Inherent Rarity in Community Restoration

Abstract: We explore the implications of an often overlooked fact in community restoration: most species, in real or synthetic communities, are infrequent or rare, a phenomenon we call "inherent rarity." Whether from long-term interactions of many factors affecting birth, death, and establishment within natural communities, or from short-term interactions of recently created communities, species abundance distributions are roughly lognormal or even more attenuated. The greater the predisturbance species diversity of remnants or the planting diversity of restorations, and the smaller the area of a habitat patch, the more likely a large proportion of species will exist in populations so small that they are acutely or chronically vulnerable to local extinction. We suggest that habitat fragmentation will inevitably favor plants and animals that are highly mobile, early successional species, including many exotic weeds among plants, or species that are historically organized as metapopulations that happen to be common enough to function as such. We further explore rationales for countering the effects of inherent rarity, including connectivity from buffers, corridors, and stepping stones, and dominance suppression from seasonally appropriate mowing, grazing, or fire.     

Should heterogeneity be the basis for conservation? Grassland bird response to fire and grazing.

In tallgrass prairie, disturbances such as grazing and fire can generate patchiness across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds evolved within the context of this shifting mosaic, with some species restricted to one or two patch types created under spatially and temporally distinct disturbance regimes. Thus, management-driven reductions in heterogeneity may be partly responsible for declines in numbers of grassland birds. We experimentally altered spatial heterogeneity of vegetation structure within a tallgrass prairie by varying the spatial and temporal extent of fire and by allowing grazing animals to move freely among burned and unburned patches (patch treatment). We contrasted this disturbance regime with traditional agricultural management of the region that promotes homogeneity (traditional treatment). We monitored grassland bird abundance during the breeding seasons of 2001-2003 to determine the influence of altered spatial heterogeneity on the grassland bird community. Focal disturbances of patch burning and grazing that shifted through the landscape over several years resulted in a more heterogeneous pattern of vegetation than uniform application of fire and grazing. Greater spatial heterogeneity in vegetation provided greater variability in the grassland bird community. Some bird species occurred in greatest abundance within focally disturbed patches, while others occurred in relatively undisturbed patches in our patch treatment. Henslow's Sparrow, a declining species, occurred only within the patch treatment. Upland Sandpiper and some other species were more abundant on recently disturbed patches within the same treatment. The patch burn treatment created the entire gradient of vegetation structure required to maintain a suite of grassland bird species that differ in habitat preferences. Our study demonstrated that increasing spatial and temporal heterogeneity of disturbance in grasslands increases variability in vegetation structure that results in greater variability at higher trophic levels. Thus, management that creates a shifting mosaic using spatially and temporally discrete disturbances in grasslands can be a useful tool in conservation. In the case of North American tallgrass prairie, discrete fires that capitalize on preferential grazing behavior of large ungulates promote a shifting mosaic of habitat types that maintain biodiversity and agricultural productivity.     

Indirect Effects of Biological Control on Plant Diversity Vary across Sites in Montana Grasslands

Abstract:  Biological control with specialist, nonindigenous, herbivorous insects is an important option for controlling invasive exotic plants in wildlands and nature reserves. It is assumed that biological control agents will reduce the dominance of the target weed, thereby increasing the native diversity of the associated plant community. However, this hypothesis has rarely been tested. We introduced Aphthona nigriscutis into grassland sites infested with the invasive exotic species Euphorbia esula L. on a nature reserve in Montana (U.S.A.). Two sites with better soil had been treated previously with herbicide, whereas two other sites had not. We measured the density and biomass of Euphorbia vegetative and flowering stems and number of native and exotic shrubs, grass-like plants, and forbs in 48 microplots in Aphthona release and control macroplots at each site. After 5 years, Aphthona release was associated with a 33–39% decline in Euphorbia aboveground biomass compared with controls at all sites. Other effects of the biocontrol depended on the site. Biocontrol slowed the recovery of species diversity at the sites previously treated with herbicide but slowed the loss of diversity at sites without a history of herbicide. Biocontrol introduction was not associated with a disproportionate increase in nontarget exotic species. Release of Aphthona caused a decline in the biomass of flowering stems relative to controls at good-soil, previous-herbicide sites but was associated with a relative increase in flower stem mass at poor-soil, no-herbicide sites. Our results suggest that biocontrol reductions in weed dominance will not always be associated with increased species diversity. More emphasis should be placed on conserving desirable communities and less on simple weed control. Monitoring of community-level effects should accompany biocontrol introductions on nature reserves.     

