A century of increasing pine density and associated shifts in understory plant strategies

We analyzed one of the longest-term ecological data sets to evaluate how forest overstory structure is related to herbaceous understory plant strategies in a ponderosa pine forest. Eighty-two permanent 1-m2 chart quadrats that were established as early as 1912 were remeasured in 2007. We reconstructed historical forest structure using dendrochronological techniques. Ponderosa pine basal area increased from an average of 4 m2/ha in the early 1900s to 29 m2/ha in 2007. Understory plant foliar cover declined by 21%, species richness declined by two species per square meter, and functional diversity also declined. The relative cover of C4 graminoids decreased by 18% and C3 graminoids increased by 19%. Herbaceous plant species with low leaf and fine root nitrogen concentrations, low specific leaf area, high leaf dry matter content, large seed mass, low specific root length, short maximum height, and early flowering date increased in relative abundance in sites where pine basal area increased the most. Overall, we observed a long-term shift in composition toward more conservative shade- and stress-tolerant herbaceous species. Our analysis of temporal changes in plant strategies provides a general framework for evaluating compositional and functional changes in terrestrial plant communities.     

Effects of Livestock Grazing on Stand Dynamics and Soils in Upland Forests of the Interior West

Many ponderosa pine and mixed-conifer forests of the western, interior United States have undergone substantial structural and compositional changes since settlement of the West by Euro-Americans. Historically, these forests consisted of widely spaced, fire-tolerant trees underlain by dense grass swards. Over the last 100 years they have developed into dense stands consisting of more fire-sensitive and disease-susceptible species. These changes, sometimes referred to as a decline in "forest health," have been attributed primarily to two factors: active suppression of low-intensity fires (which formerly reduced tree recruitment, especially of fire-sensitive, shade-tolerant species), and selective logging of larger, more fire-tolerant trees. A third factor, livestock grazing, is seldom discussed, although it may be as important as the other two factors. Livestock alter forest dynamics by (1) reducing the biomass and density of understory grasses and sedges, which otherwise outcompete conifer seedlings and prevent dense tree recruitment, and (2) reducing the abundance of fine fuels, which formerly carried low-intensity fires through forests. Grazing by domestic livestock has thereby contributed to increasingly dense western forests and to changes in tree species composition. In addition, exclosure studies have shown that livestock alter ecosystem processes by reducing the cover of herbaceous plants and litter, disturbing and compacting soils, reducing water infiltration rates, and increasing soil erosion.     

Utilisation of a coastal grassland by geese after managed re-alignment

In this study we evaluate the effect of coastal re-alignment on the utilisation of coastal grasslands by staging geese. We assessed vegetation change and utilisation by geese using repeated mapping and regular dropping counts in both the restored marsh and adjacent reference sites. All measurements were started well before the actual re-alignment. In addition, we studied the effects of livestock grazing on vegetation and geese, using exclosures. The vegetation transformed from fresh grassland into salt-marsh vegetation. A relatively large proportion of the de-embanked area became covered with secondary pioneer vegetation, and the overall cover of potential food plants for geese declined. Goose utilisation had initially dropped to low levels, both in autumn and in spring, but it recovered to a level comparable to the reference marsh after ten years. Exclosure experiments revealed that livestock grazing prevented the establishment of closed swards of grass in the poorly drained lower area of the restored marsh, and thereby negatively affected goose utilisation of these areas during spring staging. Goose grazing in the restored marsh during spring showed a positive numerical response to grass cover found during the preceding growing season. (1) The value of restored salt marsh as foraging habitat for geese initially decreased after managed re-alignment but recovered after ten years. (2) Our findings support the idea that the value of foraging habitats depends largely on the cover of forage plants and that this can be manipulated by adjusting both grazing and drainage.     

