By Suzanne Macey, Ana Luz Porzecanski, and Kimberley Landrigan

What is ecology?

In 1866, German biologist Ernst Haeckel coined the term “ecology” (from oikos, the Greek word
for “house”) to classify the branch of biology that focuses on our immediate surroundings—our
environment. A few years later, Haeckel expanded his definition, saying:

By ecology, we mean the body of knowledge concerning the economy of nature—the

investigation of the total relations of the animal both to its organic and to its inorganic
environment; including above all, its friendly and inimical relation with those animals and plants
with which it comes directly or indirectly into contact—in a word, ecology is the study of all the
complex interrelationships referred to by Darwin as the conditions of the struggle for existence.

Although the term was new, scientists were already studying natural processes and our
environment. They now began to identify themselves as ecologists. The field isn’t restricted to
professional scientists: any curious person who investigates the processes of nature—be it
tracking the blooming of plants on their local reserve, or recording the arrival of migratory birds in
their backyard—is engaged in the practice of ecology.

What are these processes of nature? They are the mechanisms by which living organisms
interact with each other (bioticinteractions) and with the physical and chemical components of
their environment (abiotic interactions). Ecologists draw on an understanding of these processes
to identify patterns—observable, repeatable phenomena—that determine the abundances and
geographic distribution of organisms, and how matter cycles and energy flows within natural
systems.

How do we study ecology?

Ecologists observe these processes and patterns at different scales of time, space, size and
complexity, from the behavior of ants to the effects of global climate systems. In 1953, Eugene
Odum emphasized and popularized an ecological hierarchy (Figure 1) as an organizing principle
in ecology. Referencing this hierarchy of ecological systems, the field is often subdivided into
organismal ecology, population ecology, community ecology, ecosystem ecology, landscape
ecology, and global ecology. Although these are considered separate fields of study, there is
considerable overlap between concepts, approaches, and principles.
Figure 1. Ecological hierarchy Visual depiction of the different levels of the ecological hierarchy.
From left to right, the smallest to largest unit of study (see text for descriptions ©Nadav
Gazit/AMNH

Compared to ecologists in the 1950s, ecologists today are more likely to identify the elements of
an ecological hierarchy in less fixed, more dynamic terms. While the traditional, and somewhat
linear, hierarchy presented above is still helpful, and forms the basic structure of this course,
systems thinking is embedded throughout in order to emphasize the interdependent, complex
relationships between all these elements in the hierarchy (further discussed in the essay
“Introduction to Systems Thinking”). We use case studies to show these relationships at work on
different scales, and trust that this will help you to see how every topic in ecology has a history
and how our understanding has changed over time.
Figure 2. Classification of biomes based on temperature and precipitation. Observed vegetation
types with respect to average temperature and precipitation as originally defined by Whittaker.
Adapted from Whittaker (1975) by Nadav Gazit/AMNH.

At the organism level

Organismal ecologists study individual organisms and their adaptations to specific habitats and
conditions. For example, in Gorongosa National Park, scientists study bats and how they use
echolocation to find their prey in the darkness and how the prey (moths) are adapted to avoid the
bats. Other questions these scientists might ask are: How do bats use echolocation for social
communication? What are the mechanics of their flight? What genes allow bats to be resistant to
diseases that are devastating to other species (e.g., Ebola, Zika)? Ecological questions like these
shed light on evolution; they can help determine how adaptations—including behavioral,
physiological, genetic, or morphological traits—provide individuals with increased ability to
survive and reproduce. If so, those traits will predominate over time in a population as the result
of evolutionary change through natural selection. Additionally, ecological research can help
scientists answer other questions in different disciplines—for instance, questions related to the
bioengineering of flying robots or the investigation of human immunity.

