Multiple resource limitation of dryland soil microbial carbon cycling on the Colorado Plateau.

Understanding interactions among biogeochemical cycles is increasingly important as anthropogenic alterations of global climate and of carbon (C), nitrogen (N), and phosphorus (P) cycles interactively affect the Earth system. Ecosystem processes in the dryland biome, which makes up over 40% of Earth's terrestrial surface, are often distinctively sensitive to small changes in resource availability, likely because levels of many resources are low. However, data also suggest that simultaneous changes in the availability of multiple resources may be necessary to affect a response in these low-resource systems, offering an opportunity to test patterns and controls of co-limitation, serial limitation, and individual limitation in soil environments. While drylands may play a governing role in key aspects of Earth's C cycle, and while an improved understanding of resource limitation could substantially improve our forecasts of dryland responses to change, our understanding of interacting controls on soil C cycle processes remains notably poor in these dry systems. Here, we address multiple fundamental hypotheses of resource controls over ecosystem function to test how water, C, N, and P regulate soil C cycling individually and interactively in a dryland ecosystem on the Colorado Plateau. Using a series of laboratory incubations, we found that, while water, C, and N limited C cycling through serial limitation, water alone resulted in an extremely small respiratory response from target organisms, whereas water + C resulted in a dramatic increase in soil C cycling, suggesting a degree of functional co-limitation. Nitrogen additions alone resulted in no changes to soil C cycling, but when N was added in concert with water and C, N greatly increased soil C cycling rates relative to additions of water and C without N. Phosphorus additions had no effect on the C cycle either alone or synergistically. These patterns were consistent with the stoichiometry of the system and interactions among resources were surprising in ways that inform our understanding of critical theories in ecology, such as the Transient Maxima Hypothesis, supporting the suggestion that multiple resource limitation explains pulse-dynamic C cycling in drylands better than water limitation alone.

Migratory goose arrival time plays a larger role in influencing forage quality than advancing springs in an Arctic coastal wetland.

With warmer springs, herbivores migrating to Arctic breeding grounds may experience phenological mismatches between their energy demands and the availability of high quality forage. Yet, how the timing of the start of the season and herbivore arrival influences forage quality is often unknown. In coastal western Alaska, approximately one million migratory geese arrive each spring to breed, where foliar %N and C:N ratios are linked to gosling survival and population growth. We conducted a three-year experiment where we manipulated the start of the growing season using warming chambers and grazing times using captive Pacific black brant (Branta bernicla nigricans) to examine how the timing of these events influences the quality of an important forage species. Our results suggest that grazing timing plays a much greater role than an advanced growing season in determining forage quality. All top models included grazing timing, and suggested that compared to typical grazing timing, early grazing significantly reduced foliar %C by 6% and C:N ratios by 16%, while late goose grazing significantly reduced foliar %N by 15% and increased foliar C:N ratios by 21%. While second-ranking top models included the effect of season, the advanced growing season effect was not significant and only reduced %N by 4%, increased %C by <1%, and increased C:N ratios by 5% compared to an ambient growing season. In summary, in years where geese arrive early, they will consume higher quality forage when they arrive and throughout the season, while in years that geese arrive late they will consume lower quality forage when they arrive and for the remainder of the season. When the growing season starts has only a minor influence on this pattern. Our findings suggest that cues determining migration and arrival times to breeding areas are important factors influencing forage quality for geese in western Alaska.

Delayed herbivory by migratory geese increases summer-long CO2 uptake in coastal western Alaska.

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to "phenological mismatches" as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three-year experiment to examine the consequences for CO2 exchange of advanced spring green-up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green-up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO2 uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green-up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m-2  s-1 toward a greater CO2 sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO2 uptake. Importantly, while the direct effect of climate warming on phenology of green-up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition.

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra- and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five-fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra- and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.