- Biotic elements of communities
- Patterns of community structure
- Interspecific interactions and the organization of communities
- Commensalism and other types of interaction
- The coevolutionary process
- The study of coevolution
- The coevolutionary “arms race” versus reduced antagonism
- Coevolution and the organization of communities
- Gene-for-gene coevolution
- The geographic mosaic theory of coevolution
- Evolution of the biosphere
- General features
- Geologic history and early life-forms
- The progression of evolution
- A period of extensive glaciation and drought: The Permian Period
- The reptilian radiation
- The diversity of Cretaceous biota
- A period of transition
- Quaternary events
Specialization in parasites
It is now evident that the parasitic lifestyle often favours extreme specialization to a single host or a small group of hosts. Living for a long period of time on a single host, a parasite must remain attached within or on its host, avoid the defenses of its host, and obtain all its nutrition from that host. Unlike grazers or predators, parasites cannot move from host to host, supplementing their diet with a variety of foods.
Estimates of the number of species worldwide have risen sharply in recent decades owing to research revealing parasitic species to be much more specific to one host species than previously realized. What once may have been considered a single parasitic species attacking many different host species has often turned out to be a group of very similar, yet distinct, parasitic species, each specialized to its own host. This speciation occurs because different parasitic populations become adapted to living on different hosts and coping with the defenses of these hosts. Over time, many of these different parasite populations evolve into genetically distinct species. It is through the specialization of individuals of a species onto different hosts, ultimately resulting in speciation, that parasitism appears to have become the most common way of life on Earth.
For example, swallowtail butterflies (Papilio) include more than 500 species worldwide. In most species an adult female lays her eggs on a host plant, and, after they hatch, the caterpillars complete their development by feeding parasitically on that plant. In North America there are two groups of these butterflies that have evolved to use different hosts: the tiger swallowtail group and the Old World swallowtail group (Papilio machaon). In the Old World swallowtail group are several species that feed on plants in the carrot family Apiaceae (also called Umbelliferae), with different populations feeding on different plant species. However, one species within this group, the Oregon swallowtail (Papilio oregonius), has become specialized to feed on tarragon sagebrush (Artemisia dracunculus), which is in the plant family Asteracaea (Compositae of some sources). Among the tiger swallowtail group, various members have become specialized to different plant hosts. The eastern tiger swallowtail (Papilio glaucus) has a long list of recorded hosts, but it is now known that the northern and southern populations are adapted to different plant species, and these populations cannot develop on the others’ hosts.
Alternation among hosts
Although many parasitic species complete all developmental stages on a single host individual, thousands of other parasitic species alternate between two or more host species, specializing on a different host species at each developmental stage. Many parasites, from a diverse array of species such as certain viruses, flatworms, nematodes, and aphids, specialize on different host species at different stages of development. Among aphids alone at least 2,700 species alternate among hosts.
Parasites have evolved by three major evolutionary routes to alternate among two or more hosts. Some parasite species have evolved to alternate between their final host and an intermediate host, or vector, that transfers the parasite from one final host to another: the malarial parasite Plasmodium falciparum alternates between a final human host and an intermediate mosquito host by which the parasite is transferred from one person to another. The parasite uses the mosquito as a mobile hypodermic syringe. Examples of a similar kind of transmission between a final host and an intermediate host with piercing mouthparts occur in many other species. Viruses, rickettsias, protozoa, and nematodes all have species that are transmitted between vertebrates through biting flies. Some viruses and other parasites are similarly transmitted between plant species by aphids, whose piercing mouthparts transmit the parasites directly into plant tissues while the aphids are feeding.
Other parasites alternate between a host and the predator that eats it. These parasites have turned an evolutionary problem (being killed along with their host) into an evolutionary opportunity (being transferred to the predator and continuing to feed). As it develops, the parasite attacks hosts higher in the food chain, alternating between herbivore and predator or between an intermediate and a top predator in the food chain. Many parasites alternate between snails or other invertebrates and vertebrate predators that feed upon these invertebrates; others alternate among vertebrate species. The pork tapeworm (Taenia solium), for example, alternates between pigs and humans in societies in which improperly cooked pork is eaten.
Still other parasites employ wings, wind, or water to alternate between hosts. Many aphid species alternate between a summer host and a winter host by producing winged individuals that fly to the new host. Rust fungi such as wheat stem rust can be carried between hosts by wind currents, and the parasite Schistosoma mansoni, which causes the disease schistosomiasis, alternates between Biomphalaria snails and humans by moving through water.
The different ways by which host species are linked to parasites contribute to the complex web of interactions that shape the structure of communities. The effect of parasitism on the dynamics of populations and the organization of communities is still one of the most underexplored topics in ecology.
The strategy of grazing
The word “grazing” conjures up images of large mammals moving through seas of grass. Grazing, however, is a form of interspecific interaction that has been adopted by a number of other groups as well. A grazer is defined as any species that moves from one victim to another, feeding on part of each victim without actually killing it outright. The “victim” is to the grazer as prey is to the predator. Hence, grasshoppers that jump from plant to plant, chewing a portion of the leaves of each one they visit, are grazers, as are caterpillars that crawl from one plant to another during development rather than remain as parasites on an individual plant. The grazing lifestyle differs from the parasitic lifestyle in a few important ways. Individuals can vary their diets with different foods, and, by not remaining attached to a single individual for long periods of time, their victims do not have time to develop induced specialized defenses, such as an immune response that a host can develop against a parasite.
Grazing is more commonly perpetrated on plants than animals because plants have a modular structure that allows a part of them to be lost without the whole individual being destroyed. In contrast, most animals that lose a part of the body to an antagonist die immediately or soon afterward, rendering the interaction an act of predation rather than grazing. An exception to this rule occurs in species that can disconnect body parts—some lizards and salamanders are able to detach their tails if they are attacked by a predator.