Plant functional trait change across a warming tundra biome.

The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

Local snow melt and temperature-but not regional sea ice-explain variation in spring phenology in coastal Arctic tundra.

The Arctic is undergoing dramatic environmental change with rapidly rising surface temperatures, accelerating sea ice decline and changing snow regimes, all of which influence tundra plant phenology. Despite these changes, no globally consistent direction of trends in spring phenology has been reported across the Arctic. While spring has advanced at some sites, spring has delayed or not changed at other sites, highlighting substantial unexplained variation. Here, we test the relative importance of local temperatures, local snow melt date and regional spring drop in sea ice extent as controls of variation in spring phenology across different sites and species. Trends in long-term time series of spring leaf-out and flowering (average span: 18 years) were highly variable for the 14 tundra species monitored at our four study sites on the Arctic coasts of Alaska, Canada and Greenland, ranging from advances of 10.06 days per decade to delays of 1.67 days per decade. Spring temperatures and the day of spring drop in sea ice extent advanced at all sites (average 1°C per decade and 21 days per decade, respectively), but only those sites with advances in snow melt (average 5 days advance per decade) also had advancing phenology. Variation in spring plant phenology was best explained by snow melt date (mean effect: 0.45 days advance in phenology per day advance snow melt) and, to a lesser extent, by mean spring temperature (mean effect: 2.39 days advance in phenology per °C). In contrast to previous studies examining sea ice and phenology at different spatial scales, regional spring drop in sea ice extent did not predict spring phenology for any species or site in our analysis. Our findings highlight that tundra vegetation responses to global change are more complex than a direct response to warming and emphasize the importance of snow melt as a local driver of tundra spring phenology.

Effects of precipitation change and neighboring plants on population dynamics of Bromus tectorum.

Shifting precipitation patterns resulting from global climate change will influence the success of invasive plant species. In the Front Range of Colorado, Bromus tectorum (cheatgrass) and other non-native winter annuals have invaded grassland communities and are becoming more abundant. As the global climate warms, more precipitation may fall as rain rather than snow in winter, and an increase in winter rain could benefit early-growing winter annuals, such as B. tectorum, to the detriment of native species. In this study we measured the effects of simulated changes in seasonal precipitation and presence of other plant species on population growth of B. tectorum in a grassland ecosystem near Boulder, Colorado, USA. We also performed elasticity analyses to identify life transitions that were most sensitive to precipitation differences. In both study years, population growth rates were highest for B. tectorum growing in treatments receiving supplemental winter precipitation and lowest for those receiving the summer drought treatment. Survival of seedlings to flowering and seed production contributed most to population growth in all treatments. Biomass of neighboring native plants was positively correlated with reduced population growth rates of B. tectorum. However, exotic plant biomass had no effect on population growth rates. This study demonstrates how interacting effects of climate change and presence of native plants can influence the population growth of an invasive species. Overall, our results suggest that B. tectorum will become more invasive in grasslands if the seasonality of precipitation shifts towards wetter winters and allows B. tectorum to grow when competition from native species is low.

The effects of black-tailed prairie dogs on plant communities within a complex urban landscape: an ecological surprise?

Historically, prairie dogs (Cynomys spp.) have been considered essential keystone species of western United States grassland ecosystems because they provide unique services and increase vegetation community richness, evenness, and diversity. However, the effects of black-tailed prairie dogs (Cynomys ludovicianus) on lands adjacent to or surrounded by urban areas may not result in the same ecosystem benefits historically associated with their presence. An urban landscape presents prairie dogs with movement challenges unparalleled in natural landscapes, as well as suites of nonnative plant species that are more common in disturbed areas. This study examined a complex ecosystem where vegetation communities are being influenced by directional environmental change, and quantified the synergistic effects resulting from the protective management of a native keystone species. The data set for this analysis was comprised of 71 paired (occupied by prairie dogs vs. unoccupied) vegetation surveys and 156 additional unpaired surveys collected from around the city of Boulder, Colorado, USA for 14 yr. Linear mixed models were used to compare data from transects occupied and unoccupied by prairie dogs, as well as to evaluate the effect of prairie dog occupation duration. In the absence of prairie dogs, vegetation in this region exhibited declines in native grasses, no changes in introduced grasses, and increases in native and nonnative forbs and bare soil over the study interval. In the presence of prairie dogs, these observed directional changes were nearly all amplified at rates four to 10 times greater than when prairie dogs were absent. Areas in Boulder occupied by prairie dogs also had significantly lower richness, evenness, and diversity of plant species, compared to unoccupied areas. Analysis of plant functional groups revealed the significant reduction of perennial native grasses, as well as a significantly higher cover of introduced forbs in occupied areas. Prairie dogs amplified the effects of low-impact environmental directional changes, creating more novel vegetation communities than the environmental factors alone. Results from this research provide insight regarding the synergistic effects of a native keystone species within a complex human-impacted 21st century landscape.

