Reduced fire severity offers near-term buffer to climate-driven declines in conifer resilience across the western United States.

Increasing fire severity and warmer, drier postfire conditions are making forests in the western United States (West) vulnerable to ecological transformation. Yet, the relative importance of and interactions between these drivers of forest change remain unresolved, particularly over upcoming decades. Here, we assess how the interactive impacts of changing climate and wildfire activity influenced conifer regeneration after 334 wildfires, using a dataset of postfire conifer regeneration from 10,230 field plots. Our findings highlight declining regeneration capacity across the West over the past four decades for the eight dominant conifer species studied. Postfire regeneration is sensitive to high-severity fire, which limits seed availability, and postfire climate, which influences seedling establishment. In the near-term, projected differences in recruitment probability between low- and high-severity fire scenarios were larger than projected climate change impacts for most species, suggesting that reductions in fire severity, and resultant impacts on seed availability, could partially offset expected climate-driven declines in postfire regeneration. Across 40 to 42% of the study area, we project postfire conifer regeneration to be likely following low-severity but not high-severity fire under future climate scenarios (2031 to 2050). However, increasingly warm, dry climate conditions are projected to eventually outweigh the influence of fire severity and seed availability. The percent of the study area considered unlikely to experience conifer regeneration, regardless of fire severity, increased from 5% in 1981 to 2000 to 26 to 31% by mid-century, highlighting a limited time window over which management actions that reduce fire severity may effectively support postfire conifer regeneration.

Conservation planning integrating natural disturbances: Estimating minimum reserve sizes for an insect disturbance in the boreal forest of eastern Canada.

Large natural disturbances such as insect outbreaks and fire are important processes for biodiversity in forest landscapes. However, few methods exist for incorporating natural disturbances into conservation planning. Intact forest landscapes, such as in the North American boreal forest, can produce large natural disturbance footprints. They also have the potential to support large reserves but size estimates based on natural disturbance are needed to guide reserve design. Historical fire data have been used to estimate minimum dynamic reserves, reserve size estimates based on maintaining natural disturbance dynamics and ensuring resilience to large natural disturbance events. While this has been a significant step towards incorporating natural disturbance into reserve design, managers currently lack guidance on how to apply these concepts in areas where fire is not the dominant natural disturbance. We generalize the minimum dynamic reserve framework to accommodate insect outbreaks and demonstrate the framework in a case study for eastern spruce budworm (Choristoneura fumiferana) in the Canadian boreal forest. Our methods use geospatial analysis to identify minimum dynamic reserves based on a set of spatially explicit initial conditions, and simulation models to test for the maintenance of a set of dynamic conditions over time. We found considerable variability in minimum dynamic reserve size depending on the size of historic budworm disturbance events and the spatial patterns of disturbance-prone vegetation types. The minimum dynamic reserve framework provides an approach for incorporating wide-ranging natural disturbances into biodiversity conservation plans for both pro-active planning in intact landscapes, and reactive planning in more developed regions.

Wildfire, Smoke Exposure, Human Health, and Environmental Justice Need to be Integrated into Forest Restoration and Management.

PURPOSE OF REVIEW: Increasing wildfire size and severity across the western United States has created an environmental and social crisis that must be approached from a transdisciplinary perspective. Climate change and more than a century of fire exclusion and wildfire suppression have led to contemporary wildfires with more severe environmental impacts and human smoke exposure. Wildfires increase smoke exposure for broad swaths of the US population, though outdoor workers and socially disadvantaged groups with limited adaptive capacity can be disproportionally exposed. Exposure to wildfire smoke is associated with a range of health impacts in children and adults, including exacerbation of existing respiratory diseases such as asthma and chronic obstructive pulmonary disease, worse birth outcomes, and cardiovascular events. Seasonally dry forests in Washington, Oregon, and California can benefit from ecological restoration as a way to adapt forests to climate change and reduce smoke impacts on affected communities. RECENT FINDINGS: Each wildfire season, large smoke events, and their adverse impacts on human health receive considerable attention from both the public and policymakers. The severity of recent wildfire seasons has state and federal governments outlining budgets and prioritizing policies to combat the worsening crisis. This surging attention provides an opportunity to outline the actions needed now to advance research and practice on conservation, economic, environmental justice, and public health interests, as well as the trade-offs that must be considered. Scientists, planners, foresters and fire managers, fire safety, air quality, and public health practitioners must collaboratively work together. This article is the result of a series of transdisciplinary conversations to find common ground and subsequently provide a holistic view of how forest and fire management intersect with human health through the impacts of smoke and articulate the need for an integrated approach to both planning and practice.

