The Fire Refuge Value of Patches of a Fire‐Sensitive Tree in Fire‐prone Savannas: Callitris intratropica in Northern Australia

Patches of fire‐sensitive vegetation often occur within fire‐prone tropical savannas, and are indicative of localized areas where fire regimes are less severe. These may act as important fire refugia for fire‐sensitive biota. The fire‐sensitive tree Callitris intratropica occurs in small patches throughout the fire‐prone northern Australian savannas, and is widely seen as an indicator of low‐severity fire regimes and of good ecosystem health. Here, we address the question: to what extent do Callitris patches act as refuges for other fire‐sensitive biota, and therefore play a broader conservation role? We contrast floral and faunal species composition between Callitris patches and surrounding eucalypt savanna, using three case studies. In the first case study, a floristic analysis of 47 Callitris patches across Western Australia's Kimberley region showed that woody species in these patches were overwhelmingly widespread, fire‐tolerant savanna taxa. No species of special conservation concern occurred disproportionately within Callitris patches. Similarly, there was no concentration of fire‐sensitive fauna or flora in five Callitris patches in the East Kimberley. Finally, there was no difference in ant species composition among 12 Callitris patches and surrounding eucalypt savannas in Kakadu National Park, Northern Territory, and there were no fire‐sensitive ant species in Callitris patches. Our three case studies from throughout the northwestern Australia provide no evidence that Callitris patches act as important refuges for fire‐sensitive flora or fauna within fire‐prone eucalypt savannas. This calls into question the notion that Callitris is a strong indicator of general ecosystem health.     

Fire-mediated effects of overstorey on plant species diversity and abundance in an eastern Australian heath

Overstorey shrub species are known to influence the composition of theunderstorey in Southern Hemisphere heathlands. Overstorey densities aresusceptible to variations in fire frequency; thus, fire regimes may influenceoverstorey/understorey interactions and overall floristic composition. Wecompared patches of Banksia heath which had supported anoverstorey during a fire interval of about 30 y with patches wherethe overstorey was absent during the same period, and tested for differences inspecies composition as a function of overstorey presence. Floristic compositionvaried significantly between overstorey patches and open patches. Most specieswere less abundant in overstorey patches, however some were more abundant. Therelative abundance of species in relation to overstorey was unrelated to theirfire response, propagule longevity or propagule storage location. There wassignificantly less biomass in overstorey patches compared with open patches.Theeffect of the overstorey varied with soil moisture. In a dry area, the numberof species was lower in overstorey patches, with fewer herb and shrub speciespresent compared with open patches. Fewer species were recorded in a wetterarea, but overstorey had no effect on the number of species recorded. Reducedintensity of competition among understorey species in overstorey patches couldbe responsible for the higher abundance of some species in these patches. Wepostulate that full diversity will be maintained when the density of overstoreyshrubs fluctuates widely over a relatively short period of time. This is mostlikely when fire frequency is highly variable.     

Comparison of the Higher-Severity Fire Regime in Historical (A.D. 1800s) and Modern (A.D. 1984–2009) Montane Forests Across 624,156?ha of the Colorado Front Range

There are concerns that recent fires, following a century of land uses, are burning in dry western forests in an uncharacteristic manner with large patches of higher-severity fire affecting long-term ecosystem dynamics. For example, it is well documented that a mixed-severity fire regime predominated over montane forests of the Colorado Front Range. However, much about the historical fire regime is unknown including the size, frequency, and distribution of higher-severity fires. We addressed these questions utilizing data from the original land surveyors who recorded locations of burned timber along survey lines resulting in a coarse-scale transect of fire occurrence across 624,156?ha. We reconstructed higher-severity burn patches, size distribution, and fire rotation for the 1800s (A.D. 1809–1883) and compared to the characteristics of modern fires over a recent 26-year period (A.D. 1984–2009) taken from remotely sensed data. We found the historical geometric mean higher-severity patch was 170.9?ha and the maximum patch size was 8,331?ha; the higher-severity fire rotation was 248.7?years. In addition, we confirmed that higher-severity fires were historically less common at elevations below 2,200?m. Modern fires had a geometric mean patch size of 90.0?ha (patches >20?ha) and a maximum size of 5,183?ha; the higher-severity fire rotation was 431?years. The distributions of higher-severity patches were only 63.5% similar, as the historical distribution had fewer small patches and more large patches. The mixed-severity fire regime, historically, included a significant portion of higher-severity fire and large burn patches; modern fires appear to be within the range of historical variability.     

