The complexity of biological disturbance agents, fuels heterogeneity, and fire in coniferous forests of the western United States.

Forest biological disturbance agents (BDAs) are insects, pathogens, and parasitic plants that affect tree decline, mortality, and forest ecosystems processes. BDAs are commonly thought to increase the likelihood and severity of fire by converting live standing trees to more flammable, dead and downed fuel. However, recent research indicates that BDAs do not necessarily increase, and can reduce, the likelihood or severity of fire. This has led to confusion regarding the role of BDAs in influencing fuels and fire in fire-prone western United States forests. Here, we review the existing literature on BDAs and their effects on fuels and fire in the western US and develop a conceptual framework to better understand the complex relationships between BDAs, fuels and fire. We ask: 1) What are the major BDA groups in western US forests that affect fuels? and 2) How do BDA-affected fuels influence fire risk and outcomes? The conceptual framework is rooted in the spatiotemporal aspects of BDA life histories, which drive forest impacts, fuel characteristics and if ignited, fire outcomes. Life histories vary among BDAs from episodic, landscape-scale outbreaks (bark beetles, defoliators), to chronic, localized disturbance effects (dwarf mistletoes, root rots). Generally, BDAs convert aboveground live biomass to dead biomass, decreasing canopy fuels and increasing surface fuels. However, the rate of conversion varies with time-since-event and among BDAs and forest types, resulting in a wide range of effects on the amount of dead fuels at any given time and place, which interacts with the structure and composition of the stand before and subsequent to BDA events. A major influence on fuels may be that BDAs have emerged as dominant agents of forest heterogeneity creation. Because BDAs play complex roles in fuels and fire heterogeneity across the western US which are further complicated by interactions with climate change, drought, and forest management (fire suppression), their impacts on fuels, fire and ecological consequences cannot be categorized simply as positive or negative but need to be evaluated within the context of BDA life histories and ecosystem dynamics.

Natural disturbance impacts on trade-offs and co-benefits of forest biodiversity and carbon.

With accelerating environmental change, understanding forest disturbance impacts on trade-offs between biodiversity and carbon dynamics is of high socio-economic importance. Most studies, however, have assessed immediate or short-term effects of disturbance, while long-term impacts remain poorly understood. Using a tree-ring-based approach, we analysed the effect of 250 years of disturbances on present-day biodiversity indicators and carbon dynamics in primary forests. Disturbance legacies spanning centuries shaped contemporary forest co-benefits and trade-offs, with contrasting, local-scale effects. Disturbances enhanced carbon sequestration, reaching maximum rates within a comparatively narrow post-disturbance window (up to 50 years). Concurrently, disturbance diminished aboveground carbon storage, which gradually returned to peak levels over centuries. Temporal patterns in biodiversity potential were bimodal; the first maximum coincided with the short-term post-disturbance carbon sequestration peak, and the second occurred during periods of maximum carbon storage in complex old-growth forest. Despite fluctuating local-scale trade-offs, forest biodiversity and carbon storage remained stable across the broader study region, and our data support a positive relationship between carbon stocks and biodiversity potential. These findings underscore the interdependencies of forest processes, and highlight the necessity of large-scale conservation programmes to effectively promote both biodiversity and long-term carbon storage, particularly given the accelerating global biodiversity and climate crises.

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.

The climate sensitivity of carbon, timber, and species richness covaries with forest age in boreal-temperate North America.

Climate change threatens the provisioning of forest ecosystem services and biodiversity (ESB). The climate sensitivity of ESB may vary with forest development from young to old-growth conditions as structure and composition shift over time and space. This study addresses knowledge gaps hindering implementation of adaptive forest management strategies to sustain ESB. We focused on a number of ESB indicators to (a) analyze associations among carbon storage, timber growth rate, and species richness along a forest development gradient; (b) test the sensitivity of these associations to climatic changes; and (c) identify hotspots of climate sensitivity across the boreal-temperate forests of eastern North America. From pre-existing databases and literature, we compiled a unique dataset of 18,507 forest plots. We used a full Bayesian framework to quantify responses of nine ESB indicators. The Bayesian models were used to assess the sensitivity of these indicators and their associations to projected increases in temperature and precipitation. We found the strongest association among the investigated ESB indicators in old forests (>170 years). These forests simultaneously support high levels of carbon storage, timber growth, and species richness. Older forests also exhibit low climate sensitivity of associations among ESB indicators as compared to younger forests. While regions with a currently low combined ESB performance benefitted from climate change, regions with a high ESB performance were particularly vulnerable to climate change. In particular, climate sensitivity was highest east and southeast of the Great Lakes, signaling potential priority areas for adaptive management. Our findings suggest that strategies aimed at enhancing the representation of older forest conditions at landscape scales will help sustain ESB in a changing world.

Intermediate-severity wind disturbance in mature temperate forests: legacy structure, carbon storage, and stand dynamics.

Wind is one of the most important natural disturbances influencing forest structure, ecosystem function, and successional processes worldwide. This study quantifies the stand-scale effects of intermediate-severity windstorms (i.e., blowdowns) on (1) live and dead legacy structure, (2) aboveground carbon storage, and (3) tree regeneration and associated stand dynamics at four mature, mixed hardwood-conifer forest sites in the northeastern United States. We compare wind-affected forests to adjacent reference conditions (i.e., undisturbed portions of the same stands) 0-8 yr post-blowdown using parametric (ANOVA) and nonparametric (NMS ordination) analyses. We supplement inventory plots and downed coarse woody detritus (DCWD) transects with hemispherical photography to capture spatial variation in the light environment. Although recent blowdowns transferred a substantial proportion of live overstory trees to DCWD, residual live tree basal area was high (19-59% of reference areas). On average, the initial post-blowdown ratio of DCWD carbon to standing live tree carbon was 2.72 in blowdown stands and 0.18 in reference stands, indicating a large carbon transfer from live to dead pools. Despite these dramatic changes, structural complexity remained high in blowdown areas, as indicated by the size and species distributions of overstory trees, abundance of sound and rotten downed wood, spatial patterns of light availability, and variability of understory vegetation. Furthermore, tree species composition was similar between blowdown and reference areas at each site, with generally shade-tolerant species dominating across multiple canopy strata. Community response to intermediate-severity blowdown at these sites suggests a dynamic in which disturbance maintains late-successional species composition rather than providing a regeneration opportunity for shade-intolerant, pioneer species. Our findings suggest that intermediate-severity wind disturbances can contribute to stand-scale structural complexity as well as development toward late-successional species composition, at least when shade-tolerant regeneration is present pre-blowdown. Advance regeneration thus enhances structural and compositional resilience to this type of disturbance. This study provides a baseline for multi-cohort silvicultural systems designed to restore heterogeneity associated with natural disturbance dynamics.