James K. Detling

Plant Response to Herbivory and Belowground Nitrogen Cycling

Plant responses to herbivory and links to belowground nitrogen cycling were investigated at Wind Cave National Park, South Dakota. Laboratory estimates of net nitrogen mineralization were highest in soils from the more altered areas of prairie dog colonies (Cynomys ludovicianus) and lowest in the adjacent, lightly grazed, uncolonized grassland. The ratio of CO2: net nitrogen mineralized, an index of immobilization, was highest in the uncolonized grassland and lowest in the altered core areas. Soil moisture was an important modifier of in situ field estimates of net nitrogen mineralization. Root biomass, an important carbon source for decomposers in perennial grasslands, was lowest in the altered core area and highest in the adjacent uncolonized grassland. Decreased nitrogen immobilization and increased net nitrogen mineralization in the laboratory in- cubations likely resulted from decreased root carbon inputs in grazed areas, which limited carbon availability to decomposers. Such increases in plant-available nitrogen may partially explain the frequently reported grazing-induced increases in shoot nitrogen concentrations. These studies suggest that carbon allocation to roots is a key link determining nitrogen- cycling responses to herbivory.

Relative growth rates and the grazing optimization hypothesis

SummaryA mathematical analysis of the changes in plant relative growth rates necessary to increase aboveground production following grazing was conducted. The equation derived gives an isoline where production of a grazed and ungrazed plant will be the same. The equation has four variables (mean shoot relative growth rate, change in relative growth rate after grazing, grazing intensity, and recovery time) and may be analyzed graphically in a number of ways.Under certain conditions, small increases in shoot relative growth rate following grazing will lead to increased aboveground production. Under other conditions, very large increases in relative growth rate after grazing can occur without production being increased over that of ungrazed plants. Plants growing at nearly their maximum potential relative growth rate have little opportunity to respond positively to grazing and potentially can sustain less grazing than plants with growth rates far below maximum. Plants with high relative growth rates at the time of grazing require large increases in growth rate while slow growing plants require only small increases. High grazing intensities are least likely to increase production and high grazing frequencies require greater responses than infrequent grazing events.

Physiological responses of plant populations to herbivory and their consequences for ecosystem nutrient flow

We explored how responses of two populations variable in grazing tolerance provide feedbacks to nutrient supply by controlling carbon supply to soil heterotrophs. The study focused on differences in production and carbon and nitrogen allocation patterns between the two populations. The grazing-tolerant population, or on-colony population, is found on intensively grazed prairie dog colonies, and a grazing-intolerant population, the off-colony population, is found in uncolonized grasslands. Equations describing the production and allocation responses to defoliation for the two ecotypes described were incorporated into CENTURY, a nutrientcycling simulation model. Simulations showed an increase in plant production that paralleled increases in net nitrogen mineralization. Production was greater with grazing and was maintained at higher grazing intensities for the on-colony than the off-colony population. Differences between the populations provided important controls over nitrogen losses. Feedbacks between plant responses to grazing and nitrogen cycling accounted for increased nitrogen availability with grazing. These feedbacks were more important determinants of ecosystem function than were fertilization effects of urine and feces deposition. The simulation results suggest that ecosystem function may be sensitive to physiological differences in population responses to periodic disturbances like herbivory.

Plant-herbivore interactions in a North American mixed-grass prairie

SummaryResearch was conducted to determine the effects of a native, sedentary rodent of North American grasslands, the black-tailed prairie dog (Cynomys ludovicianus), on seasonal aboveground plant biomass and nutrient dynamics and plant species diversity. The study was done on a northern mixed-grass prairie site at wind Cave National Park, South Dakota.Peak live plant biomass was greatest (190 g/m2) on the uncolonized part of the study area and least (95 g/m2) on a part of the prairie dog town colonized for 3 to 8 y. Peak live plant biomass (170 g/m2) of the oldest portion of the prairie dog town (colonized >26 y) was not significantly different from that of uncolonized prairie. However, where-as graminoids composed >85% of the total biomass of the latter area, forbs and dwarf shrubs (Artemisia frigida) were >95% of the total of the former. Both standing-dead plant biomass and litter declined markedly as time since colonization increased. Total plant species diversity (H) was greatest in the young prairie dog town (colonized for 3 to 8 y).Nitrogen concentration of plant shoots varied significantly as a function of time since colonization. Shoot-nitrogen was lowest in plants from the uncolonized site and greatest in plants collected from the longest-colonized areas of the prairie dog town. Shoot-nitrogen declined significantly over the growing season and tended to be higher in C3 graminoids than in C4 graminoids. In vitro digestible dry matter showed similar trends; the differences between C3 and C4 digestibilities were greatest during the last half of the growing season.We suggest that prairie dog-induced changes in plant biomass, plant species diversity, plant nutrient content, and forage digestibility may lead to further alterations of nutrient cycling and trophic dynamics in this mixed-grass prairie ecosystem.