Fire enhances litter decomposition and reduces vegetation cover influences on decomposition in a dry woodland

Dry woodlands frequently experience fire, and the heterogeneous spatial patterning of vegetation cover and fire behavior in these systems can lead to interspersed burned and unburned patches of different vegetation cover types. Biogeochemical processes may differ due to fire and vegetation cover influences on biotic and abiotic conditions, but these persistent influences of fire in the months or years following fire are not as well understood as the immediate impacts of fire. In particular, leaf litter decomposition, a process controlling nutrient availability and soil organic matter accumulation, is poorly understood in drylands but may be sensitive to vegetation cover and fire history. Decomposition is responsive to changes in abiotic drivers or interactions between abiotic conditions and biotic drivers, suggesting that decomposition rates may differ with vegetation cover and fire. The objective of this study was to assess the role of vegetation cover and fire on leaf litter decomposition in a semi-arid pinyon-juniper woodland in southern New Mexico, USA, where prescribed fire is used to combat increasing woody cover. A spatially heterogeneous prescribed burn led to closely co-located but discrete burned and unburned patches of all three dominant vegetation cover types (grass, shrub, tree). Decomposition rates of leaf litter from two species were measured in mesh litterbags deployed in factorial combination of the three vegetation cover types and two fire treatments (burned and unburned patches). For both litter types, decomposition was lower for unburned trees than for unburned grass or shrubs, perhaps due to greater soil–litter mixing and solar radiation away from tree canopies. Fire enhanced litter mass loss under trees, making decomposition rates similarly rapid in burned patches of all three vegetation cover types. Understanding decomposition dynamics in spatially heterogeneous vegetation cover of dry woodlands is critical for understanding biogeochemical process responses to fire in these systems.     

The Short-term Impact of Forest Fire on Soil Invertebrates in the Miombo

A study was conducted in eastern Zambia to assess the impact of fire on soil invertebrate communities between December 2003 and November 2004. Soil samples were collected 4 times from secondary miombo forest patches that were burnt in July–September 2003 and 2004 and patches not affected by fire. Macro-invertebrates were hand-sorted from soil samples and their population densities computed. The total number of orders per sample and the population density of Annelida, Chilopoda, Arachnida and some Hexapoda were lower under burnt forest patches compared to unburnt ones throughout the study period. Although the difference between burnt and unburnt patches in populations of other taxa such as Lepidoptera and Diptera remained below the threshold of statistical significance, fire appeared to have reduced their densities. It is concluded that fire can alter the structure of soil invertebrate communities through direct mortality or by its effect on availability of food resources. Further studies are needed to establish the linkage between fire, invertebrate community structure, ecosystem processes and floristic composition of the miombo.     

Ecological Disturbance Through Patch-Burn Grazing Influences Lesser Prairie-Chicken Space Use

Across portions of the western Great Plains in North America, natural fire has been removed from grassland ecosystems, decreasing vegetation heterogeneity and allowing woody encroachment. The loss of fire has implications for grassland species requiring diverse vegetation patches and structure or patches that have limited occurrence in the absence of fire. The lesser prairie-chicken (Tympanuchus pallidicinctus) is a declining species of prairie-grouse that requires heterogeneous grasslands throughout its life history and fire has been removed from much of its occupied range. Patch-burn grazing is a management strategy that re-establishes the fire-grazing interaction to a grassland system, increasing heterogeneity in vegetation structure and composition. We evaluated the effects of patch-burn grazing on lesser prairie-chicken space use, habitat features, and vegetation selection during a 4-year field study from 2014–2017. Female lesser prairie-chickens selected 1- and 2-year post-fire patches during the lekking season, ≥4-year post-fire patches during the nesting season, and year-of-fire and 1-year post-fire patches during post-nesting and nonbreeding seasons. Vegetation selection during the lekking season was not similar to available vegetation in selected patches, suggesting that lesser prairie-chickens cue in on other factors during the lekking season. During the nesting season, females selected nest sites with greater visual obstruction, which was available in ≥4-year post-fire patches; during the post-nesting season, females selected sites with 15–25% bare ground, which was available in the year-of-fire, 1-year post-fire, and 2-year post-fire patches; and during the nonbreeding season they selected sites with lower visual obstruction, available in the year-of-fire and 1-year post-fire patches. Because lesser prairie-chickens selected all available time-since-fire patches during their life history, patch-burn grazing may be a viable management tool to restore and maintain lesser prairie-chicken habitat on the landscape. © 2021 The Wildlife Society.     

