Contrasting fire responses to climate and management: insights from two Australian ecosystems

This study explores effects of climate change and fuel management on unplanned fire activity in ecosystems representing contrasting extremes of the moisture availability spectrum (mesic and arid). Simulation modelling examined unplanned fire activity (fire incidence and area burned, and the area burned by large fires) for alternate climate scenarios and prescribed burning levels in: (i) a cool, moist temperate forest and wet moorland ecosystem in south‐west Tasmania (mesic); and (ii) a spinifex and mulga ecosystem in central Australia (arid). Contemporary fire activity in these case study systems is limited, respectively, by fuel availability and fuel amount. For future climates, unplanned fire incidence and area burned increased in the mesic landscape, but decreased in the arid landscape in accordance with predictions based on these limiting factors. Area burned by large fires (greater than the 95th percentile of historical, unplanned fire size) increased with future climates in the mesic landscape. Simulated prescribed burning was more effective in reducing unplanned fire activity in the mesic landscape. However, the inhibitory effects of prescribed burning are predicted to be outweighed by climate change in the mesic landscape, whereas in the arid landscape prescribed burning reinforced a predicted decline in fire under climate change. The potentially contrasting direction of future changes to fire will have fundamentally different consequences for biodiversity in these contrasting ecosystems, and these will need to be accommodated through contrasting, innovative management solutions.     

Ecology of grazing lawns in Africa

Grazing lawns are a distinct grassland community type, characterised by short‐stature and with their persistence and spread promoted by grazing. In Africa, they reveal a long co‐evolutionary history of grasses and large mammal grazers. The attractiveness to grazers of a low‐biomass sward lies in the relatively high quality of forage, largely due to the low proportion of stem material in the sward; this encourages repeat grazing that concomitantly suppresses tall‐grass growth forms that would otherwise outcompete lawn species for light. Regular grazing that prevents shading and maintains sward quality is thus the cornerstone of grazing lawn dynamics. The strong interplay between abiotic conditions and disturbance factors, which are central to grazing lawn existence, can also cause these systems to be highly dynamic. Here we identify differences in growth form among grazing lawn grass species, and assess how compositional differences among lawn types, as well as environmental variables, influence their maintenance requirements (i.e. grazing frequency) and vulnerability to degradation. We also make a clear distinction between the processes of lawn establishment and lawn maintenance. Rainfall, soil nutrient status, grazer community composition and fire regime have strong and interactive influences on both processes. However, factors that concentrate grazing pressure (e.g. nutrient hotspots and sodic sites) have more bearing on where lawns establish. Similarly, we discuss the relevance of enhanced rates of nitrogen cycling and of sodium levels to lawn maintenance. Grazer community composition and density has considerable significance to grazing lawn dynamics; not all grazers are adapted to foraging on short‐grass swards, and differences in body size and relative mouth dimensions determine which species are able to convert tall‐grass swards into grazing lawns under different conditions. Hence, we evaluate the roles of different grazers in lawn dynamics, as well as the benefits that grazer populations derive from having access to grazing lawns. The effects of grazing lawns can extend well beyond their borders, due to their influence on grazer densities, behaviour and movements as well as fire spread, intensity and frequency. Variation in the area and proportion of a landscape that is grazing lawn can thus have a profound impact on system dynamics. We provide a conceptual model that summarises grazing lawn dynamics, and identify a rainfall range where we predict grazing lawns to be most prevalent. We also examine the biodiversity associated with grazing lawn systems, and consider their functional contribution to the conservation of this biodiversity. Finally, we assess the utility of grazing lawns as a resource in a rangeland context.     