Community invasion resistance is influenced by interactions between plant traits and site productivity.

Plant communities are predicted to be more resistant to invasion if they are highly productive, harbor species with similar functional traits to invaders, or support species with high competitive potential. However, the strength of competition may decrease with increasing abiotic stress if species more heavily invest in traits that confer stress tolerance over competitive ability, potentially influencing community trait-resistance relationships. Recent research examining how community traits influence invasion resistance has been predominantly focused on single vegetation types, and results between studies are often conflicting. Few studies have evaluated the extent to which abiotic factors and community traits interact to influence invasion along vegetation gradients. Here, we use an in situ seed addition experiment to examine how above- and below-ground plant traits and vegetation type interact to influence community resistance to invasion by a recently introduced annual grass, Ventenata dubia, along a productivity gradient in eastern Oregon, USA. To measure invasion resistance, we evaluated V. dubia biomass in seeded subplots with varying trait compositions across three vegetation types situated along a productivity gradient: scab-flats (sparsely vegetated dwarf-shrublands), low sage-steppe, and ephemeral wet meadows. Trait-resistance relationships were highly context dependent. In wet meadows (the most productive sites), resistance to invasion increased with increasing resident biomass and as community weighted mean trait values for specific leaf area, fine-to-total root volume, and height become more similar to V. dubia's trait values, although these relationships were relatively weak. We did not find evidence that neighboring species influenced invasion resistance in less productive vegetation types, in contrast to our expectations that facilitative interactions may increase with decreasing productivity as posited by the stress-gradient hypothesis. Unlike V. dubia, which heavily invaded all three vegetation types, introduced species with similar trait values, including Bromus tectorum, were not abundant throughout the study area demonstrating V. dubia's unique ability to take advantage of available resources. Our results illustrate how community traits and site productivity interact to influence community resistance to invasion and highlight that communities with lower overall biomass and few functionally similar species to V. dubia may be at the greatest risk for invasion.

Contrasting the efficiency of landscape versus community protection fuel treatment strategies to reduce wildfire exposure and risk.

We examined the financial efficiency and effectiveness of landscape versus community protection fuel treatments to reduce structure exposure and loss to wildfire on a large fire-prone area of central Idaho (USA). The study area contained 63,707 structures distributed in 20 rural communities and resorts, encompassing 13,804 km2. We used simulation modeling to estimate expected structure loss based on burn probability and characteristics of the home ignition zone. We then designed three fuel management strategies that targeted treatments to: 1) the surrounding areas predicted to be the source of exposure to communities from large fires, 2) the home ignition zone, and 3) a combination of the landscape and home ignition zone. We evaluated each treatment scenario in terms of exposure and expected structure loss compared to a no-treatment scenario. The potential revenue from wood products was estimated for each scenario to assess the cost-efficiency. We found that the combined landscape and home ignition zone treatment scenario which treated 5.7% of the study area resulted in the highest overall reduction in predicted exposure (47.5%, 100 structures yr-1) and predicted loss (69.1%, 57 structures yr-1). Home ignition zone treatments provided the best predicted economic and per area treated performance where exposure and loss were reduced by one structure by treating 89 and 111 ha per year, respectively, with an annual cost of $33,645 and $73,672. Revenue from thinning was the highest for landscape fuel treatments and covered 16% of the required investment. This work highlighted economic and risk tradeoffs associated with alternative fuel treatment strategies to protect developed areas from large wildland fires.

