From old fields to forests: Understanding plant successional dynamics through the lens of functional traits

Vegetative succession describes the turnover of plant species through time. This turnover enables coexistence of species temporally, but also spatially as different locations co-occur at different successional stages. Moreover, the suite of species that occupy different successional stages varies due to heterogeneous environments across both local and regional spatial scales. Understanding the processes that underlie succession as well as those that drive spatial variation in the species that comprise similar successional stages is a central goal in ecology. In order to understand these processes in this dissertation, I recast species into functional traits that connect species physiologies to their environments. Using a suite of traits thought to influence species success at various stages of succession, I examine functional trait changes through time in plant communities of the eastern US. Chapters 2 and 3 use an old field experiment to examine how soil nutrients and plant enemies influence temporal dynamics of early secondary succession by examining species-level trait responses (Chapter 2) and community-level trait responses (Chapter 3). Old fields are important and well-studied community types due to their frequency in the landscape and lend themselves well to experimental manipulation given the relatively rapid life cycles and small stature of their constituent herbaceous species. Chapters 4 and 5 use a continental-scale forest database to examine similar processes in trees, albeit at larger spatial and temporal gradients. Chapter 4 uses a space-for-time substitute approach to ask how tree community traits change along a forest age gradient, while Chapter 5 asks how traits of tree seedling communities respond to forest disturbances using resampled plots. In Chapter 6, I synthesize my findings on trait responses to successional gradients in these two distinct successional stages. Overall, I found that seed mass, indicative of dispersal strategy, and investment in structural biomass (plant height and wood density) capture plant successional strategies. Leaf traits, however, did not consistently vary with succession or the manipulated environmental gradients in the old field experiment. Rather, leaf traits displayed large, unexplained variation across space, suggesting that they are responding to processes related to spatial heterogeneity independent of succession.Doctor of Philosoph

Drought effects on montane grasslands nullify benefits of advanced flowering phenology due to warming

Abstract Warming due to climate change is generally expected to lengthen the growing season in areas of seasonal climate and to advance plant phenology, particularly the onset of leafing and flowering. However, a reduction in aboveground biomass production and reproductive output may occur when warming is accompanied by drought that crosses critical water deficit thresholds. Tracking warmer temperatures has been shown to be species‐specific with unknown impacts on community composition and productivity. The variability in species’ ability to leverage earlier leaf unfolding and flowering into increased aboveground net primary production (ANPP) or increased investments into reproductive organs has heretofore been poorly explored. We tested whether phenological sensitivity to temperature, as a result of experimental warming, directly translated into increased plant performance, as measured by ANPP and flower abundance. In order to experimentally simulate climate warming, we translocated a total of 45 intact soil–plant communities downslope along an elevational gradient of 900 m within the European Alps from 1260 to 350 m asl and weekly recorded flower abundance and total green cover as well as cumulative biomass production at peak growing season. We found that advanced phenology at lower elevations was related to increased reproductive performance and conditional on whether they experienced drought stress. While a temperature increase of +1K had positive effects on the amount of reproductive organs for species with accelerated phenology, temperature increase going along with drier conditions resulted in plants being unable to sustain early investment in reproduction as measured by flower abundance. This finding highlights that the interaction of two climate change drivers, warming and drought, can push communities’ past resistance thresholds. Moreover, we detected biotic competition mechanisms and shifts toward forb‐depressed states with graminoids best taking advantage of experimentally altered increased temperature and reduced precipitation. Our results suggest that while species may track warmer future climates, concurrent drought events post a high risk for failure of temperature‐driven improvement of reproductive performance and biomass production in the European Alps

High land-use intensity diminishes stability of forage provision of mountain pastures under future climate variability