Restoring plant species diversity and community composition in a ponderosa pine-bunchgrass ecosystem

Monitoring of ecological restoration treatments often focuses on changes in community structure and function. We suggest that long-term changes in community composition also need to be explicitly considered when evaluating the success of restoration treatments. In 1992, we initiated an experiment in a ponderosa pine-bunchgrass ecosystem to evaluate responses to restoration treatments: (a) thinning the overstory vegetation (‘thinning’), (b) thinning plus forest floor manipulation with periodic prescribed burning (‘composite’), and (c) untreated ‘control.’ Treatments were further stratified by forest patch type: presettlement tree clumps (trees that established prior to the onset of fire exclusion in 1876), patches of retained postsettlement trees, patches where all postsettlement trees were removed, and remnant grass openings. Species richness did not differ among treatments for 10 years, but was highest in the composite treatment in 11th and 12th year after initial treatment. Community composition diverged among treatments 5 years after initial treatment, and compositional changes were greatest in the composite treatment. Species richness and composition differed among patch types prior to treatment. Remnant grass patches were the most diverse and presettlement patches were the least diverse. Following treatment, species richness in the postsettlement removed and retained patches, gradually approached levels found in remnant grass patches. Compositional differences among patch types changed a little by 2005. Species richness at the 2 m² scale increased only where the overstory was thinned and the understory was burned. However, these changes may not be detectable for many years, and can vary temporally in response to events such as severe droughts. Nonnative species establishment may be reduced by scheduling longer burn intervals or by refraining from burning where fuel loads are not hazardous, though these options may hinder goals of increasing diversity. Restoring species diversity and community composition continues to be more difficult than restoring ecosystem structure and function.     

Increased searching and handling effort in tall swards lead to a Type IV functional response in small grazing herbivores

Understanding the functional response of species is important in comprehending the species’ population dynamics and the functioning of multi-species assemblages. A Type II functional response, where instantaneous intake rate increases asymptotically with sward biomass, is thought to be common in grazers. However, at tall, dense swards, food intake might decline due to mechanical limitations or if animals selectively forage on the most nutritious parts of a sward, leading to a Type IV functional response, especially for smaller herbivores. We tested the predictions that bite mass, cropping time, swallowing time and searching time increase, and bite rate decreases with increasing grass biomass for different-sized Canada geese (Branta canadensis) foraging on grass swards. Bite mass indeed showed an increasing asymptotic relationship with grass biomass. At high biomass, difficulties in handling long leaves and in locating bites were responsible for increasing cropping, swallowing, and searching times. Constant bite mass and decreasing bite rate caused the intake rate to decrease at high sward biomass after reaching an optimum, leading to a Type IV functional response. Grazer body mass affected maximum bite mass and intake rate, but did not change the shape of the functional response. As grass nutrient contents are usually highest in short swards, this Type IV functional response in geese leads to an intake rate that is maximised in these swards. The lower grass biomass at which intake rate was maximised allows resource partitioning between different-sized grazers. We argue that this Type IV functional response is of more importance than previously thought.     

Factors affecting the distribution patterns of zebra and wildebeest in a resource-stressed environment

Understanding the spatial dynamics of landscape use by free-ranging herbivores is integral for successful ecosystem management. We used binary logistic regression analyses to determine the relative importance of biotic, abiotic and human factors in influencing the distribution (presence/absence) of wild grazers on two Maasai ranches in Kenya's Amboseli-Tsavo ecosystem. Both ranches had low grass biomass and suffered from regular droughts. We found that grazers consistently located themselves where grass biomass was highest, usually irrespective of grass quality, suggesting that forage quantity may be the limiting factor where grass biomass is generally low. The availability of surface water had no significant effect on the likelihood of grazers being present, even in the dry season.     

Andropogon gayanus (Gamba Grass) Invasion Increases Fire-mediated Nitrogen Losses in the Tropical Savannas of Northern Australia

Invasive alien grasses can substantially alter fuel loads and fire regimes which could have significant consequences for fire-mediated nutrient losses. The effects of the alien grass Andropogon gayanus Kunth. (Gamba grass) on fire-mediated nutrient losses was evaluated in Australia’s tropical savannas. Losses of macronutrients during fire were determined by comparing the nutrient pools contained in the fine fuel before fire and in the ash after fire. Pre-fire grass nutrient pools were significantly higher in A. gayanus plots than in native grass plots for all nutrients measured (N, P, K, S, Ca, and Mg). Nutrient losses were substantially higher in A. gayanus plots, with 113% higher losses for N, 80% for P, 56% for K, 63 for S, 355% for Ca, and 345% for Mg. However, only losses of N and Mg varied significantly between grass types. A simplified savanna ecosystem nutrient budget estimated that A. gayanus fires led to the net N loss of 20 kg ha⁻¹ y⁻¹. This is a conservative estimate because total fuel loads were relatively low (7.85 t ha⁻¹) for A. gayanus invaded plots leading to a relatively moderate intensity fire (6,408 kW m⁻¹). Higher A. gayanus fuel loads and fire intensities could potentially lead to losses of up to 61.5 kg N ha⁻¹ from the grass fuel. Over the long term, this is likely to lead to depletion of soil nutrients, particularly N, in the already low-fertility tropical savanna soils.     

