The effect of natural disturbances on forest biodiversity: an ecological synthesis.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity-disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the ß-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes.

Testing the hypothesis of hierarchical predictability in ecological restoration and succession.

To advance predictive ecology, the hypothesis of hierarchical predictability proposes that community measures for which species are interchangeable (e.g., structure and species richness) are more predictable than measures for which species identity matters (e.g., community composition). Predictability is hypothesized to decrease for response measures in order of the following categories: structure, species richness, function, and species composition. We tested this hypothesis using a 14-year, oak savanna-prairie restoration experiment that removed non-native pine plantations at 24 sites in northwestern Ohio, USA. Based on 24 response measures, the data showed minimal support for the hypothesis, because response measures varied in predictability within categories. Half of response measures had over half their variability modeled using fixed (restoration treatment and year) and random plot effects, and these "predictable" measures occurred in all four categories. Pine basal area, environment (e.g., soil texture), and antecedent vegetation accounted for over half the variation in change within the first three post-restoration years for 77% of response measures. Change between the 3rd and 14th years was less predictable, but most restoration measures increased favorably via sites achieving them in unique ways. We propose that variation will not conform with the hypothesis of hierarchical predictability in ecosystems with vegetation dynamics driven by stochastic processes such as seed dispersal, or where vegetation structure and species richness are influenced by species composition. The ability to predict a community measure may be more driven by the number of combinations of casual factors affecting a measure than by the number of values it can have.

Rapidly restoring biological soil crusts and ecosystem functions in a severely disturbed desert ecosystem.

Restoring biological soil crusts (biocrusts) in degraded drylands can contribute to recovery of ecosystem functions that have global implications, including erosion resistance and nutrient cycling. To examine techniques for restoring biocrusts, we conducted a replicated, factorial experiment on recently abandoned road surfaces by applying biocrust inoculation (salvaged and stored dry for two years), salvaged topsoil, an abiotic soil amendment (wood shavings), and planting of a dominant perennial shrub (Ambrosia dumosa). Eighteen months after treatments, we measured biocrust abundance and species composition, soil chlorophyll a content and fertility, and soil resistance to erosion. Biocrust addition significantly accelerated biocrust recovery on disturbed soils, including increasing lichen and moss cover and cyanobacteria colonization. Compared to undisturbed controls, inoculated plots had similar lichen and moss composition, recovered 43% of total cyanobacteria density, had similar soil chlorophyll content, and exhibited recovery of soil fertility and soil stability. Inoculation was the only treatment that generated lichen and moss cover. Topsoil application resulted in partial recovery of the cyanobacteria community and soil properties. Compared to untreated disturbed plots, topsoil application without inoculum increased cyanobacteria density by 186% and moderately improved soil chlorophyll and ammonium content and soil stability. Topsoil application produced 22% and 51% of the cyanobacteria density g⁻¹ soil compared to undisturbed and inoculated plots, respectively. Plots not treated with either topsoil or inoculum had significantly lower cyanobacteria density, soil chlorophyll and ammonium concentrations, and significantly higher soil nitrate concentration. Wood shavings and Ambrosia had no influence on biocrust lichen and moss species recovery but did affect cyanobacteria composition and soil fertility. Inoculation of severely disturbed soil with native biocrusts rapidly restored biocrust communities and soil stability such that restored areas were similar to undisturbed desert within three years. Using salvaged biocrust as inoculum can be an effective tool in ecological restoration because of its efficacy and simple implementation. Although salvaging biocrust material can be technically difficult and potentially costly, utilizing opportunities to salvage material in planned future disturbance can provide additional land management tools.

Ten years of vegetation assembly after a North American mega fire.

Altered fuels and climate change are transforming fire regimes in many of Earth's biomes. Postfire reassembly of vegetation--paramount to C storage and biodiversity conservation--frequently remains unpredictable and complicated by rapid global change. Using a unique data set of pre and long-term postfire data, combined with long-term data from nearby unburned areas, we examined 10 years of understory vegetation assembly after the 2002 Hayman Fire. This fire was the largest wildfire in recorded history in Colorado, USA. Resistance (initial postfire deviance from prefire condition) and resilience (return to prefire condition) declined with increasing fire severity. However, via both resistance and resilience, 'legacy' species of the prefire community constituted >75% of total plant cover within 3 years even in severely burned areas. Perseverance of legacy species, coupled with new colonizers, created a persistent increase in community species richness and cover over prefire levels. This was driven by a first-year increase (maintained over time) in forbs with short life spans; a 2-3-year delayed surge in long-lived forbs; and a consistent increase in graminoids through the 10th postfire year. Burning increased exotic plant invasion relative to prefire and unburned areas, but burned communities always were >89% native. This study informs debate in the literature regarding whether these increasingly large fires are 'ecological catastrophes.' Landscape-scale severe burning was catastrophic from a tree overstory perspective, but from an understory perspective, burning promoted rich and productive native understories, despite the entire 10-year postfire period receiving below-average precipitation.

