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Summary: Evolutionary history plays a key role driving patterns of trait variation across plant species. For scaling and modeling purposes, grass species are typically organized into C3 vs C4 plant functional types (PFTs). Plant functional type groupings may obscure important functional differences among species. Rather, grouping grasses by evolutionary lineage may better represent grass functional diversity.We measured 11 structural and physiological traits in situ from 75 grass species within the North American tallgrass prairie. We tested whether traits differed significantly among photosynthetic pathways or lineages (tribe) in annual and perennial grass species.Critically, we found evidence that grass traits varied among lineages, including independent origins of C4 photosynthesis. Using a rigorous model selection approach, tribe was included in the top models for five of nine traits for perennial species. Tribes were separable in a multivariate and phylogenetically controlled analysis of traits, owing to coordination of important structural and ecophysiological characteristics.Our findings suggest grouping grass species by photosynthetic pathway overlooks variation in several functional traits, particularly for C4 species. These results indicate that further assessment of lineage‐based differences at other sites and across other grass species distributions may improve representation of C4 species in trait comparison analyses and modeling investigations. [ABSTRACT FROM AUTHOR]

Abstract: Nocturnal transpiration can impact water balance from the local community to earth‐atmosphere fluxes. However, the dynamics and drivers of nocturnal transpiration among coexisting plant functional groups in herbaceous ecosystems are unknown. Here, we addressed the following questions: (1) How do nocturnal (Enight) and diurnal (Eday) transpiration vary among coexisting grasses, forbs, and shrubs in a tallgrass prairie? (2) What environmental variables drive Enight and do these differ from the drivers of Eday? (3) Is Enight associated with daytime physiological processes? We measured diurnal and nocturnal leaf gas exchange on perennial grass, forb and woody species in a North American tallgrass prairie. Measurements were made periodically across two growing seasons (May–August 2014–2015) on three C4 grasses (Andropogon gerardii, Sorghastrum nutans and Panicum virgatum), two C3 forbs (Vernonia baldwinii and Solidago canadensis), one C3 sub‐shrub (Amorpha canescens) and two C3 shrubs (Cornus drummondii and Rhus glabra). By extending our study to multiple functional groups, we were able to make several key observations: (1) Enight was variable among co‐occurring plant functional groups, with the highest rates occurring in C4 grasses, (2) Enight and Eday exhibited different responses to vapour pressure deficit and other environmental drivers, and (3) rates of Enight were strongly related to predawn leaf water potential for grasses and woody species, and were likely modulated by small‐scale changes in soil moisture availability. Our results provide novel insight into an often‐overlooked portion of ecosystem water balance. Considering the high rates of Enight observed in C4 grasses, as well as the widespread global occurrence of C4 grasses, nocturnal water loss might constitute a greater proportion of global evapotranspiration than previously estimated. Additionally, future predictions of nocturnal water loss may be complicated by stomatal behaviour that differs between the day and at night. Finally, these data suggest a water‐use strategy by C4 grasses wherein the high rates of Enight occurring during wet periods may confer a competitive advantage to maximize resource consumption during periods of greater availability. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Global change drivers are altering climatic and edaphic conditions of ecosystems across the globe, and we expect novel plant communities to become more common as a result. In the Colorado Front Range, compositional changes have occurred in the mixed-grass prairie plant community in conjunction with shifts in winter precipitation and atmospheric nitrogen (N) deposition. To test whether these environmental changes have been responsible for the observed plant community change, we conducted an in situ manipulative experiment in a mixed-grass meadow near Boulder, CO. We simulated historical conditions by reducing N availability (+500 g C m year) and winter precipitation (with rainout shelters) for 2 years (2013-2014) and compared vegetation response to these treatments with that of ambient conditions. The site experienced an extreme precipitation event in autumn 2013 which allowed comparison of an exceptionally wet year with an average year. We measured pre-treatment species composition in 2012, and treatment responses in the spring and summer of 2013 and 2014. As predicted, simulating historical low N-winter dry conditions resulted in a plant community dominated by historically abundant species. Cool-season introduced species were significantly reduced in low N-winter dry plots, particularly the annual plants Bromus tectorum and Alyssum parviflorum. These same species responded strongly to the extreme precipitation event with large increases, while native grasses and forbs showed little change in productivity or composition under varying climatic or edaphic conditions. This work provides clear evidence linking on-going global change drivers to altered plant community composition in an otherwise relatively undisturbed grassland ecosystem. [ABSTRACT FROM AUTHOR]

