Land Use and Small Mammal Predation Effects on Shortgrass Prairie Birds

ABSTRACT Grassland birds endemic to the central shortgrass prairie ecoregion of the United States have experienced steep and widespread declines over the last 3 decades, and factors influencing reproductive success have been implicated. Nest predation is the major cause of nest failure in passerines, and nesting success for some shortgrass prairie birds is exceptionally low. The 3 primary land uses in the central shortgrass prairie ecoregion are native shortgrass prairie rangeland (62%), irrigated and nonirrigated cropland (29%), and Conservation Reserve Program (CRP, 8%). Because shortgrass–cropland edges and CRP may alter the community of small mammal predators of grassland bird nests, I sampled multiple sites on and near the Pawnee National Grasslands in northeast Colorado, USA, to evaluate 1) whether small mammal species richness and densities were greater in CRP fields and shortgrass prairie–cropland edges compared to shortgrass prairie habitats, and 2) whether daily survival probabilities of ground-nesting grassland bird nests were negatively correlated with densities of small mammals. Small mammal species richness and densities, estimated using trapping webs, were generally greater along edges and on CRP sites compared to shortgrass sites. Vegetation did not differ among edges and shortgrass sites but did differ among CRP and shortgrass sites. Daily survival probabilities of artificial nests at edge and CRP sites and natural nests at edge sites did not differ from shortgrass sites, and for natural nests small mammal densities did not affect nest survival. However, estimated daily survival probability of artificial nests was inversely proportional to thirteen-lined ground squirrel (Spermophilus tridecemlineatus) densities. In conclusion, these data suggest that although land-use patterns on the shortgrass prairie area in my study have substantial effects on the small mammal community, insufficient data existed to determine whether land-use patterns or small mammal density were affecting grassland bird nest survival. These findings will be useful to managers for predicting the effects of land-use changes in the shortgrass prairie on small mammal communities and avian nest success.     

Habitat selection by mountain plovers in shortgrass steppe

Much of the breeding range for the mountain plover (Charadrius montanus) occurs in shortgrass steppe and mixed-grass prairie in the western Great Plains of North America. Studies of mountain plovers in shortgrass steppe during the 1970s and 1990s were focused in Weld County, Colorado, which was considered a key breeding area for the species. These studies, however, did not include habitats influenced by black-tailed prairie dogs (Cynomys ludovicianus) or prescribed fire. The role of these 2 rangeland disturbance processes has increased substantially over the past 15 years. During 2008–2009, I used radial distance point count surveys to estimate mountain plover densities early in the nesting season in 4 habitats on public lands in Weld County, Colorado. All 4 habitats were grazed by cattle during the growing season at moderate stocking rates but had different additional disturbances consisting of 1) dormant-season prescribed burns, 2) active black-tailed prairie dog colonies, 3) black-tailed prairie dog colonies affected by epizootic plague in the past 1–2 years, and 4) rangeland with no recent history of fire or prairie dogs. Mountain plover densities were similar on active black-tailed prairie dog colonies ( = 6.8 birds/km², 95% CI = 4.3–10.6) and prescribed burns ( = 5.6 birds/km², 95% CI = 3.5–9.1). In contrast, no plovers were detected at randomly selected rangeland sites grazed by cattle but lacking recent disturbance by prairie dogs or fire, even though survey effort was highest for this rangeland habitat. Mountain plover densities were intermediate (2.0 birds/km², 95% CI = 0.8–5.0) on sites where black-tailed prairie dogs had recently been extirpated by plague. These findings suggest that prescribed burns and active black-tailed prairie dog colonies may enhance breeding habitat for mountain plovers in shortgrass steppe and illustrate the potential for suppressed or altered disturbance processes to influence habitat availability for declining wildlife species. © 2011 The Wildlife Society.     

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

Summary Studies were conducted during the 1979 growing season to examine how North American bison (Bison bison) use prairie dog (Cynomys ludovicianus) colonies in Wind Cave National Park, South Dakota. Objectives included (1) determining whether bison selected for prairie dog towns parkwide; (2) characterizing in greater detail bison use patterns of a 36-ha colony in Pringle Valley as a function of time since prairie dog colonization; and (3) relating these bison use patterns to measured changes in structure and nutritional value of vegetation on and off the dog town.During midsummer, prairie dog towns were one of the most frequently used habitats by bison parkwide. Day-long observations at Pringle Valley revealed that bison exerted strong selection (nearly 90% of all habitat use and feeding time) for the dog town, which occupied only 39% of the valley. While there, they partitioned their use of the colony by grazing in moderately affected areas (occupied <8 years by prairie dogs) and by resting in the oldest area (>26 years occupation).Prairie dogs facilitate bison habitat selection for a shortgrass successional stage in this mixed-grass community by causing a broad array of compositional, structural, and nutritional changes in the vegetation.     

