Craig D. Allen

APPLIED HISTORICAL ECOLOGY: USING THE PAST TO MANAGE FOR THE FUTURE

Applied historical ecology is the use of historical knowledge in the man- agement of ecosystems. Historical perspectives increase our understanding of the dynamic nature of landscapes and provide a frame of reference for assessing modern patterns and processes. Historical records, however, are often too brief or fragmentary to be useful, or they are not obtainable for the process or structure of interest. Even where long historical time series can be assembled, selection of appropriate reference conditions may be com- plicated by the past influence of humans and the many potential reference conditions encompassed by nonequilibrium dynamics. These complications, however, do not lessen the value of history; rather they underscore the need for multiple, comparative histories from many locations for evaluating both cultural and natural causes of variability, as well as for characterizing the overall dynamical properties of ecosystems. Historical knowledge may not simplify the task of setting management goals and making decisions, but 20th century trends, such as increasingly severe wildfires, suggest that disregarding history can be perilous. We describe examples from our research in the southwestern United States to illustrate some of the values and limitations of applied historical ecology. Paleoecological data from packrat middens and other natural archives have been useful for defining baseline conditions of vegetation communities, determining histories and rates of species range expansions and contractions, and discriminating between natural and cultural causes of environmental change. We describe a montane grassland restoration project in northern New Mexico that was justified and guided by an historical sequence of aerial photographs showing progressive tree invasion during the 20th century. Likewise, fire scar chronologies have been widely used to justify and guide fuel reduction and natural fire reintroduction in forests. A south- western network of fire histories illustrates the power of aggregating historical time series across spatial scales. Regional fire patterns evident in these aggregations point to the key role of interannual lags in responses of fuels and fire regimes to the El Nino-Southern Oscillation (wet/dry cycles), with important implications for long-range fire hazard fore- casting. These examples of applied historical ecology emphasize that detection and expla- nation of historical trends and variability are essential to informed management.

On underestimation of global vulnerability to tree mortality and forest die‐off from hotter drought in the Anthropocene

Patterns, mechanisms, projections, and consequences of tree mortality and associated broad-scale forest die-off due to drought accompanied by warmer temperatures—“hotter drought”, an emerging characteristic of the Anthropocene—are the focus of rapidly expanding literature. Despite recent observational, experimental, and modeling studies suggesting increased vulnerability of trees to hotter drought and associated pests and pathogens, substantial debate remains among research, management and policy-making communities regarding future tree mortality risks. We summarize key mortality-relevant findings, differentiating between those implying lesser versus greater levels of vulnerability. Evidence suggesting lesser vulnerability includes forest benefits of elevated [CO2] and increased water-use efficiency; observed and modeled increases in forest growth and canopy greening; widespread increases in woody-plant biomass, density, and extent; compensatory physiological, morphological, and genetic mechanisms; dampen...

Tree die‐off in response to global change‐type drought: mortality insights from a decade of plant water potential measurements

Global climate change is projected to produce warmer, longer, and more frequent droughts, referred to here as “global change-type droughts”, which have the potential to trigger widespread tree die-off. However, drought-induced tree mortality cannot be predicted with confidence, because long-term field observations of plant water stress prior to, and culminating in, mortality are rare, precluding the development and testing of mechanisms. Here, we document plant water stress in two widely distributed, co-occurring species, pinon pine (Pinus edulis) and juniper (Juniperus monosperma), over more than a decade, leading up to regional-scale die-off of pinon pine trees in response to global change-related drought. Pinon leaf water potentials remained substantially below their zero carbon assimilation point for at least 10 months prior to dying, in contrast to those of juniper, which rarely dropped below their zero-assimilation point. These data suggest that pinon mortality was driven by protracted water stress,...