Reimagine fire science for the anthropocene.

Fire is an integral component of ecosystems globally and a tool that humans have harnessed for millennia. Altered fire regimes are a fundamental cause and consequence of global change, impacting people and the biophysical systems on which they depend. As part of the newly emerging Anthropocene, marked by human-caused climate change and radical changes to ecosystems, fire danger is increasing, and fires are having increasingly devastating impacts on human health, infrastructure, and ecosystem services. Increasing fire danger is a vexing problem that requires deep transdisciplinary, trans-sector, and inclusive partnerships to address. Here, we outline barriers and opportunities in the next generation of fire science and provide guidance for investment in future research. We synthesize insights needed to better address the long-standing challenges of innovation across disciplines to (i) promote coordinated research efforts; (ii) embrace different ways of knowing and knowledge generation; (iii) promote exploration of fundamental science; (iv) capitalize on the "firehose" of data for societal benefit; and (v) integrate human and natural systems into models across multiple scales. Fire science is thus at a critical transitional moment. We need to shift from observation and modeled representations of varying components of climate, people, vegetation, and fire to more integrative and predictive approaches that support pathways toward mitigating and adapting to our increasingly flammable world, including the utilization of fire for human safety and benefit. Only through overcoming institutional silos and accessing knowledge across diverse communities can we effectively undertake research that improves outcomes in our more fiery future.

Where and why do conifer forests persist in refugia through multiple fire events?

Changing wildfire regimes are causing rapid shifts in forests worldwide. In particular, forested landscapes that burn repeatedly in relatively quick succession may be at risk of conversion when pre-fire vegetation cannot recover between fires. Fire refugia (areas that burn less frequently or severely than the surrounding landscape) support post-fire ecosystem recovery and the persistence of vulnerable species in fire-prone landscapes. Observed and projected fire-induced forest losses highlight the need to understand where and why forests persist in refugia through multiple fires. This research need is particularly acute in the Klamath-Siskiyou ecoregion of southwest Oregon and northwest California, USA, where expected increases in fire activity and climate warming may result in the loss of up to one-third of the region's conifer forests, which are the most diverse in western North America. Here, we leverage recent advances in fire progression mapping and weather interpolation, in conjunction with a novel application of satellite smoke imagery, to model the key controls on fire refugia occurrence and persistence through one, two, and three fire events over a 32-year period. Hotter-than-average fire weather was associated with lower refugia probability and higher fire severity. Refugia that persisted through three fire events appeared to be partially entrained by landscape features that offered protection from fire, suggesting that topographic variability may be an important stabilizing factor as forests pass through successive fire filters. In addition, smoke density strongly influenced fire effects, with fire refugia more likely to occur when smoke was moderate or dense in the morning, a relationship attributable to reduced incoming solar radiation resulting from smoke shading. Results from this study could inform management strategies designed to protect fire-resistant portions of biologically and topographically diverse landscapes.

Climate-change refugia: biodiversity in the slow lane.

Climate-change adaptation focuses on conducting and translating research to minimize the dire impacts of anthropogenic climate change, including threats to biodiversity and human welfare. One adaptation strategy is to focus conservation on climate-change refugia (that is, areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and sociocultural resources). In this Special Issue, recent methodological and conceptual advances in refugia science will be highlighted. Advances in this emerging subdiscipline are improving scientific understanding and conservation in the face of climate change by considering scale and ecosystem dynamics, and looking beyond climate exposure to sensitivity and adaptive capacity. We propose considering refugia in the context of a multifaceted, long-term, network-based approach, as temporal and spatial gradients of ecological persistence that can act as "slow lanes" rather than areas of stasis. After years of discussion confined primarily to the scientific literature, researchers and resource managers are now working together to put refugia conservation into practice.

Integrating Subjective and Objective Dimensions of Resilience in Fire-Prone Landscapes.

