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  • Article
    Model responses to CO(2) and warming are underestimated without explicit representation of Arctic small-mammal grazing
    (Ecological Society of America, 2021-10-17) Rastetter, Edward B. ; Griffin, Kevin L. ; Rowe, Rebecca J. ; Gough, Laura ; McLaren, Jennie ; Boelman, Natalie
    We use a simple model of coupled carbon and nitrogen cycles in terrestrial ecosystems to examine how “explicitly representing grazers” vs. “having grazer effects implicitly aggregated in with other biogeochemical processes in the model” alters predicted responses to elevated carbon dioxide and warming. The aggregated approach can affect model predictions because grazer-mediated processes can respond differently to changes in climate compared with the processes with which they are typically aggregated. We use small-mammal grazers in a tundra as an example and find that the typical three-to-four-year cycling frequency is too fast for the effects of cycle peaks and troughs to be fully manifested in the ecosystem biogeochemistry. We conclude that implicitly aggregating the effects of small-mammal grazers with other processes results in an underestimation of ecosystem response to climate change, relative to estimations in which the grazer effects are explicitly represented. The magnitude of this underestimation increases with grazer density. We therefore recommend that grazing effects be incorporated explicitly when applying models of ecosystem response to global change.
  • Preprint
    Species compositional differences on different-aged glacial landscapes drive contrasting responses of tundra to nutrient addition
    ( 2005-01-17) Hobbie, Sarah E. ; Gough, Laura ; Shaver, Gaius R.
    In the northern foothills of the Brooks Range, Alaska, moist non-acidic tundra dominates more recently deglaciated upland landscapes, whereas moist acidic tundra dominates older upland landscapes. In previous studies, experimental fertilization of moist acidic tussock tundra greatly increased the abundance and productivity of the deciduous dwarf shrub Betula nana. However, this species is largely absent from moist non-acidic tundra. These two common upland tundra community types exhibited markedly different responses to fertilization with nitrogen and phosphorus. In moist acidic tundra, cover of deciduous shrubs (primarily B. nana) increased after only 2 years, and by 4 years vascular biomass and above-ground net primary productivity (ANPP) had increased significantly, almost entirely because of Betula. In moist non-acidic tundra, both biomass and ANPP were again significantly greater, but no single species dominated the response to fertilization. Instead, the effect was due to a combination of several small, sometimes statistically non-significant responses by forbs, graminoids and prostrate deciduous shrubs. The different growth form and species' responses suggest that fertilization will cause carbon cycling through plant biomass to diverge in these two tundra ecosystems. Already, production of new stems by apical growth has increased relative to leaf production in acidic tundra, whereas the opposite has occurred in non-acidic tundra. Secondary stem growth has also increased as a component of primary production in acidic tundra, but is unchanged in non-acidic tundra. Thus, fertilization will probably increase carbon sequestration in woody biomass of B. nana in acidic tundra, while increasing carbon turnover (but not storage) of non-woody species in non-acidic tundra. These results indicate that nutrient enrichment can have very different consequences for plant communities that occur on different geological substrates, because of differences in composition, even though they share the same regional species pool. Although the specific edaphic factors that maintain compositional differences in this case are unknown, variation in soil pH and related variability in soil nutrient availability may well play a role.
