Landscapes do not exist in isolation. Landscapes are nested within larger landscapes that are nested within larger landscapes, and so on. In other words, each landscape has a context or regional setting, regardless of scale and how the landscape is defined. The landscape context may constrain processes operating within the landscape. Landscapes are “open” systems; energy, materials, and organisms move into and out of the landscape. This is especially true in practice, where landscapes are often somewhat arbitrarily delineated. That broad-scale processes act to constrain or influence finer-scale phenomena is one of the key principles of hierarchy theory (Allen and Star 1982) and ‘supply-side’ ecology (Rough garden et al. 1987). The importance of the landscape context is dependent on the phenomenon of interest, but typically varies as a function of the “openness” of the landscape. The “openness” of the landscape depends not only on the phenomenon under consideration, but on the basis used for delineating the landscape boundary. For example, from a geomorphological or hydrological perspective, the watershed forms a natural landscape, and a landscape defined in this manner might be considered relatively “closed”. Of course, energy and materials flow out of this landscape and the landscape context influences the input of energy and materials by affecting climate and so forth, but the system is nevertheless relatively closed. Conversely, from the perspective of a bird population, topographic boundaries may have little ecological relevance, and the landscape defined on the basis of watershed boundaries might be considered a relatively “open” system. Local bird abundance patterns may be produced not only by local processes or events operating within the designated landscape, but also by the dynamics of regional populations or events elsewhere in the species’ range (Wiens 1981, 1989b, Vaisanen et al. 1986, Haila et al. 1987, Ricklefs 1987).
Landscape metrics quantify the pattern of the landscape within the designated landscape boundary only. Consequently, the interpretation of these metrics and their ecological significance requires an acute awareness of the landscape context and the openness of the landscape relative to the phenomenon under consideration. These concerns are particularly important for nearest-neighbour metrics. Nearest-neighbour distances are computed solely from patches contained within the landscape boundary. If the landscape extent is small relative to the scale of the organism or ecological processes under consideration and the landscape is an “open” system relative to that organism or process, then nearest-neighbour results can be misleading. Consider a small subpopulation of a species occupying a patch near the boundary of a somewhat arbitrarily defined (from the organism’s perspective) landscape. The nearest neighbour within the landscape boundary might be quite far away, yet in reality the closest patch might be very close, but just outside the landscape boundary. The magnitude of this problem is a function of scale. Increasing the size of the landscape relative to the scale at which the organism under investigation perceives and responds to the environment will generally decrease the severity of this problem. In general, the larger the ratio of extent to grain (i.e., the larger the landscape relative to the average patch size), the less likely these and other metrics will be dominated by boundary effects.
KEY POINT The important point is that a landscape should be defined relative to both the patch mosaic within the landscape as well as the landscape context. Moreover, consideration should always be given to the landscape context and the openness of the landscape relative to the phenomenon under consideration when choosing and interpreting landscape metrics.