Resilience has become a common goal for science-based natural resource management, particularly in the context of changing climate and disturbance regimes. Integrating varying perspectives and definitions of resilience is a complex and often unrecognized challenge to applying resilience concepts to social-ecological systems (SESs) management. Using wildfire as an example, we develop a framework to expose and separate two important dimensions of resilience: the inherent properties that maintain structure, function, or states of an SES and the human perceptions of desirable or valued components of an SES. In doing so, the framework distinguishes between value-free and human-derived, value-explicit dimensions of resilience. Four archetypal scenarios highlight that ecological resilience and human values do not always align and that recognizing and anticipating potential misalignment is critical for developing effective management goals. Our framework clarifies existing resilience theory, connects literature across disciplines, and facilitates use of the resilience concept in research and land-management applications.

Adapt to more wildfire in western North American forests as climate changes.

Wildfires across western North America have increased in number and size over the past three decades, and this trend will continue in response to further warming. As a consequence, the wildland-urban interface is projected to experience substantially higher risk of climate-driven fires in the coming decades. Although many plants, animals, and ecosystem services benefit from fire, it is unknown how ecosystems will respond to increased burning and warming. Policy and management have focused primarily on specified resilience approaches aimed at resistance to wildfire and restoration of areas burned by wildfire through fire suppression and fuels management. These strategies are inadequate to address a new era of western wildfires. In contrast, policies that promote adaptive resilience to wildfire, by which people and ecosystems adjust and reorganize in response to changing fire regimes to reduce future vulnerability, are needed. Key aspects of an adaptive resilience approach are (i) recognizing that fuels reduction cannot alter regional wildfire trends; (ii) targeting fuels reduction to increase adaptation by some ecosystems and residential communities to more frequent fire; (iii) actively managing more wild and prescribed fires with a range of severities; and (iv) incentivizing and planning residential development to withstand inevitable wildfire. These strategies represent a shift in policy and management from restoring ecosystems based on historical baselines to adapting to changing fire regimes and from unsustainable defense of the wildland-urban interface to developing fire-adapted communities. We propose an approach that accepts wildfire as an inevitable catalyst of change and that promotes adaptive responses by ecosystems and residential communities to more warming and wildfire.

An analysis of controls on fire activity in boreal Canada: comparing models built with different temporal resolutions.

Fire regimes of the Canadian boreal forest are driven by certain environmental factors that are highly variable from year to year (e.g., temperature, precipitation) and others that are relatively stable (e.g., land cover, topography). Studies examining the relative influence of these environmental drivers on fire activity suggest that models making explicit use of interannual variability appear to better capture years of climate extremes, whereas those using a temporal average of all available years highlight the importance of land-cover variables. It has been suggested that fire models built at different temporal resolutions may provide a complementary understanding of controls on fire regimes, but this claim has not been tested explicitly with parallel data and modeling approaches. We addressed this issue by building two models of area burned for the period 1980­2010 using 14 explanatory variables to describe ignitions, vegetation, climate, and topography. We built one model at an annual resolution, with climate and some land-cover variables being updated annually, and the other model using 31-year fire "climatology" based on averaged variables. Despite substantial differences in the variables' contributions to the two models, their predictions were broadly similar, which suggests coherence between the spatial patterns of annually varying climate extremes and long-term climate normals. Where the models' predictions diverged, discrepancies between the annual and averaged models could be attributed to specific explanatory variables. For instance, annually updating land cover allowed us to identify a possible negative feedback between flammable biomass and fire activity. These results show that building models at more than one temporal resolution affords a deeper understanding of controls on fire activity in boreal Canada than can be achieved by examining a single model. However, in terms of spatial predictions, the additional effort required to build annual models of fire activity may not always be warranted in this study area. From a management and policy standpoint, this key finding should boost confidence in models that incorporate climatic normals, thereby providing a stronger foundation on which to make decisions on adaptation and mitigation strategies for future fire activity.

Scale-dependent controls on the area burned in the boreal forest of Canada, 1980-2005.

