E. Gene Towne

The Keystone Role of Bison in North American Tallgrass Prairie

Your use of the JSTOR archive indicates your acceptance of JSTORs Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTORs Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use.

VEGETATION TRENDS IN TALLGRASS PRAIRIE FROM BISON AND CATTLE GRAZING

Comparisons between how bison and cattle grazing affect the plant community are understood poorly because of confounding differences in how the herbivores are typically managed. This 10-year study compared vegetation changes in Kansas (USA) tallgrass prairie that was burned and grazed season-long at a moderate stocking rate by either bison or cattle. We held management practices constant between the herbivores and equalized grazing pressure by matching animals so that the total body mass in all pastures was similar each year. Trends in species cover and diversity indices in the bison and cattle pastures were compared with ungrazed prairie that also was burned annually. We found that little bluestem (Schizachyrium scoparium) cover decreased over time in bison pastures, and big bluestem (Andropogon gerardii) cover increased over time in cattle pastures. Grazing by either herbivore increased the canopy cover of annual forbs, perennial forbs, and cool-season graminoids, but both annual and perennial forb cove...

Flowering phenology as a functional trait in a tallgrass prairie

• The timing of flowering is a critical component of the ecology of plants and has the potential to structure plant communities. Yet, we know little about how the timing of flowering relates to other functional traits, species abundance, and average environmental conditions. • Here, we assessed first flowering dates (FFDs) in a North American tallgrass prairie (Konza Prairie) for 431 herbaceous species and compared them with a series of other functional traits, environmental metrics, and species abundance across ecological contrasts. • The pattern of FFDs among the species of the Konza grassland was shaped by local climate, can be linked to resource use by species, and patterns of species abundance across the landscape. Peak FFD for the community occurred when soils were typically both warm and wet, while relatively few species began flowering when soils tended to be the driest. Compared with late-flowering species, species that flowered early had lower leaf tissue density and were more abundant on uplands than lowlands. • Flowering phenology can contribute to the structuring of grassland communities, but was largely independent of most functional traits. Therefore, selection for flowering phenology may be independent of general resource strategies.

Climate controls on grass culm production over a quarter century in a tallgrass prairie

The flowering of grasses is a process critical to plant population dynamics and genetics, herbivore performance, and human health. To better understand the climate factors governing grass flowering, we analyzed the patterns of culm production over 25 years for three perennial tallgrass prairie species at Konza Prairie in Kansas, USA. The three species (Andropogon gerardii, Sorghastrum nutans, and Schizachyrium scoparium) all utilize the C4 photosynthetic pathway and were measured annually at the same locations for the past 25 years in an annually burned watershed. Culm production of all three species increased with higher growing-season soil moisture and precipitation but differed in their responses to water availability at different times during the growing season. Relative to Andropogon, Sorghastrum responded more to precipitation early in the growing season, and Schizachyrium responded more to precipitation late in the growing season. Flowering by each species also revealed a threshold relationship with late-season soil moisture at approximately 1 m depth, which likely is a proxy for season-long water balance. Although flowering can be influenced by conditions antecedent to the current growing season, neither soil moisture nor precipitation during the previous year influenced flowering over the 25-year period. Flowering culm production averaged 9% and 7% of total graminoid aboveground net primary production (ANPP) in the uplands and lowlands, respectively. Interannual variation in ANPP correlated only with Sorghastrum flowering, suggesting a predominant role of the species in ANPP responses to climate.

Maternal allocation in bison: co‐occurrence of senescence, cost of reproduction, and individual quality

Parental allocation strategies are of profound interest in life history because they directly impact offspring fitness and therefore are highly valuable for understanding population dynamics and informing management decisions. Yet, numerous questions about reproductive allocation patterns for wild populations of large mammals remain unanswered because of the challenges for measuring allocation in the wild. Using a nine-year longitudinal data set on life-history traits of mother-calf bison pairs, we identified sources of variation in relative maternal allocation (calf mass ratio on mother mass) and assessed the occurrence of reproductive costs associated with differential maternal allocation. We found that heavy mothers provided a lower allocation but still produced heavier calves than light mothers. Older females produced lighter calves and tended to decrease allocation as they aged, supporting the occurrence of reproductive senescence. Mothers that had produced a calf the previous year produced lighter calves and allocated less than mothers that did not lactate the previous year, revealing reproductive costs. However, greater maternal allocation did not reduce the probability of breeding in successive years, and the amount of allocation provided by a mother was positively correlated among the offspring she produced, illustrating individual heterogeneity. Although life-history studies are usually classified as either supporting costs of reproduction or individual quality, our study demonstrates that these contrasting evolutionary forces can shape variation within a single trait. Our work illustrates that many processes can coevolve within a population, emphasizing the need to integrate multiple concepts to better understand the evolution of life-history traits. With regard to management of bison herds, if the goal of culling programs is to select for animals with the best performance, this research suggests that managers should account for the condition and previous reproductive status of mothers when taking culling decisions on juvenile bison.

Community traitscape of foliar nitrogen isotopes reveals N availability patterns in a tallgrass prairie

Background and aimsNutrients are important determinants of community assembly, yet the roles they play in structuring plant communities are still poorly understood. One inferential approach to understanding how environmental factors structure plant communities is examining the distribution of key functional traits among species of a community—a community traitscape.MethodsTo better understand how nitrogen (N) and water availability structure grasslands, we measured N concentrations and isotope ratios for 366 herbaceous species in a mesic N-limited temperate grassland, Konza Prairie. We also compared foliar N concentrations and N isotopes between Konza species and a global dataset.ResultsSpecies that had either high foliar N concentrations or high δ15NL were not necessarily more or less abundant on the landscape nor more or less likely to be found in uplands, grazed areas, or burned areas. Apparently there are unique hot spots of high N availability at Konza and the typical non-Fabaceae Konza species occupies sites with greater N availability than found globally.ConclusionsAlthough nascent, the Konza traitscapes suggest that plant diversity in nutrient-limited communities might be strongly dependent on high-nutrient availability sites that enable high-fertility species to persist in a matrix of low nutrient availability.

