Resilience of terrestrial and aquatic fauna to historical and future wildfire regimes in western North America

Wildfires in many western North American forests are becoming more frequent, larger, and severe, with changed seasonal patterns. In response, coniferous forest ecosystems will transition toward dominance by fire‐adapted hardwoods, shrubs, meadows, and grasslands, which may benefit some faunal communities, but not others. We describe factors that limit and promote faunal resilience to shifting wildfire regimes for terrestrial and aquatic ecosystems. We highlight the potential value of interspersed nonforest patches to terrestrial wildlife. Similarly, we review watershed thresholds and factors that control the resilience of aquatic ecosystems to wildfire, mediated by thermal changes and chemical, debris, and sediment loadings. We present a 2‐dimensional life history framework to describe temporal and spatial life history traits that species use to resist wildfire effects or to recover after wildfire disturbance at a metapopulation scale. The role of fire refuge is explored for metapopulations of species. In aquatic systems, recovery of assemblages postfire may be faster for smaller fires where unburned tributary basins or instream structures provide refuge from debris and sediment flows. We envision that more‐frequent, lower‐severity fires will favor opportunistic species and that less‐frequent high‐severity fires will favor better competitors. Along the spatial dimension, we hypothesize that fire regimes that are predictable and generate burned patches in close proximity to refuge will favor species that move to refuges and later recolonize, whereas fire regimes that tend to generate less‐severely burned patches may favor species that shelter in place. Looking beyond the trees to forest fauna, we consider mitigation options to enhance resilience and buy time for species facing a no‐analog future.     

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire-making species, but 'natural' (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from 'natural' background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.     

Two fine‐resolution Pliocene charcoal records and their bearing on pre‐human fire frequency in south‐western Australia

Abstract Yallalie is a probable meteor impact crater and in the Upper Pliocene contained a substantial lake. Two Mid‐Pliocene finely laminated sediment records from Palaeolake Yallalie, from about 3 million years ago, provide evidence of fire and fire frequency in the sclerophyll woodland and heaths of south‐western Australia in the absence of humans. Fine charcoal was observed in all samples examined, and was deposited at a rate of about 0.3–0.8 cm² cm^?2 year^?1 in Palaeolake Yallalie. This evidence suggests the occurrence of annual fires occurring every year in the slightly warmer and wetter climate compared with today. The near coastal western location and the prevailing westerly winds probably carry charcoal from the near region or lake catchment scale. The data indicate that local fires occurred at a variety of time intervals between 3 and 13 years, with a typical average of 6–10 years. The results are comparable with those of Atahan et al. (2004) for the same site but from a period of about 200 000 years later in the Mid‐Pliocene. Thus, the records which differ in age by some hundreds of thousands of years have all recorded fire frequencies that are longer than for the historical period and this may have important implications for the long‐term survival of the integrity of the high biodiversity plant communities of the region.     

Unsupported inferences of high‐severity fire in historical dry forests of the western United States: response to Williams and Baker

Reconstructions of dry western US forests in the late 19th century in Arizona, Colorado and Oregon based on General Land Office records were used by Williams & Baker (2012; Global Ecology and Biogeography, 21 , 1042–1052; hereafter W&B) to infer past fire regimes with substantial moderate and high‐severity burning. The authors concluded that present‐day large, high‐severity fires are not distinguishable from historical patterns. We present evidence of important errors in their study. First, the use of tree size distributions to reconstruct past fire severity and extent is not supported by empirical age–size relationships nor by studies that directly quantified disturbance history in these forests. Second, the fire severity classification of W&B is qualitatively different from most modern classification schemes, and is based on different types of data, leading to an inappropriate comparison. Third, we note that while W&B asserted ‘surprising’ heterogeneity in their reconstructions of stand density and species composition, their data are not substantially different from many previous studies which reached very different conclusions about subsequent forest and fire behaviour changes. Contrary to the conclusions of W&B, the preponderance of scientific evidence indicates that conservation of dry forest ecosystems in the western United States and their ecological, social and economic value is not consistent with a present‐day disturbance regime of large, high‐severity fires, especially under changing climate.     