Effects of Historic Livestock Grazing on Vegetation at Chaco Culture National Historic Park, New Mexico

Abstract:   Livestock grazing is the most ubiquitous land use in western North America, yet it rarely has been studied in a controlled manner because of the lack of large areas free of grazing. We compared the ecological effects of three grazing treatments—long-term protection, short-term protection, and currently grazed—at Chaco Culture National Historic Park in northern New Mexico. Chaco has a long history of human habitation and is now one of the largest grazing exclosures in the American West. We studied the effects of livestock grazing on the cover of plants, soil crusts, and plant species richness at six sites with different potential natural vegetation. Species richness was higher under long-term protection than under current grazing at all six sites. Trends in shrub and grass response varied significantly across the six sites. Shrub cover increased with long-term protection at four upland sites, and grass cover increased with protection at four sites. The response of Chaco vegetation to release from grazing varied significantly according to each site's ecological potential, determined in part by edaphic and topographic characteristics. These nuances in vegetation response represent natural ecological variation and contrast with the notions of widespread shrub "invasion" often inferred in the past.     

Changes in Avian and Plant Communities of Aspen Woodlands over 12 Years after Livestock Removal in the Northwestern Great Basin

Abstract: Riparian and quaking aspen (Populus tremuloides) woodlands are centers of avian abundance and diversity in the western United States, but they have been affected adversely by land use practices, particularly livestock grazing. In 1990, cattle were removed from a 112,500‐ha national wildlife refuge in southeastern Oregon. Thereafter, we monitored changes in vegetation and bird abundance in years 1–3 (phase 1) and 10–12 (phase 2) in 17 riparian and 9 snow‐pocket aspen plots. On each 1.5‐ha plot, we sampled vegetation in 6 transects. Three times during each breeding season, observers recorded all birds 50 m to each side of the plot's 150‐m centerline for 25 minutes. We analyzed data with multivariate analysis of variance and paired t tests with p values adjusted for multiple comparisons. In both periods, riparian and snow‐pocket aspen produced extensive regeneration of new shoots ( stems/ha and 7079 stems/ha, respectively). By phase 2, a 64% increase in medium‐diameter trees in riparian stands indicated successful recruitment into the overstory, but this pattern was not seen in snow‐pocket stands, where the density of trees was over 2 times greater. By phase 2 in riparian and snow‐pocket stands, native forb cover had increased by 68% and 57%, respectively, mesic shrub cover had increased by 29% and 58%, and sagebrush cover had decreased by 24% and 31%. Total avian abundance increased by 33% and 39% in riparian and snow‐pocket aspen, respectively, ground or understory nesters increased by 133% and 67% and overstory nesters increased by 34% and 33%. Similarly, ground or understory foragers increased by 25% and 32%, aerial foragers by 55% and 57%, and overstory foragers by 66% and 43%. We interpreted the substantial regeneration of aspen shoots, increased densities of riparian forbs and shrubs, and increased avian abundances as a multitrophic‐level response to the total removal of livestock and as substantial movement toward recovery of biological integrity.     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

Impacts of Livestock Grazing and Tree Clearing on Birds of Woodland and Riparian Habitats

Abstract:  We investigated the impact of pastoral management on birds in subtropical grassy eucalypt woodland in southeastern Queensland, Australia, where the patterns of land management have made it possible to disentangle the effects of livestock grazing from those of tree clearing. We recorded changes in bird species composition, density, and relative abundance across two woodland habitat types (riparian and nonriparian) and two levels of clearing (wooded and nonwooded) and three levels of livestock grazing (low, moderate, and high) replicated over space (1000 km²) and time (2001–2002). We predicted that species that depend on understory vegetation would be most negatively affected by livestock grazing. A Bayesian generalized linear model showed that the level of grazing had the greatest effect when trees were present. When trees were absent, the impact of grazing was overshadowed by the effects of a lack of trees. Over 65% of species responded to different levels of grazing, and the abundance of 42% of species varied markedly with habitat and grazing. The most common response to grazing was high species relative abundance under low levels of grazing (28% of species), species absence at high levels of grazing (20%), and an increase in abundance with increasing grazing (18%). Despite having similar bird assemblages, the effect of grazing was stronger in riparian habitat than in adjacent woodland habitat. Our results suggest that any level of commercial livestock grazing is detrimental to some woodland birds, particularly the understory-dependant species, as predicted. Nevertheless, provided trees are not cleared, a rich and abundant bird fauna can coexist with moderate levels of grazing. Habitats with high levels of grazing, on the other hand, resulted in a species-poor bird assemblage dominated by birds that are increasing in abundance nationally .     

