skip to main content


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

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 can have consequences for herbivores and forage quality, developing mismatches could also influence other traits of plants, such as above‐ and below‐ground biomass and the type of reproduction, that are often not investigated.

In coastal western Alaska, we conducted a 3‐year factorial experiment that simulated scenarios of phenological mismatch by manipulating the start of the growing season (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.

After 3 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 below‐ground biomass, stem height, and the number of tillers; while late season grazing increased the number of inflorescences and standing dead biomass. Therefore, an advanced growing season and late grazing had similar directional effects on most plant traits, but a 3‐week delay in grazing had an impact on traits 3–5 times greater than a similarly timed shift in the advancement of spring. 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.

Synthesis. Factors affecting the timing of migration have a larger influence than earlier springs on an important forage species 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 below‐ground biomass and sexual reproduction of the often‐clonally reproducingC. subspathacea. Finally, the implications of mismatch may be difficult to predict because some variables required successive years of mismatch to respond.

 
more » « less
Award ID(s):
1304523 1633756
PAR ID:
10447387
Author(s) / Creator(s):
 ;  ;  ;  ;  ;  ;
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Journal of Ecology
Volume:
107
Issue:
5
ISSN:
0022-0477
Page Range / eLocation ID:
p. 2503-2518
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    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 CO2exchange 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 CO2uptake 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−1toward a greater CO2sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO2uptake. 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.

     
    more » « less
  2. Abstract

    A combination of theory and experiments predicts that increasing soil nutrients will modify herbivore and microbial impacts on ecosystem carbon cycling.

    However, few studies of herbivores and soil nutrients have measured both ecosystem carbon fluxes and carbon pools. Even more rare are studies manipulating microbes and nutrients that look at ecosystem carbon cycling responses.

    We added nutrients to a long‐term, experiment manipulating foliar fungi, soil fungi, mammalian herbivores and arthropods in a low fertility grassland. We measured gross primary production (GPP), ecosystem respiration (ER), net ecosystem exchange (NEE) and plant biomass throughout the growing season to determine how nutrients modify consumer impacts on ecosystem carbon cycling.

    Nutrient addition increased above‐ground biomass and GPP, but not ER, resulting in an increase in ecosystem carbon uptake rate. Reducing foliar fungi and arthropods increased plant biomass. Nutrients amplified consumer effects on plant biomass, such that arthropods and foliar fungi had a threefold larger impact on above‐ground biomass in fertilized plots.

    Synthesis. Our work demonstrates that throughout the growing season soil resources modify carbon uptake rates as well as animal and fungal impacts on plant biomass production. Taken together, ongoing nutrient pollution may increase ecosystem carbon uptake and drive fungi and herbivores to have larger impacts on plant biomass production.

     
    more » « less
  3. Abstract

    Given the current rates of climate change, with associated shifts in herbivore population densities, understanding the role of different herbivores in ecosystem functioning is critical for predicting ecosystem responses. Here, we examined how migratory geese and resident, non‐migratory reindeer—two dominating yet functionally contrasting herbivores—control vegetation and ecosystem processes in rapidly warming Arctic tundra.

    We collected vegetation and ecosystem carbon (C) flux data at peak plant growing season in the two longest running, fully replicated herbivore removal experiments found in high‐Arctic Svalbard. Experiments had been set up independently in wet habitat utilised by barnacle geeseBranta leucopsisin summer and in moist‐to‐dry habitat utilised by wild reindeerRangifer tarandus platyrhynchusyear‐round.

    Excluding geese induced vegetation state transitions from heavily grazed, moss‐dominated (only 4 g m−2of live above‐ground vascular plant biomass) to ungrazed, graminoid‐dominated (60 g m−2after 4‐year exclusion) and horsetail‐dominated (150 g m−2after 15‐year exclusion) tundra. This caused large increases in vegetation C and nitrogen (N) pools, dead biomass and moss‐layer depth. Alterations in plant N concentration and CN ratio suggest overall slower plant community nutrient dynamics in the short‐term (4‐year) absence of geese. Long‐term (15‐year) goose removal quadrupled net ecosystem C sequestration (NEE) by increasing ecosystem photosynthesis more than ecosystem respiration (ER).

    Excluding reindeer for 21 years also produced detectable increases in live above‐ground vascular plant biomass (from 50 to 80 g m−2; without promoting vegetation state shifts), as well as in vegetation C and N pools, dead biomass, moss‐layer depth and ER. Yet, reindeer removal did not alter the chemistry of plants and soil or NEE.