Understory vegetation indicates historic fire regimes in ponderosa pine‐dominated ecosystems in the Colorado Front Range

Question: Can current understory vegetation composition across an elevation gradient of Pinus ponderosa‐dominated forests be used to identify areas that, prior to 20th century fire suppression, were characterized by different fire frequencies and severities (i.e., historic fire regimes)? Location: P. ponderosa‐dominated forests in the montane zone of the northern Colorado Front Range, Boulder and Larimer Counties, Colorado, USA. Methods: Understory species composition and stand characteristics were sampled at 43 sites with previously determined fire histories. Indicator species analyses and indirect ordination were used to determine: (1) if stands within a particular historic fire regime had similar understory compositions, and (2) if understory vegetation was associated with the same environmental gradients that influence fire regime. Classification and regression tree analysis was used to ascertain which species could predict fire regimes. Results: Indicator species analysis identified 34 understory species as significant indicators of three distinct historic fire regimes along an elevation gradient from low‐ to high‐elevation P. ponderosa forests. A predictive model derived from a classification tree identified five species as reliable predictors of fire regime. Conclusions: P. ponderosa‐dominated forests shaped by three distinct historic fire regimes have significantly different floristic composition, and current understory compositions can be used as reliable indicators of historical differences in past fire frequency and severity. The feasibility demonstrated in the current study using current understory vegetation properties to detect different historic fire regimes, should be examined in other fire‐prone forest ecosystems.     

High‐severity fire corroborated in historical dry forests of the western United States: response to Fulé et al.

Accurate assessment of changing fire regimes is important, since climatic change and people may be promoting more wildfires. Government wildland fire policies and restoration programmes in dry western US forests are based on the hypothesis that high‐severity fire was rare in historical fire regimes, modern fire severity is unnaturally high and restoration efforts should focus primarily on thinning forests to eliminate high‐severity fire. Using General Land Office (GLO) survey data over large dry‐forest landscapes, we showed that the proportion of historical forest affected by high‐severity fire was not insignificant, fire severity has not increased as a proportion of total fire area and large areas of dense forest were present historically (Williams & Baker, Global Ecology and Biogeography, 21 , 1042–1052, 2012; W&B). In response, Fulé et al. (Global Ecology and Biogeography, 2013, doi: 10.1111/geb.12136; FE) suggest that our inferences are unsupported and land management based on our research could be damaging to native ecosystems. Here, we show that the concerns of FE are unfounded. Their criticism comes from misquoting W&B, mistaking W&B's methods, misusing evidence (e.g. from Aldo Leopold) and missing substantial available evidence. We also update corroboration for the extensive historical high‐severity fire shown by W&B. We suggest that restoration programmes are misdirected in seeking to reduce all high‐severity fire in dry forests, given findings from spatially extensive GLO data and other sources.     

Continental‐scale syntheses of Australian pyromes – misclassification of south‐western eucalypt woodlands misinforms management

Global and continental‐scale analysis of ecological phenomena can offer important insights through the identification of patterns and associations not detectable at smaller scales. However, using proxies for ecological phenomena, such as vegetation mapping for spatially projecting fire regime niches and post‐fire plant responses, require critical examination of predictions to determine utility. Using local studies in south‐western Australia, we demonstrate that while this approach has been largely successful in mallee woodland and shrubland, it has failed in eucalypt woodland, with the consequence that values for a range of fire‐related parameters from the continent‐wide approaches, if adopted in informing management, would result in undesirable conservation outcomes for the world's largest extant temperate woodland.     

Anomalies in grasstree fire history reconstructions for south‐western Australian vegetation

Abstract A new fire history for south‐western Australian sclerophyll forests was proposed recently based on grasstree (Xanthorrhoea preissii ) records that were interpreted to show a high frequency (3–5 years) ‘pre‐European burning regime’. Such a fire regime appears incompatible with the long‐term survival of many fire‐killed woody taxa. We investigated the local fire history in a small area of the northern sand‐plain shrub‐lands of south‐western Australia using 15 grasstrees, examining individual grasstree records in detail and comparing this with the decadal or averaged approach used in the original research, and with fire histories reconstructed from satellite images for the period since 1975. Results lead us to question the utility of the proposed grasstree fire history record as a tool for understanding past fire regimes for two reasons: First, inconsistencies in fire histories among individual grasstrees were considerable – some individuals were not burnt by known fires, while some apparently were burned many times during periods when others were not burned at all. Second, the grasstree record indicates a possible increase in patchiness of fires since 1930, while contemporary evidence and interpretations of the nature of Aboriginal (pre‐European) fire regimes would suggest the opposite. We believe that further research is needed to identify to what extent the grasstree method for reconstruction of fire histories can be used to re‐interpret how fire operated in many highly diverse ecosystems prior to European settlement of Australia.     

