Linda B. Brubaker

The Sustainable Biosphere Initiative: An Ecological Research Agenda: A Report from the Ecological Society of America

In this document, the Ecological Society of America proposes the Sustainable Biosphere Initiative (SBI), an initiative that focuses on the necessary role of ecological science in the wise management of Earths resources and the maintenance of Earths life support systems. This document is intended as a call to arms for all ecologists, but it will also serve as a means to communicate with individuals in other disciplines with whom ecologists must join forces to address a common predicament. This document focuses primarily on the acquisition of ecological knowledge. It identifies the ecological research programs of highest priority and recommends steps required to pursue research objectives. The document also lays the groundwork for improving the communication and application of ecological knowledge. The SBI proposes three research priorities: global change; biological diversity; and sustainable ecological systems.

SPATIAL CONTROLS OF HISTORICAL FIRE REGIMES: A MULTISCALE EXAMPLE FROM THE INTERIOR WEST, USA

Our objective was to infer the controls of spatial variation in historical fire regimes. We reconstructed a multicentury history of fire frequency, size, season, and severity from fire scars and establishment dates of 1426 trees sampled on grids in four watersheds (-64 plots, over -1620 ha each) representative of the Blue Mountains, Oregon and Wash- ington, USA. The influence of regional climate, a top-down control, was inferred from among-watershed variation in fire regimes, while the influence of local topography, a bot- tom-up control, was inferred from within-watershed variation. Before about 1900, fire regimes varied among and within watersheds, suggesting that both top-down and bottom- up controls were important. At the regional scale, dry forests (dominated by ponderosa pine), burned twice as frequently and earlier in the growing season in southern watersheds than in northern watersheds, consistent with longer and drier fire seasons to the south. Mesic forests (dominated by subalpine fir or grand fir) probably also burned more frequently to the south. At the local scale, fire frequency varied with different parameters of topography in watersheds with steep terrain, but not in the watershed with gentle terrain. Frequency varied with aspect in watersheds where topographic facets are separated by significant barriers to fire spread, but not in watersheds where such facets interfinger without fire barriers. Frequency varied with elevation where elevation and aspect interact to create gradients in snow-cover duration and also where steep talus interrupts fuel continuity. Frequency did not vary with slope within any watershed. The presence of both regional- scale and local-scale variation in the Blue Mountains suggests that top-down and bottom- up controls were both important and acted simultaneously to influence fire regimes in the past. However, an abrupt decline in fire frequency around 1900 was much greater than any regional or local variation in the previous several centuries and indicates that 20th-century fire regimes in these watersheds were dramatically affected by additional controls such as livestock grazing and fire suppression. Our results demonstrate the usefulness of examining spatial variation in historical fire regimes across scales as a means for inferring their controls.

Climate change and Arctic ecosystems: 2. Modeling, paleodata-model comparisons, and future projections

Large variations in the composition, structure, and function of Arctic ecosystems are determined by climatic gradients, especially of growing-season warmth, soil moisture, and snow cover. A unified circumpolar classification recognizing five types of tundra was developed. The geographic distributions of vegetation types north of 55degreesN, including the position of the forest limit and the distributions of the tundra types, could be predicted from climatology using a small set of plant functional types embedded in the biogeochemistry-biogeography model BIOME4. Several palaeoclimate simulations for the last glacial maximum (LGM) and mid-Holocene were used to explore the possibility of simulating past vegetation patterns, which are independently known based on pollen data. The broad outlines of observed changes in vegetation were captured. LGM simulations showed the major reduction of forest, the great extension of graminoid and forb tundra, and the restriction of low- and high-shrub tundra (although not all models produced sufficiently dry conditions to mimic the full observed change). Mid-Holocene simulations reproduced the contrast between northward forest extension in western and central Siberia and stability of the forest limit in Beringia. Projection of the effect of a continued exponential increase in atmospheric CO2 concentration, based on a transient ocean-atmosphere simulation including sulfate aerosol effects, suggests a potential for larger changes in Arctic ecosystems during the 21st century than have occurred between mid-Holocene and present. Simulated physiological effects of the CO2 increase (to >700 ppm) at high latitudes were slight compared with the effects of the change in climate.