At the population level

How many zebras need to be introduced to Gorongosa National Park in order to ensure genetic
diversity? Why hasn’t the lion population rebounded? What is the maximum number of waterbuck
that can live in Gorongosa National Park (also known as the area’s “carrying capacity”)?
Population ecologists study how and why the numbers of individuals of a species change over
time. The term “population” describes a group of individuals of one species living in the same
area at the same time. The way a specific population is defined depends on what question the
ecologist asks or what approach they use. For example, they might use genetic criteria, or
geographic boundaries, or behavioral interactions to define a population. Ecologists use tools
such as genetic analysis and demographic and mathematical modeling (e.g., determining birth
rates and death rates) to understand the changes within populations and the factors that
influence their growth, decline, regulation, and dynamics. By using the tools of ecology, we are
able to predict changes and manage species—specifically, those that humans use (e.g.,
agricultural crops, game species), seek to conserve (e.g., endangered species), or seek to
eradicate (e.g., diseases, invasive species).

At the community level

Understanding community ecology helps us answer questions like: How can so many species of
birds inhabit the same tropical rain forest on Mount Gorongosa? How do plant species respond
to the loss of certain types of herbivores? How does the presence of a termite mound change the
community of plants and animals in the nearby area? Communities are assemblages of
populations of different species that inhabit the same place at the same time.

As with populations, there are various ways to define communities. For example, a community
might be defined as the species found in a vernal pool (e.g., species of algae, salamanders,
mosquitoes), the microorganisms found in a mosquito’s gut, or the species clustered around a
hydrothermal vent on the ocean floor (e.g., species of shrimp, tube worm, bacteria). But
communities do not necessarily have strict physical boundaries, as species may move in an out
of an area. Often an area may contain so many species that it’s hard to unravel their complex
interactions. Complex communities like these include the savannas of Mozambique, the rivers of
New York State, and the seascapes surrounding the Bahamas. Researchers may focus on a
subset of species, or on the dominant species, or on the organisms that interact with their
species of interest. At the community level, ecologists can study the connections among groups
of species (e.g., predation, herbivory, competition, mutualism, parasitism) and how these
interactions affect community structure and the diversity and relative abundances of species.

At the ecosystem level

A pioneer in the field, British botanist Arthur Tansley observed that biologists were focusing
primarily on organisms rather than the flow or exchange of materials between organisms and the
physical environment. In 1935, Tansley coined the term “ecosystem,” meaning an ecological
system, to assert the importance of incorporating the movements of energy and materials into
the study of ecology. Ecologists who work at this level study the interactions of organisms within
a community and with their physical environment as an integrated system—an ecosystem. But
what are the boundaries of an ecosystem?

An ecosystem and its processes can be studied at any scale, from a tiny tidal pool up to a global
system that integrates all living organisms and their environments—the biosphere itself. As with
communities, it can be challenging to delineate boundaries between ecosystems. Ecologists
might define ecosystems according to the dominant environment (e.g., desert spring ecosystem),
the dominant species (e.g., mangrove swamp ecosystem), natural boundaries (e.g., watershed
ecosystem), or artificial boundaries (e.g., Gorongosa National Park ecosystem). Again, the scale
or definition of the ecosystem hinges on the questions the ecologist is asking. For example,
ecologists might investigate energy flow or chemical cycles: How much carbon does a cranberry
bog sequester? Or they may want to assess the effect of physical disturbance: How does a
drought affect a new urban development? Or they might ask questions about the distribution of
certain communities: Why does a specific assemblage of species inhabit tidal marshes?

BOX 1. Landscapes

Sometimes ecologists expand upon these hierarchical levels by defining groups of

ecosystems as landscapes or seascapes—ecosystems that are clustered and affect each
other. The field of landscape ecology focuses on how energy, materials, and organisms
move between a mosaic of different (heterogeneous) but connected ecosystems. As with
other levels of ecological systems, landscapes are not necessarily defined by size, but
instead by the perspective of the investigator and the nature and scope of their questions.
For example, an ecologist studying bird migration could define the landscape based on the
habitat patches a bird species uses throughout its range. Or, from a fire-management
perspective, ecologists might define a landscape based on the patchwork of fire-dependent
ecosystems.BOX 1. Landscapes

