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Karen H. Beard

Publications and source records attributed to Karen H. Beard.

13 recordsLinked to original sources

Growth form and lifespan of herbaceous species mediate the role of traits in short-term drought response

Increased climate variability is expected to intensify short-term drought events. Plants have evolved stress tolerance strategies involving trade-offs in resource conservation, mycorrhizal collaboration and plant size, yet how these strategies promote drought resistance across different herbaceous plant groups remains unknown. Leveraging 63 globally distributed grassland and shrubland sites from the International Drought Experiment, we identified plant traits linked to drought resistance in 661 populations of 421 species after 1 year of extreme drought. We assessed how traits, site precipitation and drought severity affected cover change across growth forms and lifespans, and how trait–environment interactions influenced drought resistance. Across all species, leaf N (an acquisitive trait) was associated with drought resistance, whereas in forbs, drought resistance was also associated with a conservative root trait and plant size. In addition, interactions among traits mediated drought resistance; root traits predicted performance only in concert with other traits. Environmental variables influenced trait effects on drought resistance, notably for annuals in wetter sites, suggesting that drought-escape strategies in annuals may be advantageous only under mild stress. Our study highlights variability in traits that predict drought resistance across herbaceous plant groups, emphasizing the importance of species context, environmental stress and the selection of traits in research and management.

Nature Ecology and Evolution

Drought intensity and duration interact to magnify losses in primary productivity

As droughts become longer and more intense, impacts on terrestrial primary productivity are expected to increase progressively. Yet, some ecosystems appear to acclimate to multiyear drought, with constant or diminishing reductions in productivity as drought duration increases. We quantified the combined effects of drought duration and intensity on aboveground productivity in 74 grasslands and shrublands distributed globally. Ecosystem acclimation with multiyear drought was observed overall, except when droughts were extreme (i.e., ≤1-in-100-year likelihood of occurrence). Productivity losses after four consecutive years of extreme drought increased by ~2.5-fold compared with those of the first year. These results portend a foundational shift in ecosystem behavior if drought duration and intensity increase, from maintenance of reduced functioning over time to progressive and profound losses of productivity when droughts are extreme.

Science

Aridity drives the response of soil total and particulate organic carbon to drought in temperate grasslands and shrublands

The increasing prevalence of drought events in grasslands and shrublands worldwide potentially has impacts on soil organic carbon (SOC). We leveraged the International Drought Experiment to study how SOC, including particulate organic carbon (POC) and mineral-associated organic carbon (MAOC) concentrations, responds to extreme drought treatments (1-in-100-year) for 1 to 5 years at 19 sites worldwide. In more mesic areas (aridity index > 0.65), SOC and POC concentrations decreased by 7.9% (±3.9) and 15.9% (±6.2) with drought, respectively, but there were no impacts on MAOC concentrations. However, drought had no impact on SOC, POC, or MAOC concentrations in drylands (aridity index < 0.65). The response of SOC to drought varied along an aridity gradient, concomitant with interannual precipitation variability and standing SOC concentration gradients. These findings highlight the differing response magnitudes of POC and MAOC concentrations to drought and the key regulating role of aridity.

Science Advances

Extreme drought impacts have been underestimated in grasslands and shrublands globally

Drought has well-documented societal and economic consequences. Climate change is expected to intensify drought to even more extreme levels, but because such droughts have been historically rare, their impact on ecosystem functioning is not well known. We experimentally imposed the most frequent type of intensified drought—one that is ~1 y in duration—at 100 grassland and shrubland sites distributed across six continents. We found that loss of aboveground plant growth, a key measure of ecosystem function, was 60% greater when short-term drought was extreme (≤1-in-100-y historical occurrence). This drought-induced loss in function greatly exceeds previously reported losses for grasslands and shrublands, suggesting that the global impacts of projected increases in drought severity have been substantially underestimated.

Proceedings of the National Academy of Sciences

Sex‐related differences in aging rate are associated with sex chromosome system in amphibians

Sex‐related differences in mortality are widespread in the animal kingdom. Although studies have shown that sex determination systems might drive lifespan evolution, sex chromosome influence on aging rates have not been investigated so far, likely due to an apparent lack of demographic data from clades including both XY (with heterogametic males) and ZW (heterogametic females) systems. Taking advantage of a unique collection of capture–recapture datasets in amphibians, a vertebrate group where XY and ZW systems have repeatedly evolved over the past 200 million years, we examined whether sex heterogamy can predict sex differences in aging rates and lifespans. We showed that the strength and direction of sex differences in aging rates (and not lifespan) differ between XY and ZW systems. Sex‐specific variation in aging rates was moderate within each system, but aging rates tended to be consistently higher in the heterogametic sex. This led to small but detectable effects of sex chromosome system on sex differences in aging rates in our models. Although preliminary, our results suggest that exposed recessive deleterious mutations on the X/Z chromosome (the “unguarded X/Z effect”) or repeat‐rich Y/W chromosome (the “toxic Y/W effect”) could accelerate aging in the heterogametic sex in some vertebrate clades.

