Chapter 19. Population and Community Ecology

The most accurate way to determine population size is to count all of the individuals within the area. However, this method is usually not logistically or economically feasible, especially when studying large areas. Thus, scientists usually study populations by sampling a representative portion of each habitat and use this sample to make inferences about the population as a whole. The methods used to sample populations to determine their size and density are typically tailored to the characteristics of the organism being studied. For immobile organisms such as plants, or for very small and slow-moving organisms, a quadrat may be used. A quadrat is a wood, plastic, or metal square that is randomly located on the ground and used to count the number of individuals that lie within its boundaries. To obtain an accurate count using this method, the square must be placed at random locations within the habitat enough times to produce an accurate estimate. This counting method will provide an estimate of both population size and density. The number and size of quadrat samples depends on the type of organisms and the nature of their distribution.

For smaller mobile organisms, such as mammals, a technique called mark and recapture is often used. This method involves marking a sample of captured animals in some way and releasing them back into the environment to mix with the rest of the population; then, a new sample is captured and scientists determine how many of the marked animals are in the new sample. This method assumes that the larger the population, the lower the percentage of marked organisms that will be recaptured since they will have mixed with more unmarked individuals. For example, if 80 field mice are captured, marked, and released into the forest, then a second trapping 100 field mice are captured and 20 of them are marked, the population size (N) can be determined using the following equation:

(19.1)

Using our example, the population size would be 400.

(19.2)

These results give us an estimate of 400 total individuals in the original population. The true number usually will be a bit different from this because of chance errors and possible bias caused by the sampling methods.

Life tables provide important information about the life history of an organism and the life expectancy of individuals at each age. They are modeled after actuarial tables used by the insurance industry for estimating human life expectancy. Life tables may include the probability of each age group dying before their next birthday, the percentage of surviving individuals dying at a particular age interval (their mortality rate, and their life expectancy at each interval. An example of a life table is shown in Table 19.1 from a study of Dall mountain sheep, a species native to northwestern North America. Notice that the population is divided into age intervals (column A). The mortality rate (per 1000) shown in column D is based on the number of individuals dying during the age interval (column B), divided by the number of individuals surviving at the beginning of the interval (Column C) multiplied by 1000.

(19.3)

For example, between ages three and four, 12 individuals die out of the 776 that were remaining from the original 1000 sheep. This number is then multiplied by 1000 to give the mortality rate per thousand.

(19.4)

As can be seen from the mortality rate data (column D), a high death rate occurred when the sheep were between six months and a year old, and then increased even more from 8 to 12 years old, after which there were few survivors. The data indicate that if a sheep in this population were to survive to age one, it could be expected to live another 7.7 years on average, as shown by the life-expectancy numbers in column E.

Another tool used by population ecologists is a survivorship curve, which is a graph of the number of individuals surviving at each age interval versus time. These curves allow us to compare the life histories of different populations (Figure 19.4). There are three types of survivorship curves. In a type I curve, mortality is low in the early and middle years and occurs mostly in older individuals. Organisms exhibiting a type I survivorship typically produce few offspring and provide good care to the offspring increasing the likelihood of their survival. Humans and most mammals exhibit a type I survivorship curve. In type II curves, mortality is relatively constant throughout the entire life span, and mortality is equally likely to occur at any point in the life span. Many bird populations provide examples of an intermediate or type II survivorship curve. In type III survivorship curves, early ages experience the highest mortality with much lower mortality rates for organisms that make it to advanced years. Type III organisms typically produce large numbers of offspring, but provide very little or no care for them. Trees and marine invertebrates exhibit a type III survivorship curve because very few of these organisms survive their younger years, but those that do make it to an old age are more likely to survive for a relatively long period of time.

Figure 19.4. 

Survivorship curves show the distribution of individuals in a population according to age. Humans and most mammals have a Type I survivorship curve, because death primarily occurs in the older years. Birds have a Type II survivorship curve, as death at any age is equally probable. Trees have a Type III survivorship curve because very few survive the younger years, but after a certain age, individuals are much more likely to survive.

