoplankton species. It is likely that autotoxic effects are
common among freshwater organisms, but little research has been directed toward this
phenomenon.

One consequence of intraspecific competition is stunted growth of fish in dense

populations. Fisheries managers observed long ago that fish in a crowded population
(or with low food availability) often show low (stunted) growth rates and thus are much
smaller than individuals growing in a population with few individuals (or with abundant
resources). Stunted growth has many implications. Small and large fish often rely on
different food resources, so a stunted population may have food web effects different
from those of a population with larger individuals. In addition, smaller individuals may be
more vulnerable to predators, especially other fish that are gape-limited. Stunted
populations also may be less desirable for recreational and commercial harvest.

Intraspecific competition can also lead to increased variability in body size. Competition
is often highly asymmetric, meaning that it affects some individuals much more than
others.

This could be because some individuals are inherently better competitors, or because
some individuals arrive at a site (or are born) earlier than others and thus preempt
resources. Superior or early-arriving individuals may reach a relatively large size while
inferior competitors or late arrivers suffer reduced body size. Often there is a gradient in
competitive ability or arrival times, and a population growing under intraspecific
competition displays a wide distribution of sizes among individuals of equal age. Such
asymmetries have been demonstrated in fish, amphibians, and insects. Differences in
size initiated by intraspecific competition can become magnified over time by
size-dependent competitive superiority.

An individual that gains an initial advantage (e.g., by arriving early or by having a slightly
larger initial size) will grow more rapidly than the average individual. This individual may
use a wider range of resources (e.g., larger fish can consume a wider range of prey
items), leading to a further gain in size relative to other individuals. This difference in
size may become more pronounced over time.

Interspecific competition has not been a major focus of studies investigating the
structure of lake fish communities. We are not aware of a study showing that a certain
fish species has become excluded from a local lake community as the consequence of
competition with another local fish species, but invasion by non-native species may
induce stronger negative effects. In turn, a dominance of positive co-occurrence
patterns of species or habitat-specific groups of species among lakes suggests that
facilitation and niche similarity structure richness and composition of lake fish
communities substantially (Fig. 3) (MacDougall et al., 2018).
Mutually negative effects on abundance or biomass of competing fish species are more
likely (Eloranta et al., 2016), but systematically negative abundance or biomass
correlations between species with similar niches among lakes have not been found
(MacDougall et al., 2018). It has to be noted that studies of interspecific competition
have to consider the frequent ontogenetic niche shifts in freshwater fishes (Werner and
Gilliam, 1984), resulting in a succession of different biotic interactions while fish grow.

The effect of competition among species is usually best demonstrated by niche

segregation between co-occurring fish species (Genner et al., 1999; McMeans et al.,
2020). This process is also considered as a route towards ecological speciation in lake
fish communities (Knudsen et al., 2006). However, these young speciation events can
be reversed and sympatric species can disappear if anthropogenic influences on lakes
destroy the ecological gradients along which niche segregation has been expressed
(Vonlanthen et al., 2012)

PREDATOR

- Any organism that feeds directly on another living organism, whether or not this kills
the prey.
- Herbivores, carnivores, and omnivores, which feed on live prey, are predators.
- Predation is a powerful but complex influence on species populations in communities.

We can define predation as the ecological process in which an animal (or an organism)
kills and feeds on another animal (or an organism). The animal that kills another
animal to feed on is called a “predator“.
Predation affects:

1. All stages in the life cycles of predator and prey species.

2. Many specialized food-obtaining mechanisms.

3. The evolutionary adjustments in behavior and body characteristics that help prey
avoid being eaten and help predators more efficiently catch their prey.

What counts as predation?

A predator is an organism that consumes all or part of the body of another—living or

recently killed—organism, which is its prey. "Living or recently killed" distinguishes
predators from decomposers, such as fungi and bacteria that break down the leftover
remains of organisms that have died.

If we see a lion eating a zebra, we can feel comfortable in saying that the lion is a
predator. In the broad definition, however, the zebra is too!

