DISPERSION

in wave motion, any phenomenon associated with the propagation of individual waves at speeds that depend on their wavelengths. Ocean waves, for example, move at speeds proportional to the square root of their wavelengths; these speeds vary from a few feet per second for ripples to hundreds of miles per hour for tsunamis. A wave of light has a speed in a transparent medium that varies inversely with the index of refraction (a measure of the angle by which the direction of a wave is changed as it moves from one medium into another). Any transparent mediume.g., a glass prismwill cause an incident parallel beam of light to fan out according to the refractive index of the glass for each of the component wavelengths, or colours. Dispersion is sometimes called the separation of light into colours, an effect more properly called angular dispersion. in biology, the dissemination, or scattering, of organisms over periods within a given area or over the Earth. The two disciplines most intimately intertwined with the study of dispersion are systematics and evolution. Systematics is concerned with the relationships between organisms and includes the classification of life into ordered groups, providing the detailed information essential to all biology. The study of evolution grew from a combination of systematics and dispersion, or distribution, as both Charles Darwin and Alfred Russel Wallace, pioneers in evolutionary biology, attested; and, in turn, an understanding of the process of natural selection has illuminated the reasons for changes in distribution in the history of the Earth. A specific type of organism can establish one of three possible patterns of dispersion in a given area: a random pattern; an aggregated pattern, in which organisms gather in clumps; or a uniform pattern, with a roughly equal spacing of individuals. The type of pattern often results from the nature of the relationships within the population. Social animals, such as chimpanzees, tend to gather in groups, while territorial animals, such as birds, tend to assume uniform spacing. Close attention must be paid to the scale of study in order to get an accurate reading of these patterns. If a group of monkeys occupies three widely separated trees, their spacing will obviously be aggregate; yet in each tree, their spacing may appear to be uniform. Distribution can be affected by time of day, month, or year. The most common form of distributional change occurs among migratory animals, which may be plentiful in the summer months and virtually absent in the winter. The forces governing the dispersal of organisms are either vectorial (directed motion), that is, caused by wind, water, or some other environmental motion, or stochastic (random), as in the case of the change in seasons, which gives no indication of where the dispersing organisms may ultimately settle. Dispersion may also be affected by the interrelationship of species with one another or with nutrients. Competition between species that depend on the same food types often leads to the elimination of one species, just as the extent of plant life often determines the boundaries of a species' territory. The irregularities of most distribution patterns are simplified in the case of life forms dependent upon relatively restricted habitats, like that of intertidal mollusks, which have an almost linear distribution along rocky seacoasts. A few species, most notably humans and the animals dependent upon them, have a worldwide distribution. Among both plants and animals, dispersal usually takes place at the time of reproduction. When overcrowding forces individuals to range outside the area in which they were born to find a mate or food, new populations occasionally arise. Insects often display distinctive abilities in this regard. East African locusts have been found in two forms, a bright green variety, which is sluggish and solitary, and a highly mobile, group-oriented, dark-coloured form that swarms in enormous numbers, eating all plant material in its path. It has been found that if the young of the green variety are raised in large, constricted groups, they metamorphosize into the dark form at maturity. This is called phase polymorphism. As their numbers increase and the food supply thins, the locusts undergo developmental and behavioral changes to produce the widest dispersion pattern possible. Occasionally, natural selection acts to limit the dispersal of a species. On high mountaintops and isolated islands, for example, the predominance of flightless birds and insects is notable. Organisms are also spread by passive means, such as wind, water, and by other creatures. This method is hardly less effective than active dispersal; spiders, mites, and insects have been collected by airplanes over the Pacific as much as 3,100 km (about 1,900 miles) from land. Plants regularly spread their seeds and spores by the action of the wind and water, often with morphological adaptations to increase their potential range, as in the case of milkweed seeds. Seeds are also spread by animals, often as undigested matter in the excrement of birds or mammals, or by attaching to animals via an assortment of hooks, barbs, and sticky substances. Parasites regularly use either their hosts or other creatures as distribution mechanisms. The myxoma virus, a parasite in rabbits, is carried by mosquitoes, which may travel as far as 64 km (40 miles) before infecting another rabbit. Mountains and oceans can be effective barriers to the dispersal of organisms, as can stretches of desert or other climatological extremes. Some organisms can cross these barriers; birds can cross the English Channel, while bears cannot. In such cases, the paths of the more mobile animals are called filter routes. Over geologic ages there have been many dramatic changes in climate that have affected distribution and even the survival of many life forms. Furthermore, the continents appear to have undergone large-scale displacements (see continental drift), separating many species and encouraging their independent development. But the greatest factor in the dispersal of organisms, at least during the past 10,000 years, has been human influence.