Vegetation mediated the impacts of postglacial climate change on fire regimes in the south-central Brooks Range, Alaska

We examined direct and indirect impacts of millennial-scale climate change on fire regimes in the south-central Brooks Range, Alaska, USA, using four lake sediment records and existing paleoclimate interpretations. New techniques were introduced to identify charcoal peaks semi-objectively and to detect statistical differences between fire regimes. Peaks in charcoal accumulation rates provided estimates of fire return intervals (FRIs), which were compared among vegetation zones identified by fossil pollen and stomata. Climatic warming between ca. 15 000-9000 yr BP (calendar years before Common Era (CE) 1950) coincided with shifts in vegetation from herb tundra to shrub tundra to deciduous woodlands, all novel species assemblages relative to modern vegetation. Two sites cover this period and show decreased FRIs with the transition from herb to Betula-dominated shrub tundra ca. 13 300- 14 300 yr BP (FRImean ¼ 144 yr; 95% CI ¼ 120-169 yr), when climate warmed but remained cooler than present. Although warming would have favored shorter FRIs in the shrub tundra, the shift to more continuous, flammable fuels relative to herb tundra was probably a more important cause of increased burning. Similarly, a vegetation shift to Populus-dominated deciduous woodlands overrode the influence of warmer- and drier-than-present summers, resulting in lower fire activity from ca. 10 300-8250 yr BP (FRImean ¼ 251 yr; 95% CI ¼ 156- 347 yr). Three sites record the mid-to-late Holocene, when climatic cooling and moistening allowed Picea glauca forest-tundra and P. mariana boreal forests to establish ca. 8000 and 5500 yr BP, respectively. FRIs in forest-tundra were either similar to or shorter than those in the deciduous woodlands (FRImean range ¼ 131-238 yr). The addition of P. mariana ca. 5500 yr BP increased landscape flammability, overrode the effects of climatic cooling and moistening and resulted in lower FRIs (FRImean ¼ 145 yr; 95% CI ¼ 130-163). Overall, shifts in fire regimes were strongly linked to changes in vegetation, which were responding to millennial-scale climate change. We conclude that shifts in vegetation can amplify or override the direct influence of climate change on fire regimes, when vegetation shifts significantly modify landscape flammability. Our findings emphasize the importance of biophysical feedbacks between climate, fire, and vegetation in determining the response of ecosystems to past, and by inference, future climate change.

Responses of tree populations to climatic change

The influence of climate on the population dynamics of trees must be inferred from indirect sources of information because the long lifespans of trees preclude direct observation of population growth and decline. Important insights about these processes come from 1) observations of the life histories and ecologies of trees in contemporary forests, 2) evidence of recent treeline movements in remote areas unaffected by human disturbance, and 3) results of experiments performed on forest simulation models. Each line of evidence indicates that tree population responses are influenced by many factors: including lifespans, seed productivity and dispersibility, phenotypic plasticity, genetic variability, competition, and disturbance. Some population characteristics should allow rapid changes in population sizes, while others should confer stability in times of environmental fluctuation. Interactions between controlling factors should result in a wide array of possible responses to climatic change. Interpretations of late-Quaternary forest dynamics must be based on an understanding of the biological processes involved in population responses to environmental variations.

