Environmental filtering governs consistent vertical zonation in sedimentary microbial communities across disconnected mountain lakes.

Subsurface microorganisms make up the majority of Earth's microbial biomass, but ecological processes governing surface communities may not explain community patterns at depth because of burial. Depth constrains dispersal and energy availability, and when combined with geographic isolation across landscapes, may influence community assembly. We sequenced the 16S rRNA gene of bacteria and archaea from 48 sediment cores across 36 lakes in four disconnected mountain ranges in Wyoming, USA and used null models to infer assembly processes across depth, spatial isolation, and varying environments. Although we expected strong dispersal limitations across these isolated settings, community composition was primarily shaped by environmental selection. Communities consistently shifted from domination by organisms that degrade organic matter at the surface to methanogenic, low-energy adapted taxa in deeper zones. Stochastic processes-like dispersal limitation-contributed to differences among lakes, but because these effects weakened with depth, selection processes ultimately governed subsurface microbial biogeography.

A postglacial paleoenvironmental dataset from New England.

This paleoenvironmental database features postglacial lake-sediment records from 31 study sites located across New England. The study sites span an environmental gradient from the cooler, northern and inland part of the region to the warmer, southern and coastal areas of New England. Sediment-core chronologies were determined using 14C dating, 210Pb analysis, and pollen evidence. Detailed analyses of sediment lithology, pollen, and charcoal were used to reconstruct changes in climate, vegetation, and fire at centennial temporal scales and subregional spatial scales for the last 14,000 years. Analyses of paleoenvironmental data provide insights into the rates, patterns, and drivers of ecosystem change, helping us anticipate future ecosystem dynamics and guiding present-day conservation strategies and land management.

Rocky Mountain subalpine forests now burning more than any time in recent millennia.

The 2020 fire season punctuated a decades-long trend of increased fire activity across the western United States, nearly doubling the total area burned in the central Rocky Mountains since 1984. Understanding the causes and implications of such extreme fire seasons, particularly in subalpine forests that have historically burned infrequently, requires a long-term perspective not afforded by observational records. We place 21st century fire activity in subalpine forests in the context of climate and fire history spanning the past 2,000 y using a unique network of 20 paleofire records. Largely because of extensive burning in 2020, the 21st century fire rotation period is now 117 y, reflecting nearly double the average rate of burning over the past 2,000 y. More strikingly, contemporary rates of burning are now 22% higher than the maximum rate reconstructed over the past two millennia, during the early Medieval Climate Anomaly (MCA) (770 to 870 Common Era), when Northern Hemisphere temperatures were ∼0.3 °C above the 20th century average. The 2020 fire season thus exemplifies how extreme events are demarcating newly emerging fire regimes as climate warms. With 21st century temperatures now surpassing those during the MCA, fire activity in Rocky Mountain subalpine forests is exceeding the range of variability that shaped these ecosystems for millennia.

The quest for absolute abundance: The use of internal standards for DNA-based community ecology.

To characterize microbiomes and other ecological assemblages, ecologists routinely sequence and compare loci that differ among focal taxa. Counts of these sequences convey information regarding the occurrence and relative abundances of taxa, but provide no direct measure of their absolute abundances, due to the technical limitations of the sequencing process. The relative abundances in compositional data are inherently constrained and difficult to interpret. The incorporation of internal standards (ISDs; colloquially referred to as 'spike-ins') into DNA pools can ameliorate the problems posed by relative abundance data and allow absolute abundances to be approximated. Unfortunately, many laboratory and sampling biases cause ISDs to underperform or fail. Here, we discuss how careful deployment of ISDs can avoid these complications and be an integral component of well-designed studies seeking to characterize ecological assemblages via sequencing of DNA.

Climate change, ecosystems and abrupt change: science priorities.

