Center for Northern Studies

▪ Abstract Deer have expanded their range and increased dramatically in abundance worldwide in recent decades. They inflict major economic losses in forestry, agriculture, and transportation and contribute to the transmission of several animal and human diseases. Their impact on natural ecosystems is also dramatic but less quantified. By foraging selectively, deer affect the growth and survival of many herb, shrub, and tree species, modifying patterns of relative abundance and vegetation dynamics. Cascading effects on other species extend to insects, birds, and other mammals. In forests, sustained overbrowsing reduces plant cover and diversity, alters nutrient and carbon cycling, and redirects succession to shift future overstory composition. Many of these simplified alternative states appear to be stable and difficult to reverse. Given the influence of deer on other organisms and natural processes, ecologists should actively participate in efforts to understand, monitor, and reduce the impact of deer on ecosystems.

Permafrost warming has the potential to amplify global climate change, because when frozen sediments thaw it unlocks soil organic carbon. Yet to date, no globally consistent assessment of permafrost temperature change has been compiled. Here we use a global data set of permafrost temperature time series from the Global Terrestrial Network for Permafrost to evaluate temperature change across permafrost regions for the period since the International Polar Year (2007-2009). During the reference decade between 2007 and 2016, ground temperature near the depth of zero annual amplitude in the continuous permafrost zone increased by 0.39 ± 0.15 °C. Over the same period, discontinuous permafrost warmed by 0.20 ± 0.10 °C. Permafrost in mountains warmed by 0.19 ± 0.05 °C and in Antarctica by 0.37 ± 0.10 °C. Globally, permafrost temperature increased by 0.29 ± 0.12 °C. The observed trend follows the Arctic amplification of air temperature increase in the Northern Hemisphere. In the discontinuous zone, however, ground warming occurred due to increased snow thickness while air temperature remained statistically unchanged.

Abstract Recent research using repeat photography, long-term ecological monitoring and dendrochronology has documented shrub expansion in arctic, high-latitude and alpine tundra ecosystems. Here, we (1) synthesize these findings, (2) present a conceptual framework that identifies mechanisms and constraints on shrub increase, (3) explore causes, feedbacks and implications of the increased shrub cover in tundra ecosystems, and (4) address potential lines of investigation for future research. Satellite observations from around the circumpolar Arctic, showing increased productivity, measured as changes in ‘greenness’, have coincided with a general rise in high-latitude air temperatures and have been partly attributed to increases in shrub cover. Studies indicate that warming temperatures, changes in snow cover, altered disturbance regimes as a result of permafrost thaw, tundra fires, and anthropogenic activities or changes in herbivory intensity are all contributing to observed changes in shrub abundance. A large-scale increase in shrub cover will change the structure of tundra ecosystems and alter energy fluxes, regional climate, soil–atmosphere exchange of water, carbon and nutrients, and ecological interactions between species. In order to project future rates of shrub expansion and understand the feedbacks to ecosystem and climate processes, future research should investigate the species or trait-specific responses of shrubs to climate change including: (1) the temperature sensitivity of shrub growth, (2) factors controlling the recruitment of new individuals, and (3) the relative influence of the positive and negative feedbacks involved in shrub expansion.

Climate change is generating complex responses in both natural and human ecosystems that vary in their geographic distribution, magnitude, and timing across the global landscape. One of the major issues that scientists and policy makers now confront is how to assess such massive changes over multiple scales of space and time. Lakes and reservoirs comprise a geographically distributed network of the lowest points in the surrounding landscape that make them important sentinels of climate change. Their physical, chemical, and biological responses to climate provide a variety of information‐rich signals. Their sediments archive and integrate these signals, enabling paleolimnologists to document changes over years to millennia. Lakes are also hot spots of carbon cycling in the landscape and as such are important regulators of climate change, processing terrestrial and atmospheric as well as aquatic carbon. We provide an overview of this concept of lakes and reservoirs as sentinels, integrators, and regulators of climate change, as well as of the need for scaling and modeling these responses in the context of global climate change. We conclude by providing a brief look to the future and the creation of globally networked sensors in lakes and reservoirs around the world.