A predator's prey can be an animal, but it can also be a plant or fungus. Nor does a
predator necessarily have to kill its prey. Instead, as in a grazing cow or a bloodsucking
mosquito, it may simply take a portion of the prey's body and leave it alive.

A predator-prey relationship in which an animal or insect consumes a plant is called

herbivory—herbi- means plant, and -vory means eating.

Mechanical defenses, such as the presence of thorns on plants or the hard shell on
turtles, discourage animal predation and herbivory by causing physical pain to the
predator or by physically preventing the predator from being able to eat the prey.
Chemical defenses are produced by many animals as well as plants, such as the
foxglove, which is extremely toxic when eaten. The millipede in the lower panel below
has both chemical and mechanical defenses: when threatened, it curls into a defensive
ball and makes a noxious substance that irritates eyes and skin.

Many species use their body shape and coloration to avoid being detected by predators.
For instance, the crab spider has the coloration and body shape of a flower petal, which
makes it very hard to see when it's standing still against the background of a real flower.
Can you even see it in the picture below? It took me a minute! Another famous example
is the chameleon, which can change its color to match its surroundings. Both of these
are examples of camouflage, or avoiding detection by blending in with the background

Types of Predators

Large Predators

Predators come in all shapes and sizes. They all have special adaptations that allow
them to hunt and kill their prey, with minimal damage to themselves. The most common
traits a predator has include very large teeth, sharp claws, and great strength. Many of
them are also known to have very bad attitudes (meaning they can be very aggressive)
and can cause harm to people. Polar bears, killer whales, and great white sharks all fall
into this category. All these animals are much larger than people and have the
necessary 'weapons' for killing. Also, there aren't too many people who would want to
be trapped in a room with any of them.

The polar bear is considered the largest land carnivore. It can stand over nine feet (3 m)
tall on its back legs, and it eats nothing but meat, usually seals. It has very large paws,
each tipped with strong claws.

Killer whales can reach a length of 30 (9.5m) feet and weigh up to six tons (5,443 kg).
Each of their teeth can be four inches (10 cm) long. They eat seals, sea lions, and fish.
Great white sharks (Figure 1) are the ultimate ocean predator. They are built for nothing
more than eating. They are usually 15 feet (4.6 m) feet long, weigh 5,000 pounds (2,268
kg), and have thousands of teeth. There are not too many animals on Earth that put fear
into people so well as a great white shark.

Smaller Predators

Not all predators, however, are such animals of 'mass destruction.' For example, sea
stars (Figure 2) living on the bottom of the ocean are predators. They have neither teeth
nor sharp claws, and certainly do not cause fear in humans. Yet, sea stars are
considered one of the most dangerous animals to certain types of shellfish. When on
the hunt, sea stars use their tube feet to crawl across the ocean floor. When they find a
clam, they will grab onto its shell from both sides using their arms. They then begin to
pull as hard as they can to try to separate the valves of the shell. The clam is using all of
its strength to hold the valves together, but the sea star is stronger. Once the clam tires,
and the valves open just a little, the sea star throws up its stomach through its mouth
and into the clam. The digestive enzymes in the sea star's stomach digest the clam from
the outside.

antagonistic interactions include predation, herbivory, and parasitism. These

interactions are complex, with predators consuming herbivores, plants and animals
infected by parasites, and occasional disease outbreaks. Predation, herbivory, and
parasitism exist along a continuum of severity in terms of the extent to which they
negatively affect an organism’s fitness.

Herbivorous, predatory, and parasitic interactions among organisms within communities

regulate population sizes by preventing any one population from becoming overly
abundant. Each type of interaction becomes more likely as the population size of the
food source increases. Thus these interactions are essential to maintaining the diversity
of organisms that make up an ecological community. Each of these interactions can
alter the balance of the food web, and removal of any part of the web can have a drastic
impact on the community.

Symbiosis
 - Two or more species live intimately together, with their fates linked.
- Symbiotic relationships often enhance the survival of one or both partners.