Metapopulation Dynamics of the Mistletoe and Its Host in Savanna Areas with Different Fire Occurrence

Mistletoes are aerial hemiparasitic plants which occupy patches of favorable habitat (host trees) surrounded by unfavorable habitat and may be possibly modeled as a metapopulation. A metapopulation is defined as a subdivided population that persists due to the balance between colonization and extinction in discrete habitat patches. Our aim was to evaluate the dynamics of the mistletoe Psittacanthus robustus and its host Vochysia thyrsoidea in three Brazilian savanna areas using a metapopulation approach. We also evaluated how the differences in terms of fire occurrence affected the dynamic of those populations (two areas burned during the study and one was fire protected). We monitored the populations at six-month intervals. P. robustus population structure and dynamics met the expected criteria for a metapopulation: i) the suitable habitats for the mistletoe occur in discrete patches; (ii) local populations went extinct during the study and (iii) colonization of previously non-occupied patches occurred. The ratio of occupied patches decreased in all areas with time. Local mistletoe populations went extinct due to two different causes: patch extinction in area with no fire and fire killing in the burned areas. In a burned area, the largest decrease of occupied patch ratios occurred due to a fire event that killed the parasites without, however, killing the host trees. The greatest mortality of V. thyrsoidea occurred in the area without fire. In this area, all the dead trees supported mistletoe individuals and no mortality was observed for parasite-free trees. Because P. robustus is a fire sensitive species and V. thyrsoidea is fire tolerant, P. robustus seems to increase host mortality, but its effect is lessened by periodic burning that reduces the parasite loads.     

Transient effects of corridors on polygyne fire ants over a decade

1. Although corridors are frequently regarded as a way to mitigate the negative effects of habitat fragmentation, concerns persist that corridors may facilitate the spread of invasive species to the detriment of native species.
2. The invasive fire ant, Solenopsis invicta, has two social forms. The polygyne form has limited dispersal abilities relative to the monogyne form. Our previous work in a large-scale corridor experiment showed that in landscapes dominated by the polygyne form, fire ant density was higher and native ant species richness was lower in habitat patches connected by corridors than in unconnected patches.
3. We expected that these observed corridor effects would be transient, that is, that fire ant density and native ant species richness differences between connected and unconnected patches would diminish over time as fire ants eventually fully established within patches. We tested this prediction by resampling the three landscapes dominated by polygyne fire ants 6 to 11 years after our original study.
4. Differences in fire ant density between connected and unconnected habitat patches in these landscapes decreased, as expected. Differences in native ant species richness were variable but lowest in the last 2 years of sampling.
5. These findings support our prediction of transient corridor effects on this invasive ant and stress the importance of temporal dynamics in assessing population and community impacts of habitat connectivity.

     

A native C<Subscript>3</Subscript> grass alters fuels and fire spread in montane grassland of South Africa

Although most fire research in plant ecology focuses on vegetation responses to burning, shifts in plant community composition wrought by climate change can change wildland fuelbeds and affect fire behaviour such that the nature of fire in these systems is altered. Changes that introduce substantially different fuel types can alter the spatial extent of fire, with potential impacts on community succession and biodiversity. Montane grasslands of sub-Saharan Africa are threatened by climate change because species distributions can shift with climatically determined ranges. We studied the impact of patches of the temperate C₃ grass Festuca costata in C₄-dominated grassland at the transition between their subalpine ranges in South Africa’s Drakensberg. We used empirical data on fuel moisture and fuel load across F. costata-dominated patches in a C₄-dominated matrix in fire spread models to predict the effect of larger, higher-moisture F. costata patches on the spatial extent of fire. Results indicate F. costata reduces fire spread and burn probability in F. costata patches, and the effect increases as live fuel moisture increases and patches get larger. However, as a native species, F. costata does not appear to have the extreme, fire-suppressing effect of non-native C₃ grasses in other C₄ grasslands. Instead, F. costata patches likely increase variability in the spatial extent of fire in this C₄-dominated grassland, which likely translates to spatial variability on vegetation succession.     

