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California's Native Grasslands

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Grasslands Research Highlights 

Grasslands are important subjects of ecological research and experimentation, and are studied from a wide variety of perspectives (e.g., native and invasive species; soils; management using fire, grazing, mowing, and herbicide application; ecological restoration). CNGA will regularly browse peer-reviewed books and scientific journal articles to provide highlights of recent grasslands research. Studies are organized by year of publication.


2024 Publications

Benefits of Cultural Fire Stewardship

Taylor, Annalise, Alexii Sigona, and Maggi Kelly. 2024. “Centering Amah Mutsun Voices in the Analysis of a Culturally Important, Fire-Managed Coastal Grassland.” Ecological Applications e3014. https://doi.org/10.1002/eap.3014 - Open Access.

The study explores the impact of cultural fire stewardship by Indigenous communities in California, particularly in coastal grasslands, an ecosystem known for its biodiversity and vulnerability. The research, conducted in partnership with the Amah Mutsun Tribal Band, compares three adjacent coastal grasslands with different fire histories: one with fire suppression, one that experienced wildfire after suppression, and one that has undergone regular burning every two years for over 30 years.

Key findings include:

  1. Increased Abundance of Native Grass: Danthonia californica, a native grass species, was significantly more abundant in regularly burned sites.
  2. Shrub Encroachment: Shrubs such as Baccharis pilularis, Frangula californica, and Rubus ursinus were more prevalent in areas with no recorded fire.
  3. Cultural and Ecological Benefits: The study emphasizes the importance of cultural burns for maintaining grassland health and preventing shrub encroachment, which is critical for both ecological balance and cultural practices.

The research also used Sentinel-2 satellite imagery to assess fire severity and included interviews with Amah Mutsun tribal members to understand the cultural significance of burning practices.


Melanie T. Hacopian, Sarai S. Finks, Kathleen K. Treseder. 2024. Drought mediates the response of soil fungal communities post-wildfire in a Californian grassland and coastal sage scrubland. Soil Biology and Biochemistry, Volume 196, 109511. ISSN 0038-0717. https://doi.org/10.1016/j.soilbio.2024.109511. (https://www.sciencedirect.com/science/article/pii/S0038071724002001)

Highlights

  • Drought and ambient fungal communities remained distinct post-fire.
  • Drought and ambient fungal communities followed different trends post-fire.
  • Fungal richness was stable across post-fire collection dates.
  • We observed several indicator species post-wildfire.

Abstract: There is a knowledge gap surrounding how drought and wildfire, two increasingly frequent disturbances, will alter soil fungal communities. Moreover, studies that directly compare ambient and drought-treated soil fungal communities in the context of wildfire are exceptionally scarce. We assessed the response and recovery of soil fungal communities and functional guilds in two sites – a grassland and a coastal sage shrubland – after a severe wildfire burned a long-term drought experiment. We collected soil samples at four collection dates over an eight-month period after wildfire and amplified fungal DNA. We predicted that fungal communities within the drought and ambient treatments would differ significantly across collection dates owing to differing responses to post-wildfire conditions. Richness was stable across collection dates, regardless of precipitation treatment, in both sites. Differences between treatments were significant at every collection date with respect to taxonomical community composition. Differences in community composition between collection dates within each treatment were also significant. Additionally, the monotonic trends of drought and ambient communities over time differed in strength and direction. Differences in shrubland functional guild composition across collection dates and contrasting trends suggest a drought-dependent shift after the fire. Overall, we conclude that drought mediates how soil fungal communities respond after a wildfire in the long term, however drought effects may differ across ecosystems.


Invasives: Lengthy Lag Time Phase (40 - 320 Years) Can Precede Expansion 

Invading plants remain undetected in a lag phase while they explore suitable climates.  Philipp Robeck, Franz Essl, Mark van Kleunen, Petr Pyšek, Jan Pergl, Patrick Weigelt & Mohsen B. Mesgaran

Nature Ecology & Evolution volume 8, pages 477–488 (2024) https://doi.org/10.1038/s41559-023-02313-4

Abstract. Successful alien species may experience a period of quiescence, known as the lag phase, before becoming invasive and widespread. The existence of lags introduces severe uncertainty in risk analyses of aliens as the present state of species is a poor predictor of future distributions, invasion success and impact. Predicting a species’ ability to invade and pose negative impacts requires a quantitative understanding of the commonality and magnitude of lags, environmental factors and mechanisms likely to terminate lag. Using herbarium and climate data, we analysed over 5,700 time series (species × regions) in 3,505 naturalized plant species from nine regions in temperate and tropical climates to quantify lags and test whether there have been shifts in the species’ climatic space during the transition from the lag phase to the expansion phase. Lags were identified in 35% of the assessed invasion events. We detected phylogenetic signals for lag phases in temperate climate regions and that annual self-fertilizing species were less likely to experience lags. Where lags existed, they had an average length of 40 years and a maximum of 320 years. Lengthy lags (>100 years) were more likely to occur in perennial plants and less frequent in self-pollinating species. For 98% of the species with a lag phase, the climate spaces sampled during the lag period differed from those in the expansion phase based on the assessment of centroid shifts or degree of climate space overlap. Our results highlight the importance of functional traits for the onset of the expansion phase and suggest that climate discovery may play a role in terminating the lag phase. However, other possibilities, such as sampling issues and climate niche shifts, cannot be ruled out.


Climate Change and California’s Terrestrial Biodiversity

S. Harrison, J. Franklin, R.R. Hernandez, M. Ikegami, H.D. Safford, J.H. Thorne, Climate change and California’s terrestrial biodiversity, Proc. Natl. Acad. Sci. U.S.A.: 121 (32) e2310074121, https://doi.org/10.1073/pnas.2310074121 (2024). 

Abstract: In this review and synthesis, we argue that California is an important test case for the nation and world because terrestrial biodiversity is very high, present and anticipated threats to biodiversity from climate change and other interacting stressors are severe, and innovative approaches to protecting biodiversity in the context of climate change are being developed and tested. We first review salient dimensions of California’s terrestrial physical, biological, and human diversity. Next, we examine four facets of the threat to their sustainability of these dimensions posed by climate change: direct impacts, illustrated by a new analysis of shifting diversity hotspots for plants; interactive effects involving invasive species, land-use change, and other stressors; the impacts of changing fire regimes; and the impacts of land-based renewable energy development. We examine recent policy responses in each of these areas, representing attempts to better protect biodiversity while advancing climate adaptation and mitigation. We conclude that California’s ambitious 30 × 30 Initiative and its efforts to harmonize biodiversity conservation with renewable energy development are important areas of progress. Adapting traditional suppression-oriented fire policies to the reality of new fire regimes is an area in which much progress remains to be made.

Applying Community Assembly Theory

Nguyen, M.A., Kimball, S., Burger, J.C., Freese, R., Lulow, M., Schmidt, K.T., Ta, P. and Funk, J.L. (2024), Applying community assembly theory to restoration: overcoming dispersal and abiotic filters is key to diversifying California grassland. Restor Ecol, 32: e14018. https://doi.org/10.1111/rec.14018

Summary (adapted from abstract)

In the context of restoration, ecological filters (dispersal, abiotic, and biotic) can be manipulated to alter the resulting plant community by favoring native species or disadvantaging non-native invasive species. The authors conducted two studies manipulating assembly filters at two California grassland sites previously dominated by non-native species. At Site 1, they explored how variations in sequential seeding of native grasses and forbs (to overcome dispersal and biotic filters caused by priority effects) influenced the resulting community. At Site 2, they explored how thatch removal (to overcome the abiotic filter of light limitation) and herbicide-based weed control (to overcome the biotic filter of competition from non-native species) influenced the addition of native forbs into a partially restored grassland. Native forbs at Site 1 did not suffer from arriving after grasses, but native grasses benefited when given 1 year priority over forbs. At Site 2, dethatching increased native forb cover in a high rainfall year. Herbicide application reduced non-native grass cover in dethatched plots without negatively affecting native cover. Native forb and grass cover were significant predictors of non-native grass cover. However, they accounted for only 29% of the variation observed, suggesting there are other influential factors not considered in this study. Our results suggest that forbs can be incorporated into established native grasslands more successfully after dethatching.

Implications for Practice

  • Diversifying restored grasslands requires management of dispersal, abiotic, and biotic filters to promote success of sown species.
  • There are clear benefits to utilizing mowing as a restoration tool, both to promote native forb success and as a weed management tool.
  • In years with sufficient rainfall, seeding native forbs into dethatched stands of established native bunchgrasses can achieve higher forb cover than seeding both native grasses and forbs concurrently or sequentially at the start of a restoration project.


2023 Publications

Ratcliff, Felix, Sheila Barry, Devii Rao, Rowan Peterson, Theresa Becchetti, Ermias Kebreab, Kaveh Motamed, Minju Jung, and Frank Mitloehner. 2023. "Cattle Grazing Moderates Greenhouse Gas and Particulate Matter Emissions from California Grassland Wildfires" Sustainability 15, no. 18: 13539. https://doi.org/10.3390/su151813539

Abstract

Between 2010 and 2020, an average of 36,037 hectares of grassland burned in wildfires in California each year, emitting greenhouse gasses (GHGs) and particulate matter (PM). These emissions impact climate and human health. Cattle grazing removes herbaceous fuel through the consumption of forage; however, ruminant digestion also emits GHGs. The purpose of this study was to examine the GHG and PM impact of livestock grazing in grasslands that go on to burn. We used Monte Carlo simulation to determine whether forage consumption by livestock led to reductions in grassland wildfire emissions and whether these reductions outweighed the emissions from the digestion of that forage. We estimate that between 2010 and 2020, an average of 11,590 metric tons (MT) of herbaceous fuel were removed by cattle annually from grasslands in California that went on to burn. This resulted in annual wildfire emission reductions ranging between 0.001 and 0.025 million metric tons (MMT) of CO2 equivalents (CO2e) and between 11 and 314 MT of PM2.5; a small fraction of total GHG and PM emissions from wildfires in California. We also evaluated the change in emissions if burned grasslands in California’s Central and North Coast regions—where removing grazing can lead to the encroachment of shrubs into grasslands—were instead shrublands. If the grasslands that burned in these regions in 2020 had instead been shrublands, we estimate that as much as 0.90 MMT more CO2e and 8448 MT more PM2.5 would have been emitted by wildfires, highlighting the long-term implications of livestock grazing.