  • Article
    Shallow soils are warmer under trees and tall shrubs across arctic and boreal ecosystems
    (IOP Publishing, 2020-12-18) Kropp, Heather ; Loranty, Michael M. ; Natali, Susan M. ; Kholodov, Alexander L. ; Rocha, Adrian V. ; Myers-Smith, Isla H. ; Abbott, Benjamin W. ; Abermann, Jakob ; Blanc-Betes, Elena ; Blok, Daan ; Blume-Werry, Gesche ; Boike, Julia ; Breen, Amy L. ; Cahoon, Sean M. P. ; Christiansen, Casper T. ; Douglas, Thomas A. ; Epstein, Howard E. ; Frost, Gerald V. ; Goeckede, Mathias ; Høye, Toke T. ; Mamet, Steven D. ; O’Donnell, Jonathan A. ; Olefeldt, David ; Phoenix, Gareth K. ; Salmon, Verity G. ; Sannel, A. Britta K. ; Smith, Sharon L. ; Sonnentag, Oliver ; Smith Vaughn, Lydia ; Williams, Mathew ; Elberling, Bo ; Gough, Laura ; Hjort, Jan ; Lafleur, Peter M. ; Euskirchen, Eugenie ; Heijmans, Monique M. P. D. ; Humphreys, Elyn ; Iwata, Hiroki ; Jones, Benjamin M. ; Jorgenson, M. Torre ; Grünberg, Inge ; Kim, Yongwon ; Laundre, James A. ; Mauritz, Marguerite ; Michelsen, Anders ; Schaepman-Strub, Gabriela ; Tape, Ken D. ; Ueyama, Masahito ; Lee, Bang-Yong ; Langley, Kirsty ; Lund, Magnus
    Soils are warming as air temperatures rise across the Arctic and Boreal region concurrent with the expansion of tall-statured shrubs and trees in the tundra. Changes in vegetation structure and function are expected to alter soil thermal regimes, thereby modifying climate feedbacks related to permafrost thaw and carbon cycling. However, current understanding of vegetation impacts on soil temperature is limited to local or regional scales and lacks the generality necessary to predict soil warming and permafrost stability on a pan-Arctic scale. Here we synthesize shallow soil and air temperature observations with broad spatial and temporal coverage collected across 106 sites representing nine different vegetation types in the permafrost region. We showed ecosystems with tall-statured shrubs and trees (>40 cm) have warmer shallow soils than those with short-statured tundra vegetation when normalized to a constant air temperature. In tree and tall shrub vegetation types, cooler temperatures in the warm season do not lead to cooler mean annual soil temperature indicating that ground thermal regimes in the cold-season rather than the warm-season are most critical for predicting soil warming in ecosystems underlain by permafrost. Our results suggest that the expansion of tall shrubs and trees into tundra regions can amplify shallow soil warming, and could increase the potential for increased seasonal thaw depth and increase soil carbon cycling rates and lead to increased carbon dioxide loss and further permafrost thaw.
  • Article
    Biomass offsets little or none of permafrost carbon release from soils, streams, and wildfire : an expert assessment
    (IOPScience, 2016-03-07) Abbott, Benjamin W. ; Jones, Jeremy B. ; Schuur, Edward A. G. ; Chapin, F. Stuart ; Bowden, William B. ; Bret-Harte, M. Syndonia ; Epstein, Howard E. ; Flannigan, Michael ; Harms, Tamara K. ; Hollingsworth, Teresa N. ; Mack, Michelle C. ; McGuire, A. David ; Natali, Susan M. ; Rocha, Adrian V. ; Tank, Suzanne E. ; Turetsky, Merritt R. ; Vonk, Jorien E. ; Wickland, Kimberly ; Aiken, George R. ; Alexander, Heather D. ; Amon, Rainer M. W. ; Benscoter, Brian ; Bergeron, Yves ; Bishop, Kevin ; Blarquez, Olivier ; Bond-Lamberty, Benjamin ; Breen, Amy L. ; Buffam, Ishi ; Cai, Yihua ; Carcaillet, Christopher ; Carey, Sean K. ; Chen, Jing M. ; Chen, Han Y. H. ; Christensen, Torben R. ; Cooper, Lee W. ; Cornelissen, Johannes H. C. ; de Groot, William J. ; DeLuca, Thomas Henry ; Dorrepaal, Ellen ; Fetcher, Ned ; Finlay, Jacques C. ; Forbes, Bruce C. ; French, Nancy H. F. ; Gauthier, Sylvie ; Girardin, Martin ; Goetz, Scott J. ; Goldammer, Johann G. ; Gough, Laura ; Grogan, Paul ; Guo, Laodong ; Higuera, Philip E. ; Hinzman, Larry ; Hu, Feng Sheng ; Hugelius, Gustaf ; JAFAROV, ELCHIN ; Jandt, Randi ; Johnstone, Jill F. ; Karlsson, Jan ; Kasischke, Eric S. ; Kattner, Gerhard ; Kelly, Ryan ; Keuper, Frida ; Kling, George W. ; Kortelainen, Pirkko ; Kouki, Jari ; Kuhry, Peter ; Laudon, Hjalmar ; Laurion, Isabelle ; Macdonald, Robie W. ; Mann, Paul J. ; Martikainen, Pertti ; McClelland, James W. ; Molau, Ulf ; Oberbauer, Steven F. ; Olefeldt, David ; Paré, David ; Parisien, Marc-André ; Payette, Serge ; Peng, Changhui ; Pokrovsky, Oleg ; Rastetter, Edward B. ; Raymond, Peter A. ; Raynolds, Martha K. ; Rein, Guillermo ; Reynolds, James F. ; Robards, Martin ; Rogers, Brendan ; Schädel, Christina ; Schaefer, Kevin ; Schmidt, Inger K. ; Shvidenko, Anatoly ; Sky, Jasper ; Spencer, Robert G. M. ; Starr, Gregory ; Striegl, Robert ; Teisserenc, Roman ; Tranvik, Lars J. ; Virtanen, Tarmo ; Welker, Jeffrey M. ; Zimov, Sergey A.
