Carbon uptake in Eurasian boreal forests dominates the high-latitude net ecosystem carbon budget.

Arctic-boreal landscapes are experiencing profound warming, along with changes in ecosystem moisture status and disturbance from fire. This region is of global importance in terms of carbon feedbacks to climate, yet the sign (sink or source) and magnitude of the Arctic-boreal carbon budget within recent years remains highly uncertain. Here, we provide new estimates of recent (2003-2015) vegetation gross primary productivity (GPP), ecosystem respiration (Reco ), net ecosystem CO2 exchange (NEE; Reco - GPP), and terrestrial methane (CH4 ) emissions for the Arctic-boreal zone using a satellite data-driven process-model for northern ecosystems (TCFM-Arctic), calibrated and evaluated using measurements from >60 tower eddy covariance (EC) sites. We used TCFM-Arctic to obtain daily 1-km2 flux estimates and annual carbon budgets for the pan-Arctic-boreal region. Across the domain, the model indicated an overall average NEE sink of -850 Tg CO2 -C year-1 . Eurasian boreal zones, especially those in Siberia, contributed to a majority of the net sink. In contrast, the tundra biome was relatively carbon neutral (ranging from small sink to source). Regional CH4 emissions from tundra and boreal wetlands (not accounting for aquatic CH4 ) were estimated at 35 Tg CH4 -C year-1 . Accounting for additional emissions from open water aquatic bodies and from fire, using available estimates from the literature, reduced the total regional NEE sink by 21% and shifted many far northern tundra landscapes, and some boreal forests, to a net carbon source. This assessment, based on in situ observations and models, improves our understanding of the high-latitude carbon status and also indicates a continued need for integrated site-to-regional assessments to monitor the vulnerability of these ecosystems to climate change.

Surface temperatures reveal the patterns of vegetation water stress and their environmental drivers across the tropical Americas.

Vegetation is a key component in the global carbon cycle as it stores ~450 GtC as biomass, and removes about a third of anthropogenic CO2 emissions. However, in some regions, the rate of plant carbon uptake is beginning to slow, largely because of water stress. Here, we develop a new observation-based methodology to diagnose vegetation water stress and link it to environmental drivers. We used the ratio of remotely sensed land surface to near surface atmospheric temperatures (LST/Tair ) to represent vegetation water stress, and built regression tree models (random forests) to assess the relationship between LST/Tair and the main environmental drivers of surface energy fluxes in the tropical Americas. We further determined ecosystem traits associated with water stress and surface energy partitioning, pinpointed critical thresholds for water stress, and quantified changes in ecosystem carbon uptake associated with crossing these critical thresholds. We found that the top drivers of LST/Tair , explaining over a quarter of its local variability in the study region, are (1) radiation, in 58% of the study region; (2) water supply from precipitation, in 30% of the study region; and (3) atmospheric water demand from vapor pressure deficits (VPD), in 22% of the study region. Regions in which LST/Tair variation is driven by radiation are located in regions of high aboveground biomass or at high elevations, while regions in which LST/Tair is driven by water supply from precipitation or atmospheric demand tend to have low species richness. Carbon uptake by photosynthesis can be reduced by up to 80% in water-limited regions when critical thresholds for precipitation and air dryness are exceeded simultaneously, that is, as compound events. Our results demonstrate that vegetation structure and diversity can be important for regulating surface energy and carbon fluxes over tropical regions.

Understory plant removal counteracts tree thinning effect on soil respiration in a temperate forest.