ECOHYDROLOGY OF A RESOURCE‐CONSERVING SEMIARID WOODLAND: EFFECTS OF SCALE AND DISTURBANCE

In semiarid landscapes, the linkage between runoff and vegetation is a par- ticularly close one. In this paper we report on the results of a long-term and multiple-scale study of interactions between runoff, erosion, and vegetation in a pinon-juniper woodland in New Mexico. We use our results to address three knowledge gaps: (1) the temporal scaling relationships between precipitation and runoff; (2) the effects of spatial scale on runoff and erosion, as influenced by vegetation; and (3) the influence of disturbance on these relationships. On the basis of our results, we tested three assumptions that represent current thinking in these areas (as evidenced, for example, by explicit or implicit assump- tions embedded in commonly used models). The first assumption, that aggregated precip- itation can be used as a surrogate for total runoff in semiarid environments, was not verified by our findings. We found that when runoff is generated mainly by overland flow in these systems, aggregated precipitation amounts alone (by year, season, or individual event) are a poor predictor of runoff amounts. The second assumption, that at the hillslope and smaller scales runoff and erosion are independent of spatial scale, was likewise not verified. We found that the redistribution of water and sediment within the hillslope was substantial and that there was a strong and nonlinear reduction in unit-area runoff and erosion with in- creasing scale (our scales were slope lengths ranging fro m1mt o 105 m). Thethird assumption, that disturbance-related increases in runoff and erosion remain constant with time, was partially verified. We found that for low-slope-gradient sites, disturbance led to accelerated runoff and erosion, and these conditions may persist for a decade or longer. On the basis of our findings, we further suggest that (a) disturbance alters the effects of scale on runoff and erosion in a predictable way—scale relationships in degraded areas will be fundamentally different from those in nondegraded areas because more runoff will escape off site and erosion rates will be much higher; and (b) there exists a slope threshold, below which semiarid landscapes will eventually recover following disturbance and above which there will be no recovery without mitigation or remediation.

Forest responses to increasing aridity and warmth in the southwestern United States

In recent decades, intense droughts, insect outbreaks, and wildfires have led to decreasing tree growth and increasing mortality in many temperate forests. We compared annual tree-ring width data from 1,097 populations in the coterminous United States to climate data and evaluated site-specific tree responses to climate variations throughout the 20th century. For each population, we developed a climate-driven growth equation by using climate records to predict annual ring widths. Forests within the southwestern United States appear particularly sensitive to drought and warmth. We input 21st century climate projections to the equations to predict growth responses. Our results suggest that if temperature and aridity rise as they are projected to, southwestern trees will experience substantially reduced growth during this century. As tree growth declines, mortality rates may increase at many sites. Increases in wildfires and bark-beetle outbreaks in the most recent decade are likely related to extreme drought and high temperatures during this period. Using satellite imagery and aerial survey data, we conservatively calculate that ≈2.7% of southwestern forest and woodland area experienced substantial mortality due to wildfires from 1984 to 2006, and ≈7.6% experienced mortality associated with bark beetles from 1997 to 2008. We estimate that up to ≈18% of southwestern forest area (excluding woodlands) experienced mortality due to bark beetles or wildfire during this period. Expected climatic changes will alter future forest productivity, disturbance regimes, and species ranges throughout the Southwest. Emerging knowledge of these impending transitions informs efforts to adaptively manage southwestern forests.

Viewpoint: Sustainability of piñon-juniper ecosystems - A unifying perspective of soil erosion thresholds

Many pinon-juniper ecosystems in the western U.S. are subject to accelerated erosion while others are undergoing little or no erosion. Controversy has developed over whether invading or encroaching pinon and juniper species are inherently harmful to rangeland ecosystems. We developed a conceptual model of soil erosion in pinon-juniper ecosystems that is consistent with both sides of the controversy and suggests that the diverse perspectives on this issue arise from threshold effects operating under very different site conditions. Soil erosion rate can be viewed as a function of (1) site erosion potential (SEP), determined by climate, geomorphology and soil erodibility; and (2) ground cover. Site erosion potential and cover act synergistically to determine soil erosion rates, as evident even from simple USLE predictions of erosion. In pinon-juniper ecosystems with high SEP, the erosion rate is highly sensitive to ground cover and can cross a threshold so that erosion increases dramatically in response to a small decrease in cover. The sensitivity of erosion rate to SEP and cover can be visualized as a cusp catastrophe surface on which changes may occur rapidly and irreversibly. The mechanisms associated with a rapid shift from low to high erosion rate can be illustrated using percolation theory to incorporate spatial, temporal, and scale-dependent patterns of water storage capacity on a hillslope. Percolation theory demonstrates how hillslope runoff can undergo a threshold response to a minor change in storage capacity. Our conceptual model suggests that pinon and juniper contribute to accelerated erosion only under a limited range of site conditions which, however, may exist over large areas.