Ecologists have long studied patterns, directions and tempos of change, but there is a pressing need to extend current understanding to empirical observations of abrupt changes as climate warming accelerates. Abrupt changes in ecological systems (ACES)-changes that are fast in time or fast relative to their drivers-are ubiquitous and increasing in frequency. Powerful theoretical frameworks exist, yet applications in real-world landscapes to detect, explain and anticipate ACES have lagged. We highlight five insights emerging from empirical studies of ACES across diverse ecosystems: (i) ecological systems show ACES in some dimensions but not others; (ii) climate extremes may be more important than mean climate in generating ACES; (iii) interactions among multiple drivers often produce ACES; (iv) contingencies, such as ecological memory, frequency and sequence of disturbances, and spatial context are important; and (v) tipping points are often (but not always) associated with ACES. We suggest research priorities to advance understanding of ACES in the face of climate change. Progress in understanding ACES requires strong integration of scientific approaches (theory, observations, experiments and process-based models) and high-quality empirical data drawn from a diverse array of ecosystems. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Mid-latitude net precipitation decreased with Arctic warming during the Holocene.

The latitudinal temperature gradient between the Equator and the poles influences atmospheric stability, the strength of the jet stream and extratropical cyclones1-3. Recent global warming is weakening the annual surface gradient in the Northern Hemisphere by preferentially warming the high latitudes4; however, the implications of these changes for mid-latitude climate remain uncertain5,6. Here we show that a weaker latitudinal temperature gradient-that is, warming of the Arctic with respect to the Equator-during the early to middle part of the Holocene coincided with substantial decreases in mid-latitude net precipitation (precipitation minus evapotranspiration, at 30° N to 50° N). We quantify the evolution of the gradient and of mid-latitude moisture both in a new compilation of Holocene palaeoclimate records spanning from 10° S to 90° N and in an ensemble of mid-Holocene climate model simulations. The observed pattern is consistent with the hypothesis that a weaker temperature gradient led to weaker mid-latitude westerly flow, weaker cyclones and decreased net terrestrial mid-latitude precipitation. Currently, the northern high latitudes are warming at rates nearly double the global average4, decreasing the Equator-to-pole temperature gradient to values comparable with those in the early to middle Holocene. If the patterns observed during the Holocene hold for current anthropogenically forced warming, the weaker latitudinal temperature gradient will lead to considerable reductions in mid-latitude water resources.

Predictable hydrological and ecological responses to Holocene North Atlantic variability.

Climate variations in the North Atlantic region can substantially impact surrounding continents. Notably, the Younger Dryas chronozone was named for the ecosystem effects of abrupt changes in the region at circa (ca.) 12.9-11.7 ka (millennia before 1950 AD). Holocene variations since then, however, have been hard to diagnose, and the responsiveness of terrestrial ecosystems continues to be debated. Here, we show that Holocene climate variations had spatial patterns consistent with changes in Atlantic overturning and repeatedly steepened the temperature gradient between Nova Scotia and Greenland since >8 ka. The multicentury changes correlated with hydrologic and vegetation changes in the northeast United States, including when an enhanced temperature gradient coincided with subregional droughts indicated by water-level changes at multiple coastal lakes at 4.9-4.6, 4.2-3.9, 2.8-2.1, and 1.3-1.2 ka. We assessed the variability and its effects by replicating signals across sites, using converging evidence from multiple methods, and applying forward models of the systems involved. We evaluated forest responses in the northeast United States and found that they tracked the regional climate shifts including the smallest magnitude (∼5% or 50 mm) changes in effective precipitation. Although a long-term increase in effective precipitation of >45% (>400 mm) could have prevented ecological communities from equilibrating to the continuously changing conditions, our comparisons confirm stable vegetation-climate relationships and support the use of fossil pollen records for quantitative paleoclimate reconstruction. Overall, the network of records indicates that centennial climate variability has repeatedly affected the North Atlantic region with predictable consequences.

Detecting past changes in vegetation resilience in the context of a changing climate.