Elucidating the response mechanism of soil respiration (Rs) to silvicultural practices is pivotal to evaluating the effects of management practices on soil carbon cycling in planted forest ecosystems. However, as common management practices, how thinning, understory plant removal, and their interactions affect Rs and its autotrophic and heterotrophic components (Ra and Rh) remains unclear. Therefore, we investigated Rs, Ra and Rh by the trenching method from 2011 to 2015 in a Pinus tabuliformis plantation in northern China, subjecting to four treatments (intact control plots [CK], thinning [T], understory removal [UR], and thinning with understory removal [TUR]). Mean annual Rs was significantly increased by thinning (by 15.3%), whereas decreased by UR (by 17.4%), compared with CK. These variations in Rs were mainly attributed to changes in Ra. The increments of Ra were caused by the enhanced growth of fine root biomass after thinning. However, UR led to lower Ra compared with CK (p < .05), indicating that understory growth is inadequate to compensate for the decreased respiring root biomass induced by understory removal. Rs was unchanged between TUR and the intact control plot due to the opposite effects of thinning and UR on the Ra. Changes in Rh exhibited no significant differences among the treatments, partly because of the stable microbial biomass carbon (MBC) and forest floor mass (litter and fine woody debris). No interaction effect between thinning and understory removal was detected on Rs, Ra, and Rh. The lowest temperature sensitivity (Q10 ) value of Ra was found in CK. This study highlights the necessity of incorporating understory plant effects on soil CO2 efflux in assessing forest management practices on soil carbon cycling.

Lowering water table reduces carbon sink strength and carbon stocks in northern peatlands.

Peatlands at high latitudes have accumulated >400 Pg carbon (C) because saturated soil and cold temperatures suppress C decomposition. This substantial amount of C in Arctic and Boreal peatlands is potentially subject to increased decomposition if the water table (WT) decreases due to climate change, including permafrost thaw-related drying. Here, we optimize a version of the Organizing Carbon and Hydrology In Dynamic Ecosystems model (ORCHIDEE-PCH4) using site-specific observations to investigate changes in CO2 and CH4 fluxes as well as C stock responses to an experimentally manipulated decrease of WT at six northern peatlands. The unmanipulated control peatlands, with the WT <20 cm on average (seasonal max up to 45 cm) below the surface, currently act as C sinks in most years (58 ± 34 g C m-2  year-1 ; including 6 ± 7 g C-CH4 m-2  year-1 emission). We found, however, that lowering the WT by 10 cm reduced the CO2 sink by 13 ± 15 g C m-2  year-1 and decreased CH4 emission by 4 ± 4 g CH4 m-2  year-1 , thus accumulating less C over 100 years (0.2 ± 0.2 kg C m-2 ). Yet, the reduced emission of CH4 , which has a larger greenhouse warming potential, resulted in a net decrease in greenhouse gas balance by 310 ± 360 g CO2-eq  m-2  year-1 . Peatlands with the initial WT close to the soil surface were more vulnerable to C loss: Non-permafrost peatlands lost >2 kg C m-2 over 100 years when WT is lowered by 50 cm, while permafrost peatlands temporally switched from C sinks to sources. These results highlight that reductions in C storage capacity in response to drying of northern peatlands are offset in part by reduced CH4 emissions, thus slightly reducing the positive carbon climate feedbacks of peatlands under a warmer and drier future climate scenario.

Reconciling carbon-cycle processes from ecosystem to global scales.

Understanding carbon (C) dynamics from ecosystem to global scales remains a challenge. Although expansion of global carbon dioxide (CO2) observatories makes it possible to estimate C-cycle processes from ecosystem to global scales, these estimates do not necessarily agree. At the continental US scale, only 5% of C fixed through photosynthesis remains as net ecosystem exchange (NEE), but ecosystem measurements indicate that only 2% of fixed C remains in grasslands, whereas as much as 30% remains in needleleaf forests. The wet and warm Southeast has the highest gross primary productivity and the relatively wet and cool Midwest has the highest NEE, indicating important spatial mismatches. Newly available satellite and atmospheric data can be combined in innovative ways to identify potential C loss pathways to reconcile these spatial mismatches. Independent datasets compiled from terrestrial and aquatic environments can now be combined to advance C-cycle science across the land-water interface.

Asymmetric responses of ecosystem productivity to rainfall anomalies vary inversely with mean annual rainfall over the conterminous United States.

