Climate change forecasts of more frequent climate extremes suggest that such events will become increasingly important drivers of future ecosystem dynamics and function. Because the rarity and unpredictability of naturally occurring climate extremes limits assessment of their ecological impacts, we experimentally imposed extreme drought and a mid-summer heat wave over two years in a central U.S. grassland. While the ecosystem was resistant to heat waves, it was not resistant to extreme drought, which reduced aboveground net primary productivity (ANPP) below the lowest level measured in this grassland in almost 30 years. This extreme reduction in ecosystem function was a consequence of reduced productivity of both C4 grasses and C3 forbs. However, the dominant forb was negatively impacted by the drought more than the dominant grass, and this led to a reordering of species abundances within the plant community. Although this change in community composition persisted post-drought, ANPP recovered completely t...

Resistance and resilience of a grassland ecosystem to climate extremes

Soil, through its various functions, plays a vital role in the Earth's ecosystems and provides multiple ecosystem services to humanity. Pedotransfer functions (PTFs) are simple to complex knowledge rules that relate available soil information to soil properties and variables that are needed to parameterize soil processes. In this paper, we review the existing PTFs and document the new generation of PTFs developed in the different disciplines of Earth system science. To meet the methodological challenges for a successful application in Earth system modeling, we emphasize that PTF development has to go hand in hand with suitable extrapolation and upscaling techniques such that the PTFs correctly represent the spatial heterogeneity of soils. PTFs should encompass the variability of the estimated soil property or process, in such a way that the estimation of parameters allows for validation and can also confidently provide for extrapolation and upscaling purposes capturing the spatial variation in soils. Most actively pursued recent developments are related to parameterizations of solute transport, heat exchange, soil respiration and organic carbon content, root density and vegetation water uptake. Further challenges are to be addressed in parameterization of soil erosivity and land use change impacts at multiple scales. We argue that a comprehensive set of PTFs can be applied throughout a wide range of disciplines of Earth system science, with emphasis on land surface models. Novel sensing techniques provide a true breakthrough for this, yet further improvements are necessary for methods to deal with uncertainty and to validate applications at global scale.

https://agupubs.onlinelibrary.wiley.com/doi/epdf/10.1002/2017RG000581

Pedotransfer functions in Earth system science: challenges and perspectives

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

https://hal.archives-ouvertes.fr/hal-02918834/document

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements

Poaceae (the grasses) is arguably the most successful plant family, in terms of its global occurrence in (almost) all ecosystems with angiosperms, its ecological dominance in many ecosystems, and high species richness. We suggest that the success of grasses is best understood in context of their capacity to colonize, persist, and transform environments (the "Viking syndrome"). This results from combining effective long-distance dispersal, efficacious establishment biology, ecological flexibility, resilience to disturbance and the capacity to modify environments by changing the nature of fire and mammalian herbivory. We identify a diverse set of functional traits linked to dispersal, establishment and competitive abilities. Enhanced long-distance dispersal is determined by anemochory, epizoochory and endozoochory and is facilitated via the spikelet (and especially the awned lemma) which functions as the dispersal unit. Establishment success could be a consequence of the precocious embryo and large starch reserves, which may underpin the extremely short generation times in grasses. Post-establishment genetic bottlenecks may be mitigated by wind pollination and the widespread occurrence of polyploidy, in combination with gametic self-incompatibility. The ecological competitiveness of grasses is corroborated by their dominance across the range of environmental extremes tolerated by angiosperms, facilitated by both C3and C4photosynthesis, well-developed frost tolerance in several clades, and a sympodial growth form that enabled the evolution of both annual and long-lived life forms. Finally, absence of investment in wood (except in bamboos), and the presence of persistent buds at or below ground level, provides tolerance of repeated defoliation (whether by fire, frost, drought or herbivores). Biotic modification of environments via feedbacks with herbivory or fire reinforce grass dominance leading to open ecosystems. Grasses can be both palatable and productive, fostering high biomass and diversity of mammalian herbivores. Many grasses have a suite of architectural and functional traits that facilitate frequent fire, including a tufted growth form, and tannin-like substances in leaves which slow decomposition. We mapped these traits over the phylogeny of the Poales, spanning the grasses and their relatives, and demonstrated the accumulation of traits since monocots originated in the mid-Cretaceous. Although the sympodial growth form is a monocot trait, tillering resulting in the tufted growth form most likely evolved within the grasses. Similarly, although an ovary apparently constructed of a single carpel evolved in the most recent grass ancestor, spikelets and the awned lemma dispersal units evolved within the grasses. Frost tolerance and C4photosynthesis evolved relatively late (late Palaeogene), and the last significant trait to evolve was probably the production of tannins, associated with pyrophytic savannas. This fits palaeobotanical data, suggesting several phases in the grass success story: from a late Cretaceous origin, to occasional tropical grassland patches in the later Palaeogene, to extensive C3grassy woodlands in the early-middle Miocene, to the dramatic expansion of the tropical C4grass savannas and grasslands in the Pliocene, and the C3steppe grasslands during the Pleistocene glacial periods. Modern grasslands depend heavily on strongly seasonal climates, making them sensitive to climate change.