The consequences of genetic diversity in competitive communities

Several lines of evidence suggest that the species diversity and composition of communities should depend on genetic diversity within component species, but there has been very little effort to directly assess this possibility. Here I use models of competition among genotypes and species to demonstrate a strong positive effect of the number of genotypes per species on species diversity across a range of conditions. Genetic diversity allows species to respond to selection imposed by competition, resulting in both functional convergence and divergence among species depending on their initial niche positions. This ability to respond to selection promotes species coexistence and contributes to a reduction in variation in species composition among communities. These models suggest that whenever individual fitness depends on the degree of functional similarity between a focal individual and its competitors, genetic diversity should promote species coexistence; this prediction is consistent with the few relevant empirical data collected to date. The results point to the importance of considering the genetic origin and diversity of material used in ecological experiments and in restoration efforts, in addition to highlighting potentially important community consequences of the loss of genetic diversity in natural populations.     

Incorporating differences between genetic diversity of trees and herbaceous plants in conservation strategies

Reviews that summarize the genetic diversity of plant species in relation to their life history and ecological traits show that forest trees have more genetic diversity at population and species levels than annuals or herbaceous perennials. In addition, among-population genetic differentiation is significantly lower in trees than in most herbaceous perennials and annuals. Possible reasons for these differences between trees and herbaceous perennials and annuals have not been discussed critically. Several traits, such as high rates of outcrossing, long-distance pollen and seed dispersal, large effective population sizes (N_e), arborescent stature, low population density, longevity, overlapping generations, and occurrence in late successional communities, may make trees less sensitive to genetic bottlenecks and more resistant to habitat fragmentation or climate change. We recommend that guidelines for genetic conservation strategies be designed differently for tree species versus other types of plant species. Because most tree species fit an LH scenario (low [L] genetic differentiation and high [H] genetic diversity), tree seeds could be sourced from a few populations distributed across the species’ range. For the in situ conservation of trees, translocation is a viable option to increase N_e. In contrast, rare herbaceous understory species are frequently HL (high differentiation and low diversity) species. Under the HL scenario, seeds should be taken from many populations with high genetic diversity. In situ conservation efforts for herbaceous plants should focus on protecting habitats because the typically small populations of these species are vulnerable to the loss of genetic diversity. The robust allozyme genetic diversity databases could be used to develop conservation strategies for species lacking genetic information. As a case study of reforestation with several tree species in denuded areas on the Korean Peninsula, we recommend the selection of local genotypes as suitable sources to prevent adverse effects and to insure the successful restoration in the long term.     

Predicting Plant Extinction Based on Species-Area Curves in Prairie Fragments with High Beta Richness

Abstract:  Species-area relationships and island biogeography theory are commonly used to predict how species richness will decline with fragmentation. There are a variety of largely untested assumptions in these approaches, including the assumptions that populations are distributed uniformly before fragmentation, and that local extinctions are due to effects of small population sizes. If populations are not distributed uniformly, then populations can be abundant locally but rare globally. This would cause extinction rates to be smaller than predicted. We tested extinction theory by developing estimates of the number of plant species that should be present in small tallgrass prairie fragments and then testing the uniformity assumption by partitioning species richness into α (within site) and β (among site) components in Iowa prairies. Many more native prairie plant species were present in surveys of prairie fragments (491) than was predicted based on theory (27–207). A large proportion (75%) of the total species richness was β richness. We suggest that the high proportion of β richness was responsible for the shallow species-area slopes and the lower than expected number of species losses and that a better understanding of what determines β diversity will improve predictions of fragmentation effects on richness of plants. We also suggest that plants in prairie remnants may be best conserved by protecting different prairie types rather than by protecting a few large areas containing a single prairie type.     

Differing Effects of Cattle Grazing on Native and Alien Plants

Abstract:   Habitat managers use cattle grazing to reduce alien plant cover and promote native species in California grasslands and elsewhere in the western United States. We tested the effectiveness of grazing as a restoration method by examining the effects of herbivory on native and alien plants. At Carrizo Plain National Monument, California, we surveyed native and alien species cover in adjacent grazed and ungrazed areas. We also established experimental plots in which plants were clipped or mulch (dead biomass) was removed. In addition, we clipped plants grown in pots and plants in the field that grew with and without competitors. Native species were negatively affected by clipping in 1999, 2000, and 2001, whereas alien species were unaffected. In the experimental field plots, the European annual forb Erodium cicutarium compensated in growth and reproduction following simulated herbivory. In contrast, growth and reproduction of the native perennial bunchgrass Poa secunda were reduced 1 year after clipping. In pots, E. cicutarium overcompensated and grasses undercompensated. In the field, European grasses were unaffected by the removal of competitors. It is unclear by what mechanism E. cicutarium was able to compensate, but the ability may be related to its basal rosette growth form and indeterminately growing inflorescences. The native California grassland community assembled in the absence of grazing herds, whereas invasive European species have been exposed to grazing for centuries. It may be that these invaders have adaptations that better enable them to recover from grazing. In the grassland we studied, the strategy of livestock grazing for restoration is counterproductive. It harms native species and promotes alien plant growth.     