Wildfire refugia in forests: Severe fire weather and drought mute the influence of topography and fuel age

Wildfire refugia (unburnt patches within large wildfires) are important for the persistence of fire‐sensitive species across forested landscapes globally. A key challenge is to identify the factors that determine the distribution of fire refugia across space and time. In particular, determining the relative influence of climatic and landscape factors is important in order to understand likely changes in the distribution of wildfire refugia under future climates. Here, we examine the relative effect of weather (i.e. fire weather, drought severity) and landscape features (i.e. topography, fuel age, vegetation type) on the occurrence of fire refugia across 26 large wildfires in south‐eastern Australia. Fire weather and drought severity were the primary drivers of the occurrence of fire refugia, moderating the effect of landscape attributes. Unburnt patches rarely occurred under ‘severe’ fire weather, irrespective of drought severity, topography, fuels or vegetation community. The influence of drought severity and landscape factors played out most strongly under ‘moderate’ fire weather. In mesic forests, fire refugia were linked to variables that affect fuel moisture, whereby the occurrence of unburnt patches decreased with increasing drought conditions and were associated with more mesic topographic locations (i.e. gullies, pole‐facing aspects) and vegetation communities (i.e. closed‐forest). In dry forest, the occurrence of refugia was responsive to fuel age, being associated with recently burnt areas (<5 years since fire). Overall, these results show that increased severity of fire weather and increased drought conditions, both predicted under future climate scenarios, are likely to lead to a reduction of wildfire refugia across forests of southern Australia. Protection of topographic areas able to provide long‐term fire refugia will be an important step towards maintaining the ecological integrity of forests under future climate change.     

Sonoran Desert Ecosystem transformation by a C4 grass without the grass/fire cycle

Aim Biological invasions facilitate ecosystem transformation by altering the structure and function, diversity, dominance and disturbance regimes. A classic case is the grass–fire cycle in which grass invasion increases the frequency, scale and/or intensity of wildfires and promotes the continued invasion of invasive grasses. Despite wide acceptance of the grass–fire cycle, questions linger about the relative roles that interspecific plant competition and fire play in ecosystem transformations. Location Sonoran Desert Arizona Upland of the Santa Catalina Mountains, Arizona, USA. Methods We measured species cover, density and saguaro (Carnegiea gigantea) size structure along gradients of Pennisetum ciliare invasion at 10 unburned/ungrazed P. ciliare patches. Regression models quantified differences in diversity, cover and density with respect to P. ciliare cover, and residence time and a Fisher’s exact test detected demographic changes in saguaro populations. Because P. ciliare may have initially invaded locations that were both more invasible and less diverse, we ran analyses with and without the plots in which initial infestations were located. Results Richness and diversity decreased with P. ciliare cover as did cover and density of most dominant species. Richness and diversity declined with increasing time since invasion, suggesting an ongoing transformation. The proportion of old‐to‐young Carnegiea gigantea was significantly lower in plots with dominant P. ciliare cover. Main conclusions Rich desert scrub (15–25 species per plot) was transformed into depauperate grassland (2–5 species per plot) within 20 years following P. ciliare invasion without changes to the fire regime. While the onset of a grass–fire cycle may drive ecosystem change in the later stages and larger scales of grass invasions of arid lands, competition by P. ciliare can drive small‐scale transformations earlier in the invasion. Linking competition‐induced transformation rates with spatially explicit models of spread may be necessary for predicting landscape‐level impacts on ecosystem processes in advance of a grass–fire cycle.     