Evans, A.W., Woodward, B.D., Wyckoff, A.C., Toledo, D., Duke, S., Fischer, C. et al. (2023) Livestock grazing is an effective conservation tool for Californian coastal grassland ecology: An eight-year study on vegetation dynamics. Applied Vegetation Science, 26, e12736. Available from: https://doi.org/10.1111/avsc.12736

The study aimed to assess the impact of targeted grazing on various ecological factors over an eight-year period across 17 study sites. The researchers monitored grazed and ungrazed plots and measured effects on bare ground cover, litter depth and cover, herb height, and diversity of native and non-native species. The results showed that grazing increased bare ground cover by 2.8%, reduced litter depth by 3.5 cm, lowered litter cover by 12%, and decreased herb height by 3.9 cm on average. Grazed plots exhibited higher cover of native annual forbs (+1.2%) and reduced invasive Bromus diandrus cover by 6.0%. However, there was no significant difference in diversity between native and non-native species. The study concludes that targeted grazing can be a useful conservation tool, aiding biomass management and canopy clearing while enhancing native annual forbs without posing a risk to overall native species. Future research should explore adjustments in grazing techniques to further support native species and inhibit invasive species growth.


Buisson E, S Archibald, A Fidelis, KN Suding. 2022. Ancient grasslands guide ambitious goals in grassland restoration. Science 377 (6606):594-598 (4 Aug 2022). DOI:10.1126/science.abo4605

Abstract. Grasslands, which constitute almost 40% of the terrestrial biosphere, provide habitat for a great diversity of animals and plants and contribute to the livelihoods of more than 1 billion people worldwide. Whereas the destruction and degradation of grasslands can occur rapidly, recent work indicates that complete recovery of biodiversity and essential functions occurs slowly or not at all. Grassland restoration—interventions to speed or guide this recovery—has received less attention than restoration of forested ecosystems, often due to the prevailing assumption that grasslands are recently formed habitats that can reassemble quickly. Viewing grassland restoration as long-term assembly toward old-growth endpoints, with appreciation of feedbacks and threshold shifts, will be crucial for recognizing when and how restoration can guide recovery of this globally important ecosystem.

Glassman, Sydney I., James W.J. Randolph, Sameer S. Saroa, Joia K. Capocchi, Kendra E. Walters, M. Fabiola Pulido-Chavez, Loralee Larios. 2023. “Prescribed versus wildfire impacts on exotic plants and soil microbes in California grasslands.” Applied Soil Ecology 185, 104795, ISSN 0929-1393, https://doi.org/10.1016/j.apsoil.2022.104795.

Abstract: Prescribed burns are often used as a management tool to decrease exotic plant cover and increase native plant cover in grasslands. These changes may also be mediated by fire impacts on soil microbial communities, which drive plant productivity and function. Yet, the ecological effects of prescribed burns compared to wildfires on either plant or soil microbial composition remain unclear. Grassland fires account for roughly 80 % of global annual fires, but only roughly 12 % of research on belowground impacts of fires occurs in grasslands, limiting our understanding of aboveground belowground connections in these important habitats. Here, we took advantage of the serendipitous opportunity of a wildfire burning through the same reserve where we had previously sampled a prescribed burn. This enabled us to investigate the impacts of a spring prescribed burn versus a fall wildfire on plant cover and community composition and bacterial and fungal richness, abundance, and composition. Our California grassland sites were thus within the same reserve, limiting environmental, vegetation, or climate variation between the sites. We used qPCR of 16S and 18S to assess impacts on bacterial and fungal abundance and Illumina MiSeq of 16S and ITS2 to assess impacts on bacterial and fungal richness and composition. Wildfire had stronger impacts than prescribed burns on microbial communities and both fires had similar impacts on plants with both prescribed and wildfire reducing exotic plant cover but neither reducing exotic plant richness. Fungal richness declined after the wildfire but not prescribed burn, but bacterial richness was unaffected by either. Yet, fire exposure in both fire types resulted in reduced bacterial and fungal abundance and altered bacterial and fungal composition. Plant diversity differentially impacted soil microbial diversity, with exotic plant diversity positively impacting bacterial richness and having no effect on arbuscular mycorrhizal richness. However, the remainder of the soil microbial communities were more related to aspects of soil chemistry including cation exchange capacity, organic matter, pH and phosphorous. Our coupled plant and soil community sampling allowed us to capture the sensitivity to fire of the fungal community and highlights the importance of potentially incorporating management actions such as soil or fungal amendments to promote this critical community that mediates native plant performance.


Keeley, J.E., Klinger, R.C., Brennan, T.J., Lawson, D.M., La Grange, J. and Berg, K.N. (2023), A decade-long study of repeated prescription burning in California native grassland restoration. Restoration Ecology, 31: e13939. https://doi.org/10.1111/rec.13939

This study focuses on restoring native grasslands in California dominated by Stipa pulchra, which currently coexist with non-native annual grasses. The primary aim is to shift the balance towards native grass dominance. The research examines the impact of burning on vegetation recovery.

In the first year after burning, non-native annual grass cover was significantly reduced by 70%, with Bromus diandrus experiencing the most substantial decrease. However, S. pulchra recovery was minimally affected. Over the next three years, S. pulchra levels returned to those of unburned areas, while non-native annual grasses remained below their original levels until the fifth year. Some non-native Erodium species increased in response to reduced annual grass cover but presented a lesser threat to native bunchgrasses. They returned to near control levels within three years.

The native geophyte Brodiaea kinkiensis was present throughout the grasslands and was unaffected by burning treatments. A second burn conducted five years later replicated the patterns observed after the first burn. Precipitation influenced S. pulchra cover, but non-native grass cover was more responsive to precipitation, with similar trends in both burned and unburned plots.


Rethinking local seed sourcing for the restoration of a foundational grass species in California

Nolan, M.P., Luong, J.C., Valliere, J.M., Mazer, S.J. and D'Antonio, C.M. (2023), Rethinking local seed sourcing for the restoration of a foundational grass species in California. Restor Ecol e13992. https://doi.org/10.1111/rec.13992

Madeline P. Nolan is a CNGA GRASS Recipient,  Justin C. Luong is a CNGA Director & former GRASS Recipient

This research article examines the impact of seed sourcing strategies on the success of restoration projects using the grass species Stipa pulchra. Using seeds from seven populations from Central (Santa Cruz) to Southern California (Santa Rosa Plateau), the study finds that regional admixture seed sourcing may be more beneficial for increasing resilience to environmental variation compared to restricting seed sourcing to local populations. 

The findings regarding the performance of local populations of Stipa pulchra compared to non-local populations in the common gardens showed that local populations did not consistently outperform non-local populations in the Santa Cruz common garden. The study found that there was no clear benefit to using local seed sources to establish native populations of Stipa pulchra, and that sourcing from a population close to the common garden did not improve the performance of the plants.

The distance between the seed source location and the common gardens did not have a significant effect on the growth of Stipa pulchra in 2018 or 2019 in any common garden.

The suggested seed-sourcing strategy for grassland restoration in the context of climate change, based on the study's findings, is to consider using a regional admixture provenancing approach that incorporates seed from populations adapted to different climatic conditions. This approach may help improve establishment success under climate variability, such as year-to-year variation in rainfall.


Oram, N. J., Ingrisch, J., Bardgett, R. D., Brennan, F., Dittmann, G., Gleixner, G., Illmer, P., Praeg, N., & Bahn, M. (2023). Drought intensity alters productivity, carbon allocation and plant nitrogen uptake in fast versus slow grassland communities. Journal of Ecology, 111, 1681–1699. https://doi.org/10.1111/1365-2745.14136

Summary: This research article investigates the effects of drought intensity on plant productivity, carbon allocation, and nitrogen uptake with contrasting resource acquisition strategies (fast- vs slow-strategy). The study found that slow-strategy plant communities were more drought resistant than fast-strategy communities, with different thresholds of plant productivity at peak drought. The study also observed shifts in carbon and nitrogen dynamics in the plant-soil system during and after drought, with implications for plant community recovery trajectories.

Overall, the implications of drought intensity on carbon and nitrogen dynamics in grassland ecosystems include changes in plant productivity, carbon uptake, and nitrogen uptake. Understanding these dynamics is crucial for predicting and managing the impacts of drought on grassland ecosystems.

Joseph T. Smith, Brady W. Allred, Chad S. Boyd, Kirk W. Davies, Andrew R. Kleinhesselink, Scott L. Morford, David E. Naugle. 2023.  Fire needs annual grasses more than annual grasses need fire. Biological Conservation 286. 110299. ISSN 0006-3207. https://doi.org/10.1016/j.biocon.2023.110299.