Contrasting responses of small mammals to fire and topographic refugia

Unburnt patches within burnt landscapes are expected to provide an important resource for fauna, potentially acting as a refuge from direct effects of fire and allowing animals to persist in burnt landscapes. Nevertheless, there is little information about the way refugia are used by fauna and how populations may be affected by them. Planned burns are often patchy, with unburnt areas generally associated with gully systems providing a good opportunity to study faunal use of refugia. We used a before–after control‐impact design associated with a planned burn in south eastern Australia to investigate how two small mammal species, the bush rat Rattus fuscipes and agile antechinus Antechinus agilis, used unburnt gully systems within a larger burnt area. We tested three alternative hypotheses relating to post‐fire abundance: (i) active refugia – abundance would increase in unburnt patches because of a post‐fire shift of individuals from burnt to unburnt areas; (ii) passive refugia – abundance in unburnt patches would remain similar to pre‐fire levels; and (iii) limited or no refugia – abundance would reduce in unburnt patches related to the change induced by fire in the wider landscape. We found the two species responded differently to the presence of unburnt refugia in the landscape. Relative to controls, fire had little effect on bush rat abundance in gullies, supporting hypothesis 2. In contrast, agile antechinus abundance increased in gullies immediately post‐fire consistent with a shift of individuals from burnt parts of the landscape, supporting hypothesis 1. Differences in site fidelity, habitat use and intraspecific competition between these species are suggested as likely factors influencing responses to refugia. The way unburnt patches function as faunal refugia and the subsequent influence they have on post‐fire population dynamics, will to some extent depend on the life history attributes of individual species.     

Hypericum cumulicola demography in unoccupied and occupied Florida scrub patches with different time‐since‐fire

1 Metapopulation models predict that unoccupied, but suitable, patches will exist for species subject to extinction and colonization dynamics. We compared the demographic responses of Hypericum cumulicola , a rare herbaceous species almost entirely restricted to Florida rosemary scrub, when transplanted to occupied or unoccupied patches.
2 Seedlings were transplanted and seeds buried into Florida rosemary scrub patches differing in time since last fire, and in the presence or absence of H. cumulicola. We used a replicated, factorial design to place the transplants and seeds in the field, and monitored their performance for 18 months.
3 Neither time‐since‐fire nor prior H. cumulicola site occupancy affected survival of transplants. Only time‐since‐fire affected growth. Time‐since‐fire, H. cumulicola occupancy, and their interaction affected reproductive effort, but these effects were not consistent between years.
4 Flowering and seed production led to subsequent seedling recruitment near transplants, mainly in recently burned sites. Genetic screening of transplants and seedlings showed that transplants in occupied sites could have crossed with nearby resident plants, but that offspring in sites previously unoccupied were likely to have been parented only by nearby transplants.
5 Seeds buried, and later exhumed, germinated after 1 or 2 years of burial, demonstrating a persistent soil seed bank from which populations could recover after fire. Neither time‐since‐fire nor H. cumulicola occupancy affected seed dormancy or germination.
6 Similar demography in unoccupied and occupied patches suggests that the patchy pattern of site occupancy by H. cumulicola is probably due to limited dispersal and periodic extinction, especially associated with long fire‐free intervals. Conservation measures need to protect unoccupied patches to allow metapopulation dynamics and persistence.     

A Burning Question: What are the Implications of Forest Fires for Woodland Caribou?

