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1. Spatio-temporal differences in leaf physiology are associated with fire, not drought, in a clonally integrated shrub

   https://doi.org/10.1093/aobpla/plab037  

   Wedel, Emily R; O’Keefe, Kimberly; Nippert, Jesse B; Hoch, Braden; O’Connor, Rory C (August 2021, AoB PLANTS)

   Mitchell, Patrick (Ed.)

   Abstract In highly disturbed environments, clonality facilitates plant survival via resprouting after disturbance, resource sharing among interconnected stems and vegetative reproduction. These traits likely contribute to the encroachment of deep-rooted clonal shrubs in tallgrass prairie. Clonal shrubs have access to deep soil water and are typically thought of as relatively insensitive to environmental variability. However, how leaf physiological traits differ among stems within individual clonal shrubs (hereafter ‘intra-clonal’) in response to extreme environmental variation (i.e. drought or fire) is unclear. Accounting for intra-clonal differences among stems in response to disturbance is needed to more accurately parameterize models that predict the effects of shrub encroachment on ecosystem processes. We assessed intra-clonal leaf-level physiology of the most dominant encroaching shrub in Kansas tallgrass prairie, Cornus drummondii, in response to precipitation and fire. We compared leaf gas exchange rates from the periphery to centre within shrub clones during a wet (2015) and extremely dry (2018) year. We also compared leaf physiology between recently burned shrubs (resprouts) with unburned shrubs in 2018. Resprouts had higher gas exchange rates and leaf nitrogen content than unburned shrubs, suggesting increased rates of carbon gain can contribute to recovery after fire. In areas recently burned, resprouts had higher gas exchange rates in the centre of the shrub than the periphery. In unburned areas, leaf physiology remained constant across the growing season within clonal shrubs (2015 and 2018). Results suggest single measurements within a shrub are likely sufficient to parameterize models to understand the effects of shrub encroachment on ecosystem carbon and water cycles, but model parameterization may require additional complexity in the context of fire. 

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2. Intra-canopy leaf trait variation facilitates high leaf area index and compensatory growth in a clonal woody encroaching shrub

   https://doi.org/10.1093/treephys/tpac078  

   Tooley, E Greg; Nippert, Jesse B; Bachle, Seton; Keen, Rachel M (July 2022, Tree Physiology)

   Cavaleri, Molly (Ed.)

   Abstract Leaf trait variation enables plants to utilize large gradients of light availability that exist across canopies of high leaf area index (LAI), allowing for greater net carbon gain while reducing light availability for understory competitors. While these canopy dynamics are well understood in forest ecosystems, studies of canopy structure of woody shrubs in grasslands are lacking. To evaluate the investment strategy used by these shrubs, we investigated the vertical distribution of leaf traits and physiology across canopies of Cornus drummondii, the predominant woody encroaching shrub in the Kansas tallgrass prairie. We also examined the impact of disturbance by browsing and grazing on these factors. Our results reveal that leaf mass per area (LMA) and leaf nitrogen per area (Na) varied approximately threefold across canopies of C. drummondii, resulting in major differences in the physiological functioning of leaves. High LMA leaves had high photosynthetic capacity, while low LMA leaves had a novel strategy for maintaining light compensation points below ambient light levels. The vertical allocation of leaf traits in C. drummondii canopies was also modified in response to browsing, which increased light availability at deeper canopy depths. As a result, LMA and Na increased at lower canopy depths, leading to a greater photosynthetic capacity deeper in browsed canopies compared to control canopies. This response, along with increased light availability, facilitated greater photosynthesis and resource-use efficiency deeper in browsed canopies compared to control canopies. Our results illustrate how C. drummondii facilitates high LAI canopies and a compensatory growth response to browsing—both of which are key factors contributing to the success of C. drummondii and other species responsible for grassland woody encroachment. 

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3. Woody Encroachment Modifies Subsurface Structure and Hydrological Function

   https://doi.org/10.1002/eco.2731  

   Jarecke, Karla M.; Zhang, Xi; Keen, Rachel M.; Dumont, Marc; Li, Bonan; Sadayappan, Kayalvizhi; Moreno, Victoria; Ajami, Hoori; Billings, Sharon A.; Flores, Alejandro N.; et al (November 2024, Ecohydrology)