Plant species richness and species-area relations in a shortgrass steppe in Colorado

Abstract. Plant species richness and species-area relations were examined for three landscapes (toposequences), each with a summit or upland, a midslope and a toeslope or lowland, in a shortgrass steppe in Colorado. The number of plant species in the largest plot size (0.16 ha) varied from 38 to 53. Neither the exponential relationship: s = a + b log A, nor the power function: S = cAz fit the data equally well in all situations. The processes acting upon species diversity here seem to operate at two spatial scales. The number of species in plots smaller than 3 m² was independent of the total number of species in the 0.16-ha plots and was constrained by the presence of the dominant bunchgrasses. Beyond 3 m², species number in each plot size was a function of the total number of species in the 0.16-ha plot.     

Fine-scale habitat associations of Oklahoma's longspurs

Overwintering is a key demographic stage for migratory birds but remains poorly understood, especially among multiple declining grassland bird species. The non-breeding ranges all 4 species of longspur (i.e., chestnut-collared [Calcarius ornatus], Smith's [C. pictus], Lapland [C. lapponicus], thick-billed [Rhynchophanes mccownii]) overlap in Oklahoma and the Texas Panhandle, USA, making this region ideal to study their wintering ecology. We evaluated the relationship between wintering longspur occurrence and fine-scale habitat characteristics using a combination of standardized bird surveys and vegetation plot sampling. Our study encompassed large, representative tracts of 3 prairie ecosystems (i.e., shortgrass, mixed-grass, and tallgrass prairies) that intersect within the Southern Great Plains, during winters of 2018–2019 and 2019–2020. Using randomization tests and classification trees, we characterized longspur habitats and compared these associations across the 3 prairie ecosystems. Fine-scale winter habitats (horizontal structure, vertical structure, and species compositions) varied among all 4 longspur species, varied at very fine scales, and differed between grassland types. Our findings can be applied to the management of grasslands such as decreasing vegetation height in mixed-grass prairies for chestnut-collared longspurs or removing woody vegetation in shortgrass prairies for thick-billed longspurs to help develop full-life cycle conservation for longspurs, which have experienced population declines.     

Hemlock woolly adelgid invasion affects microhabitat characteristics and small mammal communities

Hemlock woolly adelgid (HWA) invasion and preemptive logging practices alter the habitat structure of New England forests and may indirectly affect associated small mammal communities. Microhabitat structure was measured and small mammals were censused in eight large experimental plots to quantify these effects. The Harvard Forest long-term ecological research experiment is a replicated two-block design that includes four ~ 0.81-ha canopy treatments: (1) hemlock control, (2) hardwood control, (3) girdled Treatment, in which hemlock trees were killed by girdling in 2005 and left standing to simulate HWA invasion, and (4) logged treatment, in which trees were removed to simulate preemptive logging management practices. Nine microhabitat characteristics were measured from plot photos revealing differences among ground and canopy microhabitat structure. Small mammals were censused during high (2012) and low (2013) abundant years. Populations of common species were estimated with mark-recapture analysis. Peromyscus spp. were not affected by treatment in either year, but southern red-backed vole populations were greatest in the girdled treatments in 2012 and rarely captured in 2013. Between 6 and 9 mammal species were recorded in all treatments and species composition varied slightly. Estimated species richness was greater in girdled treatments than hemlock controls, but did not differ between girdled and logged treatments, which suggests preemptive logging is as detrimental to some small mammal species as HWA invasion. Overall, there is little evidence of a major shift in small mammal community structure in response to HWA invasion, with only minor changes in relative abundance both years.     