This article discusses the impact of invasive annual grasses on sagebrush ecosystems in the U.S. Great Basin. It challenges the commonly held belief that fire is necessary for the spread of invasive annual grasses, finding that the invasion, degradation, and domination of native plant communities by annual grasses can progress independently of fire.

The overemphasis on fire as the primary driver of ecosystem transformation in the Great Basin has led to a reactive management approach, where wildfire has become the default point of intervention in the grass-fire cycle. This highlights the importance of developing and implementing proactive management strategies that specifically target the invasion and spread of annual grasses, rather than solely focusing on fire management.


2022 Publications

Science Volume 377, Issue 6606, 5 August 2022

Special Section on Grasslands https://www.science.org/toc/science/377/6606

Karen D. Holl, Justin C. Luong, Pedro H.S. Brancalion, 2022. Overcoming biotic homogenization in ecological restoration. Trends in Ecology & Evolution, ISSN 0169-5347. https://doi.org/10.1016/j.tree.2022.05.002.

(https://www.sciencedirect.com/science/article/pii/S0169534722001136)

Abstract: Extensive evidence shows that regional (gamma) diversity is often lower across restored landscapes than in reference landscapes, in part due to common restoration practices that favor widespread species through selection of easily-grown species with high survival and propagation practices that reduce genetic diversity. We discuss approaches to counteract biotic homogenization, such as reintroducing species that are adapted to localized habitat conditions and are unlikely to colonize naturally; periodically reintroducing propagules from remnant populations to increase genetic diversity; and reintroducing higher trophic level fauna to restore interaction networks and processes that promote habitat heterogeneity. Several policy changes would also increase regional diversity; these include regional coordination amongst restoration groups, financial incentives to organizations producing conservation-valued species, and experimental designations for rare species introductions.

Keywords: gamma-diversity; diversity conservation; habitat heterogeneity; reintroduction; rewilding

Luong, J. C., & Loik, M. E. (2022). Adjustments in physiological and morphological traits suggest drought-induced competitive release of some California plants. Ecology and Evolution, 12, e8773. https://doi.org/10.1002/ece3.8773

Short Summary by the Author: In the greenhouse, we tested the role of drought and invasive plant competition on the growth, morphology and physiology of five native species: Bromus carinatus, Diplacus aurantiacus, Lupinus nanus, Stipa pulchra, Sidalcea malviflora. The non-native species that we used for competition were Festuca bromoides, Carduus pycnocephalus, Geranium dissectum, Medicago polymorpha, Raphanus sativus. We most species focused adjustments in plant growth and physiology for drought, and not as much for competition, implying that drought may be a stronger factor for impacting planting outcomes compared to plant invasion. We also found that certain plants potentially experienced competitive release from non-native species during drought, due to lower performance of non-native species in drought. This may indicate that they are ideal to use in invaded areas that may experience drought, and these species were Bromus carinatus and Diplacus aurantiacus. We also found that Stipa pulchra and Sidalcea malviflora were typically not strongly affected by drought. Lupinus nanus was sensitive to both drought and competition which means it may be better to sow and introduce in wetter years, areas with supplemental irrigation or those with lower competition. 

Also see this file of an awesome illustrated summary that Vicky Cheung created about this study.


Luong, JC. (2022), Nonperiodic grassland restoration management can promote native woody shrub encroachment. Restor Ecol e13650. https://doi.org/10.1111/rec.13650

Supporting information at: http://onlinelibrary.wiley.com/doi/10.1111/rec.13650/suppinfo

Short Summary from the Author:Essentially, I noticed that Baccharis pilularis was heavily recruiting in plots that were restored through one-off planting and weeding actions and no management thereafter. There was a huge flowering Baccharis pilularis equidistant to both treatment plot types but I noticed that it was only recruiting into the one that had been "restored". I took some data and found those restoration actions promoted woody invasion by Baccharis pilularis in high abundance, and there were zero recruits on those without similar restoration actions. Drought reduced the recruitment, but those that recruited still achieved similar cover. This is nothing super new, but it helps support that restored grasslands are prone to woody invasion and transformation by Baccharis pilularis if no continued management occurs, similar to what many practitioners have mentioned to me. Potential implications are that woody species may need to be monitored and targeted if the ultimate goal of the restoration project is to maintain native grasslands vs shrublands.


2021 Publications

Conservation Grazing 

Michaels, J. S., Tate, K. W., & Eviner, V. T. (2021). Vernal pool wetlands respond to livestock grazing, exclusion and reintroduction. Journal of Applied Ecology, 00, 1– 12. https://doi.org/10.1111/1365-2664.14001

Synthesis and applications. Our results show that reintroduced grazing at moderate stocking rates can have significant effects on plant communities after just 2 years and can increase native cover more quickly than overall diversity. Our findings suggest that the negative effects of long-term grazing exclusion in vernal pools may be reversible, but that land managers interested in restoring diversity should plan to monitor beyond the first two years of grazing reintroduction.

Read more about this research: Once perceived as a problem conservation grazing by cattle a boon to vernal pools, University of California Division of Agriculture and Natural Resources


Barry, Sheila & Huntsinger, Lynn. (2021). Rangeland Land-Sharing, Livestock Grazing’s Role in the Conservation of Imperiled Species. Sustainability. 13. 4466. 10.3390/su13084466.

Land sharing, conserving biodiversity on productive lands, is globally promoted. Much of the land highest in California’s biodiversity is used for livestock production, providing an opportunity to understand land sharing and species conservation. A review of United States Fish and Wildlife Service listing documents for 282 threatened and endangered species in California reveals a complex and varied relationship between grazing and conservation. According to these documents, 51% or 143 of the federally listed animal and plant species are found in habitats with grazing. While livestock grazing is a stated threat to 73% (104) of the species sharing habitat with livestock, 59% (85) of the species are said to be positively influenced, with considerable overlap between species both threatened and benefitting from grazing. Grazing is credited with benefiting flowering plants, mammals, insects, reptiles, amphibians, fish, crustaceans, and bird species by managing the state’s novel vegetation and providing and maintaining habitat structure and ecosystem functions. Benefits are noted for species across all of California’s terrestrial habitats, except alpine, and for some aquatic habitats, including riparian, wetlands, and temporary pools. Managed grazing can combat anthropomorphic threats, such as invasive species and nitrogen deposition, supporting conservation-reliant species as part of land sharing.


Nolan, M., Dewees, S. and Ma Lucero, S. (2021), Identifying effective restoration approaches to maximize plant establishment in California grasslands through a meta-analysis. Restoration Ecology, 29: e13370.  https://doi.org/10.1111/rec.13370

One reason restoration often fails is a lack of native plant establishment. Establishment failure can occur due to a lack of dispersal into the site, inappropriate abiotic conditions for plant survival, or biotic interactions that reduce germination rates or seedling survival. Understanding what is limiting establishment is critical for restoration success. Unfortunately, there has been little synthetic work to identify which restoration practices (e.g., direct seeding, weeding, irrigation) are most likely to improve plant establishment. Here we used a meta-analytical approach to determine what practices are most likely to enhance establishment success in grassland communities in California. We explored the impact of these practices on whole communities as well as on different functional groups (grass and forb). We categorized techniques by the constraint they addressed (dispersal, abiotic, or biotic) to determine which restoration practices are more likely to improve plant establishment. We found that addressing the dispersal filter is the best way to improve plant establishment regardless of the functional group. This suggests that most native plants targeted in restoration are seed limited. While we were able to identify restoration techniques that improve plant establishment, our synthesis revealed that there were insufficient numbers of studies and a lack of uniformity between treatments that hampered efforts to make broad conclusions. If we can expand research to more ecosystems, and use standardized treatments within each ecosystem, the restoration community could synthesize research more efficiently, which would inform our ultimate goal of increasing the success of restoration.


Funk, J. L. (2021). Revising the trait-based filtering framework to include interacting filters: Lessons from grassland restoration. Journal of Ecology, 109, 3466– 3472. https://doi.org/10.1111/1365-2745.13763

Abstract

  1. A trait-based framework of community assembly holds great promise for directing ecological restoration, both for selecting species with desirable traits and for manipulating community factors to enhance plant establishment and persistence. Dispersal, abiotic and biotic factors ‘filter’ species into local communities based on their traits, but interactions among these filters may complicate the use of trait-based assembly models. In this paper, I review recent studies that apply community-based theory to grassland restoration and propose a framework for incorporating interacting ecological filters into restoration design.
  2. Dispersal limits restoration success in many grassland communities while others are simultaneously limited by dispersal, environmental factors and biotic interactions. Furthermore, the relative importance of ecological filters can change over space and time. Species also differentially respond to filter manipulations which suggests that trait–environment relationships should be used to generate planting recommendations based on optimal trait values for interacting filters at a given site.
  3. Synthesis. A better understanding of how traits interact with dynamic community assembly filters will allow for site-specific management recommendations, resulting in restored communities that are resilient to a range of filter modifications including climate change, invasion by non-native species, and altered disturbance regimes.



Luong JC, Holl KD, Loik ME. 2021. Leaf traits and phylogeny explain plant survival and community dynamics in response to extreme drought in a restored coastal grassland. Journal of Applied Ecology 2021;00:1–11. https://doi. org/10.1111/1365-2664.13909


See the illustrated version of this article in Grasslands Fall 2020. Vol 30 (4):12-15. Illustrated by Leslie Goren. Grasslands


Holl, K.D., Lesage, J.C., Adams, T., Rusk, J., Schreiber, R.D. and Tang, M. (2021), Vegetative spread is key to applied nucleation success in non-native-dominated grasslands. Restor Ecol, 29: e13330. https://doi.org/10.1111/rec.13330

Abstract Excerpts: Applied nucleation (i.e. planting vegetation patches) is a restoration strategy that better recreates natural ecosystem heterogeneity and requires fewer resources compared to planting the entire area. Our results suggest that applied nucleation can be an effective restoration strategy at a small scale in non-native-dominated grasslands for species that spread vegetatively.


