Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

Elephants as agents of habitat creation for small vertebrates at the patch scale

Ecologists increasingly recognize the ability of certain species to influence ecological processes by engineering the physical environment, but efforts to develop a predictive understanding of this phenomenon are in their early stages. While many believe that the landscape-scale effects of ecosystem engineers will be to increase habitat diversity and therefore the abundance and richness of other species, few generalities exist about the effects of engineering at the scale of the engineered patch. According to one hypothesis, activities that increase structural habitat complexity within engineered patches will have positive effects on the abundance or diversity of other organisms. Here I show that, by damaging trees and increasing their structural complexity, browsing elephants create refuges used by a common arboreal lizard. Observational surveys and a lizard transplant experiment revealed that lizards preferentially occupy trees with real or simulated elephant damage. A second experiment showed that lizards vacate trees when elephant-engineered refuges are removed. Furthermore, local lizard densities increased with (and may be constrained by) local densities of elephant-damaged trees. This facilitative effect of elephants upon lizards via patch-scale habitat modification runs contrary to previously documented negative effects of the entire ungulate guild on lizards at the landscape scale, suggesting that net indirect effects of large herbivores comprise opposing trophic and engineering interactions operating at different spatial scales. Such powerful megaherbivore-initiated interactions suggest that anthropogenic changes in large-mammal densities will have important cascading consequences for ecological communities.     

Effects of species diversity on community biomass production change over the course of succession

Over the past decade an increasing number of studies have experimentally manipulated the number of species in a community and examined how this alters the aggregate production of species biomass. Many of these studies have shown that the effects of richness on biomass change through time, but we have limited understanding of the mechanisms that produce these dynamic trends. Here we report the results of an experiment in which we manipulated the richness of freshwater algae in laboratory microcosms. We used two experimental designs (additive and substitutive) that make different assumptions about how patches are initially colonized, and then tracked the development of community biomass from the point of initial colonization through a period of 6-12 generations of the focal species. We found that the effect of initial species richness on biomass production qualitatively shifted twice over the course of the experiment. The first shift occurred as species transitioned from density-independent to dependent phases of population growth. At this time, intraspecific competition caused monocultures to approach their respective carrying capacities more slowly than polycultures. As a consequence, species tended to over-yield for a brief time, generating a positive, but transient effect of diversity on community biomass. The second shift occurred as communities approached carrying capacity. At this time, strong interspecific interactions caused biomass to be dominated by the competitively superior species in polycultures. As this species had the lowest carrying capacity, a negative effect of diversity on biomass resulted in late succession. Although these two shifts produced dynamics that appeared complex, we show that the patterns can be fit to a simple Lotka-Volterra model of competition. Our results suggest that the effects of algal diversity on primary production change in a predictable sequence through successional time.     

Experimental evidence for mediation of competition by habitat succession

Habitat succession is thought to influence the importance of competition in assemblages. Competitive interactions are considered of critical importance in structuring ant assemblages, but field experiments show varied effects. I tested how succession in managed boreal forests affects the outcome of competition from dominant red wood ants, Formica aquilonia, through a removal experiment in replicated stands of three different ages (0-4, 30-40, and 80-100 years old). F. aquilonia abundance was reduced by 87%, and procedural controls showed no nontarget effects. The succession gradient revealed the full range of possible responses from ant species: decreases in 1-4-year-old stands, increases in 30-40-year-old stands, and no effects in 80-100-year-old stands, where diversity was lowest. Habitat succession thus regulates competitive interactions in this system. I propose a model for this system, where competitive effects depend on time since disturbance. In this case, soon after disturbance the dominant species facilitates increases in the abundance of other species. At intermediate times, competition reduces the abundance of some species. Finally, in long-undisturbed habitats, competitors may fail to evolve, particularly in high-stress environments. Interactions between competition and habitat succession may explain why structuring effects of ecologically dominant species appear inconsistent.     

Do natural disturbances or the forestry practices that follow them convert forests to early-successional communities?

