Ross A. Bradstock

Plant functional traits in relation to fire in crown-fire ecosystems

Disturbance is a dominant factor in many ecosystems, and the disturbance regime is likely to change over the next decades in response to land-use changes and global warming. We assume that predictions of vegetation dynamics can be made on the basis of a set of life-history traits that characterize the response of a species to disturbance. For crown-fire ecosystems, the main plant traits related to postfire persistence are the ability to resprout (persistence of individuals) and the ability to retain a persistent seed bank (persistence of populations). In this context, we asked (1) to what extent do different life- history traits co-occur with the ability to resprout and/or the ability to retain a persistent seed bank among differing ecosystems and (2) to what extent do combinations of fire- related traits (fire syndromes) change in a fire regime gradient? We explored these questions by reviewing the literature and analyzing databases compiled from different crown-fire ecosystems (mainly eastern Australia, California, and the Mediterranean basin). The review suggests that the pattern of correlation between the two basic postfire persistent traits and other plant traits varies between continents and ecosystems. From these results we predict, for instance, that not all resprouters respond in a similar way everywhere because the associated plant traits of resprouter species vary in different places. Thus, attempts to generalize predictions on the basis of the resprouting capacity may have limited power at a global scale. An example is presented for Australian heathlands. Considering the com- bination of persistence at individual (resprouting) and at population (seed bank) level, the predictive power at local scale was significantly increased.

Which mosaic? A landscape ecological approach for evaluating interactions between fire regimes, habitat and animals

The link between ‘fire mosaics’ and persistence of animal species is part of a prominent ecological/land management paradigm. This paradigm deals largely with the effects of fire on animals on the basis of individual events. The universality of the paradigm can be questioned on a variety of grounds, a major deficiency being the inability to deal with quantitative effects of recurrent fire (the fire regime). A conceptual model of fire-related habitat elements is proposed for exploration of a continuum of species/habitat/landscape/fire regime combinations. This approach predicts that the dependence of species on fire-mediated habitat heterogeneity will be highly variable and strongly context-dependent. A spatially explicit simulation model was used to examine the persistence of malleefowl (Leipoa ocellata) in a specific landscape/habitat context where dependence on fire-mosaics should be high. Results suggest that persistence of L. ocellata populations will be dependent on intervention using small patchy fires but that there is an optimum rate of intervention. Results were sensitive to spatial pattern of prescribed fire, landscape type (topography) and probability of wildfire. Underlying effects of the fire-interval distribution (the ‘invisible’ mosaic) on plant species and habitat account for these results. A management emphasis on species/landscape context and awareness of the ‘invisible’ mosaic is advocated.

Fire in Mediterranean Ecosystems: Fire in California

On the west coast of North America lies the state of California, USA (Fig. 5.1), the bulk of which is dominated by a mediterranean-type climate (MTC). Elevations range from sea level to over 4000 m. Mountain ranges are largely oriented north to south with a major valley between the coastal ranges and the interior Sierra Nevada range. In the rain shadow east of the interior mountain ranges the climate is more continental with much colder winters and increasing proportion of summer precipitation eastward. This easternmost part of the state has steppe climates in the northern portion and desert climates in the south. In Arizona and a few other parts of southwestern USA and northeastern Mexico are disjunct patches of sclerophyllous-leaved vegetation that closely resembles California MTC vegetation. These include evergreen shrublands, broadleaf woodlands and conifer forests and represent mediterranean-type vegetation (MTV) under non-MTCs. Further east at similar latitudes but under different climates are sclerophyll forests with many similarities to MTC conifer forests. The California Floristic Province (Raven & Axelrod 1978) essentially circumscribes the MTC vegetation of North America and extends across the latitudinal range of the state. On the western slopes of the major mountain ranges is a rich diversity of vegetation types that change along the elevational gradient. Ascending the coastal mountains the main vegetation types sort out along gradients of decreasing aridity in the following order: grasslands, semi-deciduous woody sage scrub, evergreen chaparral shrublands, oak woodlands and conifer forests. A similar pattern is evident on the west side of the interior Sierra Nevada except for the absence of sage scrub. These vegetation types exhibit marked differences in fire regime and tolerance to disturbance tied to the different patterns of fuel structure resulting from changes in dominant growth forms along the elevational gradient. Along this gradient there is an interaction between fires and aridity such that lower fire frequency is required to displace shrubland associations with grasslands and other herbaceous vegetation on xeric than on mesic landscapes (Keeley 2002b). Consequently there are complex local mosaics due to differences in aspect and fire history (see Fig. 1.6c).

Fire in Mediterranean Ecosystems: Fire and the Fire Regime Framework

A global view of potential vs. actual vegetation distributions points to fire as a major driver of biome distribution and determinant of community structure (Bond et al . 2005). In ecological terms, fire acts much like an herbivore, consuming biomass and competing with biotic consumers for resources, and in this sense is an important part of trophic ecology (Bond & Keeley 2005). As in other competitive interactions, not only can fire competitively exclude herbivores by temporarily eliminating resources, but intensive grazing is known to exclude fire by consuming herbaceous ground fuels (Savage & Swetnam 1990). Coexistence is often enhanced by temporal separation of trophic niches, with herbivores grazing early in the spring on green herbaceous material that is unavailable for burning, whereas later in the season the remaining dry thatch is readily consumed by fire. In many respects fire is a more potent competitor because it is not limited by either toxins or protein deficiency and readily consumes dead woody biomass, but by contrast it is often limited by ignition sources and continuity of fuels. Fire scientists have long symbolized the critical elements of fire in a triangle of fuel, oxygen and heat (Pyne et al . 1996). These are indeed necessary for fire ignition and propagation but are insufficient for predicting the global distribution of fire-prone ecosystems. The conditions both necessary and sufficient to explain the ecological distribution of fire activity can be summarized by four parameters: biomass, seasonality, ignitions and fuel structure (Fig. 2.1). In addition to biomass fuels to spread a fire there must be a dry season that converts potential fuels to available fuels. In mediterranean-type climate (MTC) ecosystems summer drought results in high fire hazard on an annual basis, in contrast to many temperate forests that are only periodically vulnerable to fire in response to decadal or longer oscillations in climate. Vegetation only burns when ignitions are present to initiate the combustion process and landscapes vary markedly in the potential for natural ignitions from lightning, and in the extent of anthropogenic ignition sources. However, understanding the ecosystem distribution of fire requires consideration of a fourth parameter, fuel structure, which is fundamental to recognizing how different fire regimes develop.