Charles Darwin, in developing his theory of natural selection, was influenced by the English clergyman Thomas Malthus. Malthus published his book in 1798 stating that populations with abundant natural resources grow very rapidly; however, they limit further growth by depleting their resources. The early pattern of accelerating population size is called exponential growth.

The best example of exponential growth in organisms is seen in bacteria. Bacteria are prokaryotes that reproduce largely by binary fission. This division takes about an hour for many bacterial species. If 1000 bacteria are placed in a large flask with an abundant supply of nutrients (so the nutrients will not become quickly depleted), the number of bacteria will have doubled from 1000 to 2000 after just an hour. In another hour, each of the 2000 bacteria will divide, producing 4000 bacteria. After the third hour, there should be 8000 bacteria in the flask. The important concept of exponential growth is that the growth rate—the number of organisms added in each reproductive generation—is itself increasing; that is, the population size is increasing at a greater and greater rate. After 24 of these cycles, the population would have increased from 1000 to more than 16 billion bacteria. When the population size, N, is plotted over time, a J-shaped growth curve is produced (Figure 19.5a).

The bacteria-in-a-flask example is not truly representative of the real world where resources are usually limited. However, when a species is introduced into a new habitat that it finds suitable, it may show exponential growth for a while. In the case of the bacteria in the flask, some bacteria will die during the experiment and thus not reproduce; therefore, the growth rate is lowered from a maximal rate in which there is no mortality. The growth rate of a population is largely determined by subtracting the death rate, D, (number organisms that die during an interval) from the birth rate, B, (number organisms that are born during an interval). The growth rate can be expressed in a simple equation that combines the birth and death rates into a single factor: r. This is shown in the following formula:

The value of r can be positive, meaning the population is increasing in size (the rate of change is positive); or negative, meaning the population is decreasing in size; or zero, in which case the population size is unchanging, a condition known as zero population growth.

In the real world, with its limited resources, exponential growth cannot continue indefinitely. Exponential growth may occur in environments where there are few individuals and plentiful resources, but when the number of individuals gets large enough, resources will be depleted and the growth rate will slow down. Eventually, the growth rate will plateau or level off (Figure 19.5b). This population size, which is determined by the maximum population size that a particular environment can sustain, is called the carrying capacity, or K. In real populations, a growing population often overshoots its carrying capacity, and the death rate increases beyond the birth rate causing the population size to decline back to the carrying capacity or below it. Most populations usually fluctuate around the carrying capacity in an undulating fashion rather than existing right at it.

The formula used to calculate logistic growth adds the carrying capacity as a moderating force in the growth rate. The expression “K – N” is equal to the number of individuals that may be added to a population at a given time, and “K – N” divided by “K” is the fraction of the carrying capacity available for further growth. Thus, the exponential growth model is restricted by this factor to generate the logistic growth equation:

(19.6)

Notice that when N is almost zero the quantity in brackets is almost equal to 1 (or K/K) and growth is close to exponential. When the population size is equal to the carrying capacity, or N = K, the quantity in brackets is equal to zero and growth is equal to zero. A graph of this equation (logistic growth) yields the S-shaped curve (Figure 19.5b). It is a more realistic model of population growth than exponential growth. There are three different sections to an S-shaped curve. Initially, growth is exponential because there are few individuals and ample resources available. Then, as resources begin to become limited, the growth rate decreases. Finally, the growth rate levels off at the carrying capacity of the environment, with little change in population number over time.

Figure 19.5. 

When resources are unlimited, populations exhibit (a) exponential growth, shown in a J-shaped curve. When resources are limited, populations exhibit (b) logistic growth. In logistic growth, population expansion decreases as resources become scarce, and it levels off when the carrying capacity of the environment is reached. The logistic growth curve is S-shaped.

