Alan J. Tepley

Interactions among spruce beetle disturbance, climate change and forest dynamics captured by a forest landscape model

The risk of bark beetle outbreaks is widely predicted to increase because of a warming climate that accelerates temperature-driven beetle population growth and drought stress that impairs host tree defenses. However, few if any studies have explicitly evaluated climatically enhanced beetle population dynamics in relation to climate-driven changes in forest composition and structure that may alter forest suitability for beetle infestation. We synthesized current understanding of the interactions among climate, spruce beetles (Dendroctonus rufipennis) and forest dynamics to parameterize and further advance the bark beetle module of a dynamic forest landscape model (LandClim) that also integrates fire and wind disturbance and climate-driven forest succession. We applied the model to a subalpine watershed in northwestern Colorado to examine the mechanisms and feedbacks that may lead to shifts in forest composition and spruce beetle disturbance under three climate change scenarios. Simulation results suggest increased drought- and beetle-induced reduction of large Engelmann spruce (Picea engelmannii) trees while Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) increased in dominance throughout the study area under all climate change scenarios. This shift in forest composition and structure counterbalances the enhancing effects of accelerated beetle population development and increased drought-induced susceptibility of spruce to beetles. As a result, we projected a long-term decrease in beetle-induced spruce mortality to below historical values under all climate scenarios at low elevations (<2800 m asl). Beetle-induced spruce mortality above 2800 m asl and under moderate climate change was slightly higher and more variable than under historical conditions but decreased to 36% and 6% of historical values under intermediate and extreme climate change, respectively. Because mechanisms driving beetle disturbance dynamics are similar across different bark beetle species, we argue that the depletion of host trees due to drought and beetle disturbance may also be important in other climate-sensitive beetle-host systems. We advocate for the consideration of climate-driven shifts in forest and disturbance dynamics in devising adaptive management strategies.

Spatiotemporal fire dynamics in mixed‐conifer and aspen forests in the San Juan Mountains of southwestern Colorado, USA

Mixed-severity fire regimes may be the most extensive yet poorly understood fire regimes of western North America. Understanding their long-term spatiotemporal dynamics is central to debates regarding altered fire regimes and the need for restoration in the context of changing climate and nearly a century of active fire suppression. However, the complexity of fire patterns and forest stand and landscape structures characteristic of mixed-severity regimes poses a substantial challenge to understanding their long-term dynamics. In this study, we develop analysis methods aimed at understanding the fire-driven forest dynamics of mixed-severity systems and apply them in the San Juan Mountains of southwestern Colorado. We sampled fire scars, stand structure, and >4300 tree ages across two 1340-ha landscapes (Williams Creek and Squaretop Mountain) that span the environmental gradient of montane mixed-conifer and aspen forests. New approaches were applied to identify pulses of tree recruitment, evaluate climate and fire as potential drivers of synchronous recruitment pulses, and combine fire scar and recruitment data to reconstruct fires. The reconstructions provided detailed fire history for each stand, which in turn was used to develop a fire-severity metric, compare fire frequency and severity by forest type, and develop a simulation procedure to evaluate the degree to which tree regeneration depended on fire by species within each forest type. Twenty fires were reconstructed since 1685 at Williams Creek and 13 fires since 1748 at Squaretop Mountain. Patterns of fire severity varied within each fire and over successive events, including high-severity patches of hundreds of hectares in both study areas. Dry mixed-conifer forests experienced relatively short fire intervals (mean 21 years) and low fire severity, and regeneration of the main conifer species was largely dependent on open conditions sustained over successive fires. Moist mixed-conifer forests experienced longer fire intervals (mean 32 years) and a broader range of severities, and fire-caused canopy openings were important for initiating pulses of tree recruitment. Most (83%) aspen stands included two or more post-fire cohorts. The methods presented here can be adapted to other mixed-severity systems to better understand their long-term spatial and temporal dynamics and develop restoration priorities.

Fire-Vegetation Feedbacks and Alternative States: Common Mechanisms of Temperate Forest Vulnerability to Fire in Southern South America and New Zealand

ABSTRACT In the context of global warming and increasing impacts of invasive plants and animals, we examine how positive fire–vegetation feedbacks are increasing the vulnerability of pyrophobic temperate forests to conversion to pyrophytic non-forest vegetation in southern South America and New Zealand. We extensively review the relevant literature to reveal how these temperate southern hemisphere floras have generated similar positive fire–vegetation feedback mechanisms resulting in increased vulnerability to anthropogenically altered fire regimens. For the two regions, we address the following questions. 1. What are the major plant species, physiognomic types and functional types characteristic of pyrophytic versus pyrophobic vegetation types and how do their traits affect flammability, resistance to fire and recovery after fire? 2. What are the roles of herbivory and microclimate in enhancing fire–vegetation feedbacks? 3. Are there similarities in trends of cover type transitions in relation to altered fire regimens? 4. How are climate change, land-use trends and the effects of introduced plants and animals affecting the vulnerability of these ecosystems to fire-induced transitions to alternative stable states? Most temperate forests of New Zealand and southern South America evolved under conditions of low fire frequencies so few taxa became adapted to recurrent fire. Current dichotomous landscapes consisting of juxtaposed pyrophobic and pyrophytic vegetation types are the outcome of the expansion of fire-resilient and fire-promoting species associated with the arrival of humans. Despite considerable differences in human history and biogeographic history, the case studies presented here show remarkable parallels in life-history traits of the key pyrophobic taxa, fire–vegetation feedback mechanisms, overall ecosystem responses to anthropogenic alteration of fire regimens, and likely vulnerability to expected global change influences on future fire regimens.

