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Observations of changes in phenology have provided some of the strongest signals of the effects of climate change on terrestrial ecosystems. The International Tundra Experiment (ITEX), initiated in the early 1990s, established a common protocol to measure plant phenology in tundra study areas across the globe. Today, this valuable collection of phenology measurements depicts the responses of plants at the colder extremes of our planet to experimental and ambient changes in temperature over the past decades. The database contains 150,434 phenology observations of 278 plant species taken at 28 study areas for periods of 1 to 26 years. Here we describe the full dataset to increase the visibility and use of these data in global analyses, and to invite phenology data contributions from underrepresented tundra locations. Portions of this tundra phenology database have been used in three recent syntheses, some datasets are expanded, others are from entirely new study areas, and the entirety of these data are now available at the Polar Data Catalogue., Arctic Science, 8 (3), ISSN:2368-7460

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.

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., Nature Communications, 12 (1), ISSN:2041-1723

National Park Service (NPS) units in the northern Great Plains (NGP) were established to preserve and interpret the history of America, protect and showcase unusual geology and paleontology, and provide a home for vanishing large wildlife. A unifying feature among these national parks, monuments, and historic sites is mixed-grass prairie, which not only provides background scenery but is the very foundation of many park missions. As recognition of the prairie’s importance to park fundamental resources and values has grown, so too has the realization that invasive plants threaten these values by reducing native species diversity, altering food webs, and marring the visitor experience. Parks manage invasive species despite uncertainties in treatment effectiveness because management cannot wait for research to provide definitive answers. Under these circumstances, adaptive management (AM) is an appropriate approach. In the NGP, we formed a collaborative adaptive vegetation management team to apply AM towards reducing invasive species (with a focus on exotic annual grasses) and improving native vegetation conditions. In our AM framework, the team uses a Bayesian model built from NPS Inventory & Monitoring and Fire Effects monitoring data and experimental results to predict the effects of management actions on park management units, according to those units’ vegetation condition and management history. These predictions inform management decisions, which are then applied.

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.

Woody plants are encroaching into many herbaceous-dominated communities across the globe, including arctic and alpine tundra. Quantifying the encroachment rate, testing which factors contribute to encroachment, and determining how encroachment is taking place and in which community types encroachment is occurring are essential for predicting shifts in tundra vegetation and carbon (C) storage. We examined willow cover changes from 1946 to 2008 in 18 ha of alpine tundra in Colorado using aerial photographs. We linked this pattern of change with experimental assessment of the effects of increasing summer temperatures, winter precipitation, and nitrogen (N) deposition—factors that this region has experienced over this period—on willow growth and survival. Shrub cover expanded by 441% over 62 years and is increasing at an exponential rate, corresponding to increases in C storage of 137 kg ha-1. Nitrogen and temperature facilitate willow growth and snow increases survival, although N and the combinatio...

Two sources of complexity make predicting plant community response to global change particularly challenging. First, realistic global change scenarios involve multiple drivers of environmental change that can interact with one another to produce non-additive effects. Second, in addition to these direct effects, global change drivers can indirectly affect plants by modifying species interactions. In order to tackle both of these challenges, we propose a novel population modeling approach, requiring only measurements of abundance and climate over time. To demonstrate the applicability of this approach, we model population dynamics of eight abundant plant species in a multifactorial global change experiment in alpine tundra where we manipulated nitrogen, precipitation, and temperature over 7 years. We test whether indirect and interactive effects are important to population dynamics and whether explicitly incorporating species interactions can change predictions when models are forecast under future climate change scenarios. For three of the eight species, population dynamics were best explained by direct effect models, for one species neither direct nor indirect effects were important, and for the other four species indirect effects mattered. Overall, global change had negative effects on species population growth, although species responded to different global change drivers, and single-factor effects were slightly more common than interactive direct effects. When the fitted population dynamic models were extrapolated under changing climatic conditions to the end of the century, forecasts of community dynamics and diversity loss were largely similar using direct effect models that do not explicitly incorporate species interactions or best-fit models; however, inclusion of species interactions was important in refining the predictions for two of the species. The modeling approach proposed here is a powerful way of analyzing readily available datasets which should be added to our toolbox to tease apart complex drivers of global change.