Below-ground carbon and nitrogen accumulation in perennial grasses: A comparison of caespitose and rhizomatous growth forms

An experiment was conducted to compare below-ground soil organic carbon and total nitrogen accumulation between caespitose and rhizomatous perennial grasses in long-term (<25 yrs) grazed and ungrazed sites in semi-arid and mesic communities in the North American Great Plains. Development of greater nutrient pools beneath than between clones occurred at minimal clone basal areas (<60 cm²) for both caespitose species. Caespitose grasses accumulated substantially greater pools of carbon (20–200 fold) and nitrogen (50–500 fold) in soils to a depth of 10 cm beneath clones than rhizomatous grasses accumulated in rhizomes in both communities. Carbon and nitrogen pools in soils beneath caespitose clones exceeded combined (soil + rhizome) pools for rhizomatous grasses for a majority of the clone basal areas (>90 cm²) in the mesic community. In contrast, both pool sizes were smaller beneath the caespitose grass at all clone basal areas than the combined pools for the rhizomatous grass in the semi-arid community. The occurrence of larger soil nutrient pools beneath the rhizomatous species in the semi-arid community was largely a consequence of niche separation for microsites characterized by soils with higher nutrient concentrations, rather than plant-induced increases in nutrient concentrations. Although nutrient islands do not occur beneath rhizomatous grasses, their distribution in the semi-arid community was restricted to microsites characterized by soils with higher SOC and N concentrations. A greater efficiency of nutrient accumulation per unit rhizome mass and the maintenance of rhizome nutrient pools of similar magnitude to those of the rhizomatous grass in the mesic community may also contribute to the distribution of rhizomatous grasses in semi-arid communities. The existence of nutrient islands beneath a wide range of clone sizes in both mesic and semi-arid communities provides circumstantial evidence to suggest that nutrient islands beneath caespitose grasses may contribute to clone fitness in this growth form.     

An Invasive Grass Increases Live Fuel Proportion and Reduces Fire Spread in a Simulated Grassland

Fire is a globally important ecosystem process, and invasive grass species generally increase fire spread by increasing the fuel load and continuity of native grassland fuelbeds. We suggest that invasive grasses that are photosynthetically active, while the native plant community is dormant reduce fire spread by introducing high-moisture, live vegetation gaps in the fuelbed. We describe the invasion pattern of a high-moisture, cool-season grass, tall fescue (Schedonorus phoenix (Scop.) Holub), in tallgrass prairie, and use spatially explicit fire behavior models to simulate fire spread under several combinations of fuel load, invasion, and fire weather scenarios. Reduced fuel load and increased extent of tall fescue invasion reduced fire spread, but high wind speed and low relative humidity can partially mitigate these effects. We attribute reduced fire spread to asynchrony in the growing seasons of the exotic, cool-season grass, tall fescue, and the native, warm-season tallgrass prairie community in this model system. Reduced fire spread under low fuel load scenarios indicate that fuel load is an important factor in fire spread, especially in invaded fuel beds. These results present a novel connection between fire behavior and asynchronous phenology between invasive grasses and native plant communities in pyrogenic ecosystems.     

Ant diversity in relation to time since fire in a mallee landscape of South-Eastern Australia

Fire is a dominant process shaping the Australian landscape and in many regions the frequency and severity of wildfires are predicted to increase under climate change. The primary impact of fire on fauna is typically indirect through habitat change. In particular, in mesic forests different animal species are favoured at different times since fire as habitat complexity increases with vegetation recovery. However, this will not necessarily be the case in habitats with low complexity such as many of those occurring in arid and semi-arid regions. Here, we investigate the relationship between fire history and ant diversity and composition in semi-arid mallee of south-eastern Australia. We surveyed ants at 11 sites in the Little Desert National Park and nearby private land that last burnt 0.5, 6 or 40 years ago. We found no relationship between time since fire and either ant diversity or composition, and this can be explained by a lack of relationship between time since fire and vegetation cover. Our findings contrast with those for mallee bird species, which show clear successional patterns following fire, but are likely to be typical of ground-foraging fauna that lack specialized habitat requirements.     