Abstract. The Arctic is a water-rich region, with freshwater systems covering about 16 % of the northern permafrost landscape. Permafrost thaw creates new freshwater ecosystems, while at the same time modifying the existing lakes, streams, and rivers that are impacted by thaw. Here, we describe the current state of knowledge regarding how permafrost thaw affects lentic (still) and lotic (moving) systems, exploring the effects of both thermokarst (thawing and collapse of ice-rich permafrost) and deepening of the active layer (the surface soil layer that thaws and refreezes each year). Within thermokarst, we further differentiate between the effects of thermokarst in lowland areas vs. that on hillslopes. For almost all of the processes that we explore, the effects of thaw vary regionally, and between lake and stream systems. Much of this regional variation is caused by differences in ground ice content, topography, soil type, and permafrost coverage. Together, these modifying factors determine (i) the degree to which permafrost thaw manifests as thermokarst, (ii) whether thermokarst leads to slumping or the formation of thermokarst lakes, and (iii) the manner in which constituent delivery to freshwater systems is altered by thaw. Differences in thaw-enabled constituent delivery can be considerable, with these modifying factors determining, for example, the balance between delivery of particulate vs. dissolved constituents, and inorganic vs. organic materials. Changes in the composition of thaw-impacted waters, coupled with changes in lake morphology, can strongly affect the physical and optical properties of thermokarst lakes. The ecology of thaw-impacted lakes and streams is also likely to change; these systems have unique microbiological communities, and show differences in respiration, primary production, and food web structure that are largely driven by differences in sediment, dissolved organic matter, and nutrient delivery. The degree to which thaw enables the delivery of dissolved vs. particulate organic matter, coupled with the composition of that organic matter and the morphology and stratification characteristics of recipient systems will play an important role in determining the balance between the release of organic matter as greenhouse gases (CO2 and CH4), its burial in sediments, and its loss downstream. The magnitude of thaw impacts on northern aquatic ecosystems is increasing, as is the prevalence of thaw-impacted lakes and streams. There is therefore an urgent need to quantify how permafrost thaw is affecting aquatic ecosystems across diverse Arctic landscapes, and the implications of this change for further climate warming.

Abstract Glacial cycles have played a dominant role in shaping the genetic structure and distribution of biota in northwestern North America. The two major ice age refugia of Beringia and the Pacific Northwest were connected by major mountain chains and bordered by the Pacific Ocean. As a result, numerous refugial options were available for the regions taxa during glacial advances. We reviewed the importance of glaciations and refugia in shaping northwestern North America’s phylogeographic history. We also tested whether ecological variables were associated with refugial history. The recurrent phylogeographic patterns that emerged were the following: (i) additional complexity, i.e. refugia within refugia , in both Beringia and the Pacific Northwest; and (ii) strong evidence for cryptic refugia in the Alexander Archipelago and Haida Gwaii, the Canadian Arctic and within the ice‐sheets. Species with contemporary ranges that covered multiple refugia, or those with high dispersal ability, were significantly more likely to have resided in multiple refugia. Most of the shared phylogeographic patterns can be attributed to multiple refugial locales during the last glacial maximum or major physiographic barriers like rivers and glaciers. However, some of the observed patterns are much older and appear connected to the orogeny of the Cascade‐Sierra chain or allopatric differentiation during historic glacial advances. The emergent patterns from this review suggest we should refine the classic Beringian‐southern refugial paradigm for northwestern North American biota and highlight the ecological and evolutionary consequences of colonization from multiple refugia.