Stand-replacing natural disturbances in mature forests are traditionally seen as events that cause forests to revert to early stages of succession and maintain species diversity. In some cases, however, such transitions could be an artifact of salvage logging and may increase biotic homogenization. We present initial (two-year) results of a study of the effects of tornado damage and the combined effects of tornado damage and salvage logging on environmental conditions and ground cover plant communities in mixed oak-pine forests in north central Mississippi. Plots were established in salvage-logged areas, adjacent to plots established before the storm in unlogged areas, spanning a gradient of storm damage intensity. Vegetation change directly attributable to tornado damage was driven primarily by a reduction in canopy cover but was not consistent with a transition to an early stage of succession. Although we observed post-storm increases of several disturbance indicators (ruderals), we also observed significant increases in the abundance of a few species indicative of upland forests. Increases in flowering were just as likely to occur in species indicative of forests as in species indicative of open woodlands. Few species declined as a result of the tornado, resulting in a net increase in species richness. Ruderals were very abundant in salvage-logged areas, which contained significantly higher amounts of bare ground and greater variance in soil penetrability than did damaged areas that were not logged. In contrast to unlogged areas severely damaged by the tornado, most upland forest indicators were not abundant in logged areas. Several of the forest and open-woodland indicators that showed increased flowering in damaged areas were absent or sparse in logged areas. Species richness was lower in salvage-logged areas than in adjacent damaged areas but similar to that in undamaged areas. These results suggest that salvage logging prevented positive responses of several forest and open-woodland species to tornado damage. Anthropogenic disturbances such as salvage logging appear to differ fundamentally from stand-level canopy-reducing disturbances in their effects on ground cover vegetation in the forests studied here and are perhaps more appropriately viewed as contributing to biotic homogenization than as events that maintain diversity.     

Biodiversity Conservation, Ecosystem Functioning, and Economic Incentives under Cocoa Agroforestry Intensification

Abstract:  World chocolate demand is expected to more than double by 2050. Decisions about how to meet this challenge will have profound effects on tropical rainforests and wild species in cocoa-producing countries. Cocoa, "the chocolate tree," is traditionally produced under a diverse and dense canopy of shade trees that provide habitat for a high diversity of organisms. The current trend to reduce or eliminate shade cover raises concerns about the potential loss of biodiversity. Nevertheless, few studies have assessed the ecological consequences and economic trade-offs under different management options in cocoa plantations. Here we describe the relationships between ant ecology (species richness, community composition, and abundance) and vegetation structure, ecosystem functions, and economic profitability under different land-use management systems in 17 traditional cocoa forest gardens in southern Cameroon. We calculated an index of profitability, based on the net annual income per hectare. We found significant differences associated with the different land-use management systems for species richness and abundance of ants and species richness and density of trees. Ant species richness was significantly higher in floristically and structurally diverse, low-intensity, old cocoa systems than in intensive young systems. Ant species richness was significantly related to tree species richness and density. We found no clear relationship between profitability and biodiversity. Nevertheless, we suggest that improving the income and livelihood of smallholder cocoa farmers will require economic incentives to discourage further intensification and ecologically detrimental loss of shade cover. Certification programs for shade-grown cocoa may provide socioeconomic incentives to slow intensification.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Facilitation drives local abundance and regional distribution of a rare plant in a harsh environment

The importance of facilitation to local community dynamics is becoming increasingly recognized. However, the predictability of positive interactions in stressful environments, the balance of competition and facilitation along environmental gradients, and the scaling of local positive interactions to regional distributions are aspects of facilitation that remain unresolved. I explored these questions in a habitat specialist, Delphinium uliginosum, and a moss, Didymodon tophaceus, both found in small serpentine wetlands. I tested three hypotheses: (1) moss facilitates germination, growth, and/or fecundity of D. uliginosum; (2) facilitation is stronger at the harsher ends of gradients in soil moisture, toxicity, and/or biomass; and (3) facilitation is reflected in positive associations at the levels of local abundance and regional occurrence. Although considerable competitive interactions occurred in later life stages, moss strongly facilitated D. uliginosum seedling emergence. There was no evidence that this facilitative effect weakened, or switched to competition, in benign environments. D. uliginosum was more locally abundant and more frequently present, across a large portion of its range, with than without moss, indicating a net facilitative effect in the face of competitive influences. Facilitated recruitment, possibly by seed retention, was found to be an important control on abundance and distribution in this rare species.     

