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Understanding how mutations arise and spread through individuals and populations is fundamental to evolutionary biology. Most organisms have a life cycle with unicellular bottlenecks during reproduction. However, some organisms like plants, fungi, or colonial animals can grow indefinitely, changing the manner in which mutations spread throughout both the individual and the population. Furthermore, clonally reproducing organisms may also achieve exceedingly long lifespans, making somatic mutation an important mechanism of creating heritable variation for Darwinian evolution by natural selection. Yet, little is known about intra-organism mutation rates and evolutionary trajectories in long-lived species. Here, we study the Pando aspen clone, the largest known quaking aspen (Populus tremuloides) clone founded by a single seedling and thought to be one of the oldest studied organisms. Aspen reproduce vegetatively via new root-borne stems forming clonal patches, sometimes spanning several hectares. To study the evolutionary history of the Pando clone, we collected and sequenced over 500 samples from Pando and neighboring clones, as well as from various tissue types within Pando, including leaves, roots, and bark. We applied a series of filters to distinguish somatic mutations from the pool of both somatic and germline mutations, incorporating a technical replicate sequencing approach to account for uncertainty in somatic mutation detection. Despite root spreading being spatially constrained, we observed only a modest positive correlation between genetic and spatial distance, suggesting the presence of a mechanism preventing the accumulation and spread of mutations across units. Phylogenetic models estimate the age of the clone to between ~16,000-80,000 years. This age is generally corroborated by the near-continuous presence of aspen pollen in a lake sediment record collected from Fish Lake near Pando. Overall, this work enhances understanding of mutation accumulation and dispersal within and between ramets of long-lived, clonally-reproducing organisms. Significance StatementThis study enhances our understanding of evolutionary processes in long-lived clonal organisms by investigating somatic mutation accumulation and dispersal patterns within the iconic Pando aspen clone. The authors estimated the clone to be between 10,000 and 80,000 years old and uncovered a modest spatial genetic structure in the 42.6-hectare clone, suggesting localized mutation build-up rather than dispersal along tissue lineages. This work sheds light on an ancient organism and how plants may evolve to preserve genetic integrity in meristems fueling indefinite growth, with implications for our comprehension of adaptive strategies in long-lived perennials. 

Abstract. Land cover governs the biogeophysical and biogeochemical feedbacks between the land surface and atmosphere. Holocene vegetation-atmosphere interactions are of particular interest, both to understand the climate effects of intensifying human land use and as a possible explanation for the Holocene Conundrum, a widely studied mismatch between simulated and reconstructed temperatures. Progress has been limited by a lack of data-constrained, quantified, and consistently produced reconstructions of Holocene land cover change. As a contribution to the Past Global Changes (PAGES) LandCover6k Working Group, we present a new suite of land cover reconstructions with uncertainty for North America, based on a network of 1445 sedimentary pollen records and the REVEALS pollen-vegetation model coupled with a Bayesian spatial model. These spatially comprehensive land cover maps are then used to determine the pattern and magnitude of North American land cover changes at continental to regional scales. Early Holocene afforestation in North America was driven by rising temperatures and deglaciation, and this afforestation likely amplified early Holocene warming via the albedo effect. A continental-scale mid-Holocene peak in summergreen trees and shrubs (8.5 to 4 ka) is hypothesized to represent a positive and understudied feedback loop among insolation, temperature, and phenology seasonality. A last-millennium decrease in summergreen trees and shrubs with corresponding increases in open land likely was driven by a spatially varying combination of intensifying land use and neoglacial cooling. Land cover trends vary within and across regions, due to individualistic taxon-level responses to environmental change. Major species-level events, such as the mid-Holocene decline of eastern hemlock, may have altered regional climates. The substantial land-cover changes reconstructed here support the importance of biogeophysical vegetation feedbacks to Holocene climate dynamics. However, recent model experiments that invoke vegetation feedbacks to explain the Holocene Conundrum may have overestimated the land cover forcing by replacing Northern Hemisphere grasslands >30° N with forests; an ecosystem state that is not supported by these land cover reconstructions. These Holocene reconstructions for North America, along with similar LandCover6k products now available for other continents, serve the Earth system modeling community by providing better-constrained land cover scenarios and benchmarks for model evaluation, ultimately making it possible to better understand the regional- to global-scale processes driving Holocene land cover dynamics. 

Abstract Climatic conditions exert an important influence on wildfire activity in the western United States; however, Indigenous farming activity may have also shaped the local fire regimes for millennia. The Fish Lake Plateau is located on the Great Basin–Colorado Plateau boundary, the only region in western North America where maize farming was adopted then suddenly abandoned. Here we integrate sedimentary archives, tree rings, and archeological data to reconstruct the past 1200 years of fire, climate, and human activity. We identify a period of high fire activity during the apex of prehistoric farming between 900 and 1400 CE, and suggest that farming likely obscured the role of climate on the fire regime through the use of frequent low-severity burning. Climatic conditions again became the dominant driver of wildfire when prehistoric populations abandoned farming around 1400 CE. We conclude that Indigenous populations shaped high-elevation mixed-conifer fire regimes on the Fish Lake Plateau through land-use practices.