We discuss hidden Markov‐type models for fitting a variety of multistate random walks to wildlife movement data. Discrete‐time hidden Markov models (HMMs) achieve considerable computational gains by focusing on observations that are regularly spaced in time, and for which the measurement error is negligible. These conditions are often met, in particular for data related to terrestrial animals, so that a likelihood‐based HMM approach is feasible. We describe a number of extensions of HMMs for animal movement modeling, including more flexible state transition models and individual random effects (fitted in a non‐Bayesian framework). In particular we consider so‐called hidden semi‐Markov models, which may substantially improve the goodness of fit and provide important insights into the behavioral state switching dynamics. To showcase the expediency of these methods, we consider an application of a hierarchical hidden semi‐Markov model to multiple bison movement paths.

Recent warming in the Arctic, which has been amplified during the winter1–3, greatly enhances microbial decomposition of soil organic matter and subsequent release of carbon dioxide (CO2)4. However, the amount of CO2 released in winter is not known and has not been well represented by ecosystem models or empirically based estimates5,6. Here we synthesize regional in situ observations of CO2 flux from Arctic and boreal soils to assess current and future winter carbon losses from the northern permafrost domain. We estimate a contemporary loss of 1,662 TgC per year from the permafrost region during the winter season (October–April). This loss is greater than the average growing season carbon uptake for this region estimated from process models (−1,032 TgC per year). Extending model predictions to warmer conditions up to 2100 indicates that winter CO2 emissions will increase 17% under a moderate mitigation scenario—Representative Concentration Pathway 4.5—and 41% under business-as-usual emissions scenario—Representative Concentration Pathway 8.5. Our results provide a baseline for winter CO2 emissions from northern terrestrial regions and indicate that enhanced soil CO2 loss due to winter warming may offset growing season carbon uptake under future climatic conditions. Winter warming in the Arctic will increase the CO2 flux from soils. A pan-Arctic analysis shows a current loss of 1,662 TgC per year over the winter, exceeding estimated carbon uptake in the growing season; projections suggest a 17% increase under RCP 4.5 and a 41% increase under RCP 8.5 by 2100.

Abstract A snapshot of the thermal state of permafrost in northern North America during the International Polar Year (IPY) was developed using ground temperature data collected from 350 boreholes. More than half these were established during IPY to enhance the network in sparsely monitored regions. The measurement sites span a diverse range of ecoclimatic and geological conditions across the continent and are at various elevations within the Cordillera. The ground temperatures within the discontinuous permafrost zone are generally above −3°C, and range down to −15°C in the continuous zone. Ground temperature envelopes vary according to substrate, with shallow depths of zero annual amplitude for peat and mineral soils, and much greater depths for bedrock. New monitoring sites in the mountains of southern and central Yukon suggest that permafrost may be limited in extent. In concert with regional air temperatures, permafrost has generally been warming across North America for the past several decades, as indicated by measurements from the western Arctic since the 1970s and from parts of eastern Canada since the early 1990s. The rates of ground warming have been variable, but are generally greater north of the treeline. Latent heat effects in the southern discontinuous zone dominate the permafrost thermal regime close to 0°C and allow permafrost to persist under a warming climate. Consequently, the spatial diversity of permafrost thermal conditions is decreasing over time. Copyright © 2010 Crown in the right of Canada and John Wiley & Sons, Ltd.

MOTIVATION: The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. MAIN TYPES OF VARIABLES INCLUDED: The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. SPATIAL LOCATION AND GRAIN: ). TIME PERIOD AND GRAIN: BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. MAJOR TAXA AND LEVEL OF MEASUREMENT: BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. SOFTWARE FORMAT: .csv and .SQL.

Arctic warming can influence tundra ecosystem function with consequences for climate feedbacks, wildlife and human communities. Yet ecological change across the Arctic tundra biome remains poorly quantified due to field measurement limitations and reliance on coarse-resolution satellite data. Here, we assess decadal changes in Arctic tundra greenness using time series from the 30 m resolution Landsat satellites. From 1985 to 2016 tundra greenness increased (greening) at ~37.3% of sampling sites and decreased (browning) at ~4.7% of sampling sites. Greening occurred most often at warm sampling sites with increased summer air temperature, soil temperature, and soil moisture, while browning occurred most often at cold sampling sites that cooled and dried. Tundra greenness was positively correlated with graminoid, shrub, and ecosystem productivity measured at field sites. Our results support the hypothesis that summer warming stimulated plant productivity across much, but not all, of the Arctic tundra biome during recent decades.