The logistic model assumes that every individual within a population will have equal access to resources and, thus, an equal chance for survival. For plants, the amount of water, sunlight, nutrients, and space to grow are the important resources, whereas in animals, important resources include food, water, shelter, nesting space, and mates.

In the real world, phenotypic variation among individuals within a population means that some individuals will be better adapted to their environment than others. The resulting competition for resources among population members of the same species is termed intraspecific competition. Intraspecific competition may not affect populations that are well below their carrying capacity, as resources are plentiful and all individuals can obtain what they need. However, as population size increases, this competition intensifies. In addition, the accumulation of waste products can reduce carrying capacity in an environment.

Yeast, a microscopic fungus used to make bread and alcoholic beverages, exhibits the classical S-shaped curve when grown in a test tube (Figure 19.6a). Its growth levels off as the population depletes the nutrients that are necessary for its growth. In the real world, however, there are variations to this idealized curve. Examples in wild populations include sheep and harbor seals (Figure 19.6b). In both examples, the population size exceeds the carrying capacity for short periods of time and then falls below the carrying capacity afterwards. This fluctuation in population size continues to occur as the population oscillates around its carrying capacity. Still, even with this oscillation, the logistic model is confirmed.

Art Connection

 

Figure 19.6. 

(a) Yeast grown in ideal conditions in a test tube shows a classical S-shaped logistic growth curve, whereas (b) a natural population of seals shows real-world fluctuation. The yeast is visualized using differential interference contrast light micrography. (credit a: scale-bar data from Matt Russell)

If the major food source of seals declines due to pollution or overfishing, which of the following would likely occur?

1. The carrying capacity of seals would decrease, as would the seal population.
2. The carrying capacity of seals would decrease, but the seal population would remain the same.
3. The number of seal deaths would increase, but the number of births would also increase, so the population size would remain the same.
4. The carrying capacity of seals would remain the same, but the population of seals would decrease.

Predation and predator avoidance are strong selective agents. Any heritable character that allows an individual of a prey population to better evade its predators will be represented in greater numbers in later generations. Likewise, traits that allow a predator to more efficiently locate and capture its prey will lead to a greater number of offspring and an increase in the commonness of the trait within the population. Such ecological relationships between specific populations lead to adaptations that are driven by reciprocal evolutionary responses in those populations. Species have evolved numerous mechanisms to escape predation and herbivory (the consumption of plants for food). Defenses may be mechanical, chemical, physical, or behavioral.

Mechanical defenses, such as the presence of armor in animals or thorns in plants, discourage predation and herbivory by discouraging physical contact (Figure 19.14a). Many animals produce or obtain chemical defenses from plants and store them to prevent predation. Many plant species produce secondary plant compounds that serve no function for the plant except that they are toxic to animals and discourage consumption. For example, the foxglove produces several compounds, including digitalis, that are extremely toxic when eaten (Figure 19.14b). (Biomedical scientists have purposed the chemical produced by foxglove as a heart medication, which has saved lives for many decades.)

Figure 19.14. 

The (a) honey locust tree uses thorns, a mechanical defense, against herbivores, while the (b) foxglove uses a chemical defense: toxins produces by the plant can cause nausea, vomiting, hallucinations, convulsions, or death when consumed. (credit a: modification of work by Huw Williams; credit b: modification of work by Philip Jägenstedt)

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 it is stationary against a background of real twigs (Figure 19.15a). In another example, the chameleon can change its color to match its surroundings (Figure 19.15b).

Figure 19.15. 