Climate change and Arctic ecosystems: 1. Vegetation changes north of 55°N between the last glacial maximum, mid-Holocene, and present

A unified scheme to assign pollen samples to vegetation types was used to reconstruct vegetation patterns north of 55°N at the last glacial maximum (LGM) and mid-Holocene (6000 years B.P.). The pollen data set assembled for this purpose represents a comprehensive compilation based on the work of many projects and research groups. Five tundra types (cushion forb tundra, graminoid and forb tundra, prostrate dwarf-shrub tundra, erect dwarf-shrub tundra, and low- and high-shrub tundra) were distinguished and mapped on the basis of modern pollen surface samples. The tundra-forest boundary and the distributions of boreal and temperate forest types today were realistically reconstructed. During the mid-Holocene the tundra-forest boundary was north of its present position in some regions, but the pattern of this shift was strongly asymmetrical around the pole, with the largest northward shift in central Siberia (∼200 km), little change in Beringia, and a southward shift in Keewatin and Labrador (∼200 km). Low- and high-shrub tundra extended farther north than today. At the LGM, forests were absent from high latitudes. Graminoid and forb tundra abutted on temperate steppe in northwestern Eurasia while prostrate dwarf-shrub, erect dwarf-shrub, and graminoid and forb tundra formed a mosaic in Beringia. Graminoid and forb tundra is restricted today and does not form a large continuous biome, but the pollen data show that it was far more extensive at the LGM, while low- and high-shrub tundra were greatly reduced, illustrating the potential for climate change to dramatically alter the relative areas occupied by different vegetation types.

HOLOCENE FIRE HISTORY OF A COASTAL TEMPERATE RAIN FOREST BASED ON SOIL CHARCOAL RADIOCARBON DATES

The long-term role of fire in coastal temperate rain forest is poorly under- stood. To determine the historical role of fire on western Vancouver Island (British Co- lumbia, Canada), we constructed a long-term spatially explicit fire history and examined the spatial and temporal distribution of fire during the Holocene. Two fire-history parameters (time-since-fire (TSF) and fire extent) were related to three landscape parameters (landform (hill slope or terrace), aspect, and forest composition) at 83 sites in a 730-ha low-elevation (less than ;200 m) area of a mountainous watershed. We dated fires using tree rings (18 sites) and 120 soil-charcoal radiocarbon dates (65 sites). Comparisons among multiple radiocarbon dates indicated a high probability that the charcoal dated at each site represented the most recent fire, though we expect greater error in TSF estimates at sites where charcoal was very old (.6000 yr) and was restricted to mineral soil horizons. TSF estimates ranged from 64 to ;12 220 yr; 45% of the sites have burned in the last 1000 yr, whereas 20% of the sites have not burned for over 6000 yr. Differences in median TSF were more significant between landform types or across aspects than among forest types. Median TSF was sig- nificantly greater on terraces (4410 yr) than on hill slopes (740 yr). On hill slopes, all south-facing and southwest-facing sites have burned within the last 1000 yr compared to only 27% of north- and east-facing sites burning over the same period. Comparison of fire dates among neighboring sites indicated that fires rarely extended .250 m. During the late Holocene, landform controls have been strong, resulting in the bias of fires to south-facing hillslopes and thus allowing late-successional forest structure to persist for thousands of years in a large portion of the watershed. In contrast, the early Holocene regional climate and forest composition likely resulted in larger landscape fires that were not strongly controlled by landform factors. The millennial-scale TSF detected in this study supports the distinction of coastal temperate rain forest as being under a fundamentally different disturbance regime than other Pacific Northwest forests to the east and south.

Vegetation history of northcentral Alaska: A mapped summary of late-quaternary pollen data

