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Ecological Communities: Networks of Interacting

Species

We wish to learn:

 What is an ecological community and what kinds of interactions take place within it?
 How important are the various categories of species interactions, including
mutualisms, commensalisms, competition and predation?
 What kinds of interactions among species become important when many species
affect one another?
 What consequences do these interactions have for biodiversity

Species Interactions, Food Webs, and Ecological Communities

An ecological community is defined as a group of actually or potentially interacting species
living in the same place. A community is bound together by the network of influences that
species have on one another. Inherent in this view is the notion that whatever affects one
species also affects many others -- the "balance of nature". We build an understanding of
communities by examining the two-way, and then the multi-way, interactions involving
pairs of species or many species.

type of interaction sign effects

mutualism +/+ both species benefit from interaction
commensalism +/0 one species benefits, one unaffected
competition -/- each species affected negatively
predation, parasitism, herbivory +/- one species benefits, one is disadvantaged

Food webs are graphical depictions of the interconnections among species based on energy
flow . Energy enters this biological web of life at the bottom of the diagram, through the
photosynthetic fixation of carbon by green plants. Many food webs also gain energy inputs
through the decomposition of organic matter, such as decomposing leaves on the forest
floor, aided by microbes. River food webs in forested headwater streams are good
examples of this.

Energy moves from lower to higher trophic (feeding) levels by consumption: herbivores
consumes plants, predators consume herbivores, and may in turn be eaten by top
predators. Some species feed at more than one tropic level, hence are termed omnivores.
Figure 1 provides a simplified model of such a food web.

Generalized food web. A food web is an assemblage

of organisms, including producers, consumers and
decomposers, through which energy and materials
may move in a community

We can look at this food web in two ways. It can be a diagram of the flow of energy
(carbon) from plants to herbivores to carnivores, and so on. We will take this approach
when we examine energy flow in ecosystems. In addition, members of a food web may
interact with one another via any of the four interaction types named above. An interaction
between two species in one part of the web can affect species some distance away,
depending on the strength and sign of the inter-connections. Often, adding a species (as
when an exotic species invades a new area) or removing a species (as in a local extinction)
has surprisingly far-reaching effects on many other species. This is due to the complex
inter-connections of species in ecological webs.

Ecologists use the following terms to describe various categories of the effects of a change
(in abundance, or presence vs absence) of one species on another.

1. Direct effects refer to the impact of the presence (or change in abundance) of
species A on species B in a two-species interaction.
2. Indirect effects refer to the impact of the presence (or change in abundance) of
species A on species C via an intermediary species (A --> B --> C).
3. Cascading effects are those which extend across three or more trophic levels, and
can be top-down (predator --> herbivore --> plant) or bottom-up (plant -->
herbivore --> predator).
4. Keystone species are those which produce strong indirect effects.

The keystone species concept is one of the best-known ideas in community ecology.
 Although it is true that many species potentially interact
with one another in a food web such as depicted in Figure
1, in nature there are big players and little players. The
biggest players of all are referred to as keystone species.
This is a species whose presence or absence, or
substantial increase or decrease in abundance, profoundly
affects other species in the community. Evidence usually
comes from experiments in which one species is added to
or removed from a community. The name derives from
the center stone in an arch supporting its weight by
inward-leaning stones. Removal of the keystone causes
the arch to collapse.

In the rocky inter-tidal zone of Washington state, and in

other, similar areas, starfish have been shown to be
keystone species The entire
community lives on relatively vertical
rock faces in the wave-swept inter-tidal
zone. The community of marine
invertebrates and algae are adapted to
cling or adhere to the rock face, where
most fed upon the small animal life
suspended in the water (plankton). A
bivalve, the mussel Mytilus, is superior
at attaching to rock faces, making it the
competitive dominant. A starfish
(Pisaster) is an effective predator of the
mussels, making space available for
other species, and consequently is
critical to maintaining a diverse
biological community.

Instances are known where a predator so strongly suppresses its prey (herbivores), that the
trophic level below (plants) benefits because it is released from the pressures of herbivory.
Such “top-down” trophic cascades, where the community looks more or less ‘green’
depending on the abundance of predators, are well-known in lakes. We also know of
examples where fertilizing a system, which increases plant growth, results in more
predators, through the increase in abundance of herbivores. This is a “bottom-up” trophic
cascade.