Oviposition Site Selection by the Regal Fritillary, Speyeria idalia, as Affected by Proximity of Violet Host Plants

Selection of oviposition sites by female regal fritillary butterflies, Speyeria idalia (Drury), in relation to the location and abundance of their larval food plant, Viola pedatifida G. Don, was studied in Kansas tallgrass prairie. To identify potential cues that females use to select oviposition microsites, we compared plots in which eggs were laid with paired control plots in terms of violet abundance, distance from plot center to the nearest violet plant, plant species composition and relative abundance, plant diversity (Shannon's index), and average plant canopy height. No significant differences occurred between control and oviposition sites for any of these variables (P > 0.15). Violets were present in only 30% of the 1-m-diameter oviposition plots, compared with 24% of the corresponding controls. Furthermore, eggs were never laid directly on violets even when they were present nearby. These data imply that S. idalia females do not preferentially orient to or oviposit on their larval food plant. Nevertheless, female S. idalia did not oviposit randomly, for they invariably chose microsites near the ground that were shaded by the grass/forb overstory and usually deposited eggs on the underside of dead vegetation. We speculate that environmental stresses such as high temperatures and intense solar insolation during the egg stage or harsh overwintering conditions experienced by first-instar larvae impact survival and influence choice of oviposition sites. High fecundity (more than 2000 eggs in some individuals) may allow this species to use a conditional, sweepstakes strategy, i.e., producing and spreading many eggs over prairie that harbors host plants to compensate for low egg and larval survivorship.     

Contribution of tree species to the biodiversity of a 1ha Old World rainforest in Brunei, Borneo

Tree biodiversity data is presented for a 1-ha permanent study plot on a low slope position of the Kuala Belalong valley in Temburong District, Brunei, Borneo. This is the first study in the region to document the tree flora of a low slope. All trees 5cm at 1.3m were mapped, measurements of dbh and height taken, and identified to species and morpho-species level. The total and relative values of each family for density, diversity, dominance and Importance Value Indices were calculated. A total of 1062 stems were recorded on the plot (1019 live, 43 dead) with a total of 278 species representing 110 genera in 49 families. The most ecologically significant family as determined by the Family Importance Value (FIV) was Euphorbiaceae with a value of 40 and'the'most significant species, using the Importance Value (IV), was morpho-speciesFordia splendissima with an IV of 29. The total basal area of all living trees was 31.43m². Forest structure was marked by a few tall (>45m) emergents, for example,Koompassia excelsa, Crypteronia griffithii and two species of the Dipterocarpaceae family, however, lower understorey and sub-canopy species dominated the plot with the average height of all individuals over the hectare being 11m.     

Impacts of ungulates on the demography and diversity of small mammals in central Kenya

The impacts of ungulates on small mammals in an East African savanna habitat were investigated by monitoring the population and community responses of small mammals on replicated 4-ha plots from which ungulates had been excluded. The dominant small mammal in this habitat is the pouched mouse, Saccostomusmearnsi, a medium-sized murid rodent. Eight other small mammal species, including Arvicanthis sp., Mus sp., Mastomys sp., Dendromus sp., Crocidura sp., and, rarely, Tatera sp., Aethomys sp., and Acomys sp., were also captured. The dominant ungulates are elephant (Loxodonta africana), giraffe (Giraffa camelopardalis), Grevy's and common zebra (Equus grevyi and E. burchelli), buffalo (Syncerus cafer), eland (Taurotragus oryx), Grant's gazelle (Gazella granti), and domestic cattle. Within 1 year, S. mearnsi populations had responded dramatically to the exclusion of large mammals by a two-fold increase in density, a difference that was maintained through pronounced seasonal fluctuations in the second year. Though individual pouched mice showed no significant differences in their use of space with and without ungulates, male S. mearnsi maintained significantly higher body weights in the absence of ungulates, indicating that habitat quality had increased. One other species, Mastomys sp., also increased in the absence of ungulates. Overall, the small mammal community maintained relatively constant species diversity on the plots to which ungulates did not have access. On the plots to which ungulates did have access, on the other hand, there was a rapid 75% decrease in diversity in the control plots during one trapping session. Ungulates are most likely affecting small mammals through their effects on food quality, since there were no detectable differences in their exposure to predators, as determined by vegetative cover, in the absence of ungulates. These results demonstrate that ungulates can have strong and rapid impacts on small mammal abundance and diversity in East African savannas, an interaction which has not previously been given serious consideration. Received: 28 December 1997 / Accepted: 11 May 1998     

Spatial structure and genetic diversity of three tropical tree species with different habitat preferences within a natural forest