Plant traits poorly predict winner and loser shrub species in a warming tundra biome.

Climate change is leading to species redistributions. In the tundra biome, shrubs are generally expanding, but not all tundra shrub species will benefit from warming. Winner and loser species, and the characteristics that may determine success or failure, have not yet been fully identified. Here, we investigate whether past abundance changes, current range sizes and projected range shifts derived from species distribution models are related to plant trait values and intraspecific trait variation. We combined 17,921 trait records with observed past and modelled future distributions from 62 tundra shrub species across three continents. We found that species with greater variation in seed mass and specific leaf area had larger projected range shifts, and projected winner species had greater seed mass values. However, trait values and variation were not consistently related to current and projected ranges, nor to past abundance change. Overall, our findings indicate that abundance change and range shifts will not lead to directional modifications in shrub trait composition, since winner and loser species share relatively similar trait spaces.

Spatial Difference of Interactive Effect Between Temperature and Daylength on Ginkgo Budburst.

Climate warming-induced shifts in spring phenology have substantially affected the structure and function of terrestrial ecosystems and global biogeochemical cycles. Spring phenology is primarily triggered by spring temperature and is also affected by daylength and winter chilling, yet the relative importance of these cues across spatial gradients remains poorly understood. Here, we conducted a manipulative experiment with two daylength and three temperature treatments to investigate spatial differences in the response of ginkgo budburst to temperature and daylength, using twigs collected at three sites across a spatial gradient: a control site at a low latitude and low elevation on Tianmu Mountain (TMlow), a low latitude and high elevation site on Tianmu Mountain (TMhigh), and a high latitude site on Jiufeng mountain (JF). The mechanisms were also tested using in situ phenological observations of ginkgo along latitudes in China. We found that, compared to TMlow individuals, budburst dates occurred 12.6 (JF) and 7.7 (TMhigh) days earlier in high-latitude and high-elevation individuals when exposed to the same temperature and daylength treatments. Importantly, daylength only affected budburst at low latitudes, with long days (16 h) advancing budburst in low-latitude individuals by, on average, 8.1 days relative to short-day (8 h) conditions. This advance was most pronounced in low-elevation/latitude individuals (TMlow = 9.6 days; TMhigh = 6.7 days; JF = 1.6 days). In addition, we found that the temperature sensitivity of budburst decreased from 3.4 to 2.4 days °C-1 along latitude and from 3.4 to 2.5 days °C-1 along elevation, respectively. The field phenological observations verified the experimental results. Our findings provide empirical evidence of spatial differences in the relative effects of spring temperature and daylength on ginkgo budburst, which improved our understanding of spatial difference in phenological changes and the responses of terrestrial ecosystem to climate change.

INHABIT: A web-based decision support tool for invasive plant species habitat visualization and assessment across the contiguous United States.

Narrowing the communication and knowledge gap between producers and users of scientific data is a longstanding problem in ecological conservation and land management. Decision support tools (DSTs), including websites or interactive web applications, provide platforms that can help bridge this gap. DSTs can most effectively disseminate and translate research results when producers and users collaboratively and iteratively design content and features. One data resource seldom incorporated into DSTs are species distribution models (SDMs), which can produce spatial predictions of habitat suitability. Outputs from SDMs can inform management decisions, but their complexity and inaccessibility can limit their use by resource managers or policy makers. To overcome these limitations, we present the Invasive Species Habitat Tool (INHABIT), a novel, web-based DST built with R Shiny to display spatial predictions and tabular summaries of habitat suitability from SDMs for invasive plants across the contiguous United States. INHABIT provides actionable science to support the prevention and management of invasive species. Two case studies demonstrate the important role of end user feedback in confirming INHABIT's credibility, utility, and relevance.

Predicted distribution of a rare and understudied forest carnivore: Humboldt marten (Martes caurina humboldtensis).