Shifts in southern Wisconsin forest canopy and understory richness, composition, and heterogeneity

We resurveyed the under- and overstory species composition of 94 upland forest stands in southern Wisconsin in 2002-2004 to assess shifts in canopy and understory richness, composition, and heterogeneity relative to the original surveys in 1949-1950. The canopy has shifted from mostly oaks (Quercus spp.) toward more mesic and shade-tolerant trees (primarily Acer spp.). Oak-dominated early-successional stands and those on coarse, nutrient-poor soils changed the most in canopy composition. Understories at most sites (80%) lost native species, with mean species density declining 25% at the 1-m2 scale and 23.1% at the 20-m2 scale. Woody species have increased 15% relative to herbaceous species in the understory despite declining in absolute abundance. Initial canopy composition, particularly the abundance of red oaks (Quercus rubra and Q. velutina), predicted understory changes better than the changes observed in the overstory. Overall rates of native species loss were greater in later-successional stands, a pattern driven by differential immigration rather than differential extirpation. However, understory species initially found in early-successional habitats declined the most, particularly remnant savanna taxa with narrow or thick leaves. These losses have yet to be offset by compensating increases in native shade-adapted species. Exotic species have proliferated in prevalence (from 13 to 76 stands) and relative abundance (from 1.2% to 8.4%), but these increases appear unrelated to the declines in native species richness and heterogeneity observed. Although canopy succession has clearly influenced shifts in understory composition and diversity, the magnitude of native species declines and failure to recruit more shade-adapted species suggest that other factors now act to limit the richness, heterogeneity, and composition of these communities.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Effects of Long-Term Ungulate Exclusion and Recent Alien Species Control on the Preservation and Restoration of a Hawaiian Tropical Dry Forest

Abstract: Although the destruction of tropical rain forests receives much attention, tropical dry forests are in general far more threatened and endangered. Eliminating grazing ungulates is often considered a key first step toward protecting these ecosystems, but few studies have investigated the long-term effects of this technique. We examined the effects of ungulate exclusion from a 2.3-ha native dry-forest preserve on the island of Hawaii by comparing its present flora to the flora of an adjacent area subjected to continuous grazing since the preserve was fenced over 40 years ago. Relative to this adjacent area, the fenced preserve contained a more diverse flora with substantially greater coverage of native overstory and understory species. Until recently, however, regeneration of native canopy trees within the preserve appears to have been thwarted by a dominant herbaceous cover of alien fountain grass (   Pennisetum setaceum ) and predation by alien rodent species. Our results indicate that although ungulate exclusion may be a necessary and critical first step, it is not sufficient to adequately preserve and maintain Hawaii's remaining tropical dry forest remnants. Our recent efforts to control the dominant alien species within the fenced preserve suggest that this practice may facilitate both the regeneration of native species and the colonization and potential invasion of new alien plants. Comparisons of seedlings of the dominant native canopy tree Diospyros sandwicensis growing in sites both dominated by and free of fountain grass suggested that fountain grass inhibits Diospyros seedling growth and photosynthesis but may increase survival if seedlings are protected from ungulates.     

Demography of northern flying squirrels informs ecosystem management of western interior forests.