Semi-natural, agriculturally used grasslands provide important ecologic and economic services, such as feed supply. In mountain regions, pastures are the dominant agricultural system and face more severe climate change impacts than lowlands. Climate change threatens ecosystem functions, such as aboveground net primary production [ANPP] and its nutrient content. It is necessary to understand the impacts of climate change and land-management on such ecosystems to develop management practices to sustainably maintain provision of ecosystem services under future climatic conditions. We studied the effect of climate change and different land-use intensities on plant-soil communities by the downslope translocation of plant-soil mesocosms along an elevation gradient in 2016, and the subsequent application of two management types (extensive vs. intensive). Communities’ response to ANPP and leaf carbon (C), nitrogen (N), and phosphorus (P) content was quantified over the subsequent two years after translocation. ANPP increased with warming in 2017 under both management intensities, but this effect was amplified by intensive land-use management. In 2018, ANPP of intensively managed communities decreased, in comparison to 2017, from 35% to 42%, while extensively managed communities maintained their production levels. The changes in ANPP are coupled with an exceptionally dry year in 2018, with up to 100 more days of drought conditions. The C:N of extensively managed communities was higher than those of intensively managed ones, and further increased in 2018, potentially indicating shifts in resource allocation strategies that may explain production stability. Our results revealed a low resistance of intensively managed communities’ ANPP under especially dry conditions. The ability to alter resource allocation likely enables a constant level of production under extensive management, but this ability is lost under intensive management. Thus, future drought events may leave intensive management as a non-sustainable farming practice, and ultimately threaten ecosystem services of montane pastures

Intensive slurry management and climate change promote nitrogen mining from organic matter-rich montane grassland soils

Aims Consequences of climate change and land use intensification on the nitrogen (N) cycle of organic-matter rich grassland soils in the alpine region remain poorly understood. We aimed to identify fates of fertilizer N and to determine the overall N balance of an organic-matter rich grassland in the European alpine region as influenced by intensified management and warming. Methods We combined 15N cattle slurry labelling with a space for time climate change experiment, which was based on translocation of intact plant-soil mesocosms down an elevational gradient to induce warming of +1 °C and + 3 °C. Mesocosms were subject to either extensive or intensive management. The fate of slurry-N was traced in the plant-soil system. Results Grassland productivity was very high (8.2 t - 19.4 t dm ha$^{-1}$ yr$^{-1}$), recovery of slurry $^{15}$N in mowed plant biomass was, however, low (9.6–14.7%), illustrating low fertilizer N use efficiency and high supply of plant available N via mineralization of soil organic matter (SOM). Higher 15N recovery rates (20.2–31.8%) were found in the soil N pool, dominated by recovery in unextractable N. Total $^{15}$N recovery was approximately half of the applied tracer, indicating substantial loss to the environment. Overall, high N export by harvest (107–360 kg N ha$^{-1}$ yr$^{-1}$) markedly exceeded N inputs, leading to a negative grassland N balance. Conclusions Here provided results suggests a risk of soil N mining in montane grasslands, which increases both under climate change and land use intensification

Nothing lasts forever: Dominant species decline under rapid environmental change in global grasslands

1. Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them. 2. Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure. 3. We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments. 4. Synthesis. Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change.EEA Santa CruzFil: Wilfahrt, Peter A. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados UnidosFil: Seabloom, Eric William. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados UnidosFil: Bakker, Jonathan D. University of Washington. School of Environmental and Forest Sciences; Estados Unidos.Fil: Biederman, Lori A. Iowa State University. Department of Ecology, Evolution, and Organismal Biology; Estados UnidosFil: Bugalho, Miguel N. University of Lisbon. Centre for Applied Ecology “Prof. Baeta Neves” (CEABN-InBIO). School of Agriculture; Portugal.Fil: Cadotte, Marc W. University of Toronto Scarborough. Department of Biological Sciences; Canadá.Fil: Caldeira, Maria C. University of Lisbon. Forest Research Centre. School of Agriculture; Portugal.Fil: Catford, Jane A. King’s College London. Department of Geography; Reino UnidoFil: Catford, Jane A. University of Melbourne. School of Agriculture, Food and Ecosystem Sciences; Australia.Fil: Chen, Qingqing. Peking University. College of Urban and Environmental Science; China.Fil: Chen, Qingqing. German Centre for Integrative Biodiversity Research (iDiv). Halle-Jena-Leipzig; AlemaniaFil: Donohue, Ian. Trinity College Dublin. School of Natural Sciences. Department of Zoology; IrlandaFil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina.Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral.; Argentina.Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina.Fil: Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unido

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Abstract Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning

The handbook for standardized field and laboratory measurements in terrestrial climate change experiments and observational studies (ClimEx)