In the boreal forest of North America, as in any fire-prone biome, three environmental factors must coincide for a wildfire to occur: an ignition source, flammable vegetation, and weather that is conducive to fire. Despite recent advances, the relative importance of these factors remains the subject of some debate. The aim of this study was to develop models that identify the environmental controls on spatial patterns in area burned for the period 1980-2005 at several spatial scales in the Canadian boreal forest. Boosted regression tree models were built to relate high-resolution data for area burned to an array of explanatory variables describing ignitions, vegetation, and long-term patterns in fire-conducive weather (i.e., fire climate) at four spatial scales (10(2) km2, 10(3) km2, 10(4) km2, and 10(5) km2). We evaluated the relative contributions of these controls on area burned, as well as their functional relationships, across spatial scales. We also assessed geographic patterns of the influence of wildfire controls. The results indicated that extreme temperature during the fire season was a top control at all spatial scales, followed closely by a wind-driven index of ease of fire spread. However, the contributions of some variables differed substantially among the spatial scales, as did their relationship to area burned. In fact, for some key variables the polarity of relationships was inverted from the finest to the broadest spatial scale. It was difficult to unequivocally attribute values of relative importance to the variables chosen to represent ignitions, vegetation, and climate, as the interdependence of these factors precluded clear partitioning. Furthermore, the influence of a variable on patterns of area burned often changed enormously across the biome, which supports the idea that fire-environment relationships in the boreal forest are complex and nonstationary.

Constraints on global fire activity vary across a resource gradient.

We provide an empirical, global test of the varying constraints hypothesis, which predicts systematic heterogeneity in the relative importance of biomass resources to burn and atmospheric conditions suitable to burning (weather/climate) across a spatial gradient of long-term resource availability. Analyses were based on relationships between monthly global wildfire activity, soil moisture, and mid-tropospheric circulation data from 2001 to 2007, synthesized across a gradient of long-term averages in resources (net primary productivity), annual temperature, and terrestrial biome. We demonstrate support for the varying constraints hypothesis, showing that, while key biophysical factors must coincide for wildfires to occur, the relative influence of resources to burn and moisture/weather conditions on fire activity shows predictable spatial patterns. In areas where resources are always available for burning during the fire season, such as subtropical/tropical biomes with mid-high annual long-term net primary productivity, fuel moisture conditions exert their strongest constraint on fire activity. In areas where resources are more limiting or variable, such as deserts, xeric shrublands, or grasslands/savannas, fuel moisture has a diminished constraint on wildfire, and metrics indicating availability of burnable fuels produced during the antecedent wet growing seasons reflect a more pronounced constraint on wildfire. This macro-scaled evidence for spatially varying constraints provides a synthesis with studies performed at local and regional scales, enhances our understanding of fire as a global process, and indicates how sensitivity to future changes in temperature and precipitation may differ across the world.

Effects of biotic feedback and harvest management on boreal forest fire activity under climate change.

Predictions of future fire activity over Canada's boreal forests have primarily been generated from climate data following assumptions that direct effects of weather will stand alone in contributing to changes in burning. However, this assumption needs explicit testing. First, areas recently burned can be less likely to burn again in the near term, and this endogenous regulation suggests the potential for self-limiting, negative biotic feedback to regional climate-driven increases in fire. Second, forest harvest is ongoing, and resulting changes in vegetation structure have been shown to affect fire activity. Consequently, we tested the assumption that fire activity will be driven by changes in fire weather without regulation by biotic feedback or regional harvest-driven changes in vegetation structure in the mixedwood boreal forest of Alberta, Canada, using a simulation experiment that includes the interaction of fire, stand dynamics, climate change, and clear cut harvest management. We found that climate change projected with fire weather indices calculated from the Canadian Regional Climate Model increased fire activity, as expected, and our simulations established evidence that the magnitude of regional increase in fire was sufficient to generate negative feedback to subsequent fire activity. We illustrate a 39% (1.39-fold) increase in fire initiation and 47% (1.47-fold) increase in area burned when climate and stand dynamics were included in simulations, yet 48% (1.48-fold) and 61% (1.61-fold) increases, respectively, when climate was considered alone. Thus, although biotic feedbacks reduced burned area estimates in important ways, they were secondary to the direct effect of climate on fire. We then show that ongoing harvest management in this region changed landscape composition in a way that led to reduced fire activity, even in the context of climate change. Although forest harvesting resulted in decreased regional fire activity when compared to unharvested conditions, forest composition and age structure was shifted substantially, illustrating a trade-off between management goals to minimize fire and conservation goals to emulate natural disturbance.