Basically symbiosis is describing any relationship or interaction between two

dissimilar organisms. The specific kind of symbiosis depends on whether either or
both organisms benefit from the relationship.

Symbiotic relationships are an important component of life in the ocean. In such

relationships, plants or animals of different species may be dependent on one another
for survival. They may share habitats or lifestyles or interact in a specific way to benefit
from the presence of another organism.

We often refer to animals living in tandem as 'associates.'' The relationship between

associates and their hosts can be described as mutualistic, commensal, or parasitic. In
a mutualistic relationship, both animals benefit from living together. Commensal
organisms cause no harm to their hosts, but receive some benefit from living with them.
Parasites actually feed off their host organism, thus causing harm to the host.

Although there are many ways organisms interact with one another, most symbioses
involve clever ways to obtain food or protection. For example, ophiuroids (brittle stars)
are often found living within the branches of corals, using their hosts to get further up
off the seafloor into the water column to feed or for protection. At hydrothermal vents,
chemosynthetic bacteria live inside of animals in a mutualistic symbiotic relationship
where the animals support the existence of the bacteria and the bacteria provide food
to the animals in an environment where light does not penetrate. And different types of
marine parasites, including worms, isopods, and copepods, infect a variety of host
species, including crabs and fishes.

Deep-sea symbioses are poorly understood and less well documented relative to
symbiotic relationships frequently encountered in shallower habitats
The five major types of species interactions are:

● Competition.
● Predation.
● Parasitism.
● Mutualism.
● Commensalism

Types of Symbiosis:

❏ Mutualism – type of symbiosis in which both members’ benefits. (e.g. Dogs and
Humans)

❏ Commensalism – type of symbiosis in which one member clearly benefits and

the other apparently is neither benefited or harmed. (e.g. a spider building a web
on a tree)

❏ Parasitism – a form of predation may also be considered symbiosis because of

the dependency of the parasite on its host. (Fleas and mosquitoes feed on blood
from other organisms.

❏ Endosymbiosis – one species living inside another one. (e.g. Protozoans that
live inside termites and help them digest wood)

❏ Ectosymbiosis – one species living on the surface of the other species. (e.g.
Lice that feed on the skin, blood, or oil secretions of the host)
KEYSTONE SPECIES

- Plays a critical role in a biological community that is out of proportion to its

abundance.

- Thought to be the top predators like lions, wolves, and tigers that limited herbivore
abundance and reduced the herbivory of plants.

Examples of Keystone Species Show

● African Elephant.
● Beaver.
● Bee.
● Grizzly Bear.
● Hummingbird.
● Ivory Tree Coral.
● Jaguar.
● Pacific Salmon.

A keystone species is an organism that helps define an entire ecosystem. Without its
keystone species, the ecosystem would be dramatically different or cease to exist
altogether.

Keystone species have low functional redundancy. This means that if the species were
to disappear from the ecosystem, no other species would be able to fill its ecological
niche. The ecosystem would be forced to radically change, allowing new and possibly
invasive species to populate the habitat.

Any organism, from plants to fungi, may be a keystone species; they are not always the
largest or most abundant species in an ecosystem. However, almost all examples of
keystone species are animals that have a huge influence on food webs. The way these
animals influence food webs varies from habitat to habitat.

Carnivores, Herbivores, and Mutualists

Predators

A keystone species is often, but not always, a predator. Just a few predators can control
the distribution and population of large numbers of prey species.

The entire concept of keystone species was founded on research surrounding the
influence of a marine predator on its environment. American zoology professor Robert
T. Paine's research showed that removing a single species, the Pisaster ochraceus sea
star, from a tidal plain on Tatoosh Island in the U.S. state of Washington, had a huge
effect on the ecosystem. Pisaster ochraceus, commonly known as purple sea stars, are
a major predator of mussels and barnacles on Tatoosh Island. With the sea stars gone,
mussels took over the area and crowded out other species, including benthic algae that
supported communities of sea snails, limpets, and bivalves. Lacking a keystone
species, the tidal plain’s biodiversity was cut in half within a year.