(a) The tropical walking stick and (b) the chameleon use their body shape and/or coloration to prevent detection by predators. (credit a: modification of work by Linda Tanner; credit b: modification of work by Frank Vassen)

Some species use coloration as a way of warning predators that they are distasteful or poisonous. For example, the monarch butterfly caterpillar sequesters poisons from its food (plants and milkweeds) to make itself poisonous or distasteful to potential predators. The caterpillar is bright yellow and black to advertise its toxicity. The caterpillar is also able to pass the sequestered toxins on to the adult monarch, which is also dramatically colored black and red as a warning to potential predators. Fire-bellied toads produce toxins that make them distasteful to their potential predators. They have bright red or orange coloration on their bellies, which they display to a potential predator to advertise their poisonous nature and discourage an attack. These are only two examples of warning coloration, which is a relatively common adaptation. Warning coloration only works if a predator uses eyesight to locate prey and can learn—a naïve predator must experience the negative consequences of eating one before it will avoid other similarly colored individuals (Figure 19.16).

Figure 19.16. 

The fire-bellied toad has bright coloration on its belly that serves to warn potential predators that it is toxic. (credit: modification of work by Roberto Verzo)

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 some cases of mimicry, a harmless species imitates the warning coloration of a harmful species. Assuming they share the same predators, this coloration then protects the harmless ones. Many insect species mimic the coloration of wasps, which are stinging, venomous insects, thereby discouraging predation (Figure 19.17).

Figure 19.17. 

One form of mimicry is when a harmless species mimics the coloration of a harmful species, as is seen with the (a) wasp (Polistes sp.) and the (b) hoverfly (Syrphus sp.). (credit: modification of work by Tom Ings)

In other cases of mimicry, multiple species share the same warning coloration, but all of them actually have defenses. The commonness of the signal improves the compliance of all the potential predators. Figure 19.18 shows a variety of foul-tasting butterflies with similar coloration.

Figure 19.18. 

Several unpleasant-tasting Heliconius butterfly species share a similar color pattern with better-tasting varieties, an example of mimicry. (credit: Joron M, Papa R, Beltrán M, Chamberlain N, Mavárez J, et al.)

Concept in Action

Go to this website to view stunning examples of mimicry.

A commensal relationship occurs when one species benefits from a close prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure 19.20). 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. Many potential commensal relationships are difficult to identify because it is difficult to prove that one partner does not derive some benefit from the presence of the other.

Figure 19.20. 

The southern masked-weaver is starting to make a nest in a tree in Zambezi Valley, Zambia. This is an example of a commensal relationship, in which one species (the bird) benefits, while the other (the tree) neither benefits nor is harmed. (credit: “Hanay”/Wikimedia Commons)

A second type of symbiotic relationship is called mutualism, in which two species benefit from their interaction. For example, termites have a mutualistic relationship with protists that live in the insect’s gut (Figure 19.21a). The termite benefits from the ability of the protists to digest cellulose. However, the protists are able to digest cellulose only because of the presence of symbiotic bacteria within their cells that produce the cellulase enzyme. The termite itself cannot do this: without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. In turn, the protists benefit from the enzymes provided by their bacterial endosymbionts, while the bacteria benefit from a doubly protective environment and a constant source of nutrients from two hosts. Lichen are a mutualistic relationship between a fungus and photosynthetic algae or cyanobacteria (Figure 19.21b). 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. The algae of lichens can live independently given the right environment, but many of the fungal partners are unable to live on their own.

Figure 19.21. 

(a) Termites form a mutualistic relationship with symbiotic protozoa in their guts, which allow both organisms to obtain energy from the cellulose the termite consumes. (b) Lichen is a fungus that has symbiotic photosynthetic algae living in close association. (credit a: modification of work by Scott Bauer, USDA; credit b: modification of work by Cory Zanker)

A parasite is an organism that feeds off another without immediately killing the organism it is feeding on. 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. Parasites may kill their hosts, but there is usually selection to slow down this process to allow the parasite time to complete its reproductive cycle before it or its offspring are able to spread to another host.

The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure 19.22). The tapeworm can live inside the intestine of the host for several years, benefiting from the host’s food, and it may grow to be over 50 feet long by adding segments. The parasite moves from one host species to a second host species in order to complete its life cycle. Plasmodium falciparum is another parasite: the protists that cause malaria, a significant disease in many parts of the world. Living inside human liver and red blood cells, the organism reproduces asexually in the human host and then sexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to mosquito and back to human, one of many arthropod-borne infectious diseases of humans.