Recovery of Aboveground Plant Biomass and Productivity After Fire in Mesic and Dry Black Spruce Forests of Interior Alaska

Plant biomass accumulation and productivity are important determinants of ecosystem carbon (C) balance during post-fire succession. In boreal black spruce (Picea mariana) forests near Delta Junction, Alaska, we quantified aboveground plant biomass and net primary productivity (ANPP) for 4 years after a 1999 wildfire in a well-drained (dry) site, and also across a dry and a moderately well-drained (mesic) chronosequence of sites that varied in time since fire (2 to ∼116 years). Four years after fire, total biomass at the 1999 burn site had increased exponentially to 160 ± 21 g m⁻² (mean ± 1SE) and vascular ANPP had recovered to 138 ± 32 g m⁻² y⁻¹, which was not different than that of a nearby unburned stand (160 ± 48 g m⁻² y⁻¹) that had similar pre-fire stand structure and understory composition. Production in the young site was dominated by re-sprouting graminoids, whereas production in the unburned site was dominated by black spruce. On the dry and mesic chronosequences, total biomass pools, including overstory and understory vascular and non-vascular plants, and lichens, increased logarithmically (dry) or linearly (mesic) with increasing site age, reaching a maximum of 2469 ± 180 (dry) and 4008 ± 233 g m⁻² (mesic) in mature stands. Biomass differences were primarily due to higher tree density in the mesic sites because mass per tree was similar between sites. ANPP of vascular and non-vascular plants increased linearly over time in the mesic chronosequence to 335 ± 68 g m⁻² y⁻¹ in the mature site, but in the dry chronosequence it peaked at 410 ± 43 g m⁻² y⁻¹ in a 15-year-old stand dominated by deciduous trees and shrubs. Key factors regulating biomass accumulation and production in these ecosystems appear to be the abundance and composition of re-sprouting species early in succession, the abundance of deciduous trees and shrubs in intermediate aged stands, and the density of black spruce across all stand ages. A better understanding of the controls over these factors will help predict how changes in climate and fire regime will affect the carbon balance of Interior Alaska. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Contrasting mechanisms of resilience at mesic and semi-arid boundaries of fynbos,a mega-diverse heathland of South Africa

Biome boundaries are expected to be sensitive to changes in climate and disturbance, because it is here that ecological communities are at environmental, ecological or disturbance limits. Using palaeoecology to study ecosystem dynamics at biome boundaries provides opportunities for understanding ecosystem resilience or sensitivity at ecologically meaningful timescales, and under varying climatic and disturbance conditions.The fynbos biome is a megadiverse Mediterranean type shrubland, found only in South Africa, that is threatened by climate change, land-use change and invasion by alien species. We used palaeoecological records from the semi-arid and mesic boundaries of the fynbos biome to test hypotheses regarding ecosystem resilience over timescales of centuries to millennia. We hypothesised that fynbos would expand at its mesic boundary at the expense of afrotemperate forest under drier and / or more fire prone conditions. In contrast, we hypothesised that at the semi-arid boundary, fynbos would expand at the expense of succulent karoo under wetter and cooler and / or more fire-prone conditions. Contrary to our expectations, the fossil pollen record at both biome boundaries showed remarkable stability at centennial - millennial timescales. To explain our results, we generated new hypotheses exploring possible mechanisms that might confer resilience.At the mesic (temperate) boundary, we suggest that decreased seasonality of rainfall during drier phases favoured fire and fynbos persistence, while in wetter periods, increased seasonality of rainfall resulted in enhanced summer drought stress, inhibiting forest expansion. At this boundary, internal reorganisation from grassy to proteoid fynbos states conferred resilience through resistance. At the succulent karoo boundary, we suggest that increased aridity was offset by less seasonality of rainfall, which enhanced biomass and allowed fire to persist, favouring persistence of fynbos. At this boundary, fynbos sensu stricto retreated during arid phases but recovered during climate amelioration, consistent with resilience through recovery. In both cases, this mega-diverse, disturbance-adapted flora provided a range of traits that enabled fynbos to persist despite environmental perturbation. Our findings agree with general observations that for ecosystems in regions of ample resource availability (i.e. at the mesic boundary), biotic interactions and disturbance tend to become more important in ecosystem dynamics, whereas in regions of scarce resources (in this case water scarcity at the semi-arid boundary) abiotic stress is more important. Our findings contribute to debates over the mechanisms that confer resistance and resilience to environmental change. Understanding and conserving the processes and mechanisms underpinning its resilience will be critical to effective conservation planning.     