Abstract Fossil pollen data, as illustrated by isopoll and isochrone maps, document the complex late Quaternary history of tundra and boreal forest development in northcentral Alaska. Major plant taxa behaved independently over time, resulting in substantial differences in the vegetation history of eastern and western regions. Major vegetation changes are in general agreement with GCM simulations and confirm the importance of the continental ice sheet and insolation variations in determining late Quaternary climatic trends. Herb-dominated tundra characterized the vegetation between 18 and 14 ka BP, with mesic graminoid tundra in lower elevations of western areas and more xeric, sparse tundra communities in the east and at higher elevations. Moist Betula tussock tundra rapidly replaced the western herb tundra ca. 14 ka BP. However, Betula shrubs expanded more slowly in the east, establishing relatively dry shrub tundra as the predominant regional vegetation by ca. 12 ka BP. River valleys and south-facing slopes supported Populus woodlands between 11 and 9 ka BP, but shrub tundra continued to dominate most upland sites. Alnus shrubs first expanded in the southwestern Brooks Range between 10 and 9 ka BP, spreading rapidly throughout the entire region between 8 and 7 ka BP. Picea glauca populations also expanded between 10 and 9 ka BP, but from source areas in northwestern Canada. The P. glauca forests were most abundant in riparian settings, but isolated stands probably also established in the shrub tundra. P. glauca reached the central Brooks Range by ca. 8 ka BP, followed by an apparent population decline between 8 and 7 ka BP. P. mariana became the dominant tree species ca. 6 ka BP, when it invaded non-riparian P. glauca forests in eastern and central areas and moist shrub tussock tundra in the west. The modern distribution of communities in northcentral Alaska was achieved between 6 and 4 ka BP.

Frequent Fires in Ancient Shrub Tundra: Implications of Paleorecords for Arctic Environmental Change

Understanding feedbacks between terrestrial and atmospheric systems is vital for predicting the consequences of global change, particularly in the rapidly changing Arctic. Fire is a key process in this context, but the consequences of altered fire regimes in tundra ecosystems are rarely considered, largely because tundra fires occur infrequently on the modern landscape. We present paleoecological data that indicate frequent tundra fires in northcentral Alaska between 14,000 and 10,000 years ago. Charcoal and pollen from lake sediments reveal that ancient birch-dominated shrub tundra burned as often as modern boreal forests in the region, every 144 years on average (+/− 90 s.d.; n = 44). Although paleoclimate interpretations and data from modern tundra fires suggest that increased burning was aided by low effective moisture, vegetation cover clearly played a critical role in facilitating the paleofires by creating an abundance of fine fuels. These records suggest that greater fire activity will likely accompany temperature-related increases in shrub-dominated tundra predicted for the 21st century and beyond. Increased tundra burning will have broad impacts on physical and biological systems as well as on land-atmosphere interactions in the Arctic, including the potential to release stored organic carbon to the atmosphere.

Recent burning of boreal forests exceeds fire regime limits of the past 10,000 years

Wildfire activity in boreal forests is anticipated to increase dramatically, with far-reaching ecological and socioeconomic consequences. Paleorecords are indispensible for elucidating boreal fire regime dynamics under changing climate, because fire return intervals and successional cycles in these ecosystems occur over decadal to centennial timescales. We present charcoal records from 14 lakes in the Yukon Flats of interior Alaska, one of the most flammable ecoregions of the boreal forest biome, to infer causes and consequences of fire regime change over the past 10,000 y. Strong correspondence between charcoal-inferred and observational fire records shows the fidelity of sedimentary charcoal records as archives of past fire regimes. Fire frequency and area burned increased ∼6,000–3,000 y ago, probably as a result of elevated landscape flammability associated with increased Picea mariana in the regional vegetation. During the Medieval Climate Anomaly (MCA; ∼1,000–500 cal B.P.), the period most similar to recent decades, warm and dry climatic conditions resulted in peak biomass burning, but severe fires favored less-flammable deciduous vegetation, such that fire frequency remained relatively stationary. These results suggest that boreal forests can sustain high-severity fire regimes for centuries under warm and dry conditions, with vegetation feedbacks modulating climate–fire linkages. The apparent limit to MCA burning has been surpassed by the regional fire regime of recent decades, which is characterized by exceptionally high fire frequency and biomass burning. This extreme combination suggests a transition to a unique regime of unprecedented fire activity. However, vegetation dynamics similar to feedbacks that occurred during the MCA may stabilize the fire regime, despite additional warming.