LaForgia, M. L., Harrison, S. P., and Latimer, A. M.. 2020. Invasive species interact with climatic variability to reduce success of natives. Ecology 101(6):e03022. 10.1002/ecy.3022

Accepted manuscript available on Google Scholar (pre final editing) https://ecoevorxiv.org/hqwk4/download?format=pdf

Abstract. Plants have evolved resource-conservative and resource-acquisitive strategies to deal with variability in rainfall, but interactions with dominant invasive species may undermine these adaptations. To investigate the relative effect of invaders on species with these two strategies, we manipulated rainfall and invasive grass presence and measured demographic rates in three resource-acquisitive and three resource-conservative native annual forbs. We found that invasive grasses were harmful to all of the target species, but especially the resource-acquisitive ones, and that these effects were stronger under experimental drought. Invasive grass presence under drought lowered per capita population growth rates of acquisitive natives through increased mortality and decreased seed set. While invasive grasses also decreased per capita growth rates of resource-conservative natives, they did so by increasing mortality under experimental watering and by limiting the production of seed under experimental drought. 

Amelia A. Wolf AA, Funk JL, Selmants PC, Morozumi CN, Hernández DL,  Pasari JR, Zavaleta ES. 2021. Trait-based filtering mediates the effects of realistic biodiversity losses on ecosystem functioning. Proceedings of the National Academy of Sciences Jun 2021, 118 (26) e2022757118; DOI: 10.1073/pnas.2022757118. Accessed July 27, 2021.

Significance

We present multiyear results from a realistic biodiversity loss experiment, examining how two key ecosystem functions (productivity and invasion resistance) responded to randomized and realistic (drought-driven) species losses across years with high yearly climatic variation. We show that realistic low-diversity communities do not always have high functioning under the conditions that drove species loss, indicating a disconnect between functional response and effect traits. Our results also suggest that the functional stability of ecosystems experiencing realistic biodiversity declines could be lower than previous studies of randomized species losses have indicated. Understanding how functional traits interact with increasingly variable climatic conditions is critical to predicting ecosystem responses to the concurrent challenges of species loss and climate change.

For more information see UC Santa Cruz Newscenter article: New study shows how loss of drought-sensitive species could affect health of California grasslands. By Allison Arteaga Soergel, June 21, 2021.


Hernandez, E, EA Shaw, L Aoyama, A Brambila, C Niederer, SB Weiss, LM Hallett. 2021. Fire versus grazing as tools to restore serpentine grasslands under global change. Restoration Ecology.

"I think the resilience of the system lies in the diversity of annuals.". -Stu Weiss

Abstract. Grassland restoration in a world of change—including nitrogen deposition and invasion—requires solutions that can be sustained and effective in the face of novel drivers. This challenge is amplified in systems characterized by high spatiotemporal variability,as management to address novel drivers may affect a system differently across its range of variability. California serpentine grasslands epitomize this challenge: they host a high diversity of native species, are characterized by temporal and spatial variability, and are experiencing atmospheric nitrogen deposition that leads to a conversion from native annual forbs to non-native annual grasses. Here, we test the interactive effects of grazing and fire to restore native serpentine species following annual grass invasion and litter accumulation. We assess management outcomes (burned-grazed, burned-ungrazed, and unburned-ungrazed) using a long-term (2004–2012) monitoring dataset. A 2004 wildfire led to a reduction in annual grasses and a transient increase in native species (forb) richness. In 2008, cattle grazing was reintroduced and crossed with the burn legacy, which sustained post-fire diversity and eventually led to native recovery in unburned areas, with a caveat that a period of high precipitation promoted the growth of annual grasses. Our study indicates that short-term management, such as fire, can promote native forb recovery in invaded serpentine grassland communities, but that ongoing treatments like grazing are necessary to maintain restoration outcomes. We speculate that this recovery may be due to the presence of a robust native seed bank, which may persist despite a period of annual grass conversion.

2020 Publications

Nerlekar AN, JW Veldman (2020) High plant diversity and slow assembly of old-growth grasslands. PNAS. Vol 117 No. 31: 18550-18556. https://www.pnas.org/doi/full/10.1073/pnas.1922266117

Full Text PDF: https://web.archive.org/web/20210427194720id_/https://www.pnas.org/content/pnas/117/31/18550.full.pdf

Earth’s ancient grasslands and savannas—hereafter old-growth grasslands—have long been viewed by scientists and environmental policymakers as early successional plant communities of low conservation value. Challenging this view, emerging research suggests that old-growth grasslands support substantial biodiversity and are slow to recover if destroyed by human land uses (e.g., tillage agriculture, plantation forestry). But despite growing interest in grassland conservation, there has been no global test of whether old-growth grasslands support greater plant species diversity than secondary grasslands (i.e., herbaceous communities that assemble after destruction of old-growth grasslands). Our synthesis of 31 studies, including 92 timepoints on six continents, found that secondary grasslands supported 37% fewer plant species than old-growth grasslands (log response ratio = −0.46) and that secondary grasslands typically require at least a century, and more often millennia (projected mean 1,400 y), to recover their former richness. Young (<29 y) secondary grasslands were composed of weedy species, and even as their richness increased over decades to centuries, secondary grasslands were still missing characteristic old-growth grassland species (e.g., long-lived perennials). In light of these results, the view that all grasslands are weedy communities, trapped by fire and large herbivores in astate of arrested succession, is untenable. Moving forward, we suggest that ecologists should explicitly consider grassland assembly time and endogenous disturbance regimes in studies of plant community structure and function. We encourage environmental policymakers to prioritize old-growth grassland conservation and work to elevate the status of old-growth grasslands, alongside old-growth forests, in the public consciousness.


Griffin SR, B Bruninga-Socolar, J Gibbs (2020). Bee communities in restored prairies are structured by landscape and management, not local floral resources. Basic and Applied Ecology 50: 144-154.

Abstract. Restored habitats require long-term management to maintain biodiversity and ensure ecosystem functions. Management strategies are often developed for plant communities, including through seeding and disturbance management, but these actions are taken with a focus on plant dynamics and with little knowledge of the effects on non-plant organisms. Wild bees are often expected to respond to such management actions via their effects on local floral resource availability, but management may also affect bees by altering survival and nesting independently of plant community responses. Working in restoration plantings within a large, actively managed tallgrass prairie preserve, we separated the effects of management and landscape context on bee community abundance and richness from the effects of these covariates on bees mediated through the abundance and richness of the local flowering plant community. We found that bees responded primarily to disturbance management (via bison) and the amount of prairie and forest habitat in the landscape, indicating that across landscapes with relatively abundant flowers and nest-sites, these landscape-level resources are more important than local floral resources for structuring bee communities. In contrast, floral communities responded to restoration age and prescribed burning. Because bees respond to different factors and at a different landscape scale than local plant communities, we conclude that management designed for plants is not sufficient for pollinators. restoration may therefore require targeted habitat design and management to successfully restore functionally important animals.


LaManna, JA, Burkle, LA, Belote, RT, Myers, JA. Biotic and abiotic drivers of plant–pollinator community assembly across wildfire gradients. J Ecol. 2020; 00: 1– 14. https://doi.org/10.1111/1365-2745.13530. OPEN ACCESS

Link to Eureka Alert! Summary article about the research, November 25, 2020. In fire-prone West, plants need their pollinators -- and vice versa. WASHINGTON UNIVERSITY IN ST. LOUIS.

Summary (condensed from Eureka Alert!)

Plant-pollinator linkages are more are important in disturbed or burned landscapes and are important to consider in restoring ecosystems where natural wildfire regimes have been altered. In burned landscapes, plant-pollinator interactions are generally as important or more important than any other factor in determining the composition of species present. The importance of flowering-plant species in determining the composition of pollinator species doubled to quadrupled following wildfire. And, the importance of pollinators in determining plant composition nearly doubled following wildfire.

This study looked at field inventories of plants and pollinators at 152 plots in Montana representing a wildfire gradient including plots with no recent wildfire (unburned), mixed-severity wildfire and high-severity wildfire.

The study found that the number of pollinators (individual bees, flies and butterflies), and the flowering plants they frequent, were higher in parts of the landscape that had burned, as opposed to those that hadn't burned.

Increases were even greater in areas that had experienced mixed-severity wildfire, which leaves some vegetation intact in a mosaic of habitat types, as opposed to high-severity wildfire, which largely removes all vegetation and can damage the soil and seed bank. For example, flowering-plant abundances increased more than 10-fold in mixed-severity

Newbold, T., Oppenheimer, P., Etard, A. et al. Tropical and Mediterranean biodiversity is disproportionately sensitive to land-use and climate change. Nat Ecol Evol (2020). https://doi.org/10.1038/s41559-020-01303-0

Abstract.