Another example of a predator acting as a keystone species is the presence of gray

wolves in the Greater Yellowstone Ecosystem. The Greater Yellowstone Ecosystem
(GYE) is an enormous and diverse temperate ecosystem stretching across the
boundaries of the U.S. states of Montana, Wyoming, and Idaho. The GYE includes
active geothermal basins, mountains, forests, meadows, and freshwater habitats.

The elk, bison, rabbit, and bird species in the Greater Yellowstone Ecosystem are at
least partly controlled by the presence of wolves. The feeding behavior of these prey
species, as well as where they choose to make their nests and burrows, are largely a
reaction to wolf activity. Scavenger species, such as vultures, are also controlled by the
wolf activity.
When the U.S. government designated land for Yellowstone National Park in the late
19th century, hundreds of wolves roamed the GYE, preying primarily on abundant herds
of elk and bison. Fearing the wolves’ impact on those herds, as well as local livestock,
governments at the local, state, and federal level worked to eradicate wolves from the
GYE. The last remaining wolf pups in Yellowstone were killed in 1924.

This started a top-down trophic cascade in the Greater Yellowstone Ecosystem. A

trophic cascade describes changes in an ecosystem due to the addition or removal of a
predator. A top-down trophic cascade describes changes that result from the removal of
an ecosystem’s top predator. (A bottom-up trophic cascade describes changes that
result from the removal of a producer or primary consumer.)

Lacking an apex predator, elk populations in Yellowstone exploded. Elk herds competed
for food resources, and plants such as grasses, sedges, and reeds did not have time or
space to grow. Overgrazing influenced the populations of other species, such as fish,
beaver, and songbirds. These animals rely on plants and their products—roots, flowers,
wood, seeds—for survival.

The physical geography of the Greater Yellowstone Ecosystem was also impacted by
the loss of wolves and subsequent elk overgrazing. Stream banks eroded as wetland
plants failed to anchor valuable soil and sediments. Lake and river temperatures
increased as trees and shrubs failed to provide shaded areas.

Starting in the 1990s, the U.S. government began reintroducing wolves to the Greater
Yellowstone Ecosystem. The results have been noteworthy. Elk populations have
shrunk, willow heights have increased, and beaver and songbird populations have
recovered.
Herbivores

Herbivores can also be keystone species. Their consumption of plants helps control the
physical and biological aspects of an ecosystem.

In African savannas such as the Serengeti plains in Tanzania, elephants are a keystone
species. Elephants eat shrubs and small trees, such as acacia, that grow on the
savanna. Even if an acacia tree grows to a height of a meter or more, elephants are
able to knock it over and uproot it. This feeding behavior keeps the savanna a grassland
and not a forest or woodland.

With elephants to control the tree population, grasses thrive and sustain grazing
animals such as antelopes, wildebeests, and zebras. Smaller animals such as mice and
shrews are able to burrow in the warm, dry soil of a savanna. Predators such as lions
and hyenas depend on the savanna for prey.

Keystone Mutualists

Keystone mutualists are two or more species that engage in mutually beneficial
interactions. A change in one species would impact the other, and change the entire
ecosystem. Keystone mutualists are often pollinators, such as bees. Pollinators often
maintain gene flow and dispersal throughout widespread ecosystems.

In the woody grasslands of Patagonia (at the southern tip of South America) a species
of hummingbird and indigenous plants act together as keystone mutualists. Local trees,
shrubs, and flowering plants have evolved to only be pollinated by Sephanoides
sephanoides, a hummingbird known as the green-backed firecrown. Green-backed
firecrowns pollinate 20% of local plant species. In turn, these plants provide the sugary
nectar that makes up most of the hummingbird’s diet.

Pockets of the existing Patagonian habitat would collapse without green-backed

firecrowns, because their functional redundancy is nearly zero—no other pollinator has
adapted to pollinate these plants.

Other Organisms Crucial to Ecosystems

In addition to keystone species, there are other categories of organisms crucial to their
ecosystems' survival.