Positive Feedbacks to Fire-Driven Deforestation Following Human Colonization of the South Island of New Zealand

Altered fire regimes in the face of climatic and land-use change could potentially transform large areas from forest to shorter-statured or open-canopy vegetation. There is growing concern that once initiated, these nonforested landscapes could be perpetuated almost indefinitely through a suite of positive feedbacks with fire. The rapid deforestation of much of New Zealand following human settlement (ca. 750 years ago) provides a rare opportunity to evaluate the feedback mechanisms that facilitated such extensive transformation and thereby help us to identify factors that confer vulnerability or resilience to similar changes in other regions. Here we evaluate the structure of living and dead vegetation (fuel loading) and microclimate (fuel moisture) in beech (Nothofagaceae) forests and adjacent stands that burned within the last 60–140 years and are dominated by mānuka (Leptospermum scoparium) or kānuka (Kunzea spp.). We show that the burning of beech forests initiates a positive feedback cycle whereby the loss of microclimatic amelioration under the dense forest canopy and the abundant fine fuels that dry readily beneath the sparse mānuka/kānuka canopy enables perpetuation of these stands by facilitating repeated burning. Beech regeneration was limited to a narrow zone along the margin of unburned stands. The high flammability of vegetation that develops after fire and the long time to forest recovery were the primary factors that facilitated extensive deforestation with the introduction of human-ignited fire. Evaluating these two characteristics may be key to determining which regions may be near a tipping point where relatively small land-use- or climatically driven changes to fire regimes could bring about extensive deforestation.

Influences of fire–vegetation feedbacks and post‐fire recovery rates on forest landscape vulnerability to altered fire regimes

1Smithsonian Conservation Biology Institute, Front Royal, VA, USA; 2Department of Mathematics, Oregon State University, Corvallis, OR, USA; 3Department of Geography, University of Colorado at Boulder, Boulder, CO, USA; 4School of Environment, University of Auckland, Auckland, New Zealand; 5Department of Geography, Portland State University, Portland, OR, USA; 6Laboratorio Ecotono, INIBIOMA, CONICET-Universidad Nacional del Comahue, Bariloche, Río Negro, Argentina; 7Departamento de Ecología, Universidad Nacional del Comahue, Bariloche, Río Negro, Argentina and 8Center for Tropical Forest Science– Forest Global Earth Observatory, Smithsonian Tropical Research Institute, Panama

Long-term growth-increment chronologies reveal diverse influences of climate forcing on freshwater and forest biota in the Pacific Northwest

Analyses of how organisms are likely to respond to a changing climate have focused largely on the direct effects of warming temperatures, though changes in other variables may also be important, particularly the amount and timing of precipitation. Here, we develop a network of eight growth-increment width chronologies for freshwater mussel species in the Pacific Northwest, United States and integrate them with tree-ring data to evaluate how terrestrial and aquatic indicators respond to hydroclimatic variability, including river discharge and precipitation. Annual discharge averaged across water years (October 1-September 30) was highly synchronous among river systems and imparted a coherent pattern among mussel chronologies. The leading principal component of the five longest mussel chronologies (1982-2003; PC1(mussel)) accounted for 47% of the dataset variability and negatively correlated with the leading principal component of river discharge (PC1(discharge); r = -0.88; P < 0.0001). PC1(mussel) and PC1(discharge) were closely linked to regional wintertime precipitation patterns across the Pacific Northwest, the season in which the vast majority of annual precipitation arrives. Mussel growth was also indirectly related to tree radial growth, though the nature of the relationships varied across the landscape. Negative correlations occurred in forests where tree growth tends to be limited by drought while positive correlations occurred in forests where tree growth tends to be limited by deep or lingering snowpack. Overall, this diverse assemblage of chronologies illustrates the importance of winter precipitation to terrestrial and freshwater ecosystems and suggests that a complexity of climate responses must be considered when estimating the biological impacts of climate variability and change.

Disequilibrium of Fire-prone Forests Sets the Stage for a Rapid Decline in Conifer Dominance during the 21st Century

As trees are long-lived organisms, the impacts of climate change on forest communities may not be apparent on the time scale of years to decades. While lagged responses to environmental change are common in forested systems, potential for abrupt transitions under climate change may occur in environments where alternative vegetation states are influenced by disturbances, such as fire. The Klamath mountains (northern California and southwest Oregon, USA) are currently dominated by carbon rich and hyper-diverse temperate conifer forests, but climate change could disrupt the mechanisms promoting forest stability– regeneration and fire tolerance— via shifts in the fire regime in conjunction with lower fitness of conifers under a hotter climate. Understanding how this landscape will respond to near-term climate change (before 2100) is critical for predicting potential climate change feedbacks and to developing sound forest conservation and management plans. Using a landscape simulation model, we estimate that 1/3 of the Klamath could transition from conifer forest to shrub/hardwood chaparral, triggered by an enhanced fire activity coupled with lower post-fire conifer establishment. Such shifts were more prevalent under higher climate change forcing (RCP 8.5) but were also simulated under the climate of 1950-2000, reflecting the joint influences of early warming trends and historical forest legacies. Our results demonstrate that there is a large potential for loss of conifer forest dominance—and associated carbon stocks and biodiversity- in the Klamath before the end of the century, and that some losses would likely occur even without the influence of climate change. Thus, in the Klamath and other forested landscapes subject to similar feedback dynamics, major ecosystem shifts should be expected when climate change disrupts key stabilizing feedbacks that maintain the dominance of long-lived, slowly regenerating trees.

Alternative stable equilibria and critical thresholds created by fire regimes and plant responses in a fire-prone community

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