    As the permafrost region warms, its large organic carbon pool will be increasingly vulnerable to decomposition, combustion, and hydrologic export. Models predict that some portion of this release will be offset by increased production of Arctic and boreal biomass; however, the lack of robust estimates of net carbon balance increases the risk of further overshooting international emissions targets. Precise empirical or model-based assessments of the critical factors driving carbon balance are unlikely in the near future, so to address this gap, we present estimates from 98 permafrost-region experts of the response of biomass, wildfire, and hydrologic carbon flux to climate change. Results suggest that contrary to model projections, total permafrost-region biomass could decrease due to water stress and disturbance, factors that are not adequately incorporated in current models. Assessments indicate that end-of-the-century organic carbon release from Arctic rivers and collapsing coastlines could increase by 75% while carbon loss via burning could increase four-fold. Experts identified water balance, shifts in vegetation community, and permafrost degradation as the key sources of uncertainty in predicting future system response. In combination with previous findings, results suggest the permafrost region will become a carbon source to the atmosphere by 2100 regardless of warming scenario but that 65%–85% of permafrost carbon release can still be avoided if human emissions are actively reduced.
  • Preprint
    Ecosystem feedbacks and cascade processes : understanding their role in the responses of Arctic and alpine ecosystems to environmental change
    ( 2008-09-11) Wookey, Philip A. ; Aerts, Rien ; Bardgett, Richard D. ; Baptist, Florence ; Bråthen, Kari Anne ; Cornelissen, Johannes H. C. ; Gough, Laura ; Hartley, Iain P. ; Hopkins, David W. ; Lavorel, Sandra ; Shaver, Gaius R.
    Global environmental change, related to climate change and the deposition of airborne N-containing contaminants, has already resulted in shifts in plant community composition among plant functional types in arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that may result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate-driven increase in shrub cover in low arctic tundra, and the contrasting transformations in plant functional composition in mid-latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes which link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate-change feedbacks, can not be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.
  • Article
    Herbivore absence can shift dry heath tundra from carbon source to sink during peak growing season
    (IOP Publishing, 2021-02-01) Min, Elizabeth ; Wilcots, Megan E. ; Naeem, Shahid ; Gough, Laura ; McLaren, Jennie ; Rowe, Rebecca J. ; Rastetter, Edward B. ; Boelman, Natalie ; Griffin, Kevin L.
    In arctic tundra, large and small mammalian herbivores have substantial impacts on the vegetation community and consequently can affect the magnitude of carbon cycling. However, herbivores are often absent from modern carbon cycle models, partly because relatively few field studies focus on herbivore impacts on carbon cycling. Our objectives were to quantify the impact of 21 years of large herbivore and large and small herbivore exclusion on carbon cycling during peak growing season in a dry heath tundra community. When herbivores were excluded, we observed a significantly greater leaf area index as well as greater vascular plant abundance. While we did not observe significant differences in deciduous dwarf shrub abundance across treatments, evergreen dwarf shrub abundance was greater where large and small herbivores were excluded. Both foliose and fruticose lichen abundance were higher in the large herbivore, but not the small and large herbivore exclosures. Net ecosystem exchange (NEE) likewise indicated the highest carbon uptake in the exclosure treatments and lowest uptake in the control (CT), suggesting that herbivory decreased the capacity of dry heath tundra to take up carbon. Moreover, our calculated NEE for average light and temperature conditions for July 2017, when our measurements were taken, indicated that the tundra was a carbon source in CT, but was a carbon sink in both exclosure treatments, indicating removal of grazing pressure can change the carbon balance of dry heath tundra. Collectively, these findings suggest that herbivore absence can lead to changes in plant community structure of dry heath tundra that in turn can increase its capacity to take up carbon.