Long-term response of a Mojave Desert winter annual plant community to a whole-ecosystem atmospheric CO2 manipulation (FACE).

Desert annuals are a critically important component of desert communities and may be particularly responsive to increasing atmospheric (CO2 ) because of their high potential growth rates and flexible phenology. During the 10-year life of the Nevada Desert FACE (free-air CO2 enrichment) Facility, we evaluated the productivity, reproductive allocation, and community structure of annuals in response to long-term elevated (CO2 ) exposure. The dominant forb and grass species exhibited accelerated phenology, increased size, and higher reproduction at elevated (CO2 ) in a wet El Niño year near the beginning of the experiment. However, a multiyear dry cycle resulted in no increases in productivity or reproductive allocation for the remainder of the experiment. At the community level, early indications of increased dominance of the invasive Bromus rubens at elevated (CO2 ) gave way to an absence of Bromus in the community during a drought cycle, with a resurgence late in the experiment in response to higher rainfall and a corresponding high density of Bromus in a final soil seed bank analysis, particularly at elevated (CO2 ). This long-term experiment resulted in two primary conclusions: (i) elevated (CO2 ) does not increase productivity of annuals in most years; and (ii) relative stimulation of invasive grasses will likely depend on future precipitation, with a wetter climate favoring invasive grasses but currently predicted greater aridity favoring native dicots.

Soil, vegetation, and seed bank of a Sonoran Desert ecosystem along an exotic plant (Pennisetum ciliare) treatment gradient.

Ecological conditions following removal of exotic plants are a key part of comprehensive environmental management strategies to combat exotic plant invasions. We examined ecological conditions following removal of the management-priority buffelgrass (Pennisetum ciliare) in Saguaro National Park of the North American Sonoran Desert. We assessed soil, vegetation, and soil seed banks on seven buffelgrass site types: five different frequencies of buffelgrass herbicide plus hand removal treatments (ranging from 5 years of annual treatment to a single year of treatment), untreated sites, and non-invaded sites, with three replicates for each of the seven site types. The 22 measured soil properties (e.g., pH) differed little among sites. Regarding vegetation, buffelgrass cover was low (≤1% median cover), or absent, across all treated sites but was high (10-70%) in untreated sites. Native vegetation cover, diversity, and composition were indistinguishable across site types. Species composition was dominated by native species (>93% relative cover) across all sites except untreated buffelgrass sites. Most (38 species, 93%) of the 41 species detected in soil seed banks were native, and native seed density did not differ significantly across sites. Results suggest that: (1) buffelgrass cover was minimal across treated sites; (2) aside from high buffelgrass cover in untreated sites, ecological conditions were largely indistinguishable across sites; (3) soil seed banks harbored ≥12 species that were frequent in the aboveground vegetation; and (4) native species dominated post-treatment vegetation composition, and removing buffelgrass did not result in replacement by other exotic species.

Thinning pine plantations to reestablish oak openings species in northwestern Ohio.

Globally the area in forest plantations is rising by 2% annually, increasing the importance of plantations for production of human goods and services and for ecological functions such as carbon storage and biodiversity conservation. Specifically in the Great Lakes states and provinces of Midwestern North America, thousands of hectares of pine plantations were established in the early and mid-1900s to revegetate abandoned agricultural fields that had replaced mixed-species forests and oak-prairie ecosystems. Plantation establishment also was intended to bolster the timber base. Management priorities have shifted, with many resource managers currently seeking to manage existing plantations for promoting mixed-species ecosystems. The purpose of this study was to assess plant succession and the reestablishment of oak savanna and prairie species after thinning 14 plantations of Pinus resinosa and strobus in northwestern Ohio, USA. Thinning reduced tree basal area by an average of 75%. Plant communities were sampled on 0.05-ha plots one and 3 years after thinning and compared to 10 unthinned control plantations. By 3 years after thinning, thinned plots contained 2-3 times more species and 14 times more plant cover than control plots. The species composition of colonizing plants was most strongly correlated with residual pine basal area and soil variables related to drainage (e.g., sand concentration, available water capacity). Although plant composition was dominated by widespread colonizers such as Erechtites hieraciifolia, the coefficient of conservatism (indicative of species of more intact, undisturbed communities) significantly increased on thinned plots from year 1 to 3. This finding, coupled with the presence of four rare, state-listed Ohio species whose eight plot occurrences all were on thinned plots, suggests that plant composition is moving towards species typifying more high-quality savanna and prairie habitats.

