Background.--This group of metrics represents a loose collection of metrics that deal with the number and size of patches and the amount of edge created by these patches. Although these metrics could easily be subdivided into separate groups or assigned to other already recognized groups, there is enough similarity in the basic patterns assessed by these metrics to include them under one umbrella.
The area of each patch comprising a landscape mosaic is perhaps the single most important and useful piece of information contained in the landscape. Not only is this information the basis for many of the patch, class, and landscape indices, but patch area has a great deal of ecological utility in its own right. For example, there is considerable evidence that bird species richness and the occurrence and abundance of some species are strongly correlated with patch size (e.g., Robbins et al. 1989). Most species have minimum area requirements: the minimum area needed to meet all life history requirements. Some of these species require that their minimum area requirements be fulfilled in contiguous habitat patches; in other words, the individual habitat patch must be larger than the species minimum area requirement for them to occupy the patch. These species are sometimes referred to as “area-sensitive” species. Thus, patch size information alone could be used to model species richness, patch occupancy, and species distribution patterns in a landscape given the appropriate empirical relationships derived from field studies.
Similarly, the size and number of patches comprising a class or the entire landscape mosaic is perhaps the most basic aspect of landscape pattern that can affect myriad processes. For example, although there are myriad effects of habitat fragmentation on individual behavior, habitat use patterns, and intra- and inter-specific interactions, many of these effects are caused by: (1) a reduction in habitat area (area effects), and (2) an increase in the proportion of edge-influenced habitat (edge effects). Briefly, as a species’ habitat is lost from the landscape (without being fragmented), at some point there will be insufficient area of habitat to support a viable population, and with continued loss eventually there will be insufficient area of habitat to support even a single individual and the species will be extirpated from the landscape. This area relationship is expected to vary among species depending on their minimum area requirements. Moreover, the area threshold for occupancy may occur when total habitat area is still much greater than the individual’s minimum area requirement. For example, an individual may not occupy available habitat unless there are other individuals of the same species occupying the same or nearby patches of habitat, or an individual’s occupancy may be influenced by what other species are occupying the patch. Similarly, as habitat is lost and simultaneously fragmented into smaller and more isolated patches, at some point there will be insufficient area of suitable habitat within a home range size area to support an individual. In either case, the effect of habitat area on the occurrence and abundance of a species (or species) is referred to as the “area effect.” This is the ultimate consequence of habitat loss and fragmentation–insufficient habitat quantity and quality to support individuals and viable populations.
Total amount of edge in a landscape is important to many ecological phenomena. In particular, a great deal of attention has been given to wildlife-edge relationships (Thomas et al. 1978 and 1979, Strelke and Dickson 1980, Morgan and Gates 1982, Logan et al. 1985). In landscape ecological investigations, much of the presumed importance of spatial pattern is related to edge effects. The forest edge effect, for example, results primarily from differences in wind and light intensity and quality reaching a forest patch that alter microclimate and disturbance rates (e.g., Gratkowski 1956, Ranney et al. 1981, Chen and Franklin 1990). These changes, in combination with changes in seed dispersal and herbivory, can influence vegetation composition and structure (Ranney et al. 1981). The proportion of a forest patch that is affected in this manner is dependent, therefore, upon patch shape and orientation, and by adjacent land cover. A large but convoluted patch, for example, could be entirely edge habitat. It is now widely accepted that edge effects must be viewed from an organism-centered perspective because edge effects influence organisms differently; some species have an affinity for edges, some are unaffected, and others are adversely affected.
One of the most dramatic and well-studied consequences of habitat fragmentation is an increase in the proportional abundance of edge-influenced habitat. Early wildlife management efforts were focused on maximizing edge habitat because it was believed that most species favored habitat conditions created by edges and that the juxtaposition of different habitats would increase species diversity (Leopold 1933). Indeed this concept of edge as a positive influence guided land management practices for most of the twentieth century. Recent studies, however, have suggested that changes in microclimate, vegetation, invertebrate populations, predation, brood parasitism, and competition along forest edges (i.e., edge effects) has resulted in the population declines of several vertebrate species dependent upon forest interior conditions (e.g., Strelke and Dickson 1980, Whitcomb et al. 1981, Kroodsma 1982, Brittingham and Temple 1983, Wilcove 1985, Temple 1986, Noss 1988, Yahner and Scott 1988, Robbins et al. 1989). In fact, many of the adverse effects of forest fragmentation on organisms seem to be directly or indirectly related to these so-called edge effects. Forest interior species, therefore, may be sensitive to patch shape because for a given patch size, the more complex the shape, the larger the edge-to-interior ratio. Total class edge in a landscape, therefore, often is the most critical piece of information in the study of fragmentation, and many of the class indices directly or indirectly reflect the amount of class edge. Similarly, the total amount of edge in a landscape is directly related to the degree of spatial heterogeneity in that landscape.