Denitrification in a semi-arid grazing ecosystem

The effect of large herbivores on gaseous N loss from grasslands, particularly via denitrification, is poorly understood. In this study, we examined the influence of native migratory ungulates on denitrification in grasslands of Yellowstone National Park in two ways, by (1) examining the effect of artificial urine application on denitrification, and (2) comparing rates inside and outside long-term exclosures at topographically diverse locations. Artificial urine did not influence denitrification 3 and 12 days after application at hilltop, mid-slope, and slope-bottom sites. Likewise, grazers had no effect on community-level denitrification at dry exclosure sites, where rates were low. At mesic sites, however, ungulates enhanced denitrification by as much as 4 kg N ha⁻¹ year⁻¹, which was double atmospheric N inputs to this ecosystem. Denitrification enzyme activity (DEA, a measure of denitrification potential) was positively associated with soil moisture at exclosure sites, and herbivores stimulated DEA when accounting for the soil moisture effect. Glucose additons to soils increased denitrification and nitrate additions had no influence, suggesting that denitrification was limited by the amount of labile soil carbon, which previously has been shown to be enhanced by ungulates in Yellowstone. These results indicate that denitrification can be an ecologically important flux in portions of semi-arid landscapes, and that there is a previously unsuspected regulation of this process by herbivores. Received: 6 March 1998 / Accepted: 28 August 1998     

Mortality of native grasses after a summer fire in natural temperate grassland suggests ecosystem instability

The exclusion of regular fire and the introduction of livestock grazing have altered native grassland composition on Victoria's volcanic plains, commonly resulting in spear‐grass and wallaby‐grass pastures replacing Kangaroo Grass grasslands. The effect of reintroducing fire to these pastures is currently unknown, although it may be an important part of restoring this ecosystem. We measured the changes in basal area of the dominant grasses in a mixed Spear‐grass/Wallaby‐grass pastures after a summer wildfire, which we assume burnt a relatively homogenous grass sward. We found a 90–95% reduction in the basal area of live spear‐grass tussocks in burnt plots compared with unburned controls, due to the mortality of tussocks. This suggests that caution and structured experimentation should be applied when using fire to manage spear‐grass‐dominated grasslands.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Nutrient resorption response to fire and nitrogen addition in a semi-arid grassland

Fire and nitrogen (N) addition, both widely used grassland restoration strategies, strongly influence community composition and ecosystem functioning. However, little is known about their effects on plant nutrient resorption from senescing leaves, especially in semi-arid ecosystems. We evaluated the effects of fire, N addition (5.25 g N m⁻² yr⁻¹) and their potential interactions on nutrient resorption in five plant species in a semi-arid grassland in northern China. Foliar nutrient concentrations and resorption proficiencies and efficiencies varied substantially among species and functional groups. Fire increased green leaf N concentration ([N]g) and decreased N resorption proficiency (N RP), P resorption proficiency (P RP) and P resorption efficiency (P RE). N addition led to higher [N]g and lower N resorption, whereas it did not affect P related responses. There was no interaction between fire and N addition to affect all response variables except for green leaf P concentration ([P]g). These results suggest that fire and N addition can influence ecosystem nutrient cycling directly by changing resorption patterns and litter quality. Given the substantial interspecific variations in nutrient content and resorption and the potentially changing community composition, both fire and N addition may have indirect impacts on ecosystem nutrient cycling in this semi-arid grassland.     