Disturbance and plant succession in the Mojave and Sonoran deserts of the American Southwest.

Disturbances such as fire, land clearing, and road building remove vegetation and can have major influences on public health through effects on air quality, aesthetics, recreational opportunities, natural resource availability, and economics. Plant recovery and succession following disturbance are poorly understood in arid lands relative to more temperate regions. This study quantitatively reviewed vegetation reestablishment following a variety of disturbances in the Mojave and Sonoran Deserts of southwestern North America. A total of 47 studies met inclusion criteria for the review. The time estimated by 29 individual studies for full reestablishment of total perennial plant cover was 76 years. Although long, this time was shorter than an estimated 215 years (among 31 individual studies) required for the recovery of species composition typical of undisturbed areas, assuming that recovery remains linear following the longest time since disturbance measurement made by the studies.

Assessing an exotic plant surveying program in the Mojave Desert, Clark County, Nevada, USA.

Exotic species can threaten native ecosystems and reduce services that ecosystems provide to humans. Early detection of incipient populations of exotic species is a key step in containing exotics before explosive population growth and corresponding impacts occur. We report the results of the first three years of an exotic plant early detection and treatment program conducted along more than 3,000 km of transportation corridors within an area >1.5 million ha in the Mojave Desert, USA. Incipient populations of 43 exotic plant species were mapped using global positioning and geographic information systems. Brassica tournefortii (Sahara mustard) infested the most soil types (47% of 256) surveyed in the study area, while Nicotiana glauca (tree tobacco) and others currently occupy less than 5% of soil types. Malcolmia africana (African mustard) was disproportionately detected on gypsum soils, occurring on 59% of gypsum soil types compared to 27% of all surveyed soils. Gypsum soils constitute unique rare plant habitat in this region, and by conventional wisdom were not previously considered prone to invasion. While this program has provided an initial assessment of the landscape-scale distribution of exotic species along transportation corridors, evaluations of both the survey methods and the effectiveness of treating incipient populations are needed. An exotic plant information system most useful to resource mangers will likely include integrating planning oriented coarse-scale surveys, more detailed monitoring of targeted locations, and research on species life histories, community invasibility, and treatment effectiveness.

A systematic review of wild burro grazing effects on Mojave Desert vegetation, USA.

Wild burros (Equus asinus), protected by the 1971 Wild Free-Roaming Horse and Burro Act on some federal lands but exotic animals many ecologists and resource mangers view as damaging to native ecosystems, represent one of the most contentious environmental management problems in American Southwest arid lands. This review synthesizes the scattered literature about burro effects on plant communities of the Mojave Desert, a center of burro management contentions. I classified 24 documents meeting selection criteria for this review into five categories of research: (i) diet analyses directly determining which plant species burros consume, (ii) utilization studies of individual species, (iii) control-impact comparisons, (iv) exclosure studies, and (v) forage analyses examining chemical characteristics of forage plants. Ten diet studies recorded 175 total species that burros consumed. However, these studies and two exclosure studies suggested that burros preferentially eat graminoid and forb groups over shrubs. One study in Death Valley National Park, for example, found that Achnatherum hymenoides (Indian ricegrass) was 11 times more abundant in burro diets than expected based on its availability. Utilization studies revealed that burros also exhibit preferences within the shrub group. Eighty-three percent of reviewed documents were produced in a 12-year period, from 1972 to 1983, with the most recent document produced in 1988. Because burros remain abundant on many federal lands and grazing may interact with other management concerns (e.g., desert wildfires fueled by exotic grasses), rejuvenating grazing research to better understand both past and present burro effects could help guide revegetation and grazing management scenarios.