Canada lynx use of burned areas: Conservation implications of changing fire regimes

A fundamental problem in ecology is forecasting how species will react to major disturbances. As the climate warms, large, frequent, and severe fires are restructuring forested landscapes at large spatial scales, with unknown impacts on imperilled predators. We use the United States federally Threatened Canada lynx as a case study to examine how predators navigate recent large burns, with particular focus on habitat features and the spatial configuration (e.g., distance to edge) that enabled lynx use of these transformed landscapes. We coupled GPS location data of lynx in Washington in an area with several recent large fires and a number of GIS layers of habitat data to develop models of lynx habitat selection in recent burns. Random Forest habitat models showed lynx‐selected islands of forest skipped by large fires, residual vegetation, and areas where some trees survived to use newly burned areas. Lynx used burned areas as early as 1 year postfire, which is much earlier than the 2–4 decades postfire previously thought for this predator. These findings are encouraging for predator persistence in the face of fires, but increasingly severe fires or management that reduces postfire residual trees or slow regeneration will likely jeopardize lynx and other predators. Fire management should change to ensure heterogeneity is retained within the footprint of large fires to enable viable predator populations as fire regimes worsen with climate change.     

Seasonality of vegetation fires as modified by human action: observing the deviation from eco‐climatic fire regimes

Aim In any region affected, fires exhibit a strong seasonal cycle driven by the dynamic of fuel moisture and ignition sources throughout the year. In this paper we investigate the global patterns of fire seasonality, which we relate to climatic, anthropogenic, land‐cover and land‐use variables. Location Global, with detailed analyses from single 1°× 1° grid cells. Methods We use a fire risk index, the Chandler burning index (CBI), as an indicator of the ‘natural’, eco‐climatic fire seasonality, across all types of ecosystems. A simple metric, the middle of the fire season, is computed from both gridded CBI data and satellite‐derived fire detections. We then interpret the difference between the eco‐climatic and observed metrics as an indicator of the human footprint on fire seasonality. Results Deforestation, shifting cultivation, cropland production or tropical savanna fires are associated with specific timings due to land‐use practices, sometimes largely decoupled from the CBI dynamics. Detailed time series from relevant locations provide comprehensive information about these practices and how they are adapted to eco‐climatic conditions. Main conclusions We find a great influence of anthropogenic activities on global patterns of fire seasonality. The specificity of the main fire practices and their easy identification from global observation is a potential tool to support land‐use monitoring efforts. Our results should also prove valuable in the development of a methodological approach for improving the representation of anthropogenic fire practices in dynamic global vegetation models.     

The response of cerambycid beetles (Coleoptera: Cerambycidae) to long‐term fire frequency regimes in subtropical eucalypt forest

Fire has a varied influence on plant and animal species through direct (e.g. fire‐induced mortality) and indirect (e.g. modification of habitat) effects. Our understanding of the influence of fire regime on invertebrates and their response to fire‐induced modifications to habitat is poor. We aimed to determine the response of a beetle family (Coleoptera: Cerambycidae) to varying fire treatments and hypothesised that the abundance of cerambycid beetles is influenced by fire frequency due to modifications in habitat associated with the fire treatments. Arthropods were sampled across 3 months in annually and triennially burnt areas (treatments starting in 1952 and 1973 respectively), an area unburnt since 1946, and a former unburnt treatment, burnt by wildfire in 2006. Eleven different cerambycid taxa were collected using flight intercept panel traps, dominated by three species (Ipomoria tillides, Adrium sp. and Bethelium signiferum) which made up 99% of individuals collected. Over the sampling period the long unburnt treatment had significantly lower species richness than the triennial and wildfire treatments. Cerambycid abundance was significantly higher in the triennially burnt treatment than in all other fire treatments. Ipomoria tillides was more abundant in both frequently burnt treatments, Adrium sp. was more common in triennially burnt areas, whereas B. signiferum, was more common in the wildfire affected treatment. Some, but not all, cerambycid beetles were more common in areas with a more open understorey (i.e. resulting from frequent burning), and lower tree basal area, as this likely influences their ability to fly easily between food sources. Cerambycid abundance was positively related to the volume of coarse woody debris and healthy tree crowns. Cerambycid beetles were clearly influenced by historic fire regime, suggesting that changes in fire regime can potentially have a profound influence on arthropod assemblages, and subsequent influences on ecosystem processes, which are currently poorly understood.     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

Anthropogenic modifications to fire regimes in the wider Serengeti‐Mara ecosystem