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.     

Evaluation of Central North American Prairie Management Based on Species Diversity, Life Form, and Individual Species Metrics

Abstract:  Reintroduction of fire and grazing, alone or in combination, has increasingly been recognized as central to the restoration of North American mixed-grass and tallgrass prairies. Although ecological studies of these systems are abundant, they have generally been observational, or if experimental, have focused on plant species diversity. Species diversity measures alone are not sufficient to inform management, which often has goals associated with life-form groups and individual species. We examined the effects of prescribed fire, light cattle grazing, and a combination of fire and grazing on three vegetation components: species diversity, groups of species categorized by life-form, and individual species. We evaluated how successful these three treatments were in achieving specific management goals for prairies in the Iowa Loess Hills (U.S.A.). The grazing treatment promoted the greatest overall species richness, whereas grazing and burning and grazing treatments resulted in the lowest cover by woody species. Burning alone best achieved the management goals of increasing the cover and diversity of native species and reducing exotic forb and (predominantly exotic) cool-season grass cover. Species-specific responses to treatments appeared idiosyncratic (i.e., within each treatment there existed a set of species attaining their highest frequency) and nearly half of uncommon species were present in only one treatment. Because all management goals were not achieved by any one treatment, we conclude that management in this region may need refining. We suggest that a mosaic of burning and grazing (alone and in combination) may provide the greatest landscape-level species richness; however, this strategy would also likely promote the persistence of exotic species. Our results support the need to consider multiple measures, including species-specific responses, when planning and evaluating management .     

Plague bacterium as a transformer species in prairie dogs and the grasslands of western North America

Invasive transformer species change the character, condition, form, or nature of ecosystems and deserve considerable attention from conservation scientists. We applied the transformer species concept to the plague bacterium Yersinia pestis in western North America, where the pathogen was introduced around 1900. Y. pestis transforms grassland ecosystems by severely depleting the abundance of prairie dogs (Cynomys spp.) and thereby causing declines in native species abundance and diversity, including threatened and endangered species; altering food web connections; altering the import and export of nutrients; causing a loss of ecosystem resilience to encroaching invasive plants; and modifying prairie dog burrows. Y. pestis poses an important challenge to conservation biologists because it causes trophic‐level perturbations that affect the stability of ecosystems. Unfortunately, understanding of the effects of Y. pestis on ecosystems is rudimentary, highlighting an acute need for continued research.     

Scale-dependent interaction of fire and grazing on community heterogeneity in tallgrass prairie

Natural disturbances affect spatial and temporal heterogeneity in plant communities, but effects vary depending on type of disturbance and scale of analysis. In this study, we examined the effects of fire frequency (1-, 4-, and 20-yr intervals) and grazing by bison on spatial and temporal heterogeneity in species composition in tallgrass prairie plant communities. Compositional heterogeneity was estimated at 10-, 50-, and 200-m2 scales. For each measurement scale, we used the average Euclidean Distance (ED) between samples within a year (2000) to measure spatial heterogeneity and between all time steps (1993-2000) for each sample to measure temporal heterogeneity. The main effects of fire and grazing were scale independent. Spatial and temporal heterogeneity were lowest on annually burned sites and highest on infrequently burned (20-yr) sites at all scales. Grazing reduced spatial heterogeneity and increased temporal heterogeneity at all scales. The rate of community change over time decreased as fire frequency increased at all scales, whereas grazing had no effect on rate of community change over time at any spatial scale. The interactive effects of fire and grazing on spatial and temporal heterogeneity differed with scale. At the 10-m2 scale, grazing increased spatial heterogeneity in annually burned grassland but decreased heterogeneity in less frequently burned areas. At the 50-m2 scale, grazing decreased spatial heterogeneity on 4-yr burns but had no effect at other fire frequencies. At the 10-m scale, grazing increased temporal heterogeneity only on 1- and 20-yr burn sites. Our results show that the individual effects of fire and grazing on spatial and temporal heterogeneity in mesic prairie are scale independent, but the interactive effects of these disturbances on community heterogeneity change with scale of measurement. These patterns reflect the homogenizing impact of fire at all spatial scales, and the different frequency, intensity, and scale of patch grazing by bison in frequently burned vs. infrequently burned areas.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Patch Size and Connectivity Thresholds for Butterfly Habitat Restoration