Caribou ( Rangifer tarandus (L., 1758)) play a central role in the ecology and culture of much of Canada, where they were once the most abundant cervid. Most populations are currently declining, and some face extirpation. In southern Canada, caribou range has retreated considerably over the past century. The ultimate reason for their decline is habitat alterations by industrial activities. The proximate causes are predation and, to a lesser extent, overharvest. The most southerly populations of “Mountain” caribou are at imminent risk of extirpation. Mountain caribou are threatened by similar industrial activities as Boreal caribou, and face increasing harassment from motorized winter recreational activities. Most populations of “Migratory Tundra” caribou are currently declining. Although these caribou fluctuate in abundance over decades, changing harvest technologies, climate change, increasing industrial development and human presence in the North raise doubts over whether recent declines will be followed by recoveries. The Peary caribou ( Rangifer tarandus pearyi J.A. Allen, 1902), a distinct subspecies endemic to Canada’s High Arctic, has suffered drastic declines caused by severe weather, hunting and predation. It faces an increasing threat from climate change. While some questions remain about the reasons for the decline of Migratory Tundra caribou, research has clearly identified several threats to the persistence of “Boreal”, Mountain, and Peary caribou. Scientific knowledge, however, has neither effectively influenced policies nor galvanized public opinion sufficiently to push governments into effective actions. The persistence of many caribou populations appears incompatible with the ongoing pace of industrial development.

Seasonal patterns of climate and vegetation growth are expected to be altered by global warming. In alpine environments, the reproduction of birds and mammals is tightly linked to seasonality; therefore such alterations may have strong repercussions on recruitment. We used the normalized difference vegetation index (NDVI), a satellite-based measurement that correlates strongly with aboveground net primary productivity, to explore how annual variations in the timing of vegetation onset and in the rate of change in primary production during green-up affected juvenile growth and survival of bighorn sheep (Ovis canadensis), Alpine ibex (Capra ibex), and mountain goats (Oreamnos americanus) in four different populations in two continents. We indexed timing of onset of vegetation growth by the integrated NDVI (INDVI) in May. The rate of change in primary production during green-up (early May to early July) was estimated as (1) the maximal slope between any two successive bimonthly NDVI values during this period and (2) the slope in NDVI between early May and early July. The maximal slope in NDVI was negatively correlated with lamb growth and survival in both populations of bighorn sheep, growth of mountain goat kids, and survival of Alpine ibex kids, but not with survival of mountain goat kids. There was no effect of INDVI in May and of the slope in NDVI between early May and early July on juvenile growth and survival for any species. Although rapid changes in NDVI during the green-up period could translate into higher plant productivity, they may also lead to a shorter period of availability of high-quality forage over a large spatial scale, decreasing the opportunity for mountain ungulates to exploit high-quality forage. Our results suggest that attempts to forecast how warmer winters and springs will affect animal population dynamics and life histories in alpine environments should consider factors influencing the rate of changes in primary production during green-up and the timing of vegetation onset.