The CONterminous United States (CONUS) presents a large range of climate conditions and biomes where terrestrial primary productivity and its inter-annual variability are controlled regionally by rainfall and/or temperature. Here, the response of ecosystem productivity to those climate variables was investigated across different biomes from 2010 to 2018 using three climate datasets of precipitation, air temperature or drought severity, combined with several proxies of ecosystem productivity: a remote sensing product of aboveground biomass, an net primary productivity (NPP) remote sensing product, an NPP model-based product and four gross primary productivity products. We used an asymmetry index (AI) where positive AI indicates a greater increase of ecosystem productivity in wet years compared to the decline in dry years, and negative AI indicates a greater decline of ecosystem productivity in dry years compared to the increase in wet years. We found consistent spatial patterns of AI across the CONUS for the different products, with negative asymmetries over the Great Plains and positive asymmetries over the southwestern CONUS. Shrubs and, to a lesser extent, evergreen forests show a persistent positive asymmetry, whilst (natural) grasslands appear to have transitioned from positive to negative anomalies during the last decade. The general tendency of dominant negative asymmetry response for ecosystem productivity across the CONUS appears to be influenced by the negative asymmetry of precipitation anomalies. AI was found to be a function of mean rainfall: more positive AIs were found in dry areas where plants are adapted to drought and take advantage of rainfall pulses, and more negative AIs were found in wet areas, with a threshold delineating the two regimes corresponding to a mean annual rainfall of 200-400 mm/year.

Increased high-latitude photosynthetic carbon gain offset by respiration carbon loss during an anomalous warm winter to spring transition.

Arctic and boreal ecosystems play an important role in the global carbon (C) budget, and whether they act as a future net C sink or source depends on climate and environmental change. Here, we used complementary in situ measurements, model simulations, and satellite observations to investigate the net carbon dioxide (CO2 ) seasonal cycle and its climatic and environmental controls across Alaska and northwestern Canada during the anomalously warm winter to spring conditions of 2015 and 2016 (relative to 2010-2014). In the warm spring, we found that photosynthesis was enhanced more than respiration, leading to greater CO2 uptake. However, photosynthetic enhancement from spring warming was partially offset by greater ecosystem respiration during the preceding anomalously warm winter, resulting in nearly neutral effects on the annual net CO2 balance. Eddy covariance CO2 flux measurements showed that air temperature has a primary influence on net CO2 exchange in winter and spring, while soil moisture has a primary control on net CO2 exchange in the fall. The net CO2 exchange was generally more moisture limited in the boreal region than in the Arctic tundra. Our analysis indicates complex seasonal interactions of underlying C cycle processes in response to changing climate and hydrology that may not manifest in changes in net annual CO2 exchange. Therefore, a better understanding of the seasonal response of C cycle processes may provide important insights for predicting future carbon-climate feedbacks and their consequences on atmospheric CO2 dynamics in the northern high latitudes.

Reducing uncertainties in decadal variability of the global carbon budget with multiple datasets.

Conventional calculations of the global carbon budget infer the land sink as a residual between emissions, atmospheric accumulation, and the ocean sink. Thus, the land sink accumulates the errors from the other flux terms and bears the largest uncertainty. Here, we present a Bayesian fusion approach that combines multiple observations in different carbon reservoirs to optimize the land (B) and ocean (O) carbon sinks, land use change emissions (L), and indirectly fossil fuel emissions (F) from 1980 to 2014. Compared with the conventional approach, Bayesian optimization decreases the uncertainties in B by 41% and in O by 46%. The L uncertainty decreases by 47%, whereas F uncertainty is marginally improved through the knowledge of natural fluxes. Both ocean and net land uptake (B + L) rates have positive trends of 29 ± 8 and 37 ± 17 Tg C⋅y-2 since 1980, respectively. Our Bayesian fusion of multiple observations reduces uncertainties, thereby allowing us to isolate important variability in global carbon cycle processes.

Limited evidence for CO2 -related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients.

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO2 may enhance photosynthesis and/or decrease stomatal conductance (gs ) thereby enhancing intrinsic water-use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO2 and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ13 C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in gs . Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing gs may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.

Tropical nighttime warming as a dominant driver of variability in the terrestrial carbon sink.

