Late Pleistocene shrub expansion preceded megafauna turnover and extinctions in eastern Beringia.

The collapse of the steppe-tundra biome (mammoth steppe) at the end of the Pleistocene is used as an important example of top-down ecosystem cascades, where human hunting of keystone species led to profound changes in vegetation across high latitudes in the Northern Hemisphere. Alternatively, it is argued that this biome transformation occurred through a bottom-up process, where climate-driven expansion of shrub tundra (Betula, Salix spp.) replaced the steppe-tundra vegetation that grazing megafauna taxa relied on. In eastern Beringia, these differing hypotheses remain largely untested, in part because the precise timing and spatial pattern of Late Pleistocene shrub expansion remains poorly resolved. This uncertainty is caused by chronological ambiguity in many lake sediment records, which typically rely on radiocarbon (14C) dates from bulk sediment or aquatic macrofossils-materials that are known to overestimate the age of sediment layers. Here, we reexamine Late Pleistocene pollen records for which 14C dating of terrestrial macrofossils is available and augment these data with 14C dates from arctic ground-squirrel middens and plant macrofossils. Comparing these paleovegetation data with a database of published 14C dates from megafauna remains, we find the postglacial expansion of shrub tundra preceded the regional extinctions of horse (Equus spp.) and mammoth (Mammuthus primigenius) and began during a period when the frequency of 14C dates indicates large grazers were abundant. These results are not consistent with a model of top-down ecosystem cascades and support the hypothesis that climate-driven habitat loss preceded and contributed to turnover in mammal communities.

Life and extinction of megafauna in the ice-age Arctic.

Understanding the population dynamics of megafauna that inhabited the mammoth steppe provides insights into the causes of extinctions during both the terminal Pleistocene and today. Our study area is Alaska's North Slope, a place where humans were rare when these extinctions occurred. After developing a statistical approach to remove the age artifacts caused by radiocarbon calibration from a large series of dated megafaunal bones, we compare the temporal patterns of bone abundance with climate records. Megafaunal abundance tracked ice age climate, peaking during transitions from cold to warm periods. These results suggest that a defining characteristic of the mammoth steppe was its temporal instability and imply that regional extinctions followed by population reestablishment from distant refugia were characteristic features of ice-age biogeography at high latitudes. It follows that long-distance dispersal was crucial for the long-term persistence of megafaunal species living in the Arctic. Such dispersal was only possible when their rapidly shifting range lands were geographically interconnected. The end of the last ice age was fatally unique because the geographic ranges of arctic megafauna became permanently fragmented after stable, interglacial climate engendered the spread of peatlands at the same time that rising sea level severed former dispersal routes.

Inland waters and their role in the carbon cycle of Alaska.

The magnitude of Alaska (AK) inland waters carbon (C) fluxes is likely to change in the future due to amplified climate warming impacts on the hydrology and biogeochemical processes in high latitude regions. Although current estimates of major aquatic C fluxes represent an essential baseline against which future change can be compared, a comprehensive assessment for AK has not yet been completed. To address this gap, we combined available data sets and applied consistent methodologies to estimate river lateral C export to the coast, river and lake carbon dioxide (CO2 ) and methane (CH4 ) emissions, and C burial in lakes for the six major hydrologic regions in the state. Estimated total aquatic C flux for AK was 41 Tg C/yr. Major components of this total flux, in Tg C/yr, were 18 for river lateral export, 17 for river CO2 emissions, and 8 for lake CO2 emissions. Lake C burial offset these fluxes by 2 Tg C/yr. River and lake CH4 emissions were 0.03 and 0.10 Tg C/yr, respectively. The Southeast and South central regions had the highest temperature, precipitation, terrestrial net primary productivity (NPP), and C yields (fluxes normalized to land area) were 77 and 42 g C·m-2 ·yr-1 , respectively. Lake CO2 emissions represented over half of the total aquatic flux from the Southwest (37 g C·m-2 ·yr-1 ). The North Slope, Northwest, and Yukon regions had lesser yields (11, 15, and 17 g C·m2 ·yr-1 ), but these estimates may be the most vulnerable to future climate change, because of the heightened sensitivity of arctic and boreal ecosystems to intensified warming. Total aquatic C yield for AK was 27 g C·m-2 ·yr-1 , which represented 16% of the estimated terrestrial NPP. Freshwater ecosystems represent a significant conduit for C loss, and a more comprehensive view of land-water-atmosphere interactions is necessary to predict future climate change impacts on the Alaskan ecosystem C balance.