Evolution

Early goose arrival increases soil nitrogen availability more than an advancing spring in coastal western Alaska

An understudied aspect of climate change-induced phenological mismatch is its effect on ecosystem functioning, such as nitrogen (N) cycling. Migratory herbivore arrival time may alter N inputs and plant–herbivore feedbacks, whereas earlier springs are predicted to increase N cycling rates through warmer temperatures. However, the relative importance of these shifts in timing and how they interact to affect N cycling are largely unknown. We conducted a 3-year factorial experiment in coastal western Alaska that simulated different timings of Pacific black brant ( Branta bernicla nigricans ) arrival (3 weeks early, typical, 3 weeks late, or no-grazing) and the growing season (ca. 3 weeks advanced and ambient) on adsorbed and mobile inorganic (NH 4 + –N, NO 3 − –N) and mobile organic N (amino acid) pools. Early grazing increased NH 4 + –N, NO 3 − –N, and amino acids by 103%, 119%, and 7%, respectively, whereas late grazing reduced adsorbed NH 4 + –N and NO 3 − –N by 16% and 17%, respectively. In comparison, the advanced growing season increased mobile NH 4 + –N by 26%. The arrival time by geese and the start of the season did not interact to influence soil N availability. While the onset of spring in our system is advancing at twice the rate of migratory goose arrival, earlier goose migration is likely to be more significant than the advances in springs in influencing soil N, although both early goose arrival and advanced springs are likely to increase N availability in the future. This increase in soil N resources can have a lasting impact on plant community composition and productivity in this N-limited ecosystem.

Coastal western Alaska

Cloud cover and delayed herbivory relative to timing of spring onset interact to dampen climate change impacts on net ecosystem exchange in a coastal Alaskan wetland

Rapid warming in northern ecosystems over the past four decades has resulted in earlier spring, increased precipitation, and altered timing of plant–animal interactions, such as herbivory. Advanced spring phenology can lead to longer growing seasons and increased carbon (C) uptake. Greater precipitation coincides with greater cloud cover possibly suppressing photosynthesis. Timing of herbivory relative to spring phenology influences plant biomass. None of these changes are mutually exclusive and their interactions could lead to unexpected consequences for Arctic ecosystem function. We examined the influence of advanced spring phenology, cloud cover, and timing of grazing on C exchange in the Yukon–Kuskokwim Delta of western Alaska for three years. We combined advancement of the growing season using passive-warming open-top chambers (OTC) with controlled timing of goose grazing (early, typical, and late season) and removal of grazing. We also monitored natural variation in incident sunlight to examine the C exchange consequences of these interacting forcings. We monitored net ecosystem exchange of C (NEE) hourly using an autochamber system. Data were used to construct daily light curves for each experimental plot and sunlight data coupled with a clear-sky model was used to quantify daily and seasonal NEE over a range of incident sunlight conditions. Cloudy days resulted in the largest suppression of NEE, reducing C uptake by approximately 2 g C m −2 d −1 regardless of the timing of the season or timing of grazing. Delaying grazing enhanced C uptake by approximately 3 g C m −2 d −1 . Advancing spring phenology reduced C uptake by approximately 1.5 g C m −2 d −1 , but only when plots were directly warmed by the OTCs; spring advancement did not have a long-term influence on NEE. Consequently, the two strongest drivers of NEE, cloud cover and grazing, can have opposing effects and thus future growing season NEE will depend on the magnitude of change in timing of grazing and incident sunlight.

Alaska

Phenological mismatch between season advancement and migration timing alters Arctic plant traits