/pdf/global-grass-poaceae-success-underpinned-by-traits-3xihgrks95.pdf

Global grass (Poaceae) success underpinned by traits facilitating colonization, persistence and habitat transformation

https://www.fs.fed.us/rm/pubs_journals/2019/rmrs_2019_ott_j002.pdf

The ecology and significance of below-ground bud banks in plants.

Because most shoot recruitment in perennial grasses occurs from belowground axillary buds, bud dynamics determine plant population dynamics and meristem limitation to population growth. Therefore, grassland vegetation responses to environmental change or disturbance may be influenced by interspecific differences in bud banks and the patterns and environmental controls of bud development and demography. We examined bud bank dynamics in Andropogon gerardii and Dichanthelium oligosanthes in tallgrass prairie by enumerating and classifying buds throughout 15 months to determine whether grass buds live for multiple years and accumulate; whether bud natality, dormancy and outgrowth are synchronous or variable; and whether bud bank dynamics differ between these co-occurring species. Andropogon gerardii (a C4 species) maintained a larger dormant bud bank, showed synchrony in bud development and transition to tiller, and its buds lived for multiple years. Thus, multiple previous years’ bud cohorts contributed to recruitment. By contrast, D. oligosanthes (a C3 species) maintained a smaller dormant bud bank, had asynchronous bud development with active buds present year-round, and its buds lived for 1 year. Buds played different roles in the dynamics of each species, allowing A. gerardii to over-winter and recruit new spring tillers and D. oligosanthes to survive and recruit new tillers following summer dormancy. These differences in bud bank age structure, phenology, and dynamics between these species suggest greater demographic buffering and time-lag effects in A. gerardii populations. Interspecific differences in bud bank structure and dynamics may explain and help predict grassland responses to environmental change.

Contrasting bud bank dynamics of two co-occurring grasses in tallgrass prairie: implications for grassland dynamics

In arid to semi-arid grasslands and savannas, plant growth, population dynamics, and productivity are consistently and strongly limited by soil water and nutrient availability. Adaptive traits of the root systems of grasses in these ecosystems are crucial to their ability to cope with strong water and/or nutrient limitation and the increasing drought stress associated with ecosystem degradation or projected climate change. We studied 18 grass species in semi-arid savanna of the Kalahari region of Botswana to quantify interspecific variation in three important root system traits including root system architecture, rhizosheath thickness and mycorrhizal colonization. Drought- tolerant species and shorter-lived species showed greater rhizosheath thickness and fine root development but lower mycorrhizal colonization compared to later successional climax grasses and those characteristic of wetter sites. In addition, there was a significant positive correlation between root fibrousness index and rhizosheath thickness among species and a weak negative correlation between root fibrousness index and mycorrhizal colonization.These patterns suggest that an extensive fine root system and rhizosheath development may be important complementary traits of grasses coping with drought conditions, the former aiding in the acquisition of water by the grass plant and the latter aiding in water uptake and retention, and reducing water loss in the rhizosphere. Within species, both rhizosheath development and mycorrhizal colonization were significantly greater in a wet year than in a year with below-average precipitation. The observed patterns suggest that the primary benefit of rhizosheath development in African savanna grasses is improved drought tolerance and that it is a plastic trait that can be adjusted annually to changing environmental conditions. The functioning of mycorrhizal symbiosis is likely to be relatively more important in infertile savannas where nutrient limitation is higher relative to water limitation.