We studied northern flying squirrel (Glaucomys sabrinus) demography in the eastern Washington Cascade Range to test hypotheses about regional and local abundance patterns and to inform managers of the possible effects of fire and fuels management on flying squirrels. We quantified habitat characteristics and squirrel density, population trends, and demography in three typical forest cover types over a four-year period. We had 2034 captures of flying squirrels over 41 000 trap nights from 1997 through 2000 and marked 879 squirrels for mark-recapture population analysis. Ponderosa pine (Pinus ponderosa) forest appeared to be poorer habitat for flying squirrels than young or mature mixed-conifer forest. About 35% fewer individuals were captured in open pine forest than in dry mixed-conifer Douglas-fir (Pseudotsuga menziesii) and grand fir (Abies grandis) forests. Home ranges were 85% larger in pine forest (4.6 ha) than in mixed-conifer forests (2.5 ha). Similarly, population density (Huggins estimator) in ponderosa pine forest was half (1.1 squirrels/ha) that of mixed-conifer forest (2.2 squirrels/ha). Tree canopy cover was the single best correlate of squirrel density (r = 0.77), with an apparent threshold of 55% canopy cover separating stands with low- from high-density populations. Pradel estimates of annual recruitment were lower in open pine (0.28) than in young (0.35) and mature (0.37) forest. High recruitment was most strongly associated with high understory plant species richness and truffle biomass. Annual survival rates ranged from 45% to 59% and did not vary among cover types. Survival was most strongly associated with understory species richness and forage lichen biomass. Maximum snow depth had a strong negative effect on survival. Rate of per capita increase showed a density-dependent response. Thinning and prescribed burning in ponderosa pine and dry mixed conifer forests to restore stable fire regimes and forest structure might reduce flying squirrel densities at stand levels by reducing forest canopy, woody debris, and the diversity or biomass of understory plants, truffles, and lichens. Those impacts might be ameliorated by patchy harvesting and the retention of large trees, woody debris, and mistletoe brooms. Negative stand-level impacts would be traded for increased resistance and resilience of dry-forest landscapes to now-common, large-scale stand replacement fires.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Interactive effects of rabbit grazing and environmental factors on plant species-richness on dunes of Norderney

On the German island Norderney rabbit grazing effects on vegetation, soil and microclimate were investigated with exclosures in different dune vegetation. Differences in species composition between grazed and ungrazed plots increased with grazing activity. A decline in growth height resulted in higher light availability and temperature on the grazed sites. One of the most visible effects was the decline in flowering plants. Most grazing effects varied between habitat types. Grey dunes were distinctly affected by rabbit grazing. In relation to the habitat type, life forms were differently affected: in closed vegetation, rabbit activities further open patches, resulting in more therophytes. In contrast, in open vegetation rabbit activities were too strong and following sand dynamics too high for many therophytes. In general, grazing diversified habitats, slightly supported the total species-richness, and enhanced the heterogeneity of species composition. Highest species-richness in total and of herbs was reached at intermediate levels of grazing pressure. The study is in line that highest species-diversity will be found at intermediate levels of disturbance and environmental stress and that grazing effects vary with the habitat type.     

Aquatic herbivores facilitate the emission of methane from wetlands

Wetlands are significant sources of atmospheric methane. Methane produced by microbes enters roots and escapes to the atmosphere through the shoots of emergent wetland plants. Herbivorous birds graze on helophytes, but their effect on methane emission remains unknown. We hypothesized that grazing on shoots of wetland plants can modulate methane emission from wetlands. Diffusive methane emission was monitored inside and outside bird exclosures, using static flux chambers placed over whole vegetation and over single shoots. Both methods showed significantly higher methane release from grazed vegetation. Surface-based diffusive methane emission from grazed plots was up to five times higher compared to exclosures. The absence of an effect on methane-cycling microbial processes indicated that this modulating effect acts on the gas transport by the plants. Modulation of methane emission by animal-plant-microbe interactions deserves further attention considering the increasing bird populations and changes in wetland vegetation as a consequence of changing land use and climate change.     

间伐对杉木人工林林下植被多样性及其营养元素现存量影响

间伐改变了林分结构,影响林下植被的生长和发育,但长期间伐对杉木Cunninghamia lanceolata林下植被多样性的影响尚缺乏系统研究。在浙江开化通过样地调查,采用Simpson指数、Shannon-Wiener指数和Pielou均匀度指数研究3种间伐处理下杉木人工林林下植被多样性的变化;分析林下植被主要营养元素含量,研究间伐对营养元素现存量的影响。以试验区传统经营方式为对照,即林木生长中期（第14年）进行1次轻度间伐（间伐强度约15%,T1）;中度（总间伐强度约35%, T2）和强度（总间伐强度约50%,T3）间伐均在第7年和第14年进行了2次间伐。间伐15年后的结果表明,3种间伐处理下,林下植物种类数分别为18种（T1）、17种（T2）和20种（T3）,林下植被总密度和盖度均随间伐强度增加而增大;林下植物Simpson指数、Shannon-Wiener指数和Pielou均匀度指数在不同间伐处理之间均没有显著差异（P＞0.05）。林下植被全碳（TC）、全磷（TP）和全钾（TK）含量在不同间伐处理之间差异不显著（P＞0.05）,全氮（TN）含量随间伐强度增加而降低,其中T3处理显著低于T1（P＜0.05）。林下植被生物量随间伐强度增大而增加,林下植被TC、TN、TP和TK现存量在不同间伐处理之间均没有显著差异（P＞0.05）。这些试验结果说明杉木人工林间伐15年后间伐强度不会显著影响林下植被多样性和营养元素的积累。     