Global biodiversity is undergoing rapid declines, driven in large part by changes to land use and climate. Global models help us to understand the consequences of environmental changes for biodiversity, but tend to neglect important geographical variation in the sensitivity of biodiversity to these changes. Here we test whether biodiversity responses to climate change and land-use change differ among biomes (geographical units that have marked differences in environment and species composition). We find the strongest negative responses to both pressures in tropical biomes and in the Mediterranean. A further analysis points towards similar underlying drivers for the sensitivity to each pressure: we find both greater reductions in species richness in the types of land use most disturbed by humans and more negative predicted responses to climate change in areas of lower climatic seasonality, and in areas where a greater proportion of species are near their upper temperature limit. Within the land most modified by humans, reductions in biodiversity were particularly large in regions where humans have come to dominate the land more recently. Our results will help to improve predictions of how biodiversity is likely to change with ongoing climatic and land-use changes, pointing toward particularly large declines in the tropics where much future agricultural expansion is expected to occur. This finding could help to inform the development of the post-2020 biodiversity framework, by highlighting the under-studied regions where biodiversity losses are likely to be greatest.:

Science Daily Summary Article: Mediterranean and tropical biodiversity most vulnerable to human pressures. Date: September 14, 2020


Stuble KL, Young TP. Priority Treatment Leaves Grassland Restoration Vulnerable to Invasion. Diversity. 2020; 12(2):71. Full Text. 

Abstract

Priority effects can be used to promote target species during restoration. Early planting can provide an advantage over later-arriving species, increasing abundance of these early-arrivers in restored communities. However, we have limited knowledge of the indirect impacts of priority effects in restoration. In particular, we do not understand how priority effects impact non-target species. Of particular conservation concern is how these priority effects influence establishment by non-native species. We use a field-based mesocosm experiment to explore the impacts of priority effects on both target and non-target species in California grasslands. Specifically, we seeded native grasses and forbs, manipulating order of arrival by planting them at the same time, planting forbs one year before grasses, planting grasses one year before forbs, or planting each functional group alone. While our study plots were tilled and weeded for the first year, the regional species pool was heavily invaded. We found that, while early-arrival of native grasses did not promote establishment of non-native species, giving priority to native forbs ultimately left our restoration mesocosms vulnerable to invasion by non-native species. This suggests that, in some cases, establishment of non-native species may be an unintended consequence of using priority treatments as a restoration tool


Supporting evidence varies for rangeland management practices that seek to improve soil properties and forage production in California. California Agriculture 74(2):101-111. https://doi.org/10.3733/ca.2020a0015

Published online June 09, 2020

Authors

Chelsea J. Carey, Point Blue Conservation Science

Kelly Gravuer, Arizona State University and The Nature Conservancy
Sasha Gennet, The Nature Conservancy
Dillon Osleger, National Center for Ecological Analysis and Synthesis
Stephen A. Wood, The Nature Conservancy and Yale University

Publication Information

PDF  |  Citation  |  Permissions


Groves, A.M., Bauer, J.T. & Brudvig, L.A. Lasting signature of planting year weather on restored grasslands. Sci Rep 10, 5953 (2020). Open Access. Published: 06 April 2020.  https://doi.org/10.1038/s41598-020-62123-7. PDF

Abstract

Ecological restoration — the rebuilding of damaged or destroyed ecosystems — is a critical component of conservation efforts, but is hindered by inconsistent, unpredictable outcomes. We investigated a source of this variation that is anecdotally suggested by practitioners, but for which empirical evidence is rare: the weather conditions during the first growing season after planting. The idea of whether natural communities face long-term consequences from conditions even many years in the past, called historical contingency, is a debated idea in ecological research. Using a large dataset (83 sites) across a wide geographic distribution (three states), we find evidence that precipitation and temperatures in the planting year (2–19 years before present) affected the relative dominance of the sown (native target species) and non-sown (mostly non-native) species. We find strong support for lasting planting year weather effects in restored tallgrass prairies, thereby supporting the historically contingent model of community assembly in a real-world setting.

Science Daily Article April 6, 2020


Buzhdygan, O.Y., Meyer, S.T., Weisser, W.W. et al. Biodiversity increases multitrophic energy use efficiency, flow and storage in grasslands. Nat Ecol Evol 4, 393–405 (2020). https://doi.org/10.1038/s41559-020-1123-8

Abstract

The continuing loss of global biodiversity has raised questions about the risk that species extinctions pose for the functioning of natural ecosystems and the services that they provide for human wellbeing. There is consensus that, on single trophic levels, biodiversity sustains functions; however, to understand the full range of biodiversity effects, a holistic and multitrophic perspective is needed. Here, we apply methods from ecosystem ecology that quantify the structure and dynamics of the trophic network using ecosystem energetics to data from a large grassland biodiversity experiment. We show that higher plant diversity leads to more energy stored, greater energy flow and higher community-energy-use efficiency across the entire trophic network. These effects of biodiversity on energy dynamics were not restricted to only plants but were also expressed by other trophic groups and, to a similar degree, in aboveground and belowground parts of the ecosystem, even though plants are by far the dominating group in the system. The positive effects of biodiversity on one trophic level were not counteracted by the negative effects on adjacent levels. Trophic levels jointly increased the performance of the community, indicating ecosystem-wide multitrophic complementarity, which is potentially an important prerequisite for the provisioning of ecosystem services.

This research is from the Jena Experiment, a long-term grassland experiment in Germany looking at biodiveristy-ecosystem functioning. 

Would you like the significance in plain words? Read the Science Daily summary of this article. 


Cristian Román-Palacios, John J. Wiens. 2020. Recent responses to climate change reveal the drivers of species extinction and survival. Proceedings of the National Academy of Sciences Feb 2020, 117 (8) 4211-4217; DOI: 10.1073/pnas.1913007117

Abstract

Climate change may be a major threat to biodiversity in the next 100 years. Although there has been important work on mechanisms of decline in some species, it generally remains unclear which changes in climate actually cause extinctions, and how many species will likely be lost. Here, we identify the specific changes in climate that are associated with the widespread local extinctions that have already occurred. We then use this information to predict the extent of future biodiversity loss and to identify which processes may forestall extinction. We used data from surveys of 538 plant and animal species over time, 44% of which have already had local extinctions at one or more sites. We found that locations with local extinctions had larger and faster changes in hottest yearly temperatures than those without. Surprisingly, sites with local extinctions had significantly smaller changes in mean annual temperatures, despite the widespread use of mean annual temperatures as proxies for overall climate change. Based on their past rates of dispersal, we estimate that 57–70% of these 538 species will not disperse quickly enough to avoid extinction. However, we show that niche shifts appear to be far more important for avoiding extinction than dispersal, although most studies focus only on dispersal. Specifically, considering both dispersal and niche shifts, we project that only 16–30% of these 538 species may go extinct by 2070. Overall, our results help identify the specific climatic changes that cause extinction and the processes that may help species to survive.


Science Daily Summary: University of Arizona. "One-third of plant and animal species could be gone in 50 years." ScienceDaily. ScienceDaily, 12 February 2020. <www.sciencedaily.com/releases/2020/02/200212150146.htm>.


Stuart Wagenius, Jared Beck, and Gretel Kiefer. 2020. Fire synchronizes flowering and boosts reproduction in a widespread but declining prairie species. PNAS. January 27, 2020. From the Echinacea Project.

Science Daily Interview with Author 

Significance

We address a critical conservation concern: the loss of native plant species in fire-dependent ecosystems. Reduced fire frequency in ecosystems such as the North American prairie contributes to local extinctions. The leading hypothesis is that woody and large herbaceous species outcompete other species in the absence of fire. However, alternative mechanisms have not been investigated. In our 21-y study of a model prairie plant, Echinacea angustifolia, we demonstrate that fires synchronize reproduction, leading to increased mating opportunities and improved reproduction. Fire synchronizes flowering both among and within years, nearly doubling seed production. These findings demonstrate a potentially widespread mechanism by which fire can enhance plant reproduction, promote population growth, and maintain plant diversity in fire-dependent ecosystems worldwide.



Bucharova, A., F Krahulec. 2020. “Native seed addition as an effective tool for post-invasion restoration.” Basic and Applied Ecology 43 (March 2020): 34-41.

https://www.sciencedirect.com/science/article/pii/S1439179119302944

Abstract

Invasive plant species reduce biodiversity, alter ecosystem processes, and cause economic losses. Control of invasive plants is therefore highly desired by land managers and policy makers. However, invasive plant control strategies frequently fail, partly because management often concentrates only on the eradication of invasive plants and not on revegetation with native species that use the available resources and prevent reinvasion. In this study, we focused on the intracontinental invader Rumex alpinus L., which was introduced by humans from the Alps to the lower mountains of Central Europe, where it has spread to semi-natural meadows, suppresses local biodiversity, and reduces the quality of hay used as cattle fodder. The species can be effectively removed using herbicide, but this leaves behind a persistent seed bank. Without further treatment, the invader rapidly regenerates and reinvades the area. We supplemented the herbicide treatment by adding the seeds of native grasses. Addition of native-seed effectively suppressed the regeneration of the invader from the seed bank, reduced its biomass, and consequently, prevented massive reinvasion. While the invader removal was successful, the restored community remained species-poor because the dense sward of native grasses blocked the regeneration of native forbs from the seed bank. Nevertheless, the addition of native seed proved to be an effective tool in preventing reinvasion after the eradication of the invasive plant.