Tree mortality from drought, insects, and their interactions in a changing climate

Climate change is expected to drive increased tree mortality through drought, heat stress, and insect attacks, with manifold impacts on forest ecosystems. Yet, climate-induced tree mortality and biotic disturbance agents are largely absent from process-based ecosystem models. Using data sets from the western USA and associated studies, we present a framework for determining the relative contribution of drought stress, insect attack, and their interactions, which is critical for modeling mortality in future climates. We outline a simple approach that identifies the mechanisms associated with two guilds of insects - bark beetles and defoliators - which are responsible for substantial tree mortality. We then discuss cross-biome patterns of insect-driven tree mortality and draw upon available evidence contrasting the prevalence of insect outbreaks in temperate and tropical regions. We conclude with an overview of tools and promising avenues to address major challenges. Ultimately, a multitrophic approach that captures tree physiology, insect populations, and tree-insect interactions will better inform projections of forest ecosystem responses to climate change.

Larger trees suffer most during drought in forests worldwide

The frequency of severe droughts is increasing in many regions around the world as a result of climate change1–3. Droughts alter the structure and function of forests4,5. Site- and region-specific studies suggest that large trees, which play keystone roles in forests6 and can be disproportionately important to ecosystem carbon storage7 and hydrology8, exhibit greater sensitivity to drought than small trees4,5,9,10. Here, we synthesize data on tree growth and mortality collected during 40 drought events in forests worldwide to see whether this size-dependent sensitivity to drought holds more widely. We find that droughts consistently had a more detrimental impact on the growth and mortality rates of larger trees. Moreover, drought-related mortality increased with tree size in 65% of the droughts examined, especially when community-wide mortality was high or when bark beetles were present. The more pronounced drought sensitivity of larger trees could be underpinned by greater inherent vulnerability to hydraulic stress11–14, the higher radiation and evaporative demand experienced by exposed crowns4,15, and the tendency for bark beetles to preferentially attack larger trees16. We suggest that future droughts will have a more detrimental impact on the growth and mortality of larger trees, potentially exacerbating feedbacks to climate change.

Rapid warming accelerates tree growth decline in semi-arid forests of Inner Asia

Forests around the world are subject to risk of high rates of tree growth decline and increased tree mortality from combinations of climate warming and drought, notably in semi-arid settings. Here, we assess how climate warming has affected tree growth in one of the worlds most extensive zones of semi-arid forests, in Inner Asia, a region where lack of data limits our understanding of how climate change may impact forests. We show that pervasive tree growth declines since 1994 in Inner Asia have been confined to semi-arid forests, where growing season water stress has been rising due to warming-induced increases in atmospheric moisture demand. A causal link between increasing drought and declining growth at semi-arid sites is corroborated by correlation analyses comparing annual climate data to records of tree-ring widths. These ring-width records tend to be substantially more sensitive to drought variability at semi-arid sites than at semi-humid sites. Fire occurrence and insect/pathogen attacks have increased in tandem with the most recent (2007-2009) documented episode of tree mortality. If warming in Inner Asia continues, further increases in forest stress and tree mortality could be expected, potentially driving the eventual regional loss of current semi-arid forests.

Extended megadroughts in the southwestern United States during Pleistocene interglacials

The potential for increased drought frequency and severity linked to anthropogenic climate change in the semi-arid regions of the southwestern United States (US) is a serious concern. Multi-year droughts during the instrumental period and decadal-length droughts of the past two millennia were shorter and climatically different from the future permanent, ‘dust-bowl-like’ megadrought conditions, lasting decades to a century, that are predicted as a consequence of warming. So far, it has been unclear whether or not such megadroughts occurred in the southwestern US, and, if so, with what regularity and intensity. Here we show that periods of aridity lasting centuries to millennia occurred in the southwestern US during mid-Pleistocene interglacials. Using molecular palaeotemperature proxies to reconstruct the mean annual temperature (MAT) in mid-Pleistocene lacustrine sediment from the Valles Caldera, New Mexico, we found that the driest conditions occurred during the warmest phases of interglacials, when the MAT was comparable to or higher than the modern MAT. A collapse of drought-tolerant C4 plant communities during these warm, dry intervals indicates a significant reduction in summer precipitation, possibly in response to a poleward migration of the subtropical dry zone. Three MAT cycles ∼2 °C in amplitude occurred within Marine Isotope Stage (MIS) 11 and seem to correspond to the muted precessional cycles within this interglacial. In comparison with MIS 11, MIS 13 experienced higher precessional-cycle amplitudes, larger variations in MAT (4–6 °C) and a longer period of extended warmth, suggesting that local insolation variations were important to interglacial climatic variability in the southwestern US. Comparison of the early MIS 11 climate record with the Holocene record shows many similarities and implies that, in the absence of anthropogenic forcing, the region should be entering a cooler and wetter phase.