Post-fire stabilization of thaw-affected permafrost terrain in northern Alaska.

In 2007, the Anaktuvuk River fire burned more than 1000 km2 of arctic tundra in northern Alaska, ~ 50% of which occurred in an area with ice-rich syngenetic permafrost (Yedoma). By 2014, widespread degradation of ice wedges was apparent in the Yedoma region. In a 50 km2 area, thaw subsidence was detected across 15% of the land area in repeat airborne LiDAR data acquired in 2009 and 2014. Updating observations with a 2021 airborne LiDAR dataset show that additional thaw subsidence was detected in < 1% of the study area, indicating stabilization of the thaw-affected permafrost terrain. Ground temperature measurements between 2010 and 2015 indicated that the number of near-surface soil thawing-degree-days at the burn site were 3 × greater than at an unburned control site, but by 2022 the number was reduced to 1.3 × greater. Mean annual ground temperature of the near-surface permafrost increased by 0.33 °C/yr in the burn site up to 7-years post-fire, but then cooled by 0.15 °C/yr in the subsequent eight years, while temperatures at the control site remained relatively stable. Permafrost cores collected from ice-wedge troughs (n = 41) and polygon centers (n = 8) revealed the presence of a thaw unconformity, that in most cases was overlain by a recovered permafrost layer that averaged 14.2 cm and 18.3 cm, respectively. Taken together, our observations highlight that the initial degradation of ice-rich permafrost following the Anaktuvuk River tundra fire has been followed by a period of thaw cessation, permafrost aggradation, and terrain stabilization.

Expanding beaver pond distribution in Arctic Alaska, 1949 to 2019.

Beavers were not previously recognized as an Arctic species, and their engineering in the tundra is considered negligible. Recent findings suggest that beavers have moved into Arctic tundra regions and are controlling surface water dynamics, which strongly influence permafrost and landscape stability. Here we use 70 years of satellite images and aerial photography to show the scale and magnitude of northwestward beaver expansion in Alaska, indicated by the construction of over 10,000 beaver ponds in the Arctic tundra. The number of beaver ponds doubled in most areas between ~ 2003 and ~ 2017. Earlier stages of beaver engineering are evident in ~ 1980 imagery, and there is no evidence of beaver engineering in ~ 1952 imagery, consistent with observations from Indigenous communities describing the influx of beavers over the period. Rapidly expanding beaver engineering has created a tundra disturbance regime that appears to be thawing permafrost and exacerbating the effects of climate change.

Summer warming explains widespread but not uniform greening in the Arctic tundra biome.

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.

Climate-driven ecological stability as a globally shared cause of Late Quaternary megafaunal extinctions: the Plaids and Stripes Hypothesis.

Controversy persists about why so many large-bodied mammal species went extinct around the end of the last ice age. Resolving this is important for understanding extinction processes in general, for assessing the ecological roles of humans, and for conserving remaining megafaunal species, many of which are endangered today. Here we explore an integrative hypothesis that asserts that an underlying cause of Late Quaternary megafaunal extinctions was a fundamental shift in the spatio-temporal fabric of ecosystems worldwide. This shift was triggered by the loss of the millennial-scale climate fluctuations that were characteristic of the ice age but ceased approximately 11700 years ago on most continents. Under ice-age conditions, which prevailed for much of the preceding 2.6 Ma, these radical and rapid climate changes prevented many ecosystems from fully equilibrating with their contemporary climates. Instead of today's 'striped' world in which species' ranges have equilibrated with gradients of temperature, moisture, and seasonality, the ice-age world was a disequilibrial 'plaid' in which species' ranges shifted rapidly and repeatedly over time and space, rarely catching up with contemporary climate. In the transient ecosystems that resulted, certain physiological, anatomical, and ecological attributes shared by megafaunal species pre-adapted them for success. These traits included greater metabolic and locomotory efficiency, increased resistance to starvation, longer life spans, greater sensory ranges, and the ability to be nomadic or migratory. When the plaid world of the ice age ended, many of the advantages of being large were either lost or became disadvantages. For instance in a striped world, the low population densities and slow reproductive rates associated with large body size reduced the resiliency of megafaunal species to population bottlenecks. As the ice age ended, the downsides of being large in striped environments lowered the extinction thresholds of megafauna worldwide, which then increased the vulnerability of individual species to a variety of proximate threats they had previously tolerated, such as human predation, competition with other species, and habitat loss. For many megafaunal species, the plaid-to-stripes transition may have been near the base of a hierarchy of extinction causes whose relative importances varied geographically, temporally, and taxonomically.