Analyses of the spatial distribution pattern, spatial genetic structure and genetic diversity were carried out using a 33-ha plot in a hill dipterocarp forest for three dipterocarps with different habitat preferences, i.e. Shorea curtisii on the ridges, Shorea leprosula in the valleys and Shorea macroptera both on the ridges and in the valleys. The significant spatial aggregation in small-diameter trees of all the three species was explained by limited seed dispersal. At the large-diameter trees, only S. macroptera showed random distribution and this might further prove that S. macroptera is habitat generalist, whilst S. curtisii and S. leprosula are habitat specific. The levels of genetic diversity estimated based on five microsatellite loci were high and comparable in all the three studied species. As the three studied species reproduced mainly through outcrossing, the observed high levels of genetic diversity might support the fact that the plant mating system can be used as guideline to infer the levels of genetic diversity, regardless of whether the species is habitat specific or habitat generalist. The lack of spatial genetic structure but significant aggregation in the small-diameter trees of all the three species might indicate limited seed dispersal but extensive pollen flow. Hence, if seed dispersal is restricted but pollen flow is extensive, significant spatial aggregation but no spatial genetic structure will be observed at the small-diameter trees, regardless of whether the species is habitat specific or habitat generalist. The inferred extensive pollen flow might indicate that energetic pollinators are involved in the pollination of Shorea species in the hill dipterocarp forests.     

Occupancy of Mountain Plover and Burrowing Owl in Colorado

Abstract: Concern over the decline of grassland birds has spurred efforts to increase understanding of grassland bird-habitat relationships. Previous studies have suggested that black-tailed prairie dogs (Cynomys ludovicianus) provide important habitat for shortgrass prairie avifauna, such as mountain plover (Charadrius montanus) and western burrowing owl (Athene cunicularia hypugaea), although such studies are lacking in Colorado (USA). We used methods to estimate occupancy (ψ) of mountain plover and burrowing owl on prairie dog colonies and other shortgrass prairie habitats in eastern Colorado. Mountain plover occupancy was higher on prairie dog colonies (ψ = 0.50, 95% CI = 0.36–0.64) than on grassland (ψ = 0.07, 95% CI = 0.03–0.15) and dryland agriculture (ψ = 0.13, 95% CI = 0.07–0.23). Burrowing owl occupancy was higher on active prairie dog colonies (ψ = 0.80, 95% CI = 0.66–0.89) compared with inactive colonies (ψ = 0.23, 95% CI = 0.07–0.53), which in turn was much higher than on grassland (ψ = 0.01, 95% CI = 0.00–0.07) and dryland agriculture (ψ = 0.00, 95% CI ψ 0.00–0.00). Mountain plover occupancy also was positively correlated with increasing amounts of prairie dog colony in the landscape. Burrowing owl occupancy was negatively correlated with increasing amounts of prairie dog colony in the surrounding landscape. Our results suggest that actions to conserve mountain plovers and burrowing owls should incorporate land management to benefit prairie dogs. Because managing for specific colony attributes is difficult, alternative management that promotes heterogeneity may ensure that suitable habitat is available for the guild of grassland inhabitants.     

Importance of topography and soil texture in the spatial distribution of two sympatric dipterocarp trees in a Bornean rainforest

Relationships between spatial distributions and site conditions, namely topography and soil texture, were analyzed for two congeneric emergent trees, Dryobalanops aromatica and Dryobalanops lanceolata (Dipterocarpaceae), in a tropical rainforest in Sarawak, East Malaysia. A 52-ha permanent plot was divided into 1300 quadrats measuring 20m×20m; for each Dryobalanops species, the number and total basal area of trees 1cm in d.b.h. were compared among groups of quadrats with different site conditions. Because spatial distributions of both Dryobalanops and site-condition variables were aggregated, Monte-Carlo permutation tests were applied to analyze the relationships. Both single and multifactor statistical tests showed that the density and basal area distributions of the two species were significantly non-random in relation to soil texture and topographic variables. D.aromatica was significantly more abundant at higher elevations, in sandy soils, and on convex and steep slopes. In contrast, D.lanceolata preferred lower elevations and less sandy soils. In the study plot, there were very few sites (3 of 1150 quadrats tested) where the models of Hayashis method predicted the co-occurrence of the two species. These results suggest that between-species differences in habitat preferences are so large that they alone explain the spatially segregated distributions of these two species within the 52-ha study plot.     