BOX 2. Biomes

Another approach to grouping ecosystems is to characterize them as aquatic zones or

terrestrial biomes. In aquatic systems, ecologists differentiate between freshwater (e.g.,
lakes, wetlands, streams and rivers, estuaries) and marine systems (e.g., intertidal zones,
pelagic zone, coral reefs, benthic zones). Aquatic environments are dispersed across the
biosphere but united by common physical characteristics (e.g., temperature, sunlight,
salinity) that shape the communities they contain. In terrestrial systems across the
biosphere, the distribution of different types of vegetation is linked to different climate zones.
In 1975, American plant ecologist Robert H. Whittaker divided the biosphere into nine
terrestrial biomes characterized by major vegetation types in consideration of the annual
temperature and precipitation (Figure 2). Today, ecologists have characterized 12 or more
terrestrial biomes across the biosphere (Figure 3), but the original nine are still often used.
Figure 3. Terrestrial biomes across the biosphere Global distribution of twelve terrestrial biomes.
Data courtesy of HHMI BiomeViewer (available at
http://www.hhmi.org/biointeractive/biomeviewer)​ . Adapted by Nadav Gazit/AMNH.

At the global level

At this level, ecologists study how global patterns and dynamics (e.g., ocean and wind currents,
natural variations in seasons such as El Niño, plate tectonics, climate change) are responsible
for the geographic distribution and abundances of populations as well as the functioning and
productivity of ecosystems. Questions in global ecology focus on interactions among the Earth’s
ecosystems, land, oceans, and atmosphere, such as: How does ocean circulation affect the
global distribution of coral reefs? In view of rising global temperatures, if the permafrost of the
tundra melts and releases carbon dioxide and other greenhouse gases into atmosphere, how will
the carbon cycle be affected? Is there a threshold for atmospheric carbon before certain
ecosystem functions collapse? Increasingly, the study of global ecology is concentrated on the
role humans play in the functioning of the biosphere.

How does ecology contribute to conservation?

There is a long human history in the savannas of Mozambique and other ecosystems around the
world. Humans have physically altered or transformed more than three-quarters of Earth’s land
surface. Even in places where no humans are present, our activities influence processes
throughout the biosphere, such as how communities are structured, nutrient cycling, energy flow,
and natural disturbance regimes (i.e., patterns of disturbance, such as fires or floods). As this
course will explore, this growing human footprint has created an environmental crisis of
unparalleled proportions, involving the mass extinction of species (see the essay “Introduction to
Biodiversity”), ocean acidification, climate change, ozone depletion, interference with the nitrogen
and phosphorus cycles, and pollution. The study of ecology plays a fundamental role in helping
us understand and address these issues at all levels, from bringing a species back from the brink
of extinction to addressing detrimental changes in the atmosphere.

REFERENCES

Campbell, N. A., Reece, J. B., Urry, L. A., Cain, M. L., Wasserman, S. A., Minorsky, P. V., &
Jackson, R. B. (2008). ​Biology​ (8th ed.) San Francisco: Pearson Benjamin Cumming.
Chapin III, F. S., Matson, P. A., & Mooney, H. A. (2002). ​Principles of Terrestrial Ecosystem
Ecology.​ New York: Springer.

Currie, W. S. (2011). Units of nature or processes across scales? The ecosystem concept at age
75. ​New Phytologist,​ 190(1), 21-34.

Ricklefs, R. E., & Relyea, R. (2014). ​Ecology the Economy of Nature.​ New York: W. H. Freeman
and Company.

Rockström, J., et al. (2009). A safe operating space for humanity. ​Nature,​ 461(7263), 472-475.

Venter, O., et al. (2016). Sixteen years of change in the global terrestrial human footprint and
implications for biodiversity conservation. ​Nature Communications,​ 7(12558).
doi:10.1038/ncomms12558

Whittaker, R. H. (1975). ​Communities and Ecosystems​ (2nd ed). New York: Macmillan.

RELATED LINKS

Moth Mimicry: Using Ultrasound to Avoid Bats​: This video from HHMI follows scientists as they
uncover the ways in which moth species in Gorongosa National Park use ultrasound to avoid
being eaten by bats.

HHMI BiomeViewer​: An interactive from HHMI that explores biomes, climate, biodiversity, and
human impacts across the globe.