Variation in root system traits among African semi-arid savanna grasses: Implications for drought tolerance

 Premise of the Study: Perennial grasses maintain aboveground tiller populations through vegetative reproduction via belowground buds and sexual reproduction via seed. The maintenance of a bud bank has important demographic consequences for perennial grasses. A tradeoff between these reproductive modes would be expected for a plant with limited resource availability. However, the ontogeny of the tiller could affect its ability to allocate between these two modes of reproduction.  Methods: Vegetative bud production and dynamics and tiller production were examined biweekly through an annual cycle on vegetative and fl owering tillers of Andropogon gerardii .  Key Results: Andropogon gerardii maintains a large reserve of dormant buds. Although vegetative and fl owering tillers had similar bud phenology, fl owering tillers produced larger numbers of buds of larger size, and transitioned a larger proportion of their buds to tiller, than did vegetative tillers. Therefore, a negative consequence of sexual reproduction on vegetative reproduction was not evident at the tiller level. A size threshold for fl oral induction likely exists that results in fltillers having more buds per tiller than vegetative tillers. The increased bud outgrowth of fltillers could be a result of their larger bud size or weaker apical dominance as compared to vegetative tillers.  Conclusions: Plant development can place signifi cant constraints on tradeoffs between the reproductive modes in perennial grasses and could affect their plasticity in plant reproductive allocation. Differences in developmental phenology and bud production between fl owering and vegetative tillers may infl uence grass responses to environmental changes such as altered precipitation regimes or resource availability.

Bud production and dynamics of flowering and vegetative tillers in Andropogon gerardii (Poaceae): the role of developmental constraints

PREMISE OF THE STUDY: Tiller recruitment from the belowground bud bank of caespitose grasses infl uences their ability to monopolize local resources and, hence, their genet fi tness. Diff erences in bud production and outgrowth among tiller types within a genet and among species may explain co-occurrence of caespitose grasses. This study aimed to characterize genet bud-bank and tiller production and dynamics in two co-occurring species and compare their vegetative reproductive strategies. METHODS: Bud-bank and tiller dynamics of Hesperostipa comata and Nassella viridula , dominant C 3 caespitose grasses in the northern mixed-grass prairie of North America, were assessed throughout an annual cycle. KEY RESULTS: The two species showed similar strategies, maintaining polycyclic tillers and thus creating mixed-age genet bud banks comprising multiple bud cohorts produced in diff erent years. Vegetative tillers produced the majority of buds, whereas fl owering tillers contributed little to the bud bank. Buds lived for at least 2 yr and were maintained in multiple developmental stages throughout the year. Because bud longevity rarely exceeded tiller longevity, tiller longevity drove turnover within the bud bank. Tiller population dynamics, more than bud production per tiller, determined the diff erential contribution of tiller types to the bud bank. Nassella viridula had higher bud production per tiller, a consistent annual tiller recruitment density, and greater longevity of buds on senesced and fl owering tillers than H. comata . CONCLUSIONS: Co-occurring C 3 caespitose grasses had similar bud-bank and tiller dynamics contributing to genet persistence but diff ered in bud characteristics that could aff ect genet longevity and species coexistence.

Bud-bank and tiller dynamics of co-occurring C3 caespitose grasses in mixed-grass prairie

Responses of plants to herbivory are dependent on the type of damage and the ontogenetic stage of the plant. We compared the effects of stem pruning and defoliation on seedlings of Colophospermum mopane, an ecologically important tree species widely distributed in southern Africa. The growth of 160 greenhouse-grown juveniles were measured for 6-mo after germination and then 6-mo after treatments including 50% defoliation, 100% defoliation, 50% stem pruning and controls. Pruning resulted in 30% reductions in total leaf area, height and biomass. Partial defoliation resulted in 30% reductions in total leaf area and plant biomass. However, complete defoliation resulted in a 30% increase in biomass production, a doubling in leaf and lateral branch number, a 45% reduction in leaf size, and no change in total leaf area. Thus, completely defoliated seedlings showed greater performance than those that were only partially defoliated, indicating that C. mopane has become adapted to the chronic and severe defoliation inflicted by Imbrasia belina caterpillars. Comparison of our results with other studies indicates that C. mopane seedlings are less herbivory-tolerant than adults and that pruning has more negative effects than defoliation. Thus, seedling browsers may constrain recruitment in C. mopane, influencing its population dynamics and abundance.

Jacqueline P. Ott

Papers

Contrasting bud bank dynamics of two co-occurring grasses in tallgrass prairie: implications for grassland dynamics

Variation in root system traits among African semi-arid savanna grasses: Implications for drought tolerance

Bud production and dynamics of flowering and vegetative tillers in Andropogon gerardii (Poaceae): the role of developmental constraints

Bud-bank and tiller dynamics of co-occurring C3 caespitose grasses in mixed-grass prairie

Coping with herbivory at the juvenile stage: responses to defoliation and stem browsing in the African savanna tree Colophospermum mopane