Net photosynthesis, root respiration, and regrowth of Bouteloua gracilis following simulated grazing

SummaryNet photosynthesis (PN), root respiration (RR), and regrowth of Bouteloua gracilis (H.B.K.) Lag. were examined in the laboratory over a 10-day period following clipping to a 4-cm height to simulate grazing by large herbivores. Net photosynthesis rates of tissue remaining immediately following defoliation were only about 40% as great as preclipping rates. Three days after clipping, PN rates of defoliated plants had increased to values about 21% greater (per unit leaf area) than those of unclipped controls and remained at that level through Day 10. No statistically significant changes in RR occurred following defoliation. Biomass of unclipped plants nearly doubled during the 10-day study period, while that of defoliated plants increased 67%. Over half the new growth of defoliated plants was allocated to new leaf blades and only 18% to new roots, while only 33% of the new growth of control plants was allocated to new leaf blades but 29% went to new roots. As a consequence of increased PN rates and increased carbon allocation to synthesis of additional photosynthetic tissue following defoliation, net CO2 uptake per plant increased from 9% to 80% of that of the controls from Day 0 through Day 10.

Grassland Patch Dynamics and Herbivore Grazing Preference Following Urine Deposition

Field experiments were performed over two growing seasons to investigate the response of Schizachyrium scoparium (C4 photosynthetic pathway) and Poa pratensis (C3) to natural and simulated bison urine deposition in a northern, mixed prairie in South Dakota. We also assessed potential feedbacks of urine deposition on herbivore grazing by monitoring grass response to defoliation and herbivore grazing preference for vegetation occupying urine patches. Total aboveground biomass and root mass were higher and root:shoot ratios were lower on urine patches than in the surrounding plant community. Higher total aboveground biomass on urine patches resulted primarily from increased aboveground P. pratensis production. Urine deposition in May had little effect on aboveground production of S. scoparium except during July when S. scoparium was most active. Urine deposition date and plant phenology appear important in determining changes in species composition. Following urine deposition, aboveground N concentrations of P. pratensis and S. scoparium were higher on patches relative to conspecifics off patches. This increase in N concentration following urine deposition was greater in P. pratensis. We suggest the large increase in P. pratensis biomass following urine deposition is related to its relatively large response to increased soil N availability and its rhizomatous habit. Root N concentrations were higher on urine patches. Poa pratensis on urine patches initiated growth earlier in the season and postponed senescence relative to plants off patches. Aboveground production following clipping was greater on urine patches and N concentrations in regrowth of both species were higher than concentrations in plants not previously clipped. Aboveground herbivore utilization was greater on urine patches than on adjacent vege- tation. Although urine patches covered only 2% of the study site, they provided 7% of the biomass and 14% of the N consumed by aboveground herbivores from June through August. Urine patches probably provided an even greater source of forage and N for herbivores earlier and later in the growing season when surrounding vegetation was mostly quiescent.