Two Eurasian invasive annual brome grasses, cheatgrass (Bromus tectorum) and Japanese brome (Bromus japonicus), are well known for their impact in steppe ecosystems of the western United States where these grasses have altered fire regimes, reduced native plant diversity and abundance, and degraded wildlife habitat. Annual bromes are also abundant in the grasslands of the Northern Great Plains (NGP), but their impact and ecology are not as well studied. It is unclear whether the lessons learned from the steppe will translate to the mixed-grass prairie where native plant species are adapted to frequent fires and grazing. Developing a successful annual brome management strategy for National Park Service units and other NGP grasslands requires better understanding of (1) the impact of annual bromes on grassland condition; (2) the dynamics of these species through space and time; and (3) the relative importance of environmental factors within and outside managers' control for these spatiotemporal dynamics. Here, we use vegetation monitoring data collected from 1998 to 2015 in 295 sites to relate spatiotemporal variability of annual brome grasses to grassland composition, weather, physical environmental characteristics, and ecological processes (grazing and fire). Concern about the impact of these species in NGP grasslands is warranted, as we found a decline in native species richness with increasing annual brome cover. Annual brome cover generally increased over the time of monitoring but also displayed a 3- to 5-yr cycle of reduction and resurgence. Relative cover of annual bromes in the monitored areas was best predicted by park unit, weather, extant plant community, slope grade, soil composition, and fire history. We found no evidence that grazing reduced annual brome cover, but this may be due to the relatively low grazing pressure in our study. By understanding the consequences and patterns of annual brome invasion, we will be better able to preserve and restore these grassland landscapes for future generations.

Thousands of years before Rocky Mountain National Park was established, people came to the area to experience its beauty and its wildlife and, for many, to re-create themselves. And it remains so today. How do we protect this special place so future generations may have the same opportunities?

Modified environmental conditions are driving phenological changes in ecosystems around the world. Many plants have already responded to warmer temperatures by flowering earlier and sustaining longer periods of growth. Changes in other environmental factors, like precipitation and atmospheric nitrogen (N) deposition, may also influence phenology but have been less studied. Alpine plants may be good predictors of phenological response patterns because environmental changes are amplified in mountain ecosystems and extreme conditions may make alpine plants particularly sensitive to changes in limiting factors like precipitation, temperature, and N. We tested the effects of increased snowpack, temperature, and N on alpine tundra plant phenology, using snow fence, open-top warming chamber, and N fertilization treatments at the Niwot Ridge Long Term Ecological Research (LTER) site. Flowering phenology of three abundant species was recorded during two growing seasons. Treatment responses varied among sp...

To understand the role biota play in resilience or vulnerability to environmental change, we investigated soil, plant, and microbial responses to a widespread environmental change, increased nitrogen (N). Our aim was to test the plant-soil threshold hypothesis: that changed biotic structure influences resilience to accumulated changes in N. For six years, we removed one of two codominant species, Geum rossii and Deschampsia caespitosa, in moist- meadow alpine tundra in Colorado, USA. We also manipulated nutrient availability by adding carbon (C) or N, separately and in combination with the species removals. Consistent with our hypothesis, Geum was associated with soil feedbacks that slowed rates of N cycling and Deschampsia with feedbacks that increased rates of N cycling. After a four- year initial resilience period, Geum dramatically declined (by almost 70%) due to increasing N availability. In contrast, Deschampsia abundance did not respond to changes in N supply; it only responded to the removal of Geum. Forbs and graminoids responded more positively to Deschampsia removal than to Geum removal, indicating stronger competitive effects by Deschampsia. The changed biotic interactions appear to have community-level consequences: after six years of Geum (but not Deschampsia) removal, evenness of the community declined by over 35%. Increased N affected the soil-microbial feedbacks, particularly in association with Geum. Microbial biomass N declined at higher N, as did the activities of two C-acquiring and one N- acquiring extracellular microbial enzymes. In the presence of Geum, N fertilization slowed the activity of phenol oxidase, a tannin-degrading enzyme, suggesting that microbes shift from degrading Geum-derived compounds. In the absence of Geum, acid phosphatase activity increased, suggesting increased phosphorus limitation in association with Deschampsia. With continued N deposition forecast for this system, these results suggest that initial resilience of Geum to increased N will be overwhelmed through elimination of microbial feedbacks. Once Geum declines, the loss will indirectly facilitate Deschampsia via competitive release. Because Deschampsia exerts strong competitive effects on subordinate species, increased Deschampsia abundance may be accompanied by a community-wide drop in diversity. We conclude that plant-soil feedbacks through the microbial community can influence vulnerability to exogenous changes in N and contribute to threshold dynamics.