Climate change is a world-wide threat to biodiversity and ecosystem structure, functioning and services. To understand the underlying drivers and mechanisms, and to predict the consequences for nature and people, we urgently need better understanding of the direction and magnitude of climate change impacts across the soil-plant-atmosphere continuum. An increasing number of climate change studies are creating new opportunities for meaningful and high-quality generalizations and improved process understanding. However, significant challenges exist related to data availability and/or compatibility across studies, compromising opportunities for data re-use, synthesis and upscaling. Many of these challenges relate to a lack of an established 'best practice' for measuring key impacts and responses. This restrains our current understanding of complex processes and mechanisms in terrestrial ecosystems related to climate change. To overcome these challenges, we collected best-practice methods emerging from major ecological research networks and experiments, as synthesized by 115 experts from across a wide range of scientific disciplines. Our handbook contains guidance on the selection of response variables for different purposes, protocols for standardized measurements of 66 such response variables and advice on data management. Specifically, we recommend a minimum subset of variables that should be collected in all climate change studies to allow data re-use and synthesis, and give guidance on additional variables critical for different types of synthesis and upscaling. The goal of this community effort is to facilitate awareness of the importance and broader application of standardized methods to promote data re-use, availability, compatibility and transparency. We envision improved research practices that will increase returns on investments in individual research projects, facilitate second-order research outputs and create opportunities for collaboration across scientific communities. Ultimately, this should significantly improve the quality and impact of the science, which is required to fulfil society's needs in a changing world.Peer reviewe

Nothing lasts forever: Dominant species decline under rapid environmental change in global grasslands

Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them. Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure. We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments. Synthesis. Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change.Fil: Wilfahrt, Peter A.. University of Minnesota; Estados UnidosFil: Seabloom, Eric. University of Minnesota; Estados UnidosFil: Bakker, Jonathan. University of Washington; Estados UnidosFil: Biederman, Lori. Iowa State University; Estados UnidosFil: Bugalho, Miguel N.. Universidade Nova de Lisboa; PortugalFil: Cadotte, Marc W.. University of Toronto–Scarborough; Estados UnidosFil: Caldeira, Maria C.. Universidade Nova de Lisboa; PortugalFil: Catford, Jane A.. University of Melbourne; AustraliaFil: Chen, Qingqing. Peking University; China. German Centre for Integrative Biodiversity Research; AlemaniaFil: Donohue, Ian. Trinity College Dublin; IrlandaFil: Ebeling, Anne. University of Jena; AlemaniaFil: Eisenhauer, Nico. German Centre for Integrative Biodiversity Research; Alemania. Leipzig University; AlemaniaFil: Haider, Sylvia. Martin Luther University Halle-Wittenberg; Alemania. Leuphana University of Lüneburg; AlemaniaFil: Heckman, Robert W.. University of Texas; Estados Unidos. United States Forest Service; Estados UnidosFil: Jentsch, Anke. University of Bayreuth; AlemaniaFil: Koerner, Sally E.. University of North Carolina Greensboro; Estados UnidosFil: Komatsu, Kimberly J.. University of North Carolina Greensboro; Estados UnidosFil: Laungani, Ramesh. Poly Prep Country Day School; Estados UnidosFil: MacDougall, Andrew. University of Guelph; CanadáFil: Smith, Nicholas G.. Texas Tech University; Estados UnidosFil: Stevens, Carly J.. Lancaster University; Reino UnidoFil: Sullivan, Lauren L.. Michigan State University; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Tedder, Michelle. University of KwaZulu-Natal; SudáfricaFil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Tecnológica Nacional. Facultad Regional Santa Cruz. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Nacional de la Patagonia Austral. Centro de Investigaciones y Transferencia de Santa Cruz; ArgentinaFil: Tognetti, Pedro Maximiliano. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Veen, Ciska. Netherlands Institute of Ecology; Países BajosFil: Wheeler, George. University of Nebraska-Lincoln; Estados UnidosFil: Young, Alyssa L.. University of North Carolina Greensboro; Estados UnidosFil: Young, Hillary. University of California; Estados UnidosFil: Borer, Elizabeth. University of Minnesota; Estados Unido