Relevance of rangeland degradation in semiarid northeastern South Africa to the nonequilibrium theory.

According to the nonequilibrium theory, livestock grazing has a limited effect on long-term vegetation productivity of semiarid rangelands, which is largely determined by rainfall. The communal lands in northeastern South Africa contain extensive degraded areas which have been mapped by the National Land Cover (NLC) program. Much evidence suggests that long-term heavy grazing is the cause of this degradation. In order to test for the prevalence of nonequilibrium dynamics, we determined the relative effects of rainfall- and grazing-induced degradation on vegetation productivity. The vegetation production in the NLC degraded areas was estimated using growth-season sums of the Normalized Difference Vegetation Index (sigmaNDVI), calculated using data from both the Advanced Very High Resolution Radiometer (AVHRR) (1985-2003) and Moderate-resolution Imaging Spectroradiometer (MODIS) (2000-2005). On average, rainfall and degradation accounted for 38% and 20% of the AVHRR sigmaNDVI variance and 50% and 33% of the MODIS sigmaNDVI variance, respectively. Thus, degradation had a significant influence on long-term vegetation productivity, and therefore the rangelands did not behave according to the nonequilibrium model, in which grazing is predicted to have a negligible long-term impact.     

Impacts of repeated timber skidding on the chemical properties of topsoil, herbaceous cover and forest floor in an eastern beech (Fagus orientalis Lipsky) stand

In this study, long-term timber skidding effects on herbaceous understory forest floor and soil were investigated on a skid road in a stand of the eastern beech (Fagus orientalis Lipsky). For this purpose, herbaceous understory forest floor and soil samples were collected from the skid road and from an undisturbed area used as a control plot. The mass (kg ha(-1)) of herbaceous and forest floor samples was determined, and soil characteristics were examined at two depths (0-5 cm and 5-10 cm). We quantified sand, silt and clay content, as well as bulk density compaction, pH, and organic carbon content in soil samples. The quantities of N, K, P, Na, Ca, Mg, Fe, Mn, Zn and Cu were determined in all herbaceous cover forest floor and soil samples. The quantities of Na, Fe, Zn, Cu and Mn in herbaceous understory samples from the skid road were considerably higher than those in the undisturbed area, while the quantity of Mg was considerably lower. These differences could have been caused by decreased herbaceous cover in addition to variations in the properties of the forest floor and soil after skidding. A lower amount of forest floor on the skid road was the result of skidding and harvesting activities. Mg and Zn contents in forest floor samples were found to be considerably lower for the skid road than for the undisturbed area. No significant differences were found in soil chemical properties (quantities of N, P, K, Na, Ca, Mg, Fe, Zn, Cu and Mn) at the 0-5 cm soil depth. Important differences exist between soil quantities of Mg at a 5-10 cm depth on the skid road and in undisturbed areas. Both 0-5 cm and 5-10 cm soil depths, the average penetrometer resistance values for the skid road was higher than for the undisturbed area. This result shows that the compaction caused by skidding is maintained to depth of 10 cm. Skid road soil showed higher bulk density values than undisturbed areas because of compaction.     