While earlier studies on the process of invasion often focused on single factors or on the general explanation of ‘disturbance,’ recent work has attempted to move towards a more mechanistic understanding of the factors that promote plant community invasion. Manipulative experiments provide a means for discerning causal relationships and interactive effects of environmental factors in promoting invasion; such experiments have been conducted in a number of grassland and shrub ecosystems. This study extends multifactor manipulative experiments into forest communities to compare factors influencing early seedling establishment for native and invasive woody plants. In Long Island, NY, invasion patterns are correlated with forest community type (pine barrens or hardwood), light availability, and soil N and Ca. We conducted manipulative field experiments in two different years to determine the relative importance and interaction of experimental gaps and N and Ca addition in pine barrens and hardwood forests in promoting invasion. We used seedlings of seven common native and invasive species in the first experiment, and 16 native and invasive species paired phylogenetically in the second experiment. Light had the strongest effect on plant growth; all plants grew more in gaps. We found no difference in the average growth rates of native and invasive species. Invasives responded more to high resources than did natives, with highest relative growth rates in gaps in the more fertile soils of the hardwood forests. Opportunities for invasion may differ from year to year, with differential success of invaders only in some years and under some environmental conditions. Clearly, to understand the complex interactions between resources and invasion in forests will require many manipulative experiments across a range of environments and using suites of invasive and native species.

Numerous hypotheses suggest that natural enemies can influence the dynamics of biological invasions. Here, we use a group of 12 related native, invasive, and naturalized vines to test the relative importance of resistance and tolerance to herbivory in promoting biological invasions. In a field experiment in Long Island, New York, we excluded mammal and insect herbivores and examined plant growth and foliar damage over two growing seasons. This novel approach allowed us to compare the relative damage from mammal and insect herbivores and whether damage rates were related to invasion. In a greenhouse experiment, we simulated herbivory through clipping and measured growth response. After two seasons of excluding herbivores, there was no difference in relative growth rates among invasive, naturalized, and native woody vines, and all vines were susceptible to damage from mammal and insect herbivores. Thus, differential attack by herbivores and plant resistance to herbivory did not explain invasion success of these species. In the field, where damage rates were high, none of the vines were able to fully compensate for damage from mammals. However, in the greenhouse, we found that invasive vines were more tolerant of simulated herbivory than native and naturalized relatives. Our results indicate that invasive vines are not escaping herbivory in the novel range, rather they are persisting despite high rates of herbivore damage in the field. While most studies of invasive plants and natural enemies have focused on resistance, this work suggests that tolerance may also play a large role in facilitating invasions.