Shrub encroachment of grasslands is a transformative ecological process by which native woody species increase in cover and frequency and replace the herbaceous community. Mechanisms of encroachment are typically assessed using temporal data or experimental manipulations, with few large spatial assessments of shrub physiology. In a mesic grassland in North America, we measured inter- and intra-annual variability in leaf δ13C in Cornus drummondii across a grassland landscape with varying fire frequency, presence of large grazers and topographic variability. This assessment of changes in individual shrub physiology is the largest spatial and temporal assessment recorded to date. Despite a doubling of annual rainfall (in 2008 versus 2011), leaf δ13C was statistically similar among and within years from 2008-11 (range of −28 to −27‰). A topography*grazing interaction was present, with higher leaf δ13C in locations that typically have more bare soil and higher sensible heat in the growing season (upland topographic positions and grazed grasslands). Leaf δ13C from slopes varied among grazing contrasts, with upland and slope leaf δ13C more similar in ungrazed locations, while slopes and lowlands were more similar in grazed locations. In 2011, canopy greenness (normalized difference vegetation index – NDVI) was assessed at the centroid of individual shrubs using high-resolution hyperspectral imagery. Canopy greenness was highest mid-summer, likely reflecting temporal periods when C assimilation rates were highest. Similar to patterns seen in leaf δ13C, NDVI was highest in locations that typically experience lowest sensible heat (lowlands and ungrazed). The ability of Cornus drummondii to decouple leaf physiological responses from climate variability and fire frequency is a likely contributor to the increase in cover and frequency of this shrub species in mesic grassland and may be generalizable to other grasslands undergoing woody encroachment. [ABSTRACT FROM AUTHOR]

Anthropogenic climate change is projected to alter precipitation patterns, resulting in novel environments for plants. The responses of dominant plant species (e.g. Panicum virgatum L. (switchgrass)) to climate changes can drive broader ecosystem processes such as primary productivity. Using a rainfall mesocosm facility, three ecotypes of P. virgatum (collected from Kansas, Oklahoma and Texas, USA) were subjected to three precipitation regimes (average, -25%, +25%) to determine the physiological and growth responses to altered precipitation in a common garden setting. Results showed mean maximum photosynthetic rates, stomatal conductance, transpiration, midday water potential and dark-adapted chlorophyll fluorescence were lowest in the Kansas ecotypes. Increased precipitation treatments raised the mean midday water potentials and lowered water-use efficiency. Aboveground biomass responded positively to changes in precipitation, but flowering initiation was later and rates were lower for Texas ecotypes. In general, ecotype origin was a better predictor of differences in physiological responses and flowering, whereas the precipitation treatments had greater control over biomass production. Depending on the growth variable measured, these results show responses for P. virgatum are under varying ecotypic or environmental control with few interactions, suggesting that future predictions to climate change need not inherently consider localised adaptations in this economically important and widely distributed species. [ABSTRACT FROM AUTHOR]

oveground biomass in grasslands varies according to landscape gradients in resource availability and seasonal patterns of growth. Using a transect spanning a topographic gradient in annually burned ungrazed tallgrass prairie, we measured changes in the height of four abundant C grass species, LAI, biomass, and cumulative carbon flux using two closely located eddy flux towers. We hypothesized that seasonal patterns of plant growth would be similar across the gradient, but the magnitude of growth and biomass accumulation would vary by topographic position, reflecting spatial differences in microclimate, slope, elevation, and soil depth. Thus, identifying and measuring local growth responses according to topographic variability should significantly improve landscape predictions of aboveground biomass. For most of the growth variables measured, classifying topography into four positions best captured the inherent spatial variability. Biomass produced, seasonal LAI and species height increased from the upland and break positions to the slope and lowland. Similarly, cumulative carbon flux in 2008 was greater in lowland versus upland tower locations (difference of 64 g m by DOY 272). Differences in growth by topographic position reflected increased production of flowering culms by Andropogon gerardii and Sorghastrum nutans in lowland. Varying growth responses by these species may be a significant driver of biomass and carbon flux differences by topographic position, at least for wet years. Using a digital elevation model to classify the watershed into topographic positions, we performed a geographically weighted regression to predict landscape biomass. The minimum and maximum predictions of aboveground biomass for this watershed had a large range (86-393 t per 40.4 ha), illustrating the drastic spatial variability in growth within this annually-burned grassland. [ABSTRACT FROM AUTHOR]