Umbrella Species

Umbrella species are often conflated with keystone species. Both terms describe a
single species on which many other species depend. The key distinction between
umbrella species and keystone species is that the value of an umbrella species is tied to
its geographic species range.

Umbrella species have large habitat needs, and the requirements of that habitat impact
many other species living there. Most umbrella species are migratory, and their range
may include different habitat types.

The identification of an umbrella species can be an important aspect of conservation.

The minimum species range of an umbrella species is often the basis for establishing
the size of a protected area.

The Siberian tiger, an endangered species, is an umbrella species with a range of more
than 1,000 kilometers (620 miles) in Russia’s far east, with territory stretching into China
and North Korea. The species range includes heavily forested ecosystems in both
temperate and boreal (subarctic) biomes. Populations of deer, boar, and moose are
under the snowy “umbrella” of the Siberian tiger range.
REFERENCES:

https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/intraspecific-c
ompetition

https://www.nature.com/scitable/knowledge/library/species-interactions-and-competition
-102131429/

https://www.khanacademy.org/science/biology/ecology/community-ecosystem-ecology/a
/predation-herbivory

https://study.com/academy/lesson/predator-in-ecosystems-definition-lesson-quiz.html

https://oceanexplorer.noaa.gov/facts/symbiosis.html

https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/intraspecific-c
ompetition

https://www.nature.com/scitable/knowledge/library/species-interactions-and-competition
-102131429/

https://www.khanacademy.org/science/biology/ecology/community-ecosystem-ecology/a
/predation-herbivory

https://study.com/academy/lesson/predator-in-ecosystems-definition-lesson-quiz.html

https://oceanexplorer.noaa.gov/facts/symbios
Dynamic and Changing Communities

Community dynamics are the changes in community structure and composition over
time. Sometimes these changes are induced by environmental disturbances such as
volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable
structure are said to be at equilibrium. Following a disturbance, the community may or
may not return to the equilibrium state.

Succession describes the sequential appearance and disappearance of species in a

community over time. In primary succession, newly exposed or newly formed land is
colonized by living things; in secondary succession, part of an ecosystem is disturbed
and remnants of the previous community remain.

Ecological Succession

"Ecological succession" is the observed process of change in the species structure of

an ecological community over time. Within any community, some species may become
less abundant over some time interval, or they may even vanish from the ecosystem
altogether. Similarly, over some time interval, other species within the community may
become more abundant, or new species may even invade into the community from
adjacent ecosystems. This observed change over time in what is living in a particular
ecosystem is "ecological succession".

Primary Succession and Pioneer Species

Primary succession occurs when new land is formed or rock is exposed: for example,
following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava
flows into the ocean, new land is continually being formed. On the Big Island,
approximately 32 acres of land are added each year. First, weathering and other natural
forces break down the substrate enough for the establishment of certain hearty plants
and lichens with few soil requirements, known as pioneer species. These species help
to further break down the mineral-rich lava into the soil where other, less hardy species
will grow and eventually replace the pioneer species. In addition, as these early species
grow and die, they add to an ever-growing layer of decomposing organic material and
contribute to soil formation. Over time the area will reach an equilibrium state, with a set
of organisms quite different from the pioneer species.

Secondary Succession

A classic example of secondary succession occurs in oak and hickory forests cleared by
wildfire. Wildfires will burn most vegetation and kill those animals unable to flee the
area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even
when areas are devoid of life due to severe fires, the area will soon be ready for new life
to take hold.

Before the fire, the vegetation was dominated by tall trees with access to the major plant
energy resource: sunlight. Their height gave them access to sunlight while also shading
the ground and other low-lying species. After the fire, though, these trees are no longer
dominant. Thus, the first plants to grow back are usually annual plants followed within a
few years by quickly growing and spreading grasses and other pioneer species. Due to,
at least in part, changes in the environment brought on by the growth of the grasses and
other species, over many years, shrubs will emerge along with small pine, oak, and
hickory trees. These organisms are called intermediate species. Eventually, over 150
years, the forest will reach its equilibrium point where species composition is no longer
changing and resembles the community before the fire. This equilibrium state is referred
to as the climax community, which will remain stable until the next disturbance.