The terrestrial biosphere is currently a strong carbon (C) sink but may switch to a source in the 21st century as climate-driven losses exceed CO2-driven C gains, thereby accelerating global warming. Although it has long been recognized that tropical climate plays a critical role in regulating interannual climate variability, the causal link between changes in temperature and precipitation and terrestrial processes remains uncertain. Here, we combine atmospheric mass balance, remote sensing-modeled datasets of vegetation C uptake, and climate datasets to characterize the temporal variability of the terrestrial C sink and determine the dominant climate drivers of this variability. We show that the interannual variability of global land C sink has grown by 50-100% over the past 50 y. We further find that interannual land C sink variability is most strongly linked to tropical nighttime warming, likely through respiration. This apparent sensitivity of respiration to nighttime temperatures, which are projected to increase faster than global average temperatures, suggests that C stored in tropical forests may be vulnerable to future warming.

The sensitivity of soil respiration to soil temperature, moisture, and carbon supply at the global scale.

Soil respiration (Rs) is a major pathway by which fixed carbon in the biosphere is returned to the atmosphere, yet there are limits to our ability to predict respiration rates using environmental drivers at the global scale. While temperature, moisture, carbon supply, and other site characteristics are known to regulate soil respiration rates at plot scales within certain biomes, quantitative frameworks for evaluating the relative importance of these factors across different biomes and at the global scale require tests of the relationships between field estimates and global climatic data. This study evaluates the factors driving Rs at the global scale by linking global datasets of soil moisture, soil temperature, primary productivity, and soil carbon estimates with observations of annual Rs from the Global Soil Respiration Database (SRDB). We find that calibrating models with parabolic soil moisture functions can improve predictive power over similar models with asymptotic functions of mean annual precipitation. Soil temperature is comparable with previously reported air temperature observations used in predicting Rs and is the dominant driver of Rs in global models; however, within certain biomes soil moisture and soil carbon emerge as dominant predictors of Rs. We identify regions where typical temperature-driven responses are further mediated by soil moisture, precipitation, and carbon supply and regions in which environmental controls on high Rs values are difficult to ascertain due to limited field data. Because soil moisture integrates temperature and precipitation dynamics, it can more directly constrain the heterotrophic component of Rs, but global-scale models tend to smooth its spatial heterogeneity by aggregating factors that increase moisture variability within and across biomes. We compare statistical and mechanistic models that provide independent estimates of global Rs ranging from 83 to 108 Pg yr-1 , but also highlight regions of uncertainty where more observations are required or environmental controls are hard to constrain.

Forest demography and biomass accumulation rates are associated with transient mean tree size vs. density scaling relations.

Linking individual and stand-level dynamics during forest development reveals a scaling relationship between mean tree size and tree density in forest stands, which integrates forest structure and function. However, the nature of this so-called scaling law and its variation across broad spatial scales remain unquantified, and its linkage with forest demographic processes and carbon dynamics remains elusive. In this study, we develop a theoretical framework and compile a broad-scale dataset of long-term sample forest stands (n = 1,433) from largely undisturbed forests to examine the association of temporal mean tree size vs. density scaling trajectories (slopes) with biomass accumulation rates and the sensitivity of scaling slopes to environmental and demographic drivers. The results empirically demonstrate a large variation of scaling slopes, ranging from -4 to -0.2, across forest stands in tropical, temperate, and boreal forest biomes. Steeper scaling slopes are associated with higher rates of biomass accumulation, resulting from a lower offset of forest growth by biomass loss from mortality. In North America, scaling slopes are positively correlated with forest stand age and rainfall seasonality, thus suggesting a higher rate of biomass accumulation in younger forests with lower rainfall seasonality. These results demonstrate the strong association of the transient mean tree size vs. density scaling trajectories with forest demography and biomass accumulation rates, thus highlighting the potential of leveraging forest structure properties to predict forest demography, carbon fluxes, and dynamics at broad spatial scales.

Global water use efficiency saturation due to increased vapor pressure deficit.

The ratio of carbon assimilation to water evapotranspiration (ET) of an ecosystem, referred to as ecosystem water use efficiency (WUEeco), is widely expected to increase because of the rising atmospheric carbon dioxide concentration (Ca). However, little is known about the interactive effects of rising Ca and climate change on WUEeco. On the basis of upscaled estimates from machine learning methods and global FLUXNET observations, we show that global WUEeco has not risen since 2001 because of the asymmetric effects of an increased vapor pressure deficit (VPD), which depressed photosynthesis and enhanced ET. An undiminished ET trend indicates that rising temperature and VPD may play a more important role in regulating ET than declining stomatal conductance. Projected increases in VPD are predicted to affect the future coupling of the terrestrial carbon and water cycles.