Our understanding of these complex species interactions gives substance to the popular
phrase, the “balance of nature”. One can also appreciate how a human-induced removal of
one species (an extinction event) or the addition on one species (invasion of a community
by a non-native species) could result in harm to many additional species, a topic we will
consider in the second semester.

We will gain a fuller appreciation of the complex, multi-way interactions among species as
we proceed through this series of lectures. However, we can fully appreciate the complexity
of these multi-way interactions, it is helpful to first understand the nuances of the various
two-way interactions. We will develop our understanding of species interactions in ecological
communities based on these building blocks.
Mutualistic Interactions

A mutualism is an interaction where both sides benefit. Pollination is a common mutualistic

interaction. The plant gains gamete transfer, the animal gets nectar (and also pollen).

Facultative mutualisms are beneficial but not essential to survival and reproduction of either
party. Obligate mutualisms are those that are essential to the life of one or both associates.
We will examine an example of each.

A fascinating facultative mutualism involves the Boran people of Africa, and a bird known as
the honey guide. According to rock paintings, humans have
collected honey in Africa for 20,000 years. Human hunting parties
are often joined by the greater honeyguide (Indicator indicator),
which leads them to bee colonies. In unfamiliar areas, the average
search time was 8.9 hr when unguided, but only 3.2 hrs when
guided by the bird. Borans use fire and smoke to drive off the bees,
break open the nest and remove the honey, but leave larvae and
wax behind. The bird gains access to larvae and wax. The use of fire
and smoke reduces the bird's risk of being stung, and humans
increase accessibility of nests. According to the Borans, the
honeyguide informs them of: direction, from the compass bearing of
bird flight; distance, from the duration of the bird's disappearance
and height of perch; and arrival, by the "indicator call". Birds and
Borans can survive without the other, but each benefits from this
facultative mutualism.

A mutualism between certain ants and a

Pseudomyrmex Ants attack a Katydid placed on an
Acacia Plant
small tree, the acacia, provides an excellent
example of an obligate mutualism. This
particular system has been extensively
studied in Costa Rica. The acacia provides a
number of benefits to the ants, including
shelter (hollow thorns), protein (beltian
bodies at tip of leaflets), nectar (secreted
near base of leaves). The ant
(Pseudomyrmex) provides several forms of
protection. It attacks and removes
herbivorous insects, It also removes vines
that might overgrow the acacia, and kills the
growing shoots of nearby plants that might
become competitors. It clears away leaf litter
from near the plant, and since the acacia
grows in a seasonally dry environment where
it occasionally is threatened by fire, the ant's
activities protect the tree from fire damage
as well.

Many other examples of mutualisms may be familiar to you.

  Gut symbionts in herbivores: mammals can't digest cellulose
 endosymbiosis and the origin of eukaryotic cells: mitochondria, flagella, chloroplasts
are thought to be derived from free-living bacteria
 pollination systems
 the coral polyp and its endosymbiont "alga" (actually a dinoflagellate)

Commensalism

When one species benefits, and the other species is neither benefited nor harmed, the
interaction is "+/0". In the southeastern US and in South America, it is common to see
egrets in cattle pastures. They follow the cattle, eating insects that are dislodged or forced
to fly as cattle graze in the field. One might suppose that egrets benefit cattle, by
consuming insects that might compete with cows for food. The interaction would be a
mutualism if this was demonstrated (but it seems a bit far-fetched). Assuming no benefit to
the cattle, this is a commensalism. It often is the case, as this example illustrates, that we
aren't sure if the interaction is "+/O" or "+/+".

The clown fish and anemone also illustrates this point. The clown fish hides from enemies
within the stinging tentacles of a sea anemone, to which the clown fish is immune. Some
report this interaction as a mutualism, arguing that the clownfish drops scraps of food into
the mouth of the anemone. Careful studies have failed to find much support for any benefit
to the anemone, so this appears to be a commensalism.

Summary

Species interactions within ecological webs include four main types of two-way interactions:
mutualism, commensalism, competition, and predation (which includes herbivory and
parasitism). Because of the many linkages among species within a food web, changes to
one species can have far-reaching effects. We will next examine competition and predation,
and then return to a consideration of more complicated indirect and cascading effects.