1. Climate change is creating phenological mismatches between herbivores and their plant resources throughout the Arctic. While advancing growing seasons and changing arrival times of migratory herbivores has been shown to have consequences for herbivores and forage quality, developing mismatches are also likely to influence other traits of plants, such as above- and belowground biomass and the type of reproduction, that are often not investigated. 2. In coastal western Alaska, we conducted a three-year factorial experiment that simulated scenarios of phenological mismatch by manipulating the start of the growing season (ca. 3-weeks early and ambient) and grazing times (3-weeks early, typical, 3-weeks late, or no-grazing) of Pacific black brant (Branta bernicla nigricans), to examine how the timing of these events influence a primary goose forage species, Carex subspathacea. 3. After three years, an advanced growing season compared to a typical growing season increased stem heights, standing dead biomass, and the number of inflorescences. Early season grazing compared to typical season grazing reduced above- and belowground biomass, stem height, and the number of tillers; while late season grazing increased the number of inflorescences and standing dead biomass by year 3. Therefore, an advanced growing season and late grazing had comparable directional effects on most plant traits, but a 3-week delay in grazing had an impact on vegetation traits 3 to 5 times greater than a similar shift in advancement of spring conditions. In addition, changes in response to treatments for some variables, such as the number of inflorescences, were not measurable until the second year of the experiment, while other variables, such as root productivity and number of tillers, changed the direction of their responses to treatments over time. 4. Synthesis: Factors influencing the timing of migration have a larger influence on an important forage species than earlier springs in the breeding and rearing habitats of Pacific black brant. The phenological mismatch prediction for this site of earlier springs and later goose arrival will likely increase above- and belowground biomass and sexual reproduction of C. subspathacea. Implications for mismatch may be difficult to predict because some variables required successive years of mismatch to respond.

Journal of Ecology

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. However, the timing of high quality forage relative to the timing of grazing 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. Both 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 the second-ranking top model included the effect of season, the advanced growing season 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.

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 CO 2 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 CO 2 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 CO 2 sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO 2 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.

Alaska

Change in dominance determines herbivore effects on plant biodiversity

Herbivores alter plant biodiversity (species richness) in many of the world’s ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis—that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.

Nature Ecology & Evolution

Phenological mismatch in coastal western Alaska may increase summer season greenhouse gas uptake

High latitude ecosystems are prone to phenological mismatches due to climate change- driven advances in the growing season and changing arrival times of migratory herbivores. These changes have the potential to alter biogeochemical cycling and contribute to feedbacks on climate change by altering greenhouse gas (GHG) emissions of carbon dioxide (CO 2 ), methane (CH 4 ) and nitrous oxide (N 2 O) through large regions of the Arctic. Yet the effects of phenological mismatches on gas fluxes are currently unexplored. We used a three-year field experiment that altered the start of the growing season and timing of grazing to investigate how phenological mismatch affects GHG exchange. We found early grazing increased mean GHG emission to the atmosphere despite lower CH 4 emissions due to grazing-induced changes in vegetation structure that increased uptake of CO 2 . In contrast, late grazing reduced GHG emissions because greater plant productivity led to an increase in CO 2 uptake that overcame the increase in CH 4 emission. Timing of grazing was an important control on both CO 2 and CH 4 emissions, and net GHG exchange was the result of opposing fluxes of CO 2 and CH 4 . N 2 O played a negligible role in GHG flux. Advancing the growing season had a smaller effect on GHG emissions than changes to timing of grazing in this study. Our results suggest that a phenological mismatch that delays timing of grazing relative to the growing season, a change which is already developing along in western coastal Alaska, will reduce GHG emissions to the atmosphere through increased CO 2 uptake despite greater CH 4 emissions.

Environmental Research Letters

Rodent-Mediated Interactions Among Seed Species of Differing Quality in a Shrubsteppe Ecosystem

Interactions among seeds, mediated by granivorous rodents, are likely to play a strong role in shrubsteppe ecosystem restoration. Past studies typically consider only pairwise interactions between preferred and less preferred seed species, whereas rangeland seedings are likely to contain more than 2 seed species, potentially leading to complex interactions. We examined how the relative proportion of seeds in a 3-species polyculture changes rodent seed selectivity (i.e., removal) and indirect interactions among seeds. We presented 2 rodent species, Peromyscus maniculatus (deer mice) and Perognathus parvus (pocket mice), in arenas with 3-species seed mixtures that varied in the proportion of a highly preferred, moderately preferred, and least preferred seed species, based on preferences determined in this study. We then conducted a field experiment in a pocket mouse—dominated ecosystem with the same 3-species seed mixtures in both “treated” (reduced shrub and increased forb cover) and “untreated” shrubsteppe. In the arena experiment, we found that rodents removed more of the highly preferred seed when the proportions of all 3 seeds were equal. Moderately preferred seeds experienced increased removal when the least preferred seed was in highest proportion. Removal of the least preferred seed increased when the highly preferred seed was in highest proportion. In the field experiment, results were similar to those from the arena experiment and did not differ between treated and untreated shrubsteppe areas. Though our results suggest that 3-species mixtures induce complex interactions among seeds, managers applying these results to restoration efforts should carefully consider the rodent community present and the potential fate of removed seeds.

Utah