   ABSTRACT Woody encroachment—the expansion of woody shrubs into grasslands—is a widely documented phenomenon with global significance for the water cycle. However, its effects on watershed hydrology, including streamflow and groundwater recharge, remain poorly understood. A key challenge is the limited understanding of how changes to root abundance, size and distribution across soil depths influence infiltration and preferential flow. We hypothesised that woody shrubs would increase and deepen coarse‐root abundance and effective soil porosity, thus promoting deeper soil water infiltration and increasing soil water flow velocities. To test this hypothesis, we conducted a study at the Konza Prairie Biological Station in Kansas, where roughleaf dogwood (Cornus drummondii) is the predominant woody shrub encroaching into native tallgrass prairie. We quantified the distribution of coarse and fine roots and leveraged soil moisture time series and electrical resistivity imaging to analyse soil water flow beneath shrubs and grasses. We observed a greater fraction of coarse roots beneath shrubs compared to grasses, which was concurrent with greater saturated hydraulic conductivity and effective porosity. Half‐hourly rainfall and soil moisture data show that the average soil water flow through macropores was 135% greater beneath shrubs than grasses at the deepest B horizon, consistent with greater saturated hydraulic conductivity. Soil‐moisture time series and electrical resistivity imaging also indicated that large rainfall events and greater antecedent wetness promoted more flow in the deeper layers beneath shrubs than beneath grasses. These findings suggest that woody encroachment alters soil hydrologic processes with cascading consequences for ecohydrological processes, including increased vertical connectivity and potential groundwater recharge. 

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4. Contrasting intra‐annual population dynamics of two codominant species are consistent across spatial and temporal scales

   https://doi.org/10.1111/1365-2745.14055  

   Gray, Jesse E.; Smith, Melinda D. (March 2023, Journal of Ecology)

   Abstract Despite asymmetric competition and a wide array of functional similarities, two ecologically important C₄perennial grasses,Andropogon gerardiiandSorghastrum nutans, frequently codominate areas of the mesic tallgrass prairie of the US Great Plains. A subtle difference in their vegetative reproduction strategies may play a role in preventing the exclusion ofS. nutans, the presumed weaker competitor in such regions.WhileA. gerardiivegetative tiller densities peak in the early growing season and decline thereafter (determinate recruitment), those ofS. nutansmay continue to increase throughout the growing season (indeterminate recruitment), providing a potential avenue for recovery from more intensive early season competition. However, until now these patterns have only been informally observed in the field.We examined the year‐to‐year consistency of growing season vegetative tiller dynamics (measured as seasonal change in tiller densities) of each grass species from an intact tallgrass prairie in Kansas – a site within the core of both species' distributions – over a period of 8 years. Then, to investigate environmental effects on these dynamics, we examined whether they differ across a Kansas landscape varying in topography, fire management regimes, and the abundances of the study species. Finally, we expanded the investigation of environmental effects on growing season tiller dynamics by observing them at the periphery of the species' distributions in central Colorado, where climatic conditions are dryer and the study species' abundances are reduced.Synthesis. We found that the tiller densities ofA. gerardiidecline within seasons with striking consistency regardless of spatio‐temporal scale or environmental factors (topography and fire regimes). In contrast, we found the seasonal dynamics ofS. nutanstiller densities were dependent on environmental factors, with seasonal tiller density increases occurring only within the Kansas populations but not consistent between years. These observations lay the groundwork for establishing differences in tiller recruitment determinacy as a potentially important yet underappreciated mechanism for promoting coexistence and codominance among perennial plant species. 

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5. Resources do not limit compensatory response of a tallgrass prairie plant community to the loss of a dominant species

   https://doi.org/10.1111/1365-2745.13741  

   Chaves, Francis A.; Smith, Melinda D. (August 2021, Journal of Ecology)

   Abstract The effect of species loss on ecosystem productivity is determined by both the functional contribution of the species lost, and the response of the remaining species in the community. According to the mass ratio hypothesis, the loss of a dominant plant species, which has a larger proportionate contribution to productivity, is expected to exert an overwhelming effect on this important ecosystem function. However, via competitive release, loss of a dominant species can provide the opportunity for other plant species to establish, thrive and become abundant in the community, potentially compensating for the function lost. Furthermore, if resource limitation is removed, then the compensatory response of function to the loss of a dominant species should be greater and more rapid than if resources are more limiting.To evaluate how resources may limit compensation of above‐ground productivity to the loss of a dominant plant species, we experimentally removed the C₄perennial tallgrass,Andropogon gerardii, from intact plant communities. We added water for 4 years, as well as nitrogen in the fourth year, to test the effect of resource limitation on the compensatory response.Overall, above‐ground biomass production increased in the remaining community with both water and nitrogen addition. However, this increase in biomass production was not sufficient to fully compensate for the loss ofA. gerardii, indicating water and nitrogen were not limiting short‐term compensation in this community.Following the removal of the dominant species, there was reordering of species abundances in the community, rather than changes in species richness. The C₄grassBouteloua curtipendulawas the most responsive species, increasing by 57.9% in abundance with water addition and 91.0% with both water and nitrogen addition. Despite this dramatic increase in abundance, its short stature and lower per capita biomass production prevented this species from compensating for the loss ofA. gerardii.Synthesis. Short‐term compensation after the loss of a dominant plant species can be hastened by increased resource availability, but ultimately full compensation appears to be limited by the presence and abundance of species in the remaining community that possess traits that allow them compensate for the species lost. 

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