Climatic warming and the future of bison as grazers

Climatic warming is likely to exacerbate nutritional stress and reduce weight gain in large mammalian herbivores by reducing plant nutritional quality. Yet accurate predictions of the effects of climatic warming on herbivores are limited by a poor understanding of how herbivore diet varies along climate gradients. We utilized DNA metabarcoding to reconstruct seasonal variation in the diet of North American bison (Bison bison) in two grasslands that differ in mean annual temperature by 6 °C. Here, we show that associated with greater nutritional stress in warmer climates, bison consistently consumed fewer graminoids and more shrubs and forbs, i.e. eudicots. Bison in the warmer grassland consumed a lower proportion of C3 grass, but not a greater proportion of C4 grass. Instead, bison diet in the warmer grassland had a greater proportion of N2-fixing eudicots, regularly comprising >60% of their protein intake in spring and fall. Although bison have been considered strict grazers, as climatic warming reduces grass protein concentrations, bison may have to attempt to compensate by grazing less and browsing more. Promotion of high-protein, palatable eudicots or increasing the protein concentrations of grasses will be critical to minimizing warming-imposed nutritional stress for bison and perhaps other large mammalian herbivores.

High leaf tissue density grassland species consistently more abundant across topographic and disturbance contrasts in a North American tallgrass prairie

Understanding the coupling between plant functional traits and abundance provides insight into the often hidden forces that structure plant communities. To better understand the coupling between leaf traits and abundance of grassland species in a mesic North American grassland, we measured specific leaf area (SLA) and its two components, tissue density and thickness for 125 grassland species. Plants with high tissue density were more abundant over a 17-year period across a range of environments: uplands, grazed and ungrazed watersheds, and frequently and infrequently burned watersheds. The consistent relationships between leaf tissue density and abundance across ecological contrasts imply that belowground resource availability constrains community composition independent of grazing and burning regimes. Leaf tissue density did not explain species abundance in lowlands, where belowground resources are the highest. Neither did it explain the differential abundance of species between grazing or fire frequency contrasts, suggesting that changes in burning or grazing select for species based on other traits. Relative to leaf tissue density, SLA was a poor predictor of abundance, reinforcing a long-observed—but often ignored—call that measurements of SLA need to be coupled with thickness measurements in order to effectively predict the performance of species. More generally, future research needs to investigate which belowground resources control community composition in the grassland and whether the importance of water or nutrients change with burning and grazing.

Ecological Consequences of Shifting the Timing of Burning Tallgrass Prairie

In the Kansas Flint Hills, grassland burning is conducted during a relatively narrow window because management recommendations for the past 40 years have been to burn only in late spring. Widespread prescribed burning within this restricted time frame frequently creates smoke management issues downwind. A potential remedy for the concentrated smoke production in late spring is to expand burning to times earlier in the year. Yet, previous research suggested that burning in winter or early spring reduces plant productivity and cattle weight gain while increasing the proportion of undesirable plant species. In order to better understand the ecological consequences of burning at different times of the year, plant production and species abundance were measured for 20 years on ungrazed watersheds burned annually in autumn, winter, or spring. We found that there were no significant differences in total grass production among the burns on either upland or lowland topographic positions, although spring burned watersheds had higher grass culm production and lower forb biomass than autumn and winter burned watersheds. Burning in autumn or winter broadened the window of grass productivity response to precipitation, which reduces susceptibility to mid-season drought. Burning in autumn or winter also increased the phenological range of species by promoting cool-season graminoids without a concomitant decrease in warm-season grasses, potentially widening the seasonal window of high-quality forage. Incorporating autumn and winter burns into the overall portfolio of tallgrass prairie management should increase the flexibility in managing grasslands, promote biodiversity, and minimize air quality issues caused by en masse late-spring burning with little negative consequences for cattle production.

A Critical Examination of Timing of Burning in the Kansas Flint Hills

ABSTRACT Frequent burning is a crucial ecological and economic component of the Kansas Flint Hills. Although burning is important for the preservation of tallgrass prairie and improving livestock production, it has become a controversial societal issue because of its potential impact on air quality standards. Over the past 80 years, recommendations on burning in Kansas have ranged from total fire exclusion to burning only in late April; and for the past 40 years, the concept that burning should only occur in late spring has become ingrained in the cultural practices of rangeland management. Yet the scientific basis for these recommendations has received little rigorous scrutiny. Herein, we critically review the research on dormant-season burning in the Flint Hills that formed the foundation for modern burn practices in Kansas. Close examination of the historical data does not support the tenet that burning must be limited to a narrow window in late spring. Many conclusions of the research that led to recommending burning only in late spring were ambiguous, not subjected to statistical analysis, or were influenced by an antiburn bias. Current research suggests that timing of a burn is not as critical as ranchers have been led to believe and burning does not have to be restricted to a narrow window in late April. There is an absence of scientific evidence that burning earlier in the spring adversely affects forage production, plant species composition, soil moisture, or cattle weight gain. Although there is a need for research on the consequences of burning grazed pastures at different times of the year, expanding the window for burning earlier in the dormant season should help alleviate air quality issues downwind of the burned areas and potentially be beneficial to ranchers.