Grass, rainfall and herbivores as determinants of Acacia erioloba (Meyer) recruitment in an African savanna

Acacia species in arid environments are thought to only establish in years of above-average rainfall, so should exhibit cohorted or pulsed recruitment. I studied population demography of Acacia erioloba Meyer in semi-arid savanna in the Kimberley area (mean annual precipitation = 425 ± 132 mm), South Africa, to establish whether they recruit episodically. This species was found to have a sapling bank at the sites, indicating that even though cohorted recruitment probably occurs, it is not the primary factor limiting recruitment to larger size classes. A. erioloba saplings given supplementary water and protection from herbivory showed significantly less height growth than saplings given water but not protected from herbivores, and grass within exclusion plots was taller and denser than outside of exclusion plots. The generalized linear model, although finding a significant difference between watered and protected and watered and unprotected saplings, explained only 16% of variation in growth, demonstrating the importance of factors other than competition from grass when rainfall is above-average. Average height increase across all treatments for a growing season was small (43.1 ± 30.81 mm), suggesting most investment is belowground. A. erioloba sapling below surface stem diameter correlated positively with aboveground growth, indicating larger, and presumably older, individuals grow faster. When grass was removed around saplings, growth rates were not significantly greater than for saplings surrounded by grass, over a growing season. Thus, in these study sites, pulsed events allow seedlings to establish, and then saplings slowly accumulate over many growing seasons within the grass sward, owing to slow aboveground growth combined with competition from grass in absence of grazers. Release events, e.g., heavy grazing combined with good rainfall, may allow mass release, giving the impression of cohorted or pulsed recruitment.     

The allometry of patch selection in ruminants

An axiomatic feature of food consumption by animals is that intake rate and prey abundance are positively related. While this has been demonstrated rigorously for large herbivores, it is apparent from patch selection trials that grazers paradoxically tend to prefer short, sparse swards to tall, dense swards. Indeed, migratory herbivores often shift from areas of high to low sward biomass during the growing season. As nutritional quality is an inverse function of grass abundance, herbivores appear to sacrifice short-term intake for nutritional gains obtainable by eating sparse forage of higher quality. Explicit models of this trade-off suggest that individual ruminants maximize daily rates of energy gain by choosing immature swards of intermediate biomass. As body mass is related positively to both ruminant cropping rates and digestibility, there should be an allometric link between grass abundance and energy maximization, providing a tool for predicting patterns of herbivore habitat selection. We used previously published studies to develop a synthetic model of trade-offs between forage abundance and quality predicting that optimal sward biomass should scale allometrically with body size. The model predicts size-related variation in habitat selection observed in a guild of grazing ungulates in the Serengeti ecosystem.     

Contributions of Ignitions,Fuels, and Weather to the Spatial Patterns of Burn Probability of a Boreal Landscape

The spatial pattern of fire observed across boreal landscapes is the outcome of complex interactions among components of the fire environment. We investigated how the naturally occurring patterns of ignitions, fuels, and weather generate spatial pattern of burn probability (BP) in a large and highly fire-prone boreal landscape of western Canada, Wood Buffalo National Park. This was achieved by producing a high-resolution map of BP using a fire simulation model that models the ignition and spread of individual fires for the current state of the study landscape (that is, the ‘control’). Then, to extract the effect of the variability in ignitions, fuels, and weather on spatial BP patterns, we subtracted the control BP map to those produced by “homogenizing” a single environmental factor of interest (that is, the ‘experimental treatments’). This yielded maps of spatial residuals that represent the spatial BP patterns for which the heterogeneity of each factor of interest is responsible. Residuals were analyzed within a structural equation modeling framework. The results showed unequal contributions of fuels (67.4%), weather (29.2%), and ignitions (3.4%) to spatial BP patterning. The large contribution of fuels reflects how substantial heterogeneity of land cover on this landscape strongly affects BP. Although weather has a chiefly temporal control on fire regimes, the variability in fire-conducive weather conditions exerted a surprisingly large influence on spatial BP patterns. The almost negligible effect of spatial ignition patterns was surprising but explainable in the context of this area’s fire regime. Similar contributions of fuels, weather, and ignitions could be expected in other parts of the boreal forest that lack a strong anthropogenic imprint, but are likely to be altered in human-dominated fire regimes.