Author(s): Farrer, EC; Ashton, IW; Spasojevic, MJ; Fu, S; Gonzalez, DJX; Suding, KN | Abstract: Summary: Environmental change can affect species directly by altering their physical environment and indirectly by altering the abundance of interacting species. A key challenge at the interface of community ecology and conservation biology is to predict how direct and indirect effects combine to influence response in a changing environment. In particular, little is known about how direct and indirect effects on biodiversity develop over time or their potential to influence ecosystem function. We studied how nitrogen (N), winter precipitation (snow) and warming influenced diversity and ecosystem function over 6 years in alpine tundra. We used path analyses to partition direct effects of environmental manipulations from indirect effects due to changes in the abundance of two dominant plants. We hypothesize that (i) indirect effects will develop more slowly but will become stronger than direct effects over time and (ii) after 6 years, indirect effects will more strongly influence diversity while direct effects will influence ecosystem function. Indirect effects of N on diversity were consistently stronger than direct effects and actually developed quickly, prior to direct effects. Direct effects of snow on diversity were detected in year 2 but then subsequently were reversed, while indirect effects were detected in year 4 and grew stronger over time. Overall in year 6, indirect effects were much stronger than direct effects on diversity. Direct effects predominated for three of four ecosystem functions we measured (productivity, N mineralization, winter N availability). The only indirect effects we found were that N and snow indirectly affected microbial biomass N by influencing Geum abundance. Across all four ecosystem measures, indirect effects were infrequent and weaker than direct effects. Synthesis. Increasing indirect effects on diversity over time indicate that short-term experiments or monitoring of natural systems may underestimate the full magnitude of global change effects on plant communities. Moreover, explicitly accounting for changes in dominant plant abundance may be necessary for forecasting plant community response to environmental change. Conversely, weak indirect effects for ecosystem processes suggest that predicting ecosystem function without knowledge of plant responses to global change may be possible. Increasing indirect effects on diversity over time indicate that short-term experiments or monitoring of natural systems may underestimate the full magnitude of global change effects on plant communities. Explicitly accounting for changes in dominant plant abundance may be necessary for forecasting plant community response to environmental change. Conversely, weak indirect effects for ecosystem processes suggest that predicting ecosystem function without knowledge of plant responses to global change may be possible.

Invasive species can change decomposition rates within an ecosystem by changing the quality of the litter entering a system. It is not known, however, whether or not invasions can also change rates of decomposition irrespective of litter quality. We conducted an experiment to explore the differences in decomposition between leaf litter of native and exotic invasive woody plants and between invaded and uninvaded mesic hard- wood forests on Long Island, New York, USA. We evaluated the mass and nitrogen loss rates from leaf litter of four pairs of native and exotic woody species. Litter from the exotic species decomposed and released nitrogen significantly faster than litter from the native species. The largest differences in decomposition and nitrogen loss occurred between the invaded and uninvaded sites rather than between native and exotic species, with litter of all species types decomposing substantially faster in invaded sites. These results suggest that the invasion of exotic species into hardwood forests alters decomposition and nutrient cycling, irrespective of species-specific litter quality differences between natives and ex- otics.

Ecological stoichiometry, the study of the balance of energy and materials in living systems, may serve as a useful synthetic framework for evolutionary biology. Here, we review recent work that illustrates the power of a stoichiometric approach to evolution across multiple scales, and then point to important open questions that may chart the way forward in this new field. At the molecular level, stoichiometry links hereditary changes in the molecular composition of organisms to key phenotypic functions. At the level of evolutionary ecology, a simultaneous focus on the energetic and material underpinnings of evolutionary tradeoffs and transactions highlights the relationship between the cost of resource acquisition and the functional consequences of biochemical composition. At the macroevolutionary level, a stoichiometric perspective can better operationalize models of adaptive radiation and escalation, and elucidate links between evolutionary innovation and the development of global biogeochemical cycles. Because ecological stoichiometry focuses on the interaction of energetic and multiple material currencies, it should provide new opportunities for coupling evolutionary dynamics across scales from genomes to the biosphere.