Canada's federal recovery strategy for boreal woodland caribou (Rangifer tarandus caribou) classifies areas burned by forest fire as disturbed habitat. This assignment of fire as a disturbance has potential economic and social implications across Canada, and influences plans and actions to achieve caribou conservation and recovery. Previous researchers have reported caribou avoid burned habitat, but these studies did not typically consider unburned residual patches within fire perimeters. Additionally, the implications of burned habitat on individual caribou survival is unclear. We examined resource selection by boreal woodland caribou of burns, and unburned residual patches, using global positioning system (GPS) locations for 201 caribou across 6 caribou populations in Alberta, Canada. We also examined if burned habitat affected the survival of adult female caribou. Caribou avoided burns and unburned residual patches. Increased use of burned habitats, however, did not lower the survival of adult caribou. Collectively, these results provide evidence to support current assertions that burns, and the embedded unburned residual patches are not preferred caribou habitat and increase our understanding of the implications of forest fire for caribou vital rates. Our investigation offers important information about the role of forest fire in caribou ecology and enhances the identification of disturbed habitat under recovery strategy guidelines to effectively address caribou population declines. © 2021 The Wildlife Society.     

Fine-scale genetic structure and fire-created habitat patchiness in the Australian allodapine bee, Exoneura nigrescens (Hymenoptera: Apidae)

Fire promotes an abundance of nest sites for the stem nesting bee Exoneura nigrescens, which remain viable for approximately 10 years. The finite duration of nesting substrate and localized fire events suggest that migration should minimize genetic structure among suitable habitat patches. Exoneura nigrescens was sampled from 7 localities with a known fire history in southwestern Victoria, Australia. Individual bees were genotyped at 8 microsatellite loci and genic and genotypic analyses applied to examine genetic structure among burn patch localities, within burn patches, and within colonies. Despite relatively short-term availability of nesting substrates, remarkably fine-scale genetic structure was observed both among burn patches and within burn patches. The spatial distribution of relatedness shows a strong pattern of isolation-by-distance at geographic distances to 35 km, suggesting that genetic partitioning among burn patches is, at least in part, a result of dispersal ability. Genetic structure within burn patches includes colonies consisting of close kin with genic partitioning among nests. Relatedness structure within colonies suggests that polygamy, multiple breeding pairs, and a lack of inbreeding typifies the mating system.     

Short‐term effects of low intensity fire on soil macroinvertebrate assemblages in different vegetation patch types in an Australian tropical savanna

Abstract Early dry season fires are a common land management regime employed across the tropical savannas of northern Australia. The rationale is that this reduces fuel loads and so reduces fire risk in the latter part of the dry season. Despite the acceptance of fire as a major management tool the ecological effects of fire remain uncertain. Vegetation patches and their associated macroinvertebrates play a critical role in the capture and recycling of water and nutrients. The aim of this study was to examine the responses of soil macroinvertebrates, within different types of vegetation patches, to early dry season fires in tropical savanna woodland in northern Australia. The abundance of major macroinvertebrate taxa and functional groups, and taxon richness were quantified in three vegetation patch types 2 weeks before and 2 weeks after burning. Termites dominated the soil macroinvertebrate assemblage sampled. Fire led to significant decreases in ant and spider abundances and overall taxon richness. Functional group analyses showed significant overall declines in the abundances of macropredators and litter transformers. There were also interactions between fire and patch type; in tree patches, fire significantly reduced total macroinvertebrate abundance, as well as the abundance of termites and ecosystem engineers. These changes in soil macroinvertebrates will potentially influence patch functionality, with important implications for soil processes and landscape health.     

Edge fires drive the shape and stability of tropical forests

In tropical regions, fires propagate readily in grasslands but typically consume only edges of forest patches. Thus, forest patches grow due to tree propagation and shrink by fires in surrounding grasslands. The interplay between these competing edge effects is unknown, but critical in determining the shape and stability of individual forest patches, as well the landscape‐level spatial distribution and stability of forests. We analyze high‐resolution remote‐sensing data from protected Brazilian Cerrado areas and find that forest shapes obey a robust perimeter–area scaling relation across climatic zones. We explain this scaling by introducing a heterogeneous fire propagation model of tropical forest‐grassland ecotones. Deviations from this perimeter–area relation determine the stability of individual forest patches. At a larger scale, our model predicts that the relative rates of tree growth due to propagative expansion and long‐distance seed dispersal determine whether collapse of regional‐scale tree cover is continuous or discontinuous as fire frequency changes.     