The oasis effect: response of birds to exurban development in a southwestern savanna

Ranches are being converted to exurban housing developments in the southwestern United States, with potentially significant but little-studied impacts on biological diversity. We counted birds in grasslands and savannas in southeastern Arizona that were grazed by livestock, embedded in low-density exurban housing developments, or both, or neither. Species richness and bird abundance were higher in exurban neighborhoods than in undeveloped landscapes, independent of livestock grazing. The positive response to development was particularly evident among doves, quail, hummingbirds, aerial insectivores, and some but not all ground-foraging sparrows. Effects of livestock grazing were comparatively minor and mostly involved birds with requirements for tall ground cover or the lack of it. The average rank correlation between counts of individual species and housing density was positive across all transects. However, this relationship disappeared among the exurban transects alone, and bird species richness on the exurban transects was negatively correlated with the number of homes nearby. These results suggest that the positive influence of exurban development on avian abundance and variety was greatest at the lowest housing densities. We attribute the attraction of many birds to exurban development to an oasis effect, in which resources otherwise scarce in arid southwestern environments (shade, nectar, nest sites, and especially water) are relatively abundant around exurban home sites. This finding is consistent with the hypothesis that exurban home sites represented resource supply points inside birds' home ranges otherwise consisting mostly of natural vegetation.     

Nitrogen spatial heterogeneity influences diversity following restoration in a ponderosa pine forest, Montana.

The resource heterogeneity hypothesis (RHH) is frequently cited in the ecological literature as an important mechanism for maintaining species diversity. The RHH has rarely been evaluated in the context of restoration ecology in which a commonly cited goal is to restore diversity. In this study we focused on the spatial heterogeneity of total inorganic nitrogen (TIN) following restoration treatments in a ponderosa pine (Pinus ponderosa)/Douglas-fir (Pseudotsuga menziesii) forest in western Montana, USA. Our objective was to evaluate relationships between understory species richness and TIN heterogeneity following mechanical thinning (thin-only), prescribed burning (burn-only), and mechanical thinning with prescribed burning (thin/burn) to discern the ecological and management implications of these restoration approaches. We employed a randomized block design, with three 9-ha replicates of each treatment and an untreated control. Within each treatment, we randomly established a 20 x 50 m (1000 m2) plot in which we measured species richness across the entire plot and in 12 1-m(2) quadrats randomly placed within each larger plot. Additionally, we measured TIN from a grid consisting of 112 soil samples (0-5 cm) in each plot and computed standard deviations as a measure of heterogeneity. We found a correlation between the net increase in species richness and the TIN standard deviations one and two years following restoration treatments, supporting RHH. Using nonmetric multidimensional scaling ordination and chi-squared analysis, we found that high and low TIN quadrats contained different understory communities in 2003 and 2004, further supporting RHH. A comparison of restoration treatments demonstrated that thin/burn and burn-only treatments created higher N heterogeneity relative to the control. We also found that within prescribed burn treatments, TIN heterogeneity was positively correlated with fine-fuel consumption, a variable reflecting burn severity. These findings may lead to more informed restoration decisions that consider treatment effects on understory diversity in ponderosa pine/Douglas-fir ecosystems.     

Wetlands as an alternative stable state in desert streams

Historically, desert drainages of the American southwest supported productive riverine wetlands (ciénegas). Region-wide erosion of ciénegas during the late 19th and early 20th century dramatically reduced the abundance of these ecosystems, but recent reestablishment of wetlands in Sycamore Creek, Arizona, USA, provides an opportunity to evaluate the mechanisms underlying wetland development. A simple model demonstrates that density-dependent stabilization of channel substrate by vegetation results in the existence of alternative stable states in desert streams. A two-year (October 2004-September 2006) field survey of herbaceous cover and biomass at 26 sites located along Sycamore Creek is used to test the underlying assumption of this model that vegetation cover loss during floods is density dependent, as well as the prediction that the distribution of vegetation abundance should shift toward bimodality in response to floods. Observations of nonlinear, negative relationships between herbaceous biomass prior to flood events and the proportion of persistent vegetation cover were consistent with the alternative stable state model. In further support of the alternative-state hypothesis, vegetation cover diverged from an approximately normal distribution toward a distinctly bimodal distribution during the monsoon flood season of 2006. These results represent the first empirically supported example of alternative-state behavior in stream ecosystems. Identification of alternative stable states in desert streams supports recent hypotheses concerning the importance of strong abiotic-disturbance regimes and biogeomorphic mechanisms in multiple-state ecosystems.