Source–sink dynamics

Although the seeds of a source–sink model had been planted earlier, Pulliam is often recognized as the first to present a fully developed source–sink model. He defined source and sink patches in terms of their demographic parameters, or BIDE rates (birth, immigration, death, and emigration rates). In the source patch, birth rates were greater than death rates, causing the population to grow. The excess individuals were expected to leave the patch, so that emigration rates were greater than immigration rates. In other words, sources were a net exporter of individuals. In contrast, in a sink patch, death rates were greater than birth rates, resulting in a population decline toward extinction unless enough individuals emigrated from the source patch. Immigration rates were expected to be greater than emigration rates, so that sinks were a net importer of individuals. As a result, there would be a net flow of individuals from the source to the sink (see Table 1). Pulliam's work was followed by many others who developed and tested the source–sink model. Watkinson and Sutherland presented a phenomenon in which high immigration rates could cause a patch to appear to be a sink by raising the patch's population above its carrying capacity (the number of individuals it can support). However, in the absence of immigration, the patches are able to support a smaller population. Since true sinks cannot support any population, the authors called these patches "pseudo-sinks". Definitively distinguishing between true sinks and pseudo-sinks requires cutting off immigration to the patch in question and determining whether the patch is still able to maintain a population. Thomas et al. were able to do just that, taking advantage of an unseasonable frost that killed off the host plants for a source population of Edith's checkerspot butterfly (Euphydryas editha). Without the host plants, the supply of immigrants to other nearby patches was cut off. Although these patches had appeared to be sinks, they did not become extinct without the constant supply of immigrants. They were capable of sustaining a smaller population, suggesting that they were in fact pseudo-sinks. Watkinson and Sutherland's who argued that differentiating between sources and sinks themselves may be difficult. She asserted that a long-term study of the demographic parameters of the populations in each patch is necessary. Otherwise, temporary variations in those parameters, perhaps due to climate fluctuations or natural disasters, may result in a misclassification of the patches. For example, Johnson described periodic flooding of a river in Costa Rica which completely inundated patches of the host plant for a rolled-leaf beetle (Cephaloleia fenestrata). During the floods, these patches became sinks, but at other times they were no different from other patches. If researchers had not considered what happened during the floods, they would not have understood the full complexity of the system. Dias described a source—pseudo-sink inversion in butterfly populations of E. editha. who examined wood thrush (Hylocichla mustelina) survey data for evidence of source and sink populations on a large scale. The authors reasoned that emigrants from sources would likely be the juveniles produced in one year dispersing to reproduce in sinks in the next year, producing a one-year time lag between population changes in the source and in the sink. Using data from the Breeding Bird Survey, an annual survey of North American birds, they looked for relationships between survey sites showing such a one-year time lag. They found several pairs of sites showing significant relationships 60–80 km apart. Several appeared to be sources to more than one sink, and several sinks appeared to receive individuals from more than one source. In addition, some sites appeared to be a sink to one site and a source to another (see Figure 1). The authors concluded that source–sink dynamics may occur on continental scales. One of the more confusing issues involves identifying sources and sinks in the field. Runge et al. and ecological statistics in order to differentiate sources and sinks. ==Modes of dispersal==

Why would individuals ever leave high quality source habitat for a low quality sink habitat? This question is central to source–sink theory. Ultimately, it depends on the organisms and the way they move and distribute themselves between habitat patches. For example, plants disperse passively, relying on other agents such as wind or water currents to move seeds to another patch. Passive dispersal can result in source–sink dynamics whenever the seeds land in a patch that cannot support the plant's growth or reproduction. Winds may continually deposit seeds there, maintaining a population even though the plants themselves do not successfully reproduce. Another good example for this case are soil protists. Soil protists also disperse passively, relying mainly on wind to colonize other sites. As a result, source–sink dynamics can arise simply because external agents dispersed protist propagules (e.g., cysts, spores), forcing individuals to grow in a poor habitat. In contrast, many organisms that disperse actively should have no reason to remain in a sink patch, provided the organisms are able to recognize it as a poor quality patch (see discussion of ecological traps). The reasoning behind this argument is that organisms are often expected to behave according to the "ideal free distribution", which describes a population in which individuals distribute themselves evenly among habitat patches according to how many individuals the patch can support. When there are patches of varying quality available, the ideal free distribution predicts a pattern of "balanced dispersal". For example, the dominant, older individuals in a population may occupy all of the best territories in the source so that the next best territory available may be in the sink. As the subordinate, younger individuals age, they may be able to take over territories in the source, but new subordinate juveniles from the source will have to move to the sink. Pulliam argued that such a pattern of dispersal can maintain a large sink population indefinitely. Furthermore, if good breeding sites in the source are rare and poor breeding sites in the sink are common, it is even possible that the majority of the population resides in the sink. ==Importance in ecology==

The source–sink model of population dynamics has made contributions to many areas in ecology. For example, a species' niche was originally described as the environmental factors required by a species to carry out its life history, and a species was expected to be found only in areas that met these niche requirements. This concept of a niche was later termed the "fundamental niche", and described as all of the places a species could successfully occupy. In contrast, the "realized niche", was described as all of the places a species actually did occupy, and was expected to be less than the extent of the fundamental niche as a result of competition with other species. However, the source–sink model demonstrated that the majority of a population could occupy a sink which, by definition, did not meet the niche requirements of the species, Though some patches may go extinct, the regional persistence of the metapopulation depends on the ability of patches to be re-colonized. As long as there are source patches present for successful reproduction, sink patches may allow the total number of individuals in the metapopulation to grow beyond what the source could support, providing a reserve of individuals available for re-colonization. Source–sink dynamics also has implications for studies of the coexistence of species within habitat patches. Because a patch that is a source for one species may be a sink for another, coexistence may actually depend on immigration from a second patch rather than the interactions between the two species. Finally, the source–sink model has greatly influenced ecological trap theory, a model in which organisms prefer sink habitat over source habitat. Besides being ecological trap sink habitat may vary in their response i major disturbance and colonization of sink habitat may allow species survival even if population in source habitat extinct due to some catastrophic event which may substantially increase metapopulational stability. ==Conservation==

Land managers and conservationists have become increasingly interested in preserving and restoring high quality habitat, particularly where rare, threatened, or endangered species are concerned. As a result, it is important to understand how to identify or create high quality habitat, and how populations respond to habitat loss or change. Because a large proportion of a species' population could exist in sink habitat, conservation efforts may misinterpret the species' habitat requirements. Similarly, without considering the presence of a trap, conservationists might mistakenly preserve trap habitat under the assumption that an organism's preferred habitat was also good quality habitat. Simultaneously, source habitat may be ignored or even destroyed if only a small proportion of the population resides there. Degradation or destruction of the source habitat will, in turn, impact the sink or trap populations, potentially over large distances. Either way, determining which areas are sources or sinks for any one species may be very difficult, and an area that is a source for one species may be unimportant to others. Finally, areas that are sources or sinks currently may not be in the future as habitats are continually altered by human activity or climate change. Few areas can be expected to be universal sources, or universal sinks. While the presence of source, sink, or trap patches must be considered for short-term population survival, especially for very small populations, long-term survival may depend on the creation of networks of reserves that incorporate a variety of habitats and allow populations to interact. == See also ==