We conducted single- and mixed-litter experiments in a hardwood forest in Long Island, New York, using leaf litter from phylogenetically paired native and invasive species. We selected long-established, abundant invasive species with wide-ranging distributions in the eastern United States that likely make substantial contributions to the litter pool of invaded areas. Overall, leaf litter from invasive species differed from native litter, though differences varied by phylogenetic grouping. Invasive litter had lower carbon:nitrogen ratios (30.9 ± 1.96 SE vs. 32.8 ± 1.36, P = 0.034) and invasive species lost 0.03 ± 0.007 g of nitrogen and had 23.4 ± 4.9 % of their starting mass remaining at the end of 1 year compared with a loss of 0.02 ± 0.003 g nitrogen and 31.1 ± 2.6 % mass remaining for native species. Mixing litter from two species did not alter decomposition rates when native species were mixed with other native species, or when invasive species were mixed with other invasive species. However, mixing litter of native and invasive species resulted in significantly less mass and nitrogen loss than was seen in unmixed invasive litter. Mixtures of native and invasive litter lost all but 47 ± 2.2 % of initial mass, compared to 37 ± 5.8 % for invasive litter and 50 ± 5.1 % for native litter. This non-additive effect of mixing native and invasive litter suggests that an additive model of metabolic characteristics may not suffice for predicting invasion impacts in a community context, particularly as invasion proceeds over time. Because the more rapid decomposition of invasive litter tends to slow to rates typical of native species when native and invasive litters are mixed together, there may be little impact of invasive species on nutrient cycling early in an invasion, when native leaf litter is abundant (providing litter deposition is the dominant control on nutrient cycling).

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.

Plant resource partitioning of chemical forms of nitrogen (N) may be an important factor promoting species coexistence in N-limited ecosystems. Since the microbial community regulates N-form transformations, plant partitioning of N may be related to plant-soil feedbacks. We conducted a (15)N tracer addition experiment to study the ability of two alpine plant species, Acomastylis rossii and Deschampsia caespitosa, to partition organic and inorganic forms of N. The species are codominant and associated with strong plant-soil feedbacks that affect N cycling. We manipulated interspecific interactions by removing Acomastylis or Deschampsia from areas where the species were codominant to test if N uptake patterns varied in the presence of the other species. We found that Deschampsia acquired organic and inorganic N more rapidly than Acomastylis, regardless of neighbor treatment. Plant N uptake-specifically ammonium uptake-increased with plant density and the presence of an interspecific neighbor. Interestingly, this change in N uptake was not in the expected direction to reduce niche overlap and instead suggested facilitation of ammonium use. To test if N acquisition patterns were consistent with plant-soil feedbacks, we also compared microbial rhizosphere extracellular enzyme activity in patches dominated by one or the other species and in areas where they grew together. The presence of both species was generally associated with increased rhizosphere extracellular enzyme activity (five of ten enzymes) and a trend towards increased foliar N concentrations. Taken together, these results suggest that feedbacks through the microbial community, either in response to increased plant density or specific plant neighbors, could facilitate coexistence. However, coexistence is promoted via enhanced resource uptake rather than reduced niche overlap. The importance of resource partitioning to reduce the intensity of competitive interactions might vary across systems, particularly as a function of plant-soil feedbacks.

Tramp ants are invading disturbed ecosystems worldwide, exterminating untold numbers of native species. They have even invaded Biosphere 2, a 1.28-hectare closed greenhouse structure built in the Arizona desert as a microcosm for studying ecological interactions and global change. Invertebrate surveys within Biosphere 2 from 1990 to 1997 have revealed dramatic changes in faunal composition, including an almost complete replacement of the ant fauna by a single tramp ant species. In 1990-91, surveys in Biosphere 2 found no one ant species dominant. By 1993, populations of the crazy ant, Paratrechina longicornis (Latreille), a tramp species not found in 1990-91, had increased to extremely high levels. In 1996, virtually all ants (>99.9%) coming to bait were P. longicornis. We observed P. longicornis foragers feeding almost exclusively on the sugary excretions (honeydew) produced by vast numbers of Homoptera, primarily scale insects and mealybugs, found on many of the plants. High densities of ants were associated with high densities of homopterans. In 1997, soil and litter surveys found that the only invertebrates thriving in Biosphere 2, besides P. longicornis and homopterans, were either species with effective defenses against ants (well-armored isopods and millipedes) or tiny subterranean species that can escape ant predation (mites, thief ants, and springtails). A convergent pattern of biodiversity occurs in disturbed tropical and subtropical ecosystems dominated by tramp ants.