Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.

Predation and Herbivory

Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure 20.20). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.

The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.

Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.

Defense Mechanisms against Predation and Herbivory

The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.

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. Figure 20.21 shows some organisms’ defenses against predation and herbivory.

Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure 20.22a). In another example, the chameleon can change its color to match its surroundings (Figure 20.22b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.

Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure 20.23).

While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure 20.24).

In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure 20.25 shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.

Competitive Exclusion Principle

Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure 20.26, with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.

This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.

Symbiosis

Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.

Commensalism

A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure 20.27). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.

Mutualism

A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure 20.28a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure 20.28b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.

Parasitism

A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.

The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure 20.29). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.

Characteristics of Communities

Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.

Foundation Species

Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.

Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure 20.30). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.

Biodiversity, Species Richness, and Relative Species Abundance

Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure 20.31). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.

Keystone Species

A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure 20.32). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.

Community Dynamics

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.

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 is 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 (Figure 20.34). These species help to further break down the mineral rich lava into 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 (Figure 20.35). 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.