Zirbel, C. R., and Brudvig, L. A.. 2020. "Trait–environment interactions affect plant establishment success during restoration." Ecology 101(3):e02971. 10.1002/ecy.2971


Abstract [Full text is available for this article]

Establishment and persistence are central to community assembly and are determined by how traits interact with the environment to determine performance (trait–environment interactions). Community assembly studies have rarely considered such trait–environment interactions, however, which can lead to incorrect inferences about how traits affect assembly. We evaluated how functional traits, environmental conditions, and trait–environment interactions structure plant establishment, as a measure of performance. Within 12 prairie restorations created by sowing 70 species, we quantified environmental conditions and counted individuals of each seeded species to quantify first‐year establishment. Three trait–environment interactions structured establishment. Leaf nitrogen interacted with herbivore pressure, as low leaf nitrogen species established relatively better under higher herbivory than species with high leaf nitrogen. Soil moisture interacted with root mass fraction (RMF), with low‐RMF species establishing better with low soil moisture and higher‐RMF species better on wetter soils. Specific leaf area (SLA) interacted with light availability, as low‐SLA species established better under high light conditions and high‐SLA species under low light conditions. Our work illustrates how community assembly can be better described by trait–environment interactions than correlating traits or environment with performance. This knowledge can assist species selection to maximize restoration success.



2019 Publications

Valliere, J.M., Balch, S., Bell, C., Contreras, C. and Hilbig, B.E. (2019), Repeated mowing to restore remnant native grasslands invaded by nonnative annual grasses: upsides and downsides above and below ground. Restoration Ecology, 27: 261 268. https://doi.org/10.1111/rec.12873

Abstract

California grasslands have been severely impacted by the invasion of nonnative annual grasses, which often limit restoration of this important ecosystem. In this study, we explored the use of mowing as a restoration tool for native perennial grasslands at the Santa Rosa Plateau Ecological Reserve in southern California. We sought to evaluate if, over time, mowing would reduce nonnative annual grass cover and benefit native species, especially the native bunchgrass Stipa pulchra. We hypothesized that repeated mowing, carefully timed to target nonnative annual grasses prior to seed maturation, would reduce nonnative seed inputs into the soil and eventually lead to diminished abundance of these species. We monitored vegetation in mowed and unmowed plots for 4 years, and conducted a seed bank study after 5 years to better understand the cumulative effects of mowing on native and nonnative seed inputs. Consistent with our hypotheses, we found that mowing successfully reduced nonnative annual grass cover and benefitted some native species, including S. pulchra. However, we also found that nonnative forb species showed progressive increases in mowed plots over time. We observed similar patterns of species composition in the soil seed bank. Together, these results suggest that mowing can be used to control nonnative annual grasses and increase the abundance of native bunchgrasses, but that this method may also have the unintended consequence of increasing nonnative forb species.


Beaury, E.M., Finn, J.T., Corbin, J.D., Barr, V. and Bradley, B.A. (2019), Biotic resistance to invasion is ubiquitous across ecosystems of the United States.Ecol Lett. doi:10.1111/ele.13446

https://onlinelibrary.wiley.com/doi/abs/10.1111/ele.13446

Abstract

The biotic resistance hypothesis predicts that diverse native communities are more resistant to invasion. However, past studies vary in their support for this hypothesis due to an apparent contradiction between experimental studies, which support biotic resistance, and observational studies, which find that native and non‐native species richness are positively related at broad scales (small‐scale studies are more variable). Here, we present a novel analysis of the biotic resistance hypothesis using 24 456 observations of plant richness spanning four community types and seven ecoregions of the United States. Non‐native plant occurrence was negatively related to native plant richness across all community types and ecoregions, although the strength of biotic resistance varied across different ecological, anthropogenic and climatic contexts. Our results strongly support the biotic resistance hypothesis, thus reconciling differences between experimental and observational studies and providing evidence for the shared benefits between invasive species management and native biodiversity conservation.


Kimberly J. Komatsu et al. 2019. Global change effects on plant communities are magnified by time and the number of global change factors imposed. PNAS. DOI: 10.1073/pnas.1819027116. Science Daily News Article 


Luong JC, Turner PL, Phillipson CN, Seltmann KC. 2019. Local grassland restoration affects insect communities. Ecological Entomology. https://doi.org/10.1111/een.12721

Abstract. 1. It is hypothesised that ecological restoration in grasslands can induce an

alternative stable state shift in vegetation. The change in vegetation influences insect

community assemblages and allows for greater functional redundancy in pollination and

refuge for native insect species.

2. Insect community assemblages at eight coastal California grassland sites were

evaluated. Half of these sites had undergone restoration through active revegetation

of native grassland flora and half were non-restored. Insects were collected from

Lupinus bicolor (Fabaceae) within 2 × 2-m2 plots in spring 2017. Lupinus bicolor is

a common native species that is used in California restoration projects, and home and

state landscaping projects.

3. Ordination demonstrated that insect community assemblages were different between

restored and non-restored sites. These differences were seen in insect functional groups

as well as taxa-specific differences and were found to be driven by environmental

characteristics such as non-native forb cover.

4. Functional redundancy of herbivores decreased at restored sites, while pollinators

became more redundant compared with non-restored sites. The assemblages of the

common species found at restoration sites contained more native insects than those found

at non-restored sites, including species such as Bombus vosnesenskii.

5. Local grassland restoration has the potential to induce an alternative stable state

change and affect insect community assemblages. Additionally, it was found that grassland

restoration can be a potential conservation tool to provide refugia for bumblebees

(Bombus), but additional studies are required to fully understand its broader applicability.


Uricchio, Lawrence H.  S. Caroline Daws, Erin R. Spear, and Erin A. Mordecai. 2019. Priority effects of nonhierarchical competition shape species composition in a complex grassland community. The American Naturalist 193 (2). 

Abstract. Niche and fitness differences control the outcome of competition, but determining their relative importance in invaded communities—which may be far from equilibrium—remains a pressing concern. Moreover, it is unclear whether classic approaches for studying competition, which were developed predominantly for pairs of interacting species, will fully capture dynamics in complex species assemblages. We parameterized a population-dynamic model using competition experiments of two native and three exotic species from a grassland community. We found evidence for minimal fitness differences or niche differences between the native species, leading to slow replacement dynamics and priority effects, but large fitness advantages allowed exotics to unconditionally invade natives. Priority effects driven by strong interspecific competition between exotic species drove single-species dominance by one of two exotic species in 80% of model outcomes, while a complex mixture of nonhierarchical competition and coexistence between native and exotic species occurred in the remaining 20%. Fungal infection, a commonly hypothesized coexistence mechanism, had weak fitness effects and is unlikely to substantially affect coexistence. In contrast to previous work on pairwise outcomes in largely native-dominated communities, our work supports a role for nearly neutral dynamics and priority effects as drivers of species composition in invaded communities.

More about this research: Knowing how invasive grasses compete could help preserve native grasslands, according to Stanford researchers. Stanford News, January 15, 2019, by Sophie Bates 

2018 Publications

Dass, Pawlok, Benjanim Z Houlton, Yingping Want and David Warlind. 2018. Grasslands more reliable carbon sink than trees. Environmental Research Letters, Volume 13, Number 7. PDF. 


LaForgia, Marina L.,  Marko J. Spasojevic,  Erica J. Case,  Andrew M. Latimer,  Susan P. Harrison. Seed banks of native forbs, but not exotic grasses, increase during extreme drought. (2018). Ecology 99 (4):896-903. 

Abstract: Extreme droughts such as the one that affected California in 2012–2015 have been linked to severe ecological consequences in perennial‐dominated communities such as forests. In annual communities, drought impacts are difficult to assess because many species persist through facultative multiyear seed dormancy, which leads to the development of seed banks. Impacts of extreme drought on the abundance and composition of the seed banks of whole communities are little known. In 80 heterogeneous grassland plots where cover is dominated by ~15 species of exotic annual grasses and diversity is dominated by ~70 species of native annual forbs, we grew out seeds from soil cores collected early in the California drought (2012) and later in the multiyear drought (2014), and analyzed drought‐associated changes in the seed bank. Over the course of the study we identified more than 22,000 seedlings to species. We found that seeds of exotic annual grasses declined sharply in abundance during the drought while seeds of native annual forbs increased, a pattern that resembled but was even stronger than the changes in aboveground cover of these groups. Consistent with the expectation that low specific leaf area (SLA) is an indicator of drought tolerance, we found that the community‐weighted mean SLA of annual forbs declined both in the seed bank and in the aboveground community, as low‐SLA forbs increased disproportionately. In this system, seed dormancy reinforces the indirect benefits of extreme drought to the native forb community.

Eviner, Valarie and Carolyn Malmstrom.  and Carolyn Malmstrom. 2018. California's native perennial grasses provide strong suppression of goatgrass and medusahead. Grasslands 28(1): 3-6. 

California's native grasses have the ability to suppress noxious weeds and prevent them from dominating grasslands when naturalized annual exotics, which can out-compete native grasses, are controlled. 


Maresh Nelson, S.B., Coon, J.J., Duchardt, C.J. et al. Contrasting impacts of invasive plants and human-altered landscape context on nest survival and brood parasitism of a grassland bird. Landscape Ecol 33, 1799–1813 (2018). https://doi.org/10.1007/s10980-018-0703-3

Significance

Controlling tall fescue within grasslands could benefit birds by increasing nest success.

This study found that nest survival declined with increasing tall fescue cover at nest sites for a Midwestern small grassland bird called a dickcissel. Tall fescue changes the vegetation structure making it less diverse to mat-like.


The researchers identified all the plants within a 16-foot radius of every dickcissel nest they found - all 477 of them - and the more tall fescue there was, the greater the chances of nest failure. No other plant in their surveys had significant predictive power.

Read the Eureka Alert Article Summary: Invasive forage grass leads to grassland bird decline/ October 19, 2018.


2017 Publications

Balachowski, Jennifer A. and Florence A. Volaire. 2017. Implications of plant functional traits and drought survival strategies for ecological restoration. Journal of Applied Ecology (online) DOI: 10.1111/1365-2664.12979.