Abstract:  Recovery of endangered species in highly fragmented habitats often requires habitat restoration. Selection of restoration sites typically involves too many options and too much uncertainty to reach a decision based on existing reserve design methods. The Fender's blue butterfly (  Icaricia icarioides fenderi ) survives in small, isolated patches of remnant prairie in Oregon's Willamette Valley—a habitat for which <0.5% of the original remains. Recovery of this species will require considerable habitat restoration. We investigated the potential of biologically based rules of thumb and more complex models to serve as tools in making land acquisitions. Based on Fender's blue dispersal behavior and demography, we have estimated that restored patches should be <1 km from existing habitat and at least 2 ha. We compared these rules to the results of two modeling approaches: an incidence function model and a spatially explicit simulation of demography and dispersal behavior. Not surprisingly, the simple rules and complex models all conclude that large (>2 ha) connected (<1 km) patches have the highest restoration value. The dispersal model, however, suggests that small, connected patches have more restoration value than large, isolated patches, whereas the incidence function model suggests that size and connectivity are equally important. These differences stem from model assumptions. We used incidence functions to predict long-term, stochastic, steady-state conditions and dispersal simulations to predict short-term (25-year) colonization dynamics. To apply our results in the context of selecting restoration sites on the ground, we recommend selecting nearby sites when short-term colonization dynamics are expected to be an important aspect of a species' biology.     

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.     

Impact of Grazing Intensity during Drought in an Arizona Grassland

Abstract:  The ecological benefits of changing cattle grazing practices in the western United States remain controversial, due in part to a lack of experimentation. In 1997 we initiated an experimental study of two rangeland alternatives, cattle removal and high-impact grazing, and compared grassland community responses with those with more conventional, moderate grazing practices. The study was conducted in a high-elevation, semiarid grassland near Flagstaff, Arizona (U.S.A.). We conducted annual plant surveys of modified Whittaker plots for 8 years and examined plant composition shifts among treatments and years. High-impact grazing had strong directional effects that led to a decline in perennial forb cover and an increase in annual plants, particularly the exotic cheatgrass ( Bromus tectorum L.). A twofold increase in plant cover by exotic species followed a severe drought in the sixth year of the study, and this increase was greatest in the high-impact grazing plots, where native cover declined by one-half. Cattle removal resulted in little increase in native plant cover and reduced plant species richness relative to the moderate grazing control. Our results suggest that some intermediate level of cattle grazing may maintain greater levels of native plant diversity than the alternatives of cattle removal or high-density, short-duration grazing practices. Furthermore, episodic drought interacts with cattle grazing, leading to infrequent, but biologically important shifts in plant communities. Our results demonstrate the importance of climatic variation in determining ecological effects of grazing practices, and we recommend improving conservation efforts in arid rangelands by developing management plans that anticipate this variation.     

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.     

Direct and indirect effects of CO2, nitrogen, and community diversity on plant-enemy interactions

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

Biotic habitat complexity controls species diversity and nutrient effects on net biomass production

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.     

Environmental variability and allocation trade-offs maintain species diversity in a process-based model of succulent plant communities

Ecological theory suggests that environmental variability can promote coexistence, provided that species occupy differential niches. In this study, we focus on two questions: (1) Do allocation trade-offs provide a sufficient basis for niche differentiation in succulent plant communities? (2) What is the relative importance of different forms of environmental variability on species diversity and community composition? We approach these questions with a generic, individual-based simulation model. In our model, plants compete for water in a spatially explicit environment. Species differ in their size at maturity and in the allocation of carbon to roots, leaves and storage tissue. The model was fully specified with independent literature data. Model output was compared to characteristics of a species-rich community in the semi-arid Richtersveld (South Africa). The model reproduced the coexistence of plants with different sizes at maturity, the dominance of succulent shrubs, and the level of vegetation cover. We analyzed the effects of three forms of environmental variability: (a) temporal fluctuations in precipitation (rain and fog), (b) spatial heterogeneity of water supply due to run-on and run-off processes and (c) ‘rock pockets’ that limit root competition in space. The three types of variability had differential effects on diversity: diversity exhibited a strong hump-shaped response to temporal variation. Spatial variability increased diversity, with the strongest increase occurring at intermediate levels of temporal variability. Finally, rock pockets had the weakest effect, but contributed to diversity by providing refuges for small species, particularly at low temporal variability. The model thus shows that spatio-temporal variation of resource supply can maintain diversity over long time scales even in small systems, as is the case in the Richtersveld succulent communities. Trade-offs in allocation provide the basis for necessary niche differentiation. By describing resource competition between individual plants, our model provides a mechanistic basis for the link from species traits to community composition at given environmental conditions. It thereby contributes to an understanding of the forces shaping plant communities. Such an understanding is critical to reduce the threats environmental change poses to biodiversity and ecosystem services.