Anthropogenic climate change is continuously altering ecological responses to disturbance and must be accounted for when examining ecological resilience. One way to measure resilience in ecological datasets is by considering the amount and duration of change from a baseline created by perturbations, such as disturbances like wildfire. Recovery occurs when ecological conditions return to equilibrium, meaning that no subsequent changes can be attributed to the effects of the disturbance, but climate change often causes the recovered state to differ from the previous baseline. The palaeoecological record provides an opportunity to examine these expectations because palaeoclimates changed continuously; few periods existed when environmental conditions were stationary. Here we demonstrate a framework for examining resilience in palaeoecological records against the backdrop of a non-stationary climate by considering resilience as two components of (i) resistance (magnitude of change) and (ii) recovery (time required to return) to predicted equilibrium values. Measuring these components of resilience in palaeoecological records requires high-resolution fossil (e.g. pollen) records, local palaeoclimate reconstructions, a model to predict ecological change in response to climate change, and disturbance records measured at the same spatial scale as the ecological (e.g. vegetation history) record. Resistance following disturbance is measured as the deviation of the fossil record from the ecological state predicted by the palaeoclimate records, and recovery time is measured as the time required for the fossil record to return to predicted values. We show that some cases may involve nearly persistent equilibrium despite large climate changes, but that others can involve a shift to a new state without any complete recovery.

Reconciling divergent trends and millennial variations in Holocene temperatures.

Cooling during most of the past two millennia has been widely recognized and has been inferred to be the dominant global temperature trend of the past 11,700 years (the Holocene epoch). However, long-term cooling has been difficult to reconcile with global forcing, and climate models consistently simulate long-term warming. The divergence between simulations and reconstructions emerges primarily for northern mid-latitudes, for which pronounced cooling has been inferred from marine and coastal records using multiple approaches. Here we show that temperatures reconstructed from sub-fossil pollen from 642 sites across North America and Europe closely match simulations, and that long-term warming, not cooling, defined the Holocene until around 2,000 years ago. The reconstructions indicate that evidence of long-term cooling was limited to North Atlantic records. Early Holocene temperatures on the continents were more than two degrees Celsius below those of the past two millennia, consistent with the simulated effects of remnant ice sheets in the climate model Community Climate System Model 3 (CCSM3). CCSM3 simulates increases in 'growing degree days'-a measure of the accumulated warmth above five degrees Celsius per year-of more than 300 kelvin days over the Holocene, consistent with inferences from the pollen data. It also simulates a decrease in mean summer temperatures of more than two degrees Celsius, which correlates with reconstructed marine trends and highlights the potential importance of the different subseasonal sensitivities of the records. Despite the differing trends, pollen- and marine-based reconstructions are correlated at millennial-to-centennial scales, probably in response to ice-sheet and meltwater dynamics, and to stochastic dynamics similar to the temperature variations produced by CCSM3. Although our results depend on a single source of palaeoclimatic data (pollen) and a single climate-model simulation, they reinforce the notion that climate models can adequately simulate climates for periods other than the present-day. They also demonstrate that amplified warming in recent decades increased temperatures above the mean of any century during the past 11,000 years.

Extensive wildfires, climate change, and an abrupt state change in subalpine ribbon forests, Colorado.

Ecosystems may shift abruptly when the effects of climate change and disturbance interact, and landscapes with regularly patterned vegetation may be especially vulnerable to abrupt shifts. Here we use a fossil pollen record from a regularly patterned ribbon forest (alternating bands of forests and meadows) in Colorado to examine whether past changes in wildfire and climate produced abrupt vegetation shifts. Comparing the percentages of conifer pollen with sedimentary δ18 O data (interpreted as an indicator of temperature or snow accumulation) indicates a first-order linear relationship between vegetation composition and climate change with no detectable lags over the past 2,500 yr (r = 0.55, P < 0.001). Additionally, however, we find that the vegetation changed abruptly within a century of extensive wildfires, which were recognized in a previous study to have burned approximately 80% of the surrounding 1,000 km2 landscape 1,000 yr ago when temperatures rose ~0.5°C. The vegetation change was larger than expected from the effects of climate change alone. Pollen assemblages changed from a composition associated with closed subalpine forests to one similar to modern ribbon forests. Fossil pollen assemblages then remained like those from modern ribbon forests for the following ~1,000 yr, providing a clear example of how extensive disturbances can trigger persistent new vegetation states and alter how vegetation responds to climate.

Medieval warming initiated exceptionally large wildfire outbreaks in the Rocky Mountains.