Contents Summary 652 I. Introduction 652 II. Discrepancy in predicting the effects of rising [CO 2 ] on the terrestrial C sink 655 III. Carbon and nutrient storage in plants and its modelling 656 IV. Modelling the source and the sink: a plant perspective 657 V. Plant‐scale water and Carbon flux models 660 VI. Challenges for the future 662 Acknowledgements 663 Authors contributions 663 References 663 Summary The increase in atmospheric CO 2 in the future is one of the most certain projections in environmental sciences. Understanding whether vegetation carbon assimilation, growth, and changes in vegetation carbon stocks are affected by higher atmospheric CO 2 and translating this understanding in mechanistic vegetation models is of utmost importance. This is highlighted by inconsistencies between global‐scale studies that attribute terrestrial carbon sinks to CO 2 stimulation of gross and net primary production on the one hand, and forest inventories, tree‐scale studies, and plant physiological evidence showing a much less pronounced CO 2 fertilization effect on the other hand. Here, we review how plant carbon sources and sinks are currently described in terrestrial biosphere models. We highlight an uneven representation of complexity between the modelling of photosynthesis and other processes, such as plant respiration, direct carbon sinks, and carbon allocation, largely driven by available observations. Despite a general lack of data on carbon sink dynamics to drive model improvements, ways forward toward a mechanistic representation of plant carbon sinks are discussed, leveraging on results obtained from plant‐scale models and on observations geared toward model developments.

Although cyanobacteria are often thought of as warm water organisms. they are the predominant biota in cold polar environments such as ice shelves. glaciers. glacial meltwater streams and ice-capped lakes. Cyanobacteria are the primary colonizers of glacial moraines after the retreat of ice sheets. and they play an important role in the carbon and nitrogen economy of tundra and polar desert soils. Various communities dominated by cyanobacteria inhabit exposed rock surfaces. while others occur within fissures and the interstitial spaces between crystals in certain Arctic and Antarctic rock types (See Chapter 13). Highly pigmented microbial mats dominated by Nostoc or oscillatorians (Oscillatoriaceae) are a feature of streams. lakes and ponds in both polar regions. with extreme accumulations up to 90 cm thick and >40 μg Chla cm-2 at some sites. Picocyanobacteria often dominate the phytoplankton of polar and subpolar lakes. In the coastal saline lakes of Antarctica picocyanobacteria achieve some of the highest natural concentrations on record. up to 8 x 106 cells mL-1. However. picocyanobacteria are conspicuously absent or rare in the adjacent polar oceans. The ecophysiological characteristics of high-latitude cyanobacteria that contribute to their success and dominance include: an ability to grow over a wide temperature range (but at slow rates); tolerance of desiccation, freezing and salinity stress; a variety of adaptive strategies against high levels of solar radiation (including ultraviolet radiation) in exposed habitats; and acclimation to shade allowing net growth in protected dim light environments. In many polar habitats, the large standing stocks of cyanobacterial biomass are the result of gradual accumulation over many seasons, with only minor losses via biotic and abiotic removal processes. Cyanobacteria are not successful in the polar oceans where slow, temperature-depressed and light-limited growth rates are unable to keep pace with the continuous losses due to grazing, advection and mixing.

A comprehensive database of paleoclimate records is needed to place recent warming into the longer-term context of natural climate variability. We present a global compilation of quality-controlled, published, temperature-sensitive proxy records extending back 12,000 years through the Holocene. Data were compiled from 679 sites where time series cover at least 4000 years, are resolved at sub-millennial scale (median spacing of 400 years or finer) and have at least one age control point every 3000 years, with cut-off values slackened in data-sparse regions. The data derive from lake sediment (51%), marine sediment (31%), peat (11%), glacier ice (3%), and other natural archives. The database contains 1319 records, including 157 from the Southern Hemisphere. The multi-proxy database comprises paleotemperature time series based on ecological assemblages, as well as biophysical and geochemical indicators that reflect mean annual or seasonal temperatures, as encoded in the database. This database can be used to reconstruct the spatiotemporal evolution of Holocene temperature at global to regional scales, and is publicly available in Linked Paleo Data (LiPD) format.