Carbon isotope ratios and crop analyses of Arphia (Orthoptera: Acrididae) species in southeastern Wyoming Grassland

Summary Food habits of Arphia conspersa Scudder and Arphia pseudonietana (Thomas) were studied along an altitudinal transect in southeastern Wyoming shortgrass mixed prairie. Stable carbon isotope ratios indicated that diets were significantly different between study sites, between species, and between sexes. These differences were found to be primarily related to the availability of different food plants along the transect, although species with the C₃ pathway of photosynthesis were consumed in greater proportion than their availability in the habitat. The preference for C₃ species is presumably related to their higher nutritional value and digestibility, in spite of the fact that more time and energy must be spent to locate these food plants in some of the habitats studied. This study demonstrates the utility of the carbon isotope method in studying plant-animal interactions in habitats containing both C₃ and C₄ plants.     

A Test of Diversity–Productivity Models in Natural,Degraded, and Restored Wet Prairies

Conceptual restoration models depict strong correlations between structure and function, with both decreasing as an ecosystem is degraded and increasing during restoration. We evaluated the “linear” and “asymptotic” models by measuring diversity and annual net primary productivity (NPP) within four states of a southern Wisconsin floodplain: a remnant (unplowed) wet prairie, two degraded sites (soybean field and invaded prairie), and a restored prairie. Neither model fit our data for aboveground (ANPP), belowground (BNPP), or total (TNPP) productivity. ANPP declined as species richness increased (r = 0.998, df = 2), with highest values for soybeans (1,024 g/m²; two species in 30 0.25‐m² plots) and invaded prairie (937 g/m²; nine species, 99% cover of Phalaris arundinacea), intermediate for restored prairie (712 g/m²; 28 species), and lowest for diverse remnant prairie (571 g/m²; 36 species). In contrast, BNPP was lowest for soybeans (225 g/m²) and highest for remnant prairie (571 g/m²). TNPP in restored prairie (990 g/m²) matched that of the remnant (1,147 g/m²) within 7 years, but root:shoot NPP ratios were quite different (0.39 and 0.99, respectively). Overall, results suggest that the relationship between species diversity and productivity can differ with the component measured (ANPP, BNPP, or TNPP) and that diversity does not ensure high productivity. Because measuring ANPP does not fully test ecosystem‐function theory, we recommend assessing BNPP and additional ecosystem processes in future attempts to determine whether adding species will restore more function to degraded ecosystems.     

Plant community responses to resource availability and heterogeneity during restoration

Availability and heterogeneity of resources have a strong influence on plant community structure in undisturbed systems, as well as those recovering from disturbance. Less is known about the role of resource availability and heterogeneity in restored communities, although restoration provides a valuable opportunity to test our understanding of factors that influence plant community assembly. We altered soil nitrogen (N) availability and soil depth during a prairie restoration to determine if the availability and/or heterogeneity of soil resources influenced plant community composition in restored grassland communities. Plant community responses to three levels of N availability (ambient, enriched by fertilization, and reduced by carbon amendment) and two levels of soil depth (deep and shallow) were evaluated. In addition, we evaluated plant community responses to four whole plot heterogeneity treatments created from the six possible combinations of soil N availability and soil depth. The soil depth treatment had little influence on community structure during the first 3 years of restoration. Total diversity and richness declined over time under annual N enrichment, whereas diversity was maintained and richness increased over time in soil with reduced N availability. Non-native species establishment was lowest in reduced-N soil in the initial year, but their presence was negligible in all of the soil N treatments by the second year of restoration. Panicum virgatum, a native perennial C₄ grass, was the dominant species in all soil N treatments by year three, but the magnitude of its dominance was lowest in the reduced-N soil and highest in enriched-N soil. Consequently, the relative cover of P. virgatum was strongly correlated with community dominance and inversely related to diversity. The differential growth response of P. virgatum to soil N availability led to a higher degree of community similarity to native prairie in the reduced-N treatment than in the enriched-N treatment. There were no differences in plant community structure among the four whole plot-level heterogeneity treatments, which all exhibited the same degree of similarity to native prairie. Diversity and community heterogeneity in the whole-plot treatments appeared to be regulated by the dominant species effect on light availability, rather than soil N heterogeneity per se. Our results indicate that a strong differential response of a dominant species to resource availability in a restored community can regulate community structure, diversity, and similarity to the native (or target) community, but the importance of resource heterogeneity in restoring diversity may be dampened in systems where a dominant species can successfully establish across a range of resource availability.     