BACKGROUND: Many mammalian species have experienced range contractions. Following a reduction in distribution that has resulted in apparently small and disjunct populations, the Humboldt marten (Martes caurina humboldtensis) was recently designated as federally Threatened and state Endangered. This subspecies of Pacific marten occurring in coastal Oregon and northern California, also known as coastal martens, appear unlike martens that occur in snow-associated regions in that vegetation associations appear to differ widely between Humboldt marten populations. We expected current distributions represent realized niches, but estimating factors associated with long-term occurrence was challenging for this rare and little-known species. Here, we assessed the predicted contemporary distribution of Humboldt martens and interpret our findings as hypotheses correlated with the subspecies' niche to inform strategic conservation actions. METHODS: We modeled Humboldt marten distribution using a maximum entropy (Maxent) approach. We spatially-thinned 10,229 marten locations collected from 1996-2020 by applying a minimum distance of 500-m between locations, resulting in 384 locations used to assess correlations of marten occurrence with biotic and abiotic variables. We independently optimized the spatial scale of each variable and focused development of model variables on biotic associations (e.g., hypothesized relationships with forest conditions), given that abiotic factors such as precipitation are largely static and not alterable within a management context. RESULTS: Humboldt marten locations were positively associated with increased shrub cover (salal (Gautheria shallon)), mast producing trees (e.g., tanoak, Notholithocarpus densiflorus), increased pine (Pinus sp.) proportion of total basal area, annual precipitation at home-range spatial scales, low and high amounts of canopy cover and slope, and cooler August temperatures. Unlike other recent literature, we found little evidence that Humboldt martens were associated with old-growth structural indices. This case study provides an example of how limited information on rare or lesser-known species can lead to differing interpretations, emphasizing the need for study-level replication in ecology. Humboldt marten conservation would benefit from continued survey effort to clarify range extent, population sizes, and fine-scale habitat use.

Experimental warming differentially affects vegetative and reproductive phenology of tundra plants.

Rapid climate warming is altering Arctic and alpine tundra ecosystem structure and function, including shifts in plant phenology. While the advancement of green up and flowering are well-documented, it remains unclear whether all phenophases, particularly those later in the season, will shift in unison or respond divergently to warming. Here, we present the largest synthesis to our knowledge of experimental warming effects on tundra plant phenology from the International Tundra Experiment. We examine the effect of warming on a suite of season-wide plant phenophases. Results challenge the expectation that all phenophases will advance in unison to warming. Instead, we find that experimental warming caused: (1) larger phenological shifts in reproductive versus vegetative phenophases and (2) advanced reproductive phenophases and green up but delayed leaf senescence which translated to a lengthening of the growing season by approximately 3%. Patterns were consistent across sites, plant species and over time. The advancement of reproductive seasons and lengthening of growing seasons may have significant consequences for trophic interactions and ecosystem function across the tundra.

Loss of foundation species increases population growth of exotic forbs in sagebrush steppe.

The invasion and spread of exotic plants following land disturbance threatens semiarid ecosystems. In sagebrush steppe, soil water is scarce and is partitioned between deep-rooted perennial shrubs and shallower-rooted native forbs and grasses. Disturbances commonly remove shrubs, leaving grass-dominated communities, and may allow for the exploitation of water resources by the many species of invasive, tap-rooted forbs that are increasingly successful in this habitat. We hypothesized that exotic forb populations would benefit from increased soil water made available by removal of sagebrush, a foundation species capable of deep-rooting, in semiarid shrub-steppe ecosystems. To test this hypothesis, we used periodic matrix models to examine effects of experimental manipulations of soil water on population growth of two exotic forb species, Tragopogon dubius and Lactuca serriola, in sagebrush steppe of southern Idaho, USA. We used elasticity analyses to examine which stages in the life cycle of T. dubius and L. serriola had the largest relative influence on population growth. We studied the demography of T. dubius and L. serriola in three treatments: (1) control, in which vegetation was not disturbed, (2) shrubs removed, or (3) shrubs removed but winter-spring recharge of deep-soil water blocked by rainout shelters. The short-term population growth rate (Lambda) of T. dubius in the shrub-removal treatment was more than double that of T. dubius in either sheltered or control treatments, both of which had limited soil water. All L. serriola individuals that emerged in undisturbed sagebrush plots died, whereas Lambda of L. serriola was high (Lambda > 2.5) in all shrub-removal plots, whether they had rainout shelters or not. Population growth of both forbs in all treatments was most responsive to flowering and seed production, which are life stages that should be particularly reliant on deep-soil water, as well as seedling establishment, which is important to most plant populations, especially during invasion. These data indicate the importance of native species, in this case the dominant shrub, in influencing soil resources and restricting population growth of exotic plants. These results argue that management of invasive plants should focus not only on removal of nonnatives, but also on reestablishment of important native species.

Climate Change: Flowering Time May Be Shifting in Surprising Ways.