  • Preprint
    Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance
    ( 2017-09) Asmus, Ashley ; Koltz, Amanda ; McLaren, Jennie ; Shaver, Gaius R. ; Gough, Laura
    A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.
  • Article
    Time lags: insights from the U.S. Long Term Ecological Research Network
    (Ecological Society of America, 2021-05-17) Rastetter, Edward B. ; Ohman, Mark D. ; Elliott, Katherine J. ; Rehage, Jennifer S. ; Rivera-Monroy, Victor H. ; Boucek, Ross E. ; Castaneda-Moya, Edward ; Danielson, Tess M. ; Gough, Laura ; Groffman, Peter M. ; Jackson, C. Rhett ; Ford Miniat, Chelcy
    Ecosystems across the United States are changing in complex ways that are difficult to predict. Coordinated long-term research and analysis are required to assess how these changes will affect a diverse array of ecosystem services. This paper is part of a series that is a product of a synthesis effort of the U.S. National Science Foundation’s Long Term Ecological Research (LTER) network. This effort revealed that each LTER site had at least one compelling scientific case study about “what their site would look like” in 50 or 100 yr. As the site results were prepared, themes emerged, and the case studies were grouped into separate papers along five themes: state change, connectivity, resilience, time lags, and cascading effects and compiled into this special issue. This paper addresses the time lags theme with five examples from diverse biomes including tundra (Arctic), coastal upwelling (California Current Ecosystem), montane forests (Coweeta), and Everglades freshwater and coastal wetlands (Florida Coastal Everglades) LTER sites. Its objective is to demonstrate the importance of different types of time lags, in different kinds of ecosystems, as drivers of ecosystem structure and function and how these can effectively be addressed with long-term studies. The concept that slow, interactive, compounded changes can have dramatic effects on ecosystem structure, function, services, and future scenarios is apparent in many systems, but they are difficult to quantify and predict. The case studies presented here illustrate the expanding scope of thinking about time lags within the LTER network and beyond. Specifically, they examine what variables are best indicators of lagged changes in arctic tundra, how progressive ocean warming can have profound effects on zooplankton and phytoplankton in waters off the California coast, how a series of species changes over many decades can affect Eastern deciduous forests, and how infrequent, extreme cold spells and storms can have enduring effects on fish populations and wetland vegetation along the Southeast coast and the Gulf of Mexico. The case studies highlight the need for a diverse set of LTER (and other research networks) sites to sort out the multiple components of time lag effects in ecosystems.
  • Article
    Shrub encroachment in Arctic tundra : Betula nana effects on above- and belowground litter decomposition
    (John Wiley & Sons, 2017-04-07) McLaren, Jennie ; Buckeridge, Kate M. ; van de Weg, Martine J. ; Shaver, Gaius R. ; Schimel, Joshua P. ; Gough, Laura
    Rapid arctic vegetation change as a result of global warming includes an increase in the cover and biomass of deciduous shrubs. Increases in shrub abundance will result in a proportional increase of shrub litter in the litter community, potentially affecting carbon turnover rates in arctic ecosystems. We investigated the effects of leaf and root litter of a deciduous shrub, Betula nana, on decomposition, by examining species-specific decomposition patterns, as well as effects of Betula litter on the decomposition of other species. We conducted a 2-yr decomposition experiment in moist acidic tundra in northern Alaska, where we decomposed three tundra species (Vaccinium vitis-idaea, Rhododendron palustre, and Eriophorum vaginatum) alone and in combination with Betula litter. Decomposition patterns for leaf and root litter were determined using three different measures of decomposition (mass loss, respiration, extracellular enzyme activity). We report faster decomposition of Betula leaf litter compared to other species, with support for species differences coming from all three measures of decomposition. Mixing effects were less consistent among the measures, with negative mixing effects shown only for mass loss. In contrast, there were few species differences or mixing effects for root decomposition. Overall, we attribute longer-term litter mass loss patterns to patterns created by early decomposition processes in the first winter. We note numerous differences for species patterns between leaf and root decomposition, indicating that conclusions from leaf litter experiments should not be extrapolated to below-ground decomposition. The high decomposition rates of Betula leaf litter aboveground, and relatively similar decomposition rates of multiple species below, suggest a potential for increases in turnover in the fast-decomposing carbon pool of leaves and fine roots as the dominance of deciduous shrubs in the Arctic increases, but this outcome may be tempered by negative litter mixing effects during the early stages of encroachment.
