Diversity and Intermediate Disturbance

The intermediate disturbance hypothesis (Connell 1978) and the diversity-disturbance hypothesis (Huston 1979) are often used to explain the ecological mechanisms determining the high diversity of species found in traditional landscapes (e.g. Blondel 2006): species diversity peaks when communities are exposed to moderate disturbance, in terms of frequency, extent and intensity. This occurs because moderate disturbance (e.g., moderate grazing) creates discontinuities in the ecosystem that allow the maintenance of early successional species while preventing dominance of more competitive species, hence keeping the ecosystem in a transitional state between early and steady-state communities. The management of traditional landscape mosaics (Fig. 8.3), with low intensity farming, moderate grazing and maintenance of forest patches, is often described as an example of intermediate disturbance, and therefore as a promoter of species diversity (Ostermann 1998; Henle et al. 2008). Nonetheless, peaked relationships between species richness and disturbance are not the rule across ecology studies (Mackey and Currie 2001). Peaked curves are more commonly reported by studies covering small spatial scales and in the presence of natural disturbance regimes (Mackey and Currie 2001). In

Fig. 8.3  Minifundia system in a mountain landscape in Northwestern Portugal. (Photo credit: Vânia Proença)

addition, the relationship between taxa richness and the intensity of anthropogenic disturbance regimes is often non-significant (Mackey and Currie 2001), increasing the challenge of predicting the impacts of altered regimes of disturbances on biodiversity.

Effects of Land-Use Change on Disturbance Regimes

Land-use changes caused by rural abandonment can create the conditions for an increase in the frequency and intensity of disturbance events, in particular higher fire risk due to fuel accumulation and shrub encroachment, but may also result in fewer disturbances if disturbance agents, such as domestic grazers or browsers, become residual or even disappear. The trajectory of secondary succession after abandonment depends on several interacting factors and ecological filters, such as the pool of colonizer species in the surrounding landscape, their ability to colonize abandoned patches, soil quality, and, of course, disturbance regime (Cramer 2007). Disturbances will not only exert a selective pressure on community assembly, but will also respond to community structure and composition.

In landscapes where tree density is very low, such as some Mediterranean landscapes, there is a high probability of shrub encroachment after farmland abandonment due to seed limitation, predatory pressure over oak acorns and deficient abiotic conditions, such as poor soils (Acácio et al. 2007). Wildfire will further promote shrub dominance, due to many shrubs' resprouting ability. Wildfires may hence establish a reinforcing feedback loop, leading to community homogenization and a decline in diversity at all scales (Proença and Pereira 2010).

A different trajectory can be anticipated in landscapes with a higher tree density, such as semi-natural grasslands in northern Europe (Eriksson et al. 2002). There, seed availability and dispersal are not limiting factors and forest is able to colonize and regenerate in relatively short time. With an expected low disturbance regime, forest can expand, which would decline habitat heterogeneity. Some species, such as grassland specialists, will show strong reductions in abundance or even go locally extinct. Impacts at the landscape level will depend on species ability to persist in alternative habitats such as forest edges or heathlands (Proença and Pereira 2013).

The above examples describe abandoned patches in a fairly homogenous landscape matrix with either a low or high tree density. In a heterogeneous landscape with a more balanced cover of different habitats and a variety of edaphic-topographic conditions, scenarios would be different given the diversity of local responses to changes in disturbance regime. Habitat diversity will not only counteract landscape homogenization, but also provide alternative habitats for species affected by farmland abandonment, thus reducing the impact of land-use change on species diversity. The persistence of those species in the landscape will then depend on the maintenance of those alternative habitats, either by natural processes, such as herbivory by wild ungulates, or through assisted processes, such as prescribed fire or herbivore re-introduction. Such restoration approaches, either passive or assisted, are an important open question in rewilding research.

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