Productivity and resistance to weed invasion in four prairie biomass feedstocks with different diversity

High‐diversity mixtures of native tallgrass prairie vegetation should be effective biomass feedstocks because of their high productivity and low input requirements. These diverse mixtures should also enhance several of the ecosystem services provided by the traditional monoculture feedstocks used for bioenergy. In this study, we compared biomass production, year‐to‐year variation in biomass production, and resistance to weed invasion in four prairie biomass feedstocks with different diversity: one species – a switchgrass monoculture; five species – a mix of C₄ grasses; 16 species – a mix of grasses, forbs, and legumes; and 32 species – a mix of grasses, forbs, legumes, and sedges. Each diversity treatment was replicated four times on three soil types for a total of 48 research plots (0.33–0.56 ha each). We measured biomass production by harvesting all plant material to ground level in ten randomly selected quadrats per plot. Weed biomass was measured as a subset of total biomass. We replicated this design over a five‐year period (2010–2014). Across soil types, the one‐, 16‐, and 32‐species treatments produced the same amount of biomass, but the one‐species treatment produced significantly more biomass than the five‐species treatment. The rank order of our four diversity treatments differed between soil types suggesting that soil type influences treatment productivity. Year‐to‐year variation in biomass production did not differ between diversity treatments. Weed biomass was higher in the one‐species treatment than the five‐, 16‐, and 32‐species treatments. The high productivity and low susceptibility to weed invasion of our 16‐ and 32‐species treatments supports the hypothesis that high‐diversity prairie mixtures would be effective biomass feedstocks in the Midwestern United States. The influence of soil type on relative feedstock performance suggests that seed mixes used for biomass should be specifically tailored to site characteristics for maximum productivity and stand success.     

Helianthus maximiliani and species fine‐scale spatial pattern affect diversity interactions in reconstructed tallgrass prairies

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Low genetic differentiation between populations of an endemic prairie katydid despite habitat loss and fragmentation

Tallgrass prairie habitats within North America have suffered severe fragmentation and habitat loss as land has been converted for agricultural purposes. Habitat loss and fragmentation can affect gene flow and the genetic structure of insect populations. Neoconocephalus bivocatus is a prairie obligate katydid found only in isolated prairie patches. We compared genetic diversity and population differentiation using AFLP markers in N. bivocatus and N. robustus, a grassland generalist that is not isolated to prairie fragments and occupies a more contiguous range. Similar levels of genetic diversity were present within populations of both species. While population genetic structure was found in both species, there was no relationship between assigned genotypes and sampling localities. This genetic structure may instead be evidence of a past barrier to gene flow that has since been removed. Genetic differentiation within both species was low, with no evidence of a correlation with geographic distance, indicating neither species is dispersal limited at these distances. We see no significant reduction in genetic diversity or genetic differentiation within N. bivocatus when compared to N. robustus. We therefore conclude that while N. bivocatus utilizes a fragmented landscape, long-distance dispersal likely maintains gene flow between isolated prairie patches.     

Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community

Summary We examined the impact of pocket gopher disturbances on the dynamics of a shortgrass prairie community. Through their burrowing activity, pocket gophers (Thomomys bottae) cast up mounds of soil which both kill existing vegetation and create sites for colonization by competitively-inferior plant species. Three major patterns emerge from these disturbances: First, we show that 10 of the most common herbaceous perennial dicots benefit from pocket gopher disturbance; that is, a greater proportion of seedlings are found in the open space created by pocket gopher disturbance than would be expected based on the availability of disturbed habitat. Additionally, these seedlings exhibited higher growth rates than adjacent seedlings of the same species growing in undisturbed habitat. Second, we tested two predictions of the Intermediate Disturbance Hypothesis and found that species diversity was greatest for plots characterized by disturbances of intermediate age. However, we did not detect significant differences in diversity between plots characterized by intermediate and high levels of disturbance, indicating that many species are adapted to or at least tolerant of high levels of disturbance. Third, we noted that the abundance of grasses decreased with increasing disturbance, while the abundance of dicots increased with increasing disturbance.