  • Article
    Differential physiological responses to environmental change promote woody shrub expansion
    (John Wiley & Sons, 2013-03-13) Heskel, Mary ; Greaves, Heather ; Kornfeld, Ari ; Gough, Laura ; Atkin, Owen K. ; Turnbull, Matthew H. ; Shaver, Gaius R. ; Griffin, Kevin L.
    Direct and indirect effects of warming are increasingly modifying the carbon-rich vegetation and soils of the Arctic tundra, with important implications for the terrestrial carbon cycle. Understanding the biological and environmental influences on the processes that regulate foliar carbon cycling in tundra species is essential for predicting the future terrestrial carbon balance in this region. To determine the effect of climate change impacts on gas exchange in tundra, we quantified foliar photosynthesis (Anet), respiration in the dark and light (RD and RL, determined using the Kok method), photorespiration (PR), carbon gain efficiency (CGE, the ratio of photosynthetic CO2 uptake to total CO2 exchange of photosynthesis, PR, and respiration), and leaf traits of three dominant species – Betula nana, a woody shrub; Eriophorum vaginatum, a graminoid; and Rubus chamaemorus, a forb – grown under long-term warming and fertilization treatments since 1989 at Toolik Lake, Alaska. Under warming, B. nana exhibited the highest rates of Anet and strongest light inhibition of respiration, increasing CGE nearly 50% compared with leaves grown in ambient conditions, which corresponded to a 52% increase in relative abundance. Gas exchange did not shift under fertilization in B. nana despite increases in leaf N and P and near-complete dominance at the community scale, suggesting a morphological rather than physiological response. Rubus chamaemorus, exhibited minimal shifts in foliar gas exchange, and responded similarly to B. nana under treatment conditions. By contrast, E. vaginatum, did not significantly alter its gas exchange physiology under treatments and exhibited dramatic decreases in relative cover (warming: −19.7%; fertilization: −79.7%; warming with fertilization: −91.1%). Our findings suggest a foliar physiological advantage in the woody shrub B. nana that is further mediated by warming and increased soil nutrient availability, which may facilitate shrub expansion and in turn alter the terrestrial carbon cycle in future tundra environments.
  • Article
    Using structure to model function: incorporating canopy structure improves estimates of ecosystem carbon flux in arctic dry heath tundra
    (IOP Publishing, 2023-05-12) Min, Elizabeth ; Naeem, Shahid ; Gough, Laura ; McLaren, Jennie R. ; Rowe, Rebecca J. ; Rastetter, Edward ; Boelman, Natalie ; Griffin, Kevin L.
    Most tundra carbon flux modeling relies on leaf area index (LAI), generally estimated from measurements of canopy greenness using the normalized difference vegetation index (NDVI), to estimate the direction and magnitude of fluxes. However, due to the relative sparseness and low stature of tundra canopies, such models do not explicitly consider the influence of variation in tundra canopy structure on carbon flux estimates. Structure from motion (SFM), a photogrammetric method for deriving three-dimensional (3D) structure from digital imagery, is a non-destructive method for estimating both fine-scale canopy structure and LAI. To understand how variation in 3D canopy structure affects ecosystem carbon fluxes in Arctic tundra, we adapted an existing NDVI-based tundra carbon flux model to include variation in SFM-derived canopy structure and its interaction with incoming sunlight to cast shadows on canopies. Our study system consisted of replicate plots of dry heath tundra that had been subjected to three herbivore exclosure treatments (an exclosure-free control [CT], large mammals exclosure), and a large and small mammal exclosure [ExLS]), providing the range of 3D canopy structures employed in our study. We found that foliage within the more structurally complex surface of CT canopies received significantly less light over the course of the day than canopies within both exclosure treatments. This was especially during morning and evening hours, and was reflected in modeled rates of net ecosystem exchange (NEE) and gross primary productivity (GPP). We found that in the ExLS treatment, SFM-derived estimates of GPP were significantly lower and NEE significantly higher than those based on LAI alone. Our results demonstrate that the structure of even simple tundra vegetation canopies can have significant impacts on tundra carbon fluxes and thus need to be accounted for.