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.

Fire in the Earth system.

Fire is a worldwide phenomenon that appears in the geological record soon after the appearance of terrestrial plants. Fire influences global ecosystem patterns and processes, including vegetation distribution and structure, the carbon cycle, and climate. Although humans and fire have always coexisted, our capacity to manage fire remains imperfect and may become more difficult in the future as climate change alters fire regimes. This risk is difficult to assess, however, because fires are still poorly represented in global models. Here, we discuss some of the most important issues involved in developing a better understanding of the role of fire in the Earth system.

Global pyrogeography: the current and future distribution of wildfire.

Climate change is expected to alter the geographic distribution of wildfire, a complex abiotic process that responds to a variety of spatial and environmental gradients. How future climate change may alter global wildfire activity, however, is still largely unknown. As a first step to quantifying potential change in global wildfire, we present a multivariate quantification of environmental drivers for the observed, current distribution of vegetation fires using statistical models of the relationship between fire activity and resources to burn, climate conditions, human influence, and lightning flash rates at a coarse spatiotemporal resolution (100 km, over one decade). We then demonstrate how these statistical models can be used to project future changes in global fire patterns, highlighting regional hotspots of change in fire probabilities under future climate conditions as simulated by a global climate model. Based on current conditions, our results illustrate how the availability of resources to burn and climate conditions conducive to combustion jointly determine why some parts of the world are fire-prone and others are fire-free. In contrast to any expectation that global warming should necessarily result in more fire, we find that regional increases in fire probabilities may be counter-balanced by decreases at other locations, due to the interplay of temperature and precipitation variables. Despite this net balance, our models predict substantial invasion and retreat of fire across large portions of the globe. These changes could have important effects on terrestrial ecosystems since alteration in fire activity may occur quite rapidly, generating ever more complex environmental challenges for species dispersing and adjusting to new climate conditions. Our findings highlight the potential for widespread impacts of climate change on wildfire, suggesting severely altered fire regimes and the need for more explicit inclusion of fire in research on global vegetation-climate change dynamics and conservation planning.

Application of random effects to the study of resource selection by animals.

1. Resource selection estimated by logistic regression is used increasingly in studies to identify critical resources for animal populations and to predict species occurrence. 2. Most frequently, individual animals are monitored and pooled to estimate population-level effects without regard to group or individual-level variation. Pooling assumes that both observations and their errors are independent, and resource selection is constant given individual variation in resource availability. 3. Although researchers have identified ways to minimize autocorrelation, variation between individuals caused by differences in selection or available resources, including functional responses in resource selection, have not been well addressed. 4. Here we review random-effects models and their application to resource selection modelling to overcome these common limitations. We present a simple case study of an analysis of resource selection by grizzly bears in the foothills of the Canadian Rocky Mountains with and without random effects. 5. Both categorical and continuous variables in the grizzly bear model differed in interpretation, both in statistical significance and coefficient sign, depending on how a random effect was included. We used a simulation approach to clarify the application of random effects under three common situations for telemetry studies: (a) discrepancies in sample sizes among individuals; (b) differences among individuals in selection where availability is constant; and (c) differences in availability with and without a functional response in resource selection. 6. We found that random intercepts accounted for unbalanced sample designs, and models with random intercepts and coefficients improved model fit given the variation in selection among individuals and functional responses in selection. Our empirical example and simulations demonstrate how including random effects in resource selection models can aid interpretation and address difficult assumptions limiting their generality. This approach will allow researchers to appropriately estimate marginal (population) and conditional (individual) responses, and account for complex grouping, unbalanced sample designs and autocorrelation.