Figure 19.22. 

This diagram shows the life cycle of the tapeworm, a human worm parasite. (credit: modification of work by CDC)

Concept in Action

To learn more about “Symbiosis in the Sea,” watch this webisode of Jonathan Bird’s Blue World.

Ecologists have extensively studied one of the fundamental characteristics of communities: biodiversity. One measure of biodiversity used by ecologists is the number of different species in a particular area and their relative abundance. The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term used to describe the number of species living in a habitat or other unit. Species richness varies across the globe (Figure 19.23). Ecologists have struggled to understand the determinants of biodiversity. Species richness is related to latitude: the greatest species richness occurs near the equator and the lowest richness occurs near the poles. Other factors influence species richness as well. Island biogeography attempts to explain the great species richness found in isolated islands, and has found relationships between species richness, island size, and distance from the mainland.

Relative species abundance is the number individuals in a species relative to the total number of individuals in all species within a system. Foundation species, described below, often have the highest relative abundance of species.

Figure 19.23. 

The greatest species richness for mammals in North America is associated in the equatorial latitudes. (credit: modification of work by NASA, CIESIN, Columbia University)

Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are often primary producers, and they are typically an abundant organism. For example, kelp, a species of brown algae, is a foundation species that forms 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. Examples include the kelp described above or tree species found in a forest. The photosynthetic corals of the coral reef also provide structure by physically modifying the environment (Figure 19.24). The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.

Figure 19.24. 

Coral is the foundation species of coral reef ecosystems. (credit: Jim E. Maragos, USFWS)

A keystone species is one whose presence has inordinate influence in maintaining the prevalence of various species in an ecosystem, the ecological community’s structure, and sometimes its biodiversity. Pisaster ochraceus, the intertidal sea star, is a keystone species in the northwestern portion of the United States (Figure 19.25). Studies have shown that when this organism is removed from communities, mussel populations (their natural prey) increase, which completely alters the species composition and reduces 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. The banded tetra feeds largely on insects from the terrestrial ecosystem and then excretes phosphorus into the aquatic ecosystem. The relationships between populations in the community, and possibly the biodiversity, would change dramatically if these fish were to become extinct.

Figure 19.25. 

The Pisaster ochraceus sea star is a keystone species. (credit: Jerry Kirkhart)

Primary succession occurs when new land is formed, 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 to it its size each year. Weathering and other natural forces break down the rock enough for the establishment of hearty species such as lichens and some plants, known as pioneer species (Figure 19.26). These species help to further break down the mineral-rich lava into soil where other, less hardy but more competitive species, such as grasses, shrubs, and trees, will grow and eventually replace the pioneer species. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.

Figure 19.26. 

During primary succession in lava on Maui, Hawaii, succulent plants are the pioneer species. (credit: Forest and Kim Starr)

A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure 19.27). Wildfires will burn most vegetation, and unless the animals can flee the area, they are killed. Their nutrients, however, are returned to the ground in the form of ash. Thus, although the community has been dramatically altered, there is a soil ecosystem present that provides a foundation for rapid recolonization.

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, at least in part, to changes in the environment brought on by the growth of grasses and forbs, over many years, shrubs 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 and resemble the community before the fire. This equilibrium state is referred to as the climax community, which will remain until the next disturbance. The climax community is typically characteristic of a given climate and geology. Although the community in equilibrium looks the same once it is attained, the equilibrium is a dynamic one with constant changes in abundance and sometimes species identities. The return of a natural ecosystem after agricultural activities is also a well-documented secondary succession process.

Figure 19.27. 

Secondary succession is seen in an oak and hickory forest after a forest fire. A sequence of the community present at three successive times at the same location is depicted.