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.     

Threats to monsoon rainforest habitat in northern Australia: The case of Ptychosperma bleeseri Burret (Arecaceae)

Abstract Rainforest patches in the Australian wet-dry tropics occur as habitat islands. Recent studies show that many patches are severely damaged by feral animals, weed invasion and contemporary fire regimes. This paper documents the status of an endangered palm species, Ptychosperma bleeseri, which occurs within a narrow geographical range in seven rainforest patches. Results of multivariate statistical analysis of quadrats with and without P. bleeseri show that the species does not appear to occupy all suitable habitat available at either a regional scale or within single rainforest patches. Evidence, based on demographic surveys, is presented for the current decline of P. bleeseri populations and possible causes for this decline are discussed; in particular, the impacts of wildfire and feral animal activity. Recommendations for a recovery plan for P. bleeseri are discussed, and include fencing, fire protection and public education.     

Tree Patches Show Greater N Losses but Maintain Higher Soil N Availability than Grassland Patches in a Frequently Burned Oak Savanna

Long-term prescribed fires have increased woody canopy openness and reduced nitrogen (N) cycling (that is, net N mineralization) in an oak savanna in Minnesota, USA. It is unclear how fire-induced shifts from oak-dominated to C4 grass-dominated vegetation contribute to this decline in N cycling compared to direct effects of increasing fire frequency promoting greater N losses. We determined (1) the magnitude of decline in net N mineralization in oak versus grass-dominated patches with increasing fire frequency and (2) if differences in net N mineralization between oak and grass patches in frequently burned oak savanna (burned 8 out of 10 years on average during the last 40 years) could be attributed to differences in N losses through volatilization and leaching or to plant traits affecting decomposition and mineralization. In situ net N mineralization declined with increasing fire frequency overall, but this decline was less in oak- than in grass-dominated patches, with oak-dominated patches having more than two times higher net N mineralization than grass-dominated patches. Greater net N mineralization in oak-dominated patches occurred despite greater N losses through volatilization and leaching (on average 1.8 and 1.4 g m⁻² y⁻¹ for oak- and grass-dominated patches, respectively), likely because of higher plant litter N concentration in the oak-dominated patches. As total soil N pools in the first 15 cm did not differ between oak- and grass-dominated patches (on average 83 g N m⁻²), N inputs from atmospheric deposition and uptake from deep soil layers may offset higher N losses. Our results further show that net N mineralization rates decline within 5 years after tree death and subsequent colonization by C4 grasses to levels observed in grass-dominated patches. Although long-term prescribed fires often directly reduce N stocks and cycling because of increased N losses, this study has shown that fire-induced shifts in vegetation composition can strongly contribute to the declines in N cycling in systems that are frequently disturbed by fires with potential feedbacks to plant productivity.     

Fire may mediate effects of landscape connectivity on plant community richness in prairie remnants

Following the predictions of island biogeography and metapopulation theory, efforts to understand and maintain plant communities have focused on spatial attributes such as patch size and connectivity. However, despite a large body of literature, the effects of these spatial attributes remain uncertain, and accumulating evidence suggests that they may interact with local patch characteristics. Here, we assess the role of spatial attributes (patch size and connectivity at two time periods) and of patch quality (indicated by number of years since last fire, as this is a fire‐dependent system) in determining plant species richness at 63 remnant prairie sites in Wisconsin, USA. Our results demonstrate significant main effects of patch size, with larger patches being richer than small patches, and of fire, with recent‐fire sites being richer than fire‐excluded sites. We also report a novel interaction between number of years since fire and patch connectivity. Historical (1950) connectivity was negatively related to species richness at recent‐fire sites, but positively related to species richness at fire‐excluded sites, while current (2000) connectivity and species richness were positively correlated at recent‐fire sites, but unrelated at fire‐excluded sites. Our findings indicate that measures of patch quality such as time since last fire can alter the association between spatial attributes such as patch size or connectivity and species richness.     

Moderate patchiness optimizes heterogeneity,stability, and beta diversity in mesic grassland

Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire–grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%–33% of area burned) and three‐ to four‐year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.