Non-symmetric responses of leaf onset date to natural warming and cooling in northern ecosystems.

The northern hemisphere has experienced regional cooling, especially during the global warming hiatus (1998-2012) due to ocean energy redistribution. However, the lack of studies about the natural cooling effects hampers our understanding of vegetation responses to climate change. Using 15,125 ground phenological time series at 3,620 sites since the 1950s and 31-year satellite greenness observations (1982-2012) covering the warming hiatus period, we show a stronger response of leaf onset date (LOD) to natural cooling than to warming, i.e. the delay of LOD caused by 1°C cooling is larger than the advance of LOD with 1°C warming. This might be because cooling leads to larger chilling accumulation and heating requirements for leaf onset, but this non-symmetric LOD response is partially offset by warming-related drying. Moreover, spring greening magnitude, in terms of satellite-based greenness and productivity, is more sensitive to LOD changes in the warming area than in the cooling. These results highlight the importance of considering non-symmetric responses of spring greening to warming and cooling when predicting vegetation-climate feedbacks.

Field-based tree mortality constraint reduces estimates of model-projected forest carbon sinks.

Considerable uncertainty and debate exist in projecting the future capacity of forests to sequester atmospheric CO2. Here we estimate spatially explicit patterns of biomass loss by tree mortality (LOSS) from largely unmanaged forest plots to constrain projected (2015-2099) net primary productivity (NPP), heterotrophic respiration (HR) and net carbon sink in six dynamic global vegetation models (DGVMs) across continents. This approach relies on a strong relationship among LOSS, NPP, and HR at continental or biome scales. The DGVMs overestimated historical LOSS, particularly in tropical regions and eastern North America by as much as 5 Mg ha-1 y-1. The modeled spread of DGVM-projected NPP and HR uncertainties was substantially reduced in tropical regions after incorporating the field-based mortality constraint. The observation-constrained models show a decrease in the tropical forest carbon sink by the end of the century, particularly across South America (from 2 to 1.4 PgC y-1), and an increase in the sink in North America (from 0.8 to 1.1 PgC y-1). These results highlight the feasibility of using forest demographic data to empirically constrain forest carbon sink projections and the potential overestimation of projected tropical forest carbon sinks.

Respiratory loss during late-growing season determines the net carbon dioxide sink in northern permafrost regions.

Warming of northern high latitude regions (NHL, > 50 °N) has increased both photosynthesis and respiration which results in considerable uncertainty regarding the net carbon dioxide (CO2) balance of NHL ecosystems. Using estimates constrained from atmospheric observations from 1980 to 2017, we find that the increasing trends of net CO2 uptake in the early-growing season are of similar magnitude across the tree cover gradient in the NHL. However, the trend of respiratory CO2 loss during late-growing season increases significantly with increasing tree cover, offsetting a larger fraction of photosynthetic CO2 uptake, and thus resulting in a slower rate of increasing annual net CO2 uptake in areas with higher tree cover, especially in central and southern boreal forest regions. The magnitude of this seasonal compensation effect explains the difference in net CO2 uptake trends along the NHL vegetation- permafrost gradient. Such seasonal compensation dynamics are not captured by dynamic global vegetation models, which simulate weaker respiration control on carbon exchange during the late-growing season, and thus calls into question projections of increasing net CO2 uptake as high latitude ecosystems respond to warming climate conditions.

Decadal variability in land carbon sink efficiency.

BACKGROUND: The climate mitigation target of limiting the temperature increase below 2 °C above the pre-industrial levels requires the efforts from all countries. Tracking the trajectory of the land carbon sink efficiency is thus crucial to evaluate the nationally determined contributions (NDCs). Here, we define the instantaneous land sink efficiency as the ratio of natural land carbon sinks to emissions from fossil fuel and land-use and land-cover change with a value of 1 indicating carbon neutrality to track its temporal dynamics in the past decades. RESULTS: Land sink efficiency has been decreasing during 1957-1990 because of the increased emissions from fossil fuel. After the effect of the Mt. Pinatubo eruption diminished (after 1994), the land sink efficiency firstly increased before 2009 and then began to decrease again after 2009. This reversal around 2009 is mostly attributed to changes in land sinks in tropical regions in response to climate variations. CONCLUSIONS: The decreasing trend of land sink efficiency in recent years reveals greater challenges in climate change mitigation, and that climate impacts on land carbon sinks must be accurately quantified to assess the effectiveness of regional scale climate mitigation policies.