The authors measured seasonal growth and foliar and root functional traits under non-limiting water conditions, followed by recovery after severe drought in four populations of Elymus glaucus.  Dehydration tolerance and summer dormancy were associated with a more conservative strategy (lower productivity, lower Specific Leaf Area and Specific Root Length), as well as earlier reproductive phenology. The authors discuss the implication that restored populations with superior drought survival may therefore be less competitive, and recommend further investigation to inform plant materials selection protocols and management practices

Gennet, Sasha, Erica Spotswood, Michele Hammond, James W. Bartolome. 2017. Livestock grazing supports native plants and songbirds in a California annual grassland. PLoS ONE 12(6) (June 14, 2017) : e0176367. https://doi.org/10.1371/journal.pone.0176367.

An 8-year study in central California finds livestock grazing can be compatible with or support grassland bird conservation. --Michele Hammond is a current serving on the CNGA Board of Directors. 


Hungate, Bruce A. , Edward B. Barbier, Amy W. Ando, Samuel P. Marks, Peter B. Reich, Natasja Van Gestel, David Tilman, Johannes M. H. Knops, David U. Hooper, Bradley J. Butterfield, Bradley J. Cardinale. 2017.  The economic value of grassland species for carbon storage. Science Advances: E1601880. 

Biodiversity confers economic value by enhancing carbon storage in grasslands, an economic argument for biodiversity conservation.

Vaughn, K.J., Biel, C., Clary, J.J. et al. 2011. California perennial grasses are physiologically distinct from both Mediterranean annual and perennial grasses.  Plant Soil (2011) 345: 37. https://doi.org/10.1007/s11104-011-0757-3


Gornish, Elise and Julea Shaw. 2017. Restoration manual for annual grassland systems in California. University of California ANR Publication 8575.  

Techniques to provide general direction to effectively improve grassland conditions in monetary and logistically feasible ways. Developed for practitioners of any experience level to inform grassland restoration, design and application. --Elise Gornish served on the CNGA Board of Directors in 2017. 


Kidd J, Manning P, Simkin J, Peacock S, Stockdale E. 2017. Impacts of 120 years of fertilizer addition on a temperate grassland ecosystem. PLoS ONE 12(3): e0174632. https://doi.org/10.1371/journal.pone.0174632

Abstract Except: Here, we examined the effects of long-term fertilizer addition on key ecosystem properties in a long-term grassland experiment (Palace Leas Hay Meadow) in which farmyard manure (FYM) and inorganic fertilizer treatments have been applied consistently for 120 years in order to characterize the experimental site more fully and compare ecosystem responses with those observed at other long-term and short-term experiments. FYM inputs increased soil organic carbon (SOC) stocks, hay yield, nutrient availability and acted as a buffer against soil acidification (>pH 5). In contrast, N-containing inorganic fertilizers strongly acidified the soil (<pH 4.5) and increased surface SOC stocks by increasing the C stored in the coarse (2.8 mm-200 μm) and fine (200–50 μm) fractions. Application of N fertilizers also reduced plant species richness and the abundance of forbs and legumes.


2015 Publications

Veldman, J.W., Buisson, E., Durigan, G., Fernandes, G.W., Le Stradic, S., Mahy, G., Negreiros, D., Overbeck, G.E., Veldman, R.G., Zaloumis, N.P., Putz, F.E. and Bond, W.J. (2015), Toward an old‐growth concept for grasslands, savannas, and woodlands. Frontiers in Ecology and the Environment, 13: 154-162. doi:10.1890/140270
ABSTRACT:
We expand the concept of “old growth” to encompass the distinct ecologies and conservation values of the world's ancient grass‐dominated biomes. Biologically rich grasslands, savannas, and open‐canopy woodlands suffer from an image problem among scientists, policy makers, land managers, and the general public, that fosters alarming rates of ecosystem destruction and degradation. These biomes have for too long been misrepresented as the result of deforestation followed by arrested succession. We now know that grassy biomes originated millions of years ago, long before humans began deforesting. We present a consensus view from diverse geographic regions on the ecological characteristics needed to identify old‐growth grasslands and to distinguish them from recently formed anthropogenic vegetation. If widely adopted, the old‐growth grassland concept has the potential to improve scientific understanding, conservation policies, and ecosystem management.

2013 Publications
Azzari, G., M.L. Goulden, and R.B. Rusu. 2013. Rapid characterization of vegetation structure with a Microsoft Kinect sensor. Sensors 13(2): 2384–98. doi:10.3390/s130202384.


Bai, S., Q. Xu, and Y. Qin. 2013. Vibration driven vehicle inspired from grass spike. Scientific Reports 3:1851.

ABSTRACT: Searching and detecting in some harsh environments such as collapsed buildings, pipes, small cracks are crucial for human rescue and industrial detection, military surveillance etc. However, the drawbacks of traditional moving modes of current vehicles make them difficult to perform such tasks. So developing some new vehicles is urgent. Here, we report aSetaria viridis spike's interesting behavior on a vibrating track, and inspired by that phenomena we develop a concept for cargo delivery, and give a detailed discussion about its working mechanism. This vehicle can move on a wide range of smooth and rough surfaces. Moreover, its climbing capability in tilted and even vertical smooth pipe is also outstanding. These features make it suitable for search-rescue, military reconnaissance, etc. Finally, this vehicle can be reduced into micro/nano-scale, which makes it would play an important role in target-drug delivery, micro-electromechanical systems (MEMS).


Chaplin-Kramer, R., and M.R. George. 2013. Effects of climate change on range forage production in the San Francisco Bay Area. PloS One, 8(3): e57723. doi:10.1371/journal.pone.0057723

Concilio, A. L., M.E. Loik, and J. Belnap. 2013. Global change effects on Bromus tectorum L. (Poaceae) at its high-elevation range margin. Global Change biology, 19(1): 161–72. doi:10.1111/gcb.12032

Evett, R, & J.W. Bartolome. (2013). Phytolith evidence for the extent and nature of prehistoric California grasslands. The Holocene, 23(11) 1-6. doi:10.1177/0959683613499056

Californian grasslands have been radically transformed by cultivation and exotic species invasions that began with European settlement in 1769. Because native species contribute only a small percentage of total vegetation cover at most grassland sites in California today, there has been a long-standing controversy regarding the extent and composition of prehistoric grasslands. We used phytolith analysis of soil samples from 153 spatially diverse grassland sites currently cultivated or dominated by exotic annual grasses to provide the first direct evidence that, except for areas near the coast, grass was generally a minor component of the vegetation. Most of the arid, non-wetland Central Valley and surrounding foothills probably had very low grass cover. Combining our data with historical accounts of vegetation encountered by early explorers in California and relict analysis of native species in existing grassland patches, it appears that while the extent of prehistoric grassland was probably similar to the current extent of exotic annual grassland, most areas were likely dominated by a highly diverse assemblage of herbaceous species, composed largely of annual forbs adapted to exploit local environmental heterogeneity. Phytolith analysis should be routinely performed in California to guide grassland restoration efforts. The technique can also be used to reconstruct prehistoric vegetation of highly altered grassland ecosystems worldwide.


Fernandez-Going, B. M., and S. Harrison. 2013. Effects of experimental water addition depend on grassland community characteristics. Plant Ecology 214:777–786.

ABSTRACT: Plant community biomass and composition on low-productivity soils, such as serpentine, may be more resistant to climate change because they host stress-tolerant species that may respond slowly to change. These communities also host a number of endemic taxa that are of special interest because of their narrow distributions. In a 3-year study, we experimentally tested the response of serpentine and non-serpentine communities to water addition in spring. We also compared the responses of endemics and generalists to water addition, with and without biomass (competitor) removal. In the non-serpentine grassland, peak biomass was significantly greater in the water addition plots compared with control plots, but this effect depended on the year. In the serpentine grassland, there was no effect of water addition on biomass. Survival, biomass, growth rates, and seed production of soil endemics and generalists were all significantly reduced by competition, but were unaffected by water addition. Overall, endemics tended to perform better in serpentine soil and generalists in non-serpentine soil, suggesting that soil is an important factor for the establishment and survival of endemics and generalists. For endemics, the effect of biomass removal was stronger on non-serpentine soil, but for generalists this effect was similar on both soils, indicating that competition can be important in low-resource habitats. In conclusion, our results suggest that low-fertility plant communities may be slow to respond to changes in precipitation compared to communities on more fertile soil.


Mariotte, P., C. Vandenberghe, P. Kardol, F. Hagedorn, and A. Buttler. 2013. Subordinate plant species enhance community resistance against drought in semi-natural grasslands. Journal of Ecology 101:763–773.

ABSTRACT: According to the insurance hypothesis, more diverse plant communities are more likely to be resistant to drought. Whilst many experiments have been carried out to determine the effects of plant diversity on plant community insurance, the results are still contradictory. Here, we conducted a drought experiment where we tested whether the presence of subordinate species increases plant community insurance. In Swiss Jura grassland, we combined a removal experiment of subordinate species with a summer drought event using rainout shelters. Plant community composition was determined after the drought and based on biomass measurements; we estimated resistance, recovery and resilience of the plant community for each combination of treatments. Moreover, to assess drought impacts on water-use efficiency (WUE), we analysed carbon isotope ratios (13C values) in plant leaves of two dominants and two subordinates collected at the end of the drought period. We showed that subordinate species are more resistant to drought and increased community resistance by enhancing their above-ground biomass production during the imposed drought. These patterns were associated with decreased competitiveness of dominant species whose biomass decreased during drought. Significant increase in 13C values in plant tissue under drought indicated a better WUE for the measured species. Interestingly, the WUE was significantly higher in plots where subordinates were removed. Recovery and resilience were not affected by the summer drought, but the absence of subordinates reduced overall above-ground biomass in both watered and drought plots. Synthesis. We demonstrated that, independent of plant diversity, the presence of drought-resistant subordinate species increases plant community insurance against drought and, hence, is important for the functioning of grassland ecosystems.