Many of the largest wildfires in US history burned in recent decades, and climate change explains much of the increase in area burned. The frequency of extreme wildfire weather will increase with continued warming, but many uncertainties still exist about future fire regimes, including how the risk of large fires will persist as vegetation changes. Past fire-climate relationships provide an opportunity to constrain the related uncertainties, and reveal widespread burning across large regions of western North America during past warm intervals. Whether such episodes also burned large portions of individual landscapes has been difficult to determine, however, because uncertainties with the ages of past fires and limited spatial resolution often prohibit specific estimates of past area burned. Accounting for these challenges in a subalpine landscape in Colorado, we estimated century-scale fire synchroneity across 12 lake-sediment charcoal records spanning the past 2,000 y. The percentage of sites burned only deviated from the historic range of variability during the Medieval Climate Anomaly (MCA) between 1,200 and 850 y B.P., when temperatures were similar to recent decades. Between 1,130 and 1,030 y B.P., 83% (median estimate) of our sites burned when temperatures increased ∼0.5 °C relative to the preceding centuries. Lake-based fire rotation during the MCA decreased to an estimated 120 y, representing a 260% higher rate of burning than during the period of dendroecological sampling (360 to -60 y B.P.). Increased burning, however, did not persist throughout the MCA. Burning declined abruptly before temperatures cooled, indicating possible fuel limitations to continued burning.

Model systems for a no-analog future: species associations and climates during the last deglaciation.

As the earth system moves to a novel state, model systems (experimental, observational, paleoecological) are needed to assess and improve the predictive accuracy of ecological models under environments with no contemporary analog. In recent years, we have intensively studied the no-analog plant associations and climates in eastern North America during the last deglaciation to better constrain their spatiotemporal distribution, test hypotheses about climatic and megaherbivory controls, and assess the accuracy of species- and community-level models. The formation of no-analog plant associations was asynchronous, beginning first in the south-central United States; at sites in the north-central United States, it is linked to declining megafaunal abundances. Insolation and temperature were more seasonal than present, creating climates currently nonexistent in North America, and shifting species-climate relationships for some taxa. These shifts pose a common challenge to empirical paleoclimatic reconstructions, species distribution models (SDMs), and conservation-optimization models based on SDMs. Steps forward include combining recent and paleoecological data to more fully describe species' fundamental niches, employing community-level models to model shifts in species interactions under no-analog climates, and assimilating paleoecological data with mechanistic ecosystem models. Accurately modeling species interactions under novel environments remains a fundamental challenge for all forms of ecological models.

A continuous climatic impact on Holocene human population in the Rocky Mountains.

Ancient cultural changes have often been linked to abrupt climatic events, but the potential that climate can exert a persistent influence on human populations has been debated. Here, independent population, temperature, and moisture history reconstructions from the Bighorn Basin in Wyoming (United States) show a clear quantitative relationship spanning 13 ka, which explains five major periods of population growth/decline and ~45% of the population variance. A persistent ~300-y lag in the human demographic response conforms with either slow (~0.3%) intrinsic annual population growth rates or a lag in the environmental carrying capacity, but in either case, the population continuously adjusted to changing environmental conditions.

Woodland-to-forest transition during prolonged drought in Minnesota after ca. AD 1300.

Interactions among multiple causes of ecological perturbation, such as climate change and disturbance, can produce "ecological surprises." Here, we examine whether climate-fire-vegetation interactions can produce ecological changes that differ in direction from those expected from the effects of climate change alone. To do so, we focus on the "Big Woods" of central Minnesota, USA, which was shaped both by climate and fire. The deciduous Big Woods forest replaced regional woodlands and savannas after the severity of regional fire regimes declined at ca. AD 1300. A trend toward wet conditions has long been assumed to explain the forest expansion, but we show that water levels at two lakes within the region (Wolsfeld Lake and Bufflehead Pond) were low when open woodlands were transformed into the Big Woods. Water levels were high instead at ca. 2240-795 BC when regional fire regimes were most severe. Based on the correlation between water levels and fire-regime severity, we infer that prolonged or repeated droughts after ca. AD 1265 reduced the biomass and connectivity of fine fuels (grasses) within the woodlands. As a result, regional fire severity declined and allowed tree populations to expand. Tree-ring data from the region show a peak in the recruitment of key Big Woods tree species during the AD 1930s drought and suggest that low regional moisture balance need not have been a limiting factor for forest expansion. The regional history, thus, demonstrates the types of counterintuitive ecosystem changes that may arise as climate changes in the future.