It can be argued that although fragmentation disrupts adaptation, it may partially compensate in promoting biodiversity by facilitating the evolutionary process of speciation (ref. 1, p. 75). This idea is based on the idea of founder-induced speciation (36-40). Of these models, the theory of genetic transilience is not just a theory of how founder events can induce speciation but rather primarily of why the vast majority of founder events do not induce speciation (40). Very restrictive conditions must hold before a founder event is likely to trigger speciation (40), conditions of: innate properties (e.g., genomic recombination size, system of mating), historical properties (e.g., the nature of the ancestral population structure, founder numbers, the manner in which the founders were sampled), and ecological factors (the requirement for a rapid increase in population size shortly after the founder event). Recently there have been empirical tests of genetic transilience (41, 42), and the results have supported the predictions of genetic transilience theory, in its predictions both in factors favoring founder-induced speciation and those preventing such speciation (43, 44).
When conditions are favorable for genetic transilience, they can lead not only to explosive speciation rates but also to major adaptive breakthroughs and innovations and to the evolution of higher taxa. For example, the Hawaiian Drosophila have the right combination of innate and historical properties in an appropriate ecological context for genetic transilience (38). The Hawaiian Drosophila not only represent the most speciose group of Drosophila; they also display an extraordinary range of morphological, developmental, and ecological diversity for the genus as a whole and have led to the creation of new genera (45). The ecological context in this case consisted of the regular creation of new volcanic islands to serve as sites of colonization from the older islands. This ecological context creates a situation in which rare interisland founder events to newer islands should lead to explosive population growth after the founder event because of open ecological niches. Such rapid population growth shortly after the founder event is a critical and essential element to speciation via genetic transilience (40).
The requirement of rapid population growth immediately after the founder event means that founder events are likely to induce speciation only in environmental contexts of open or expanding ecological opportunities. However, the founder events induced by human fragmentation are often characterized by diminished, not enhanced, ecological opportunity. Consequently, we expect most human-induced fragmentation events to reduce genetic diversity and increase local extinction with no compensating facilitation of speciation. We know of no compelling examples, in either nature or the laboratory, of speciation via founder events without the flush phase of rapid population growth after the founder event.
Studies of the eastern collared lizard illustrate a fate of rapid local extinction after founder events induced by fragmentation. As noted above in our work on Stegall Mountain, under a fire regime, collared lizards successfully exploit small glade habitats as feeding and breeding territories. Once isolated (as they were when fires were suppressed), these small glade populations must inevitably go extinct. Since 1981, we have surveyed 130 glades in the northeastern Ozarks that had open areas that were as large as or larger than other nearby glades that had a population of collared lizards. Of these larger glades, collared lizard populations were still on 42 of them, indicating that 68% of these glades have experienced local extinction with no subsequent recolonization under the extreme fragmentation induced by fire suppression. This calculation assumes that all 130 glades had collared lizards before fire suppression occurred. In light of the fact that all these glades are close to a currently inhabited glade, this seems to be a reasonable assumption, given the results obtained at the Peck Ranch that lizards readily disperse to nearby glades when frequent fires occur. Indeed, we feel that this percentage is undoubtedly an underestimate of local extinction on larger glades, because we primarily surveyed areas with prior reports that collared lizards were present.
This local extinction process was directly observed for one glade, Victoria Glade. Because of its proximity to St. Louis, this glade has been included in a large number of scientific studies and is a common destination of field trips sponsored by Washington University and other local universities. As a consequence, there is excellent documentation of the plants and animals on this glade since the early 1950s. In the 1950s, this glade had a healthy population of collared lizards, but because of a lack of fire, eastern red cedars began to encroach on the glade, thereby destroying the open microhabitat essential for collared lizards. By 1962, the lizards had become extinct (O. Sexton, personal communication). In the 1980s, this glade was purchased in part by the MDOC and in part by The Nature Conservancy, both agencies initiating a management regime of clearing and burning (only the glade proper was initially burned and not the surrounding oak-hickory forest). By 1990, the glade had been returned to excellent condition (as judged by the plant community), but no collared lizards had recolonized the glade despite the existence of nearby natural populations on private property. Hence, fragmentation of the collared lizards in the eastern Ozarks has resulted in much local extinction without compensatory recolonization events. (This glade was subsequently recolonized by collared lizards, but only after fire management included the surrounding forest.) This situation resulted in an "extinction ratchet" (16, 46), in which each local extinction brings the total population closer and closer to global extinction. An extinction ratchet, not speciation, is the primary impact of human-induced fragmentation.