    Synthesis. Although both herbivores were key drivers of ecosystem structure and function, the control exerted by geese in their main habitat (wet tundra) was much more pronounced than that exerted by reindeer in their main habitat (moist‐to‐dry tundra). Importantly, these herbivore effects are scale dependent, because geese are more spatially concentrated and thereby affect a smaller portion of the tundra landscape compared to reindeer. Our results highlight the substantial heterogeneity in how herbivores shape tundra vegetation and ecosystem processes, with implications for ongoing environmental change.

     
    more » « less
  4. Abstract

    Changes in seasonality associated with climate warming (e.g. temperature, growing season duration) are likely to alter invertebrate prey biomass and availability in aquatic ecosystems through direct and indirect influences on physiology and phenology, particularly in arctic lakes. However, despite warmer thermal regimes, photoperiod will remain unchanged such that potential shifts resulting from longer and warmer growing seasons could be limited by availability of sunlight, especially at lower trophic levels. Thus, a better understanding of warming effects on invertebrate prey throughout the growing season (e.g. early, peak, late) is important to understand arctic lake food‐web dynamics in a changing climate.

    Here, we use a multifaceted approach to evaluate prey availability to predators in lakes of arctic Alaska. In a laboratory mesocosm experiment, we measured different metrics of abundance for snails (Lymnaea elodes) and zooplankton (Daphnia middendorffiana) across three time periods (early, mid‐ and late growing season) and across three temperature and photoperiod treatments (control, increased temperature and increased temperature × photoperiod). Additionally, we used generalised additive models and generalised additive mixed‐effects models to relate long‐term empirical observations of zooplankton biomass (1983–2015) to observed temperature regimes in an arctic lake. We then simulated zooplankton biomass for the warmest temperature observations across the growing season to inform likely zooplankton biomass regimes under future change.

    We observed variable responses by snails and zooplankton across experiments and treatments. Early in the growing season, snail development was accelerated at multiple life stages (e.g. egg and juvenile). In mid‐season, in accordance with warmer temperatures, we observed significantly increasedDaphniaabundances. However, in the late season,Daphniaappeared to be limited by photoperiod. Confirming our experimental results, our models of zooplankton biomass showed an increase of nearly 20% in warmer years. Further, these model estimates could be conservative as the consumptive demand of fishes may increase in warmer years as well.

    Overall, our results highlight the importance of interactive effects of temperature and seasonality. Based primarily on temperature, we can readily predict the response of fish metabolism in warmer temperatures. However, in this context, we generally require a better understanding of climate‐driven responses of important invertebrate prey resources. Our results suggest invertebrate prey biomass and availability are likely to respond positively with climate change based on temperature and seasonality, as well as proportionally to the metabolic requirements of fish predators. While further research is necessary to understand how other food‐web components will respond climate change, our findings suggest that the fish community at the top of arctic lake food webs will have adequate prey base in a warming climate.

     
    more » « less
  5. Abstract

    Herbivory is a key process structuring vegetation in savannas, especially in Africa where large mammal herbivore communities remain intact. Exclusion experiments consistently show that herbivores impact savanna vegetation, but effect size variation has resisted explanation, limiting our understanding of the past, present and future roles of herbivory in savanna ecosystems.

    Synthesis of vegetation responses to herbivore exclusion shows that herbivory decreased grass abundance by 57.0% and tree abundance by 30.6% across African savannas.

    The magnitude of herbivore exclusion effects scaled with herbivore abundance: more grazing herbivores resulted in larger grass responses and more browsing herbivores in larger tree responses. However, existing experiments are concentrated in semi‐arid savannas (400–800‐mm rainfall) and soils data are mostly lacking, which makes disentangling environmental constraints a challenge and priority for future research.

    Observed herbivore impacts were ~2.1× larger than existing estimates modelled based on consumption. Wildlife metabolic rates may be higher than are usually used for estimating consumption, which offers one clear avenue for reconciling estimated herbivore consumption with observed herbivore impacts. Plant‐soil feedbacks, plant community composition, and the phenological or demographic timing of herbivory may also influence vegetation productivity, thereby magnifying herbivore impacts.

    Because herbivore abundance so closely predicts vegetation impact, changes in herbivore abundance through time are likely predictive of the past and future of their impacts. Grazer diversity in Africa has declined from its peak 1 million years ago and wild grazer abundance has declined historically, suggesting that grazing likely had larger impacts in the past than it does today.

    Current wildlife impacts are dominated by small‐bodied mixed feeders, which will likely continue into the future, but the magnitude of top‐down control may also depend on changing climate, fire and atmospheric CO2.

    Synthesis. Herbivore biomass determines the magnitude of their impacts on savanna vegetation, with effect sizes based on direct observation that outstrip existing modelled estimates across African savannas. Findings suggest substantial ecosystem impacts of herbivory and allow us to generate evidence‐based hypotheses of the past and future impacts of herbivores on savanna vegetation.

     
    more » « less