Defoliation responses of western wheatgrass populations with diverse histories of prairie dog grazing

SummaryPhotosynthesis and regrowth were compared over a 10-day period following defoliation of about 75% of the tillers of western wheatgrass (Agropyron smithii) plants collected from a black-tailed prairie dog (Cynomys ludovicianus) town and a grazing exclosure at Wind Cave National Park, South Dakota. Prior to defoliation, dog town plants had more tillers, but fewer leaves per tiller, shorter and narrower leaf blades, more horizontal leaves, and higher leaf blade/leaf sheath ratios than plants from the grazing exclosure. Rates of net photosynthesis (PN) did not differ significantly among plants of the two populations, either prior to or following defoliation. From Days 2–10 following defoliation, PN of remaining undamaged leaves averaged 104% of predefoliation rates while PN of similar leaves on non-defoliated plants declined steadily with time. averaging only 79% predefoliation rates during this period. Following defoliation, transpiration rates followed similar trends to CO2 exchange, and rates did not differ between plants of the two populations. Absolute rates of leaf elongation and shoot production were greater in plants from the exclosure. However, defoliation of plants from the exclosure population resulted in a 20% reduction in their cumulative shoot dry weight, while cumulative shoot dry weight of plants from the prairie dog town was not significantly affected by defoliation. This apparent ability of plants from the prairie dog town population to withstand defoliation better than plants from the exclosure was atributed to factors such as the higher leaf blade/leaf sheath ratio and more horizontal leaf angles of plants from the former population.

Grazing History, Defoliation, and Competition: Effects on Shortgrass Production and Nitrogen Accumulation

Plants of Bouteloua gracilis were collected from heavily grazed prairie dog (Cynomys ludovicianus) colonies (ON—colony) and lightly grazed uncolonized sites (OFF—colony) in Wind Cave National Park, South Dakota. Our objectives were to determine (1) how grazing history affected production, nitrogen uptake, and biomass and nitrogen allocation following defoliation and (2) how such responses were affected by competing neighbors of the same population. The growth chamber experiment was factorial with three main treatments in 12 possible combinations. Main treatments were: population (plants from lightly and heavily grazed sites), defoliation (three intensities), and competition (with or without neighbors of the same population). There were no significant interactions among treatments. OFF—colony plants produced 121% as much biomass and their N. yield was 203% as great as ON—colony plants. ON—colony plants allocated a higher percentage of biomass and N to the roots from OFF—colony plants, while the latter allo...

Ecological roles and conservation challenges of social, burrowing, herbivorous mammals in the world's grasslands

The worlds grassland ecosystems are shaped in part by a key functional group of social, burrowing, herbivorous mammals. Through herbivory and ecosystem engineering they create distinctive and important habitats for many other species, thereby increasing biodiversity and habitat heterogeneity across the landscape. They also help maintain grassland presence and serve as important prey for many predators. However, these burrowing mammals are facing myriad threats, which have caused marked decreases in populations of the best-studied species, as well as cascading declines in dependent species and in grassland habitat. To prevent or mitigate such losses, we recommend that grasslands be managed to promote the compatibility of burrowing mammals with human activities. Here, we highlight the important and often overlooked ecological roles of these burrowing mammals, the threats they face, and future management efforts needed to enhance their populations and grassland ecosystems.

Rates of vegetation change associated with prairie dog (Cynomys ludovicianus) grazing in North American mixed-grass prairie

A prairie dog (Cynomys ludovicianus) colony with a known history of habitation was studied to quantify the effects of herbivory on plant species composition, dominance, stature and diversity in a North American mixedgrass prairie. Gradient analysis was used to quantify the relationship between plant community structure, prairie dog density, burrow density and habitation history and to document community-level responses of plants subjected to heavy grazing pressure. The results quantify the type, rate and extent of change which plant populations and communities may undergo in response to the differential grazing of plants variously tolerant of defoliation. Detrended correspondence analysis indicated that 69% of the between-sample floristic variance on the site was attributable to prairie dog habitation. Perennial grasses were rapidly displaced from the site within 3 yr of colonization and were replaced by annual forbs. The net result was an increase in species richness and diversity on the prairie dog colony. Within the colony, however, the number of species was more a function of stand size than colonization history. Significant decreases in canopy stature after 2 yr of habitation resulted from replacement of mid-height grass species by shortgrass species and forbs. In addition, there was a shift from tall growth forms of off-colony species to dwarf growth forms of the same species on the colony. Decreases in litter and increases in bare soil cover were substantial during the first 2 yr of habitation but changed little thereafter.