Fire is a key driver in savannah systems and widely used as a land management tool. Intensifying human land uses are leading to rapid changes in the fire regimes, with consequences for ecosystem functioning and composition. We undertake a novel analysis describing spatial patterns in the fire regime of the Serengeti‐Mara ecosystem, document multidecadal temporal changes and investigate the factors underlying these patterns. We used MODIS active fire and burned area products from 2001 to 2014 to identify individual fires; summarizing four characteristics for each detected fire: size, ignition date, time since last fire and radiative power. Using satellite imagery, we estimated the rate of change in the density of livestock bomas as a proxy for livestock density. We used these metrics to model drivers of variation in the four fire characteristics, as well as total number of fires and total area burned. Fires in the Serengeti‐Mara show high spatial variability—with number of fires and ignition date mirroring mean annual precipitation. The short‐term effect of rainfall decreases fire size and intensity but cumulative rainfall over several years leads to increased standing grass biomass and fuel loads, and, therefore, in larger and hotter fires. Our study reveals dramatic changes over time, with a reduction in total number of fires and total area burned, to the point where some areas now experience virtually no fire. We suggest that increasing livestock numbers are driving this decline, presumably by inhibiting fire spread. These temporal patterns are part of a global decline in total area burned, especially in savannahs, and we caution that ecosystem functioning may have been compromised. Land managers and policy formulators need to factor in rapid fire regime modifications to achieve management objectives and maintain the ecological function of savannah ecosystems.     

Influences of climate on fire regimes in montane forests of north-western Mexico

Aim To identify the influence of interannual and interdecadal climate variation on the occurrence and extent of fires in montane conifer forests of north‐western Mexico. Location This study was conducted in Jeffrey pine (Pinus jeffreyi Grev. & Balf.)‐dominated mixed‐conifer forests in the central and northern plateau of the Sierra San Pedro Mártir, Baja California, Mexico. Methods Fire occurrence was reconstructed for 12 dispersed sites for a 290‐year period (1700–1990) from cross‐dated fire‐scarred samples extracted from live trees, snags and logs. Superposed epoch analysis was used to examine the relationships of tree‐ring reconstructions of drought, the El Niño/Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO) with fire occurrence and extent. Results Years with no recorded fire scars were wetter than average. In contrast, years of widespread fires were dry and associated with phase changes of the PDO, usually from positive (warm) to negative (cold). The influence of the PDO was most evident during the La Niña phase of the ENSO. Widespread fires were also associated with warm/wet conditions 5 years before the fire. We hypothesize that the 5‐year lag between warm/wet conditions and widespread fires may be associated with the time necessary to build up sufficient quantity and continuity of needle litter to support widespread fires. Two periods of unusually high fire activity (1770–1800 and 1920–1950) were each followed by several decades of unusually low fire activity. The switch in each case was associated with strong phase changes in both PDO and ENSO. Main conclusions Climate strongly influences fire regimes in the mountains of north‐western Mexico. Wet/warm years are associated with little fire activity. However, these years may contribute to subsequent fire years by encouraging the production of sufficient needle litter to support more widespread fires that occur in dry/cool years.     

Shifting season of fire and its interaction with fire severity: Impacts on reproductive effort in resprouting plants

Fire regimes shape plant communities but are shifting with changing climate. More frequent fires of increasing intensity are burning across a broader range of seasons. Despite this, impacts that changes in fire season have on plant populations, or how they interact with other fire regime elements, are still relatively understudied. We asked (a) how does the season of fire affect plant vigor, including vegetative growth and flowering after a fire event, and (b) do different functional resprouting groups respond differently to the effects of season of fire? We sampled a total of 887 plants across 36 sites using a space‐for‐time design to assess resprouting vigor and reproductive output for five plant species. Sites represented either a spring or autumn burn, aged one to three years old. Season of fire had the clearest impacts on flowering in Lambertia formosa with a 152% increase in the number of plants flowering and a 45% increase in number of flowers per plant after autumn compared with spring fires. There were also season × severity interactions for total flowers produced for Leptospermum polygalifolium and L. trinervium with both species producing greater flowering in autumn, but only after lower severity fires. Severity of fire was a more important driver in vegetative growth than fire season. Season of fire impacts have previously been seen as synonymous with the effects of fire severity; however, we found that fire season and severity can have clear and independent, as well as interacting, impacts on post‐fire vegetative growth and reproductive response of resprouting species. Overall, we observed that there were positive effects of autumn fires on reproductive traits, while vegetative growth was positively related to fire severity and pre‐fire plant size.