Ecological Disturbance

In the ecological context, the disturbance is regarded as an event of intense

environmental stress occurring over a relatively short period of time and causing large
changes in the affected ecosystem. Disturbance can result from natural causes or from
the activities of humans.
Disturbance can be caused by physical stressors such as volcanic eruptions,
hurricanes, tornadoes, earthquakes, and over geological time, glacial advance, and
retreat. Humans can also cause physical disturbances, for example, through
construction activities. Wildfire is a type of chemical disturbance caused by the rapid
combustion of much of the biomass of an ecosystem and often causing mortality of the
dominant species of the community such as trees in the case of a forest fire. Wildfires
can ignite naturally, usually through a lightning strike, or humans can start the blaze.
Sometimes fires are set deliberately as a management activity in forestry or agriculture.
Events of unusually severe pollution by toxic chemicals, nutrients, or heat may also be
regarded as a type of disturbance if they are severe enough to result in substantial
ecological damages. Disturbance can also be biological, as when a severe infestation of
defoliating insects causes substantial mortality of trees in a forest, or of crops in
agriculture. The harvesting of forests and other ecosystems by humans is another type
of biological disturbance.
Ecological disturbance can occur at a variety of spatial scales. The most extensive
disturbances involve landscape-scale events, such as glaciation, which can affect entire
continents. Tornadoes, hurricanes, and wildfires can also affect very large areas;
sometimes wildfires extend over millions of acres.
Some disturbances, however, are much more local in their effects. For example, the
primary disturbance regime in old-growth forests is associated with the death of
individuals, large trees caused by disease, insect attack, or a lightning strike. This sort
of micro disturbance event results in a gap in the otherwise closed forest canopy, which
lets direct light reach the forest floor. This encourages the development of a different set
of plant and animal communities than those usually found on the dark, moist forest floor.
Further ecological changes occur when the dead tree falls to the ground and slowly rots.
Diverse processes of ecological recovery occur in response to the within-stand patch
dynamics associated with the deaths of large trees in old-growth forests.
Whenever an ecosystem is affected by a substantial disturbance event, individuals and
even entire species may be weakened or killed off. Other ecological damages can also
occur, such as changes in hydrologic processes or soil contamination. However, once
the actual disturbance event is finished, a process known as succession begins, which
may eventually produce a similar ecosystem to the one that existed prior to the
disturbance.
In a number of regions around the world, human activities are producing dramatic
ecological disturbances. Clear-cutting of tropical rainforests, the damming or polluting of
rivers and streams, the introduction of various chemicals and particulates into the
atmosphere from industrial facilities are all human processes that have major effects on
many ecosystems. In cases where the ecological disturbance is ongoing, succession is
forestalled, and the damaged ecosystems may fail to recover their complex and
sophisticated functions.

Introduced species

Introduced species, also called alien species, are those that have been moved by
humans to an environment where they didn't occur naturally. The term can refer to
animals, plants, fungi, or microorganisms that are non-native to an area. Species
introduction can be accidental or intentional.

Many accidental introductions of species involve boats, which travel between

continents. For example, the black rat, which is native to South Asia, has been hitching
rides in ship cellars since the first century, spreading initially to Europe and
subsequently throughout the rest of the world during the Age of Discovery. Species
have also been accidentally introduced through water discharged by cruise ships and
tankers, stuck to travelers' clothes and shoes, or in imported wood and food.

Sometimes species that don't occur naturally in a habitat are deliberately introduced by
humans for conservation efforts, population control of native species, or for boosting
agriculture and fisheries. For example, although the U.S. is today the largest producer
of corn in the world, and corn is grown in every continent, corn didn't use to exist
anywhere but the Mexican mountains until Indians and explorers started taking it to
other places. Sometimes species are introduced seemingly on a whim for human
enjoyment, like the water hyacinth, which is originally from South America and has been
used to beautify ponds in North America, Africa, Asia, Australia, and New Zealand.
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