McAdoo, J. K., B. W. Schultz, and S.R.  Swanson. 2013. Aboriginal precedent for active management of sagebrush-perennial grass communities in the Great Basin. Rangeland Ecology & Management, 66(3), 241-253.


ABSTRACT: Until recently, most contemporary ecologists have ignored or diminished anecdotal historical accounts and anthropologists' reports about aboriginal fire in the Great Basin. Literature review shows that Indians practiced regular use of fire for many purposes, including the obvious reasons of increasing the availability of desired plants, maintaining habitats for animals used as food, and driving game during hunts. Historical accounts of prehistoric anthropogenic firing, inferences from fire-scar data, and data regarding annual production capability of representative sagebrush (Artemisia spp.)-perennial grass ecological sites indicate that prehistoric conditions were neither fuel- nor ignition-limited. According to many sources, this "active management" by Indians was widespread, significant, and more common than lightning-caused fires, resulting in mosaic vegetation patterns that subsequently moderated the behavior of "natural fires." This interaction between Indian-burning and lightning fires may have strongly influenced the pre-Euro-American settlement vegetation of the Great Basin. At the very least, the landscape was a patchwork of areas altered by aboriginal people and areas shaped primarily by bio-physical processes. Based on this prehistoric precedent, current historically unprecedented conditions (fuel load and exotic weed invasion threats), and predicted climate change, contemporary active management of sagebrush-perennial grass communities is paramount. Restoration measures should be scientifically based and tailored to achieve ecological resilience and functionality in specific sites. Prescribed fire is not always ecologically appropriate or judicious, especially in Wyoming big sagebrush (A. tridentata spp. wyomingensis) communities, so managers should consider using other alternatives where an intentional low severity disturbance is deemed necessary. Properly planned active management would disrupt fuel continuity for lightning fires, ensure ecological process and successional integrity, and benefit multiple uses on a landscape scale.


McSherry, M. E., and M. E. Ritchie. 2013. Effects of grazing on grassland soil carbon: a global review. Global change biology 19:1347–57.

ABSTRACT: Soils of grasslands represent a large potential reservoir for storing CO2, but this potential likely depends on how grasslands are managed for large mammal grazing. Previous studies found both strong positive and negative grazing effects on soil organic carbon (SOC) but explanations for this variation are poorly developed. Expanding on previous reviews, we performed a multifactorial meta-analysis of grazer effects on SOC density on 47 independent experimental contrasts from 17 studies. We explicitly tested hypotheses that grazer effects would shift from negative to positive with decreasing precipitation, increasing fineness of soil texture, transition from dominant grass species with C3 to C4 photosynthesis, and decreasing grazing intensity, after controlling for study duration and sampling depth. The six variables of soil texture, precipitation, grass type, grazing intensity, study duration, and sampling depth explained 85% of a large variation (+/- 150gm2yr1) in grazing effects, and the best model included significant interactions between precipitation and soil texture (P=0.002), grass type, and grazing intensity (P=0.012), and study duration and soil sampling depth (P=0.020). Specifically, an increase in mean annual precipitation of 600mm resulted in a 24% decrease in grazer effect size on finer textured soils, while on sandy soils the same increase in precipitation produced a 22% increase in grazer effect on SOC. Increasing grazing intensity increased SOC by 67% on C4-dominated and C4C3 mixed grasslands, but decreased SOC by an average 18% in C3-dominated grasslands. We discovered these patterns despite a lack of studies in natural, wildlife-dominated ecosystems, and tropical grasslands. Our results, which suggest a future focus on why C3 vs. C4-dominated grasslands differ so strongly in their response of SOC to grazing, show that grazer effects on SOC are highly context-specific and imply that grazers in different regions might be managed differently to help mitigate greenhouse gas emissions. 

Ryals, R., & Silver, W. L. 2013. Effects of organic matter amendments on net primary productivity and greenhouse gas emissions in annual grasslands. Ecological Applications 23(1): 46–59.

Setterfield, S. A, N. A Rossiter-Rachor, M. M. Douglas, L. Wainger, A. M. Petty, P. Barrow, I. J. Shepherd, and K. B. Ferdinands. 2013. Adding fuel to the fire: the impacts of non-native grass invasion on fire management at a regional scale. PloS One 8:e59144.

ABSTRACT: Widespread invasion by non-native plants has resulted in substantial change in fire-fuel characteristics and fire-behaviour in many of the world's ecosystems, with a subsequent increase in the risk of fire damage to human life, property and the environment. Models used by fire management agencies to assess fire risk are dependent on accurate assessments of fuel characteristics but there is little evidence that they have been modified to reflect landscape-scale invasions. There is also a paucity of information documenting other changes in fire management activities that have occurred to mitigate changed fire regimes. This represents an important limitation in information for both fire and weed risk management. Methodology/Principal Findings: We undertook an aerial survey to estimate changes to landscape fuel loads in northern Australia resulting from invasion by Andropogon gayanus (gamba grass). Fuel load within the most densely invaded area had increased from 6 to 10 t ha(-1) in the past two decades. Assessment of the effect of calculating the Grassland Fire Danger Index (GFDI) for the 2008 and 2009 fire seasons demonstrated that an increase from 6 to 10 t ha(-1) resulted in an increase from five to 38 days with fire risk in the 'severe' category in 2008 and from 11 to 67 days in 2009. The season of severe fire weather increased by six weeks. Our assessment of the effect of increased fuel load on fire management practices showed that fire management costs in the region have increased markedly (similar to 9 times) in the past decade due primarily to A. gayanus invasion. Conclusions/Significance: This study demonstrated the high economic cost of mitigating fire impacts of an invasive grass. This study demonstrates the need to quantify direct and indirect invasion costs to assess the risk of further invasion and to appropriately fund fire and weed management strategies.

Skaer, M. J., D. J. Graydon, and J. H. Cushman. 2013. Community-level consequences of cattle grazing for an invaded grassland: variable responses of native and exotic vegetation. Journal of Vegetation Science 24:332-343.

Vourlitis, G. L., & J.L. Kroon. 2013. Growth and resource use of the invasive grass, Pampasgrass (Cortaderia selloana), in response to nitrogen and water availability. Weed Science, 61(1): 117–125. doi:10.1614/WS-D-11-00220.1


2012

Bagne, K.; Ford, P.; Reeves, M. (November 2012). Grasslands and Climate Change. U.S. Department of Agriculture, Forest Service, Climate Change Resource Center. https://www.fs.usda.gov/ccrc/topics/grasslands-and-climate-change

From article text:

Model projections for grasslands are at a broad scale (1km) and give a sense of likely changes summarized below [6]:

Climate suitable for California valley grasslands is likely to have significant declines, shifting towards oak woodlands and desert scrub, along with a high proportion of no analog climates (i.e., projected climates do not match any contemporary biomes).


2009

Krausman PR, DE Naugle, MR Frisina, er al. 2009. "Livestock Grazing, Wildlife Habitat, and Rangeland Values". Rangelands 31 (5), 15-19. http://doi.org/10.2111/1551-501X-31.5.15


2007

K. Blake Suttle, Meredith A. Thomsen "CLIMATE CHANGE AND GRASSLAND RESTORATION IN CALIFORNIA: LESSONS FROM SIX YEARS OF RAINFALL MANIPULATION IN A NORTH COAST GRASSLAND," Madroño 54(3), 225-233, (5 July 2007). https://doi.org/10.3120/0024-9637(2007)54[225:CCAGRI]2.0.CO;2

Abstract

Native perennial bunchgrasses have undergone steep declines across much of California but persist in sizable populations along the northern coast. The longer rainy season and less severe summer drought in this region are thought to facilitate bunchgrass persistence in the face of extensive invasion by exotic annual species. Changes in the seasonality and intensity of precipitation that accompany global climate change could critically influence efforts to conserve and restore these plants in California grasslands. We established a large-scale manipulation of rainfall in a protected Mendocino County grassland to investigate how predicted shifts in precipitation affect the performance of three native perennial bunchgrass species in exotic-dominated stands. We added seeds, plugs, and mature tussocks of Danthonia californica, Elymus glaucus, and Elymus multisetus into replicate plots of exotic annual grassland and subjected the plots to one of three experimental precipitation regimes: increased winter rainfall, increased spring rainfall, and ambient rainfall. Responses to rainfall addition varied widely by age class and species and depended heavily on seasonal timing of the increase. Establishment from seed was rare for all three species and showed little response to water addition, likely due to concomitant changes in the surrounding communities. Production of exotic annual grasses rose markedly following repeated extensions of the rainy season, and while established bunchgrasses benefited despite this change, new plants could not establish into thickening stands of exotic vegetation. In contrast, survival was high for transplanted plugs and tussocks of all three species across all three rainfall treatments, suggesting that plugs and tussocks can survive a wide range of climatic conditions and high local densities of exotic annual grasses. Restoration approaches focused on these life stages may be most robust to changing climate. Transplanted individuals can provide a continual source of propagules to surrounding areas that then recruit during years in which conditions in the physical and biological environment are amenable to seedling establishment.



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