Future climate change is likely to reduce the floristic diversity of grasslands. Yet the potential consequences of climate-induced plant species losses for the functioning of these ecosystems are poorly understood. We investigated how climate change might alter the functional composition of grasslands for Konza Prairie, a diverse tallgrass prairie in central North America. With species-specific climate envelopes, we show that a reduction in mean annual precipitation would preferentially remove species that are more abundant in the more productive lowland positions at Konza. As such, decreases in precipitation could reduce productivity not only by reducing water availability but by also removing species that inhabit the most productive areas and respond the most to climate variability. In support of this prediction, data on species abundance at Konza over 16 years show that species that are more abundant in lowlands than uplands are preferentially reduced in years with low precipitation. Climate change is likely to also preferentially remove species from particular functional groups and clades. For example, warming is forecast to preferentially remove perennials over annuals as well as Cyperaceae species. Despite these predictions, climate change is unlikely to unilaterally alter the functional composition of the tallgrass prairie flora, as many functional traits such as physiological drought tolerance and maximum photosynthetic rates showed little relationship with climate envelope parameters. In all, although climatic drying would indirectly alter grassland productivity through species loss patterns, the insurance afforded by biodiversity to ecosystem function is likely to be sustained in the face of climate change. [ABSTRACT FROM AUTHOR]

The majority of tallgrass prairie root biomass is located in the upper soil layers (0–25 cm), but species differences exist in reliance on soil water at varying depths. These differences have led to the hypothesis that resource partitioning belowground facilitates species co-existence in this mesic grassland. To determine if plant water relations can be linked to soil water partitioning as a potential mechanism allowing C3 species to persist among the more dominant C4 grasses, we measured differences in the source of water-use using the isotopic signature of xylem water, volumetric soil water content at 4 depths, and leaf water potentials. Data were collected for seven species representing C4 grasses, C3 forbs and C3 shrubs over three growing seasons at the Konza Prairie (Kansas, USA) to encompass a range of natural climatic conditions. C4 grasses relied on shallow soil water (5 cm) across the growing season and had midday leaf water potentials that were highly correlated with shallow soil water regardless of soil water availability at other portions of the soil profile (20, 40 and 90 cm). In contrast, C3 species only used shallow soil water when plentiful at this depth; these species increased their dependence on soil water from greater depths as the upper soil layers dried. Structural equation models describing plant water relations were very similar for the three C4 species, whereas a unique set of models and drivers were identified for each of the C3 species. These results support soil water partitioning as a mechanism for species coexistence, as C4 species in this grassland have relatively consistent dependence on water in shallow soil layers, whereas C3 species show niche differentiation in water use strategies to avoid competition with C4 grasses for water in shallow soil layers when this resource is limiting and leaf water stress is high. [ABSTRACT FROM AUTHOR]

Ecosystem process models are often used to predict carbon flux on a landscape or on a global scale. Such models must be aggregate and canopies are often treated as a uniform unit of foliage. Parameters that are known to vary within the canopy, e.g., nitrogen content and leaf mass per area, are often estimated by a mean value for the canopy. Estimating appropriate means is complicated, especially in mixed-species stands and in complex terrain.We analyzed sources of variation in specific parameters with the goal of testing various simplifying assumptions. The measurements came from mixed-species forests in the northern Rocky Mountains. We found that, for three important parameters (nitrogen concentration and content, and leaf mass per area), a sample taken near the vertical center of the crown provided a good estimate of the mean values for the crown. Altitude (700–1700 m), solar insolation (4200–5400 MJ m−2 year−1) and leaf area index (1–11) had negligible effects on the parameters; only species differences were consistently detected. The correlation between mass-based photosynthetic rates and mass-based nitrogen concentrations was much weaker than the correlation between area-based photosynthetic rates and area-based nitrogen concentration. Comparison of photosynthesis–nitrogen relationships for a wide variety of conifer species and sites revealed a broad general trend that can be used in models.These results suggest important potential simplifications in model parameterization, most notably that canopy means can be estimated with ease, that complex terrain is a minor source of variation in these parameters and that use of one photosynthesis–nitrogen relationship for conifer species does not result in large errors. Species-to-species variation, however, was large and needs to be accounted for when parameterizing process models. [ABSTRACT FROM AUTHOR]