A new study examined how flowering phenology has changed over the past three decades along an elevational gradient. These findings indicate that climate change is shifting flowering time in complex ways, even across local spatial gradients.

Projected impacts of climate change on the range and phenology of three culturally-important shrub species.

Climate change is shifting both the habitat suitability and the timing of critical biological events, such as flowering and fruiting, for plant species across the globe. Here, we ask how both the distribution and phenology of three food-producing shrubs native to northwestern North America might shift as the climate changes. To address this question, we compared gridded climate data with species location data to identify climate variables that best predicted the current bioclimatic niches of beaked hazelnut (Corylus cornuta), Oregon grape (Mahonia aquifolium), and salal (Gaultheria shallon). We also developed thermal-sum models for the timing of flowering and fruit ripening for these species. We then used multi-model ensemble future climate projections to estimate how species range and phenology may change under future conditions. Modelling efforts showed extreme minimum temperature, climate moisture deficit, and mean summer precipitation were predictive of climatic suitability across all three species. Future bioclimatic niche models project substantial reductions in habitat suitability across the lower elevation and southern portions of the species' current ranges by the end of the 21st century. Thermal-sum phenology models for these species indicate that flowering and the ripening of fruits and nuts will advance an average of 25 days by the mid-21st century, and 36 days by the late-21st century under a high emissions scenario (RCP 8.5). Future changes in the climatic niche and phenology of these important food-producing species may alter trophic relationships, with cascading impacts on regional ecosystems.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Effectiveness of winter temperatures for satisfying chilling requirements for reproductive budburst of red alder (Alnus rubra).

BACKGROUND: Experiencing an adequate amount of cold temperatures over winter is necessary for many temperate tree species to break dormancy and flower in spring. Thus, changes in winter and spring temperatures associated with climate change may influence when trees break dormancy and flower in the future. There have been several experimental studies that have quantified the effectiveness of cold temperatures for chilling requirements for vegetative budburst of temperate trees; however, there are few experimental studies addressing the chilling requirements for reproductive budburst of trees, as it is difficult to place reproductively mature trees in temperature-controlled environments. METHODS: To identify how changing temperatures associated with climate change may impact reproductive phenology, we completed a temperature-controlled growth chamber experiment using cuttings of reproductive branches of red alder (Alnus rubra), one of the most widespread hardwood tree species of the Pacific Northwest, USA. The purpose of this study was to examine how colder (4 °C) and warmer (9 °C) winter temperature regimes influenced the timing of reproductive budburst of red alder cuttings in spring. We also compared the date of budburst of cuttings to that of branches from intact trees. RESULTS: We found that cuttings flowered earlier after pretreatment with a 4 °C winter temperature regime than after a 9 °C winter temperature regime. We found no significant differences between the timing of male budburst of cuttings exposed to ambient conditions compared to male budburst of branches from intact trees. We used our experimental data to estimate a "possibility-line" that shows the accumulated chilling and forcing temperatures necessary prior to reproductive budburst of red alder. DISCUSSION: This study provides a preliminary indication that warmer winters with climate change may not be as effective as colder winters for satisfying chilling temperature requirements of a Northwest hardwood tree species.

Interspecific Plant Interactions Reflected in Soil Bacterial Community Structure and Nitrogen Cycling in Primary Succession.

Past research demonstrating the importance plant-microbe interactions as drivers of ecosystem succession has focused on how plants condition soil microbial communities, impacting subsequent plant performance and plant community assembly. These studies, however, largely treat microbial communities as a black box. In this study, we sought to examine how emblematic shifts from early successional Alnus viridus ssp. sinuata (Sitka alder) to late successional Picea sitchensis (Sitka spruce) in primary succession may be reflected in specific belowground changes in bacterial community structure and nitrogen cycling related to the interaction of these two plants. We examined early successional alder-conditioned soils in a glacial forefield to delineate how alders alter the soil microbial community with increasing dominance. Further, we assessed the impact of late-successional spruce plants on these early successional alder-conditioned microbiomes and related nitrogen cycling through a leachate addition microcosm experiment. We show how increasingly abundant alder select for particular bacterial taxa. Additionally, we found that spruce leachate significantly alters the composition of these microbial communities in large part by driving declines in taxa that are enriched by alder, including bacterial symbionts. We found these effects to be spruce specific, beyond a general leachate effect. Our work also demonstrates a unique influence of spruce on ammonium availability. Such insights bolster theory relating the importance of plant-microbe interactions with late-successional plants and interspecific plant interactions more generally.