Empirical estimates of regional carbon budgets imply reduced global soil heterotrophic respiration.

Resolving regional carbon budgets is critical for informing land-based mitigation policy. For nine regions covering nearly the whole globe, we collected inventory estimates of carbon-stock changes complemented by satellite estimates of biomass changes where inventory data are missing. The net land-atmospheric carbon exchange (NEE) was calculated by taking the sum of the carbon-stock change and lateral carbon fluxes from crop and wood trade, and riverine-carbon export to the ocean. Summing up NEE from all regions, we obtained a global 'bottom-up' NEE for net land anthropogenic CO2 uptake of -2.2 ± 0.6 PgC yr-1 consistent with the independent top-down NEE from the global atmospheric carbon budget during 2000-2009. This estimate is so far the most comprehensive global bottom-up carbon budget accounting, which set up an important milestone for global carbon-cycle studies. By decomposing NEE into component fluxes, we found that global soil heterotrophic respiration amounts to a source of CO2 of 39 PgC yr-1 with an interquartile of 33-46 PgC yr-1-a much smaller portion of net primary productivity than previously reported.

Unsaturated fatty acid content in seston and tropho-dynamic coupling in lakes.

Determining the factors that control food web interactions is a key issue in ecology. The empirical relationship between nutrient loading (total phosphorus) and phytoplankton standing stock (chlorophyll a) in lakes was described about 30 years ago and is central for managing surface water quality. The efficiency with which biomass and energy are transferred through the food web and sustain the production of higher trophic levels (such as fish) declines with nutrient loading and system productivity, but the underlying mechanisms are poorly understood. Here we show that in seston (fine particles in water) during summer, specific omega3-polyunsaturated fatty acids (omega3-PUFAs), which are important for zooplankton, are significantly correlated to the trophic status of the lake. The omega3-PUFAs octadecatetraenoic acid, eicosapentaenoic acid (EPA) and docosahexaenoic acid, but not alpha-linolenic acid, decrease on a double-logarithmic scale with increasing total phosphorus. By combining the empirical relationship between EPA-to-carbon content and total phosphorus with functional models relating EPA-to-carbon content to the growth and egg production of daphnids, we predict secondary production for this key consumer. Thus, the decreasing efficiency in energy transfer with increasing lake productivity can be explained by differences in omega3-PUFA-associated food quality at the plant-animal interface.

Evolution of woodcutting behaviour in Early Pliocene beaver driven by consumption of woody plants.

Modern beavers (Castor) are prolific ecosystem engineers and dramatically alter the landscape through tree harvesting and dam building. Little is known, however, about the evolutionary drivers of their woodcutting behaviour. Here we investigate if early woodcutting behaviour in Castoridae was driven by nutritional needs. We measured stable carbon and nitrogen isotopes (δ13C and δ15N) of coeval subfossil plants and beaver collagen (Dipoides sp.) from the Early Pliocene, High Arctic Beaver Pond fossil locality (Ellesmere Island), in order to reconstruct Dipoides sp. diet. Isotopic evidence indicates a diet of woody plants and freshwater macrophytes, supporting the hypothesis that this extinct semiaquatic beaver engaged in woodcutting behaviour for feeding purposes. In a phylogenetic context, the isotopic evidence implies that woodcutting and consumption of woody plants can be traced back to a small-bodied, semiaquatic Miocene castorid, suggesting that beavers have been consuming woody plants for over 20 million years. We propose that the behavioural complex (swimming, woodcutting, and consuming woody plants) preceded and facilitated the evolution of dam building. Dam building and food caching behaviours appear to be specializations for cold winter survival and may have evolved in response to late Neogene northern cooling.