Cell formation in growth rings of balsam fir (Abies balsamea (L.) Mill) in the boreal forest was studied to describe the timing of ring formation and the development patterns of earlywood and latewood. Wood micro-cores were extracted during the growing season from 1998 to 2000. The micro-cores were stained with cresyl fast violet to facilitate counting the number of cells in the radial enlargement, wall thickening phases, and mature cell phases. The periods required to complete these various phases were then estimated. Variations in the beginning of the growing season (May 7 – June 7), the earlywood–latewood transition (July 2 – July 19), and the end of the growing season ( August 20 – September 20) were observed. Short cell enlargement durations of less than a week for earlywood and 5–10 days for latewood were observed. Time required for cell wall thickening was about 20 days for earlywood and longer than 10–15 days for latewood. A certain flexibility was observed in the ring formation patterns and in the cell development rate, providing an advantage in the boreal forest where optimal growth conditions change from year to year. These findings on the spatial and temporal patterns of ring development may be useful for understanding tree relationships with climate or other environmental parameters.

Abstract Numerous international scientific assessments and related articles have, during the last decade, described the observed and potential impacts of climate change as well as other related environmental stressors on Arctic ecosystems. There is increasing recognition that observed and projected changes in freshwater sources, fluxes, and storage will have profound implications for the physical, biogeochemical, biological, and ecological processes and properties of Arctic terrestrial and freshwater ecosystems. However, a significant level of uncertainty remains in relation to forecasting the impacts of an intensified hydrological regime and related cryospheric change on ecosystem structure and function. As the terrestrial and freshwater ecology component of the Arctic Freshwater Synthesis, we review these uncertainties and recommend enhanced coordinated circumpolar research and monitoring efforts to improve quantification and prediction of how an altered hydrological regime influences local, regional, and circumpolar‐level responses in terrestrial and freshwater systems. Specifically, we evaluate (i) changes in ecosystem productivity; (ii) alterations in ecosystem‐level biogeochemical cycling and chemical transport; (iii) altered landscapes, successional trajectories, and creation of new habitats; (iv) altered seasonality and phenological mismatches; and (v) gains or losses of species and associated trophic interactions. We emphasize the need for developing a process‐based understanding of interecosystem interactions, along with improved predictive models. We recommend enhanced use of the catchment scale as an integrated unit of study, thereby more explicitly considering the physical, chemical, and ecological processes and fluxes across a full freshwater continuum in a geographic region and spatial range of hydroecological units (e.g., stream‐pond‐lake‐river‐near shore marine environments).

Abstract Geophysical techniques can be used to examine the spatial distribution of subsurface geophysical properties to delineate horizontally and vertically the active layer, permafrost and taliks. Spatial and temporal changes in subsurface geophysical properties due to permafrost cooling, warming, aggradation or degradation can also be assessed through geophysical monitoring. This paper reviews the geophysical methods most frequently applied in mountain and arctic/subarctic lowland permafrost investigations. Key results and recommendations based on the analysis of the applicability and reliability of different geophysical techniques for permafrost studies are summarised. Emphasis is put on the tomographic capabilities of geophysical methods. Recent advances in application and data interpretation are shown in relation to five case studies, and future perspectives are highlighted. Copyright © 2008 John Wiley & Sons, Ltd.

As the permafrost region warms, its large organic carbon pool will be increasingly vulnerable to decomposition, combustion, and hydrologic export. Models predict that some portion of this release will be offset by increased production of Arctic and boreal biomass; however, the lack of robust estimates of net carbon balance increases the risk of further overshooting international emissions targets. Precise empirical or model-based assessments of the critical factors driving carbon balance are unlikely in the near future, so to address this gap, we present estimates from 98 permafrost-region experts of the response of biomass, wildfire, and hydrologic carbon flux to climate change. Results suggest that contrary to model projections, total permafrost-region biomass could decrease due to water stress and disturbance, factors that are not adequately incorporated in current models. Assessments indicate that end-of-the-century organic carbon release from Arctic rivers and collapsing coastlines could increase by 75% while carbon loss via burning could increase four-fold. Experts identified water balance, shifts in vegetation community, and permafrost degradation as the key sources of uncertainty in predicting future system response. In combination with previous findings, results suggest the permafrost region will become a carbon source to the atmosphere by 2100 regardless of warming scenario but that 65%-85% of permafrost carbon release can still be avoided if human emissions are actively reduced.