Contrasting effects of habitat loss and fragmentation on coral-associated reef fishes

Disturbance can result in the fragmentation and/or loss of suitable habitat, both of which can have important consequences for survival, species interactions, and resulting patterns of local diversity. However, effects of habitat loss and fragmentation are typically confounded during disturbance events, and previous attempts to determine their relative significance have proved ineffective. Here we experimentally manipulated live coral habitats to examine the potential independent and interactive effects of habitat loss and fragmentation on survival, abundance, and species richness of recruitment-stage, coral-associated reef fishes. Loss of 75% of live coral from experimental reefs resulted in low survival of a coral-associated damselfish and low abundance and richness of other recruits 16 weeks after habitat manipulations. In contrast, fragmentation had positive effects on damselfish survival and resulted in greater abundance and species richness of other recruits. We hypothesize that spacing of habitat through fragmentation weakens competition within and among species. Comparison of effect sizes over the course of the study period revealed that, in the first six weeks following habitat manipulations, the positive effects of fragmentation were at least four times stronger than the effects of habitat loss. This initial positive effect of fragmentation attenuated considerably after 16 weeks, whereas the negative effects of habitat loss increased in strength over time. There was little indication that the amount of habitat influenced the magnitude of the habitat fragmentation effect. Numerous studies have reported dramatic declines in coral reef fish abundance and diversity in response to disturbances that cause the loss and fragmentation of coral habitats. Our results suggest that these declines occur as a result of habitat loss, not habitat fragmentation. Positive fragmentation effects may actually buffer against the negative effects of habitat loss and contribute to the resistance of reef fish populations to declines in coral cover.     

Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

Associations of Forest Cover,Fragment Area,and Connectivity with Neotropical Understory Bird Species Richness and Abundance

Theoretical and empirical studies demonstrate that the total amount of forest and the size and connectivity of fragments have nonlinear effects on species survival. We tested how habitat amount and configuration affect understory bird species richness and abundance. We used mist nets (almost 34,000 net hours) to sample birds in 53 Atlantic Forest fragments in southeastern Brazil. Fragments were distributed among 3 10,800‐ha landscapes. The remaining forest in these landscapes was below (10% forest cover), similar to (30%), and above (50%) the theoretical fragmentation threshold (approximately 30%) below which the effects of fragmentation should be intensified. Species‐richness estimates were significantly higher (F= 3715, p = 0.00) where 50% of the forest remained, which suggests a species occurrence threshold of 30–50% forest, which is higher than usually occurs (<30%). Relations between forest cover and species richness differed depending on species sensitivity to forest conversion and fragmentation. For less sensitive species, species richness decreased as forest cover increased, whereas for highly sensitive species the opposite occurred. For sensitive species, species richness and the amount of forest cover were positively related, particularly when forest cover was 30–50%. Fragment size and connectivity were related to species richness and abundance in all landscapes, not just below the 30% threshold. Where 10% of the forest remained, fragment size was more related to species richness and abundance than connectivity. However, the relation between connectivity and species richness and abundance was stronger where 30% of the landscape was forested. Where 50% of the landscape was forested, fragment size and connectivity were both related to species richness and abundance. Our results demonstrated a rapid loss of species at relatively high levels of forest cover (30–50%). Highly sensitive species were 3‐4 times more common above the 30–50% threshold than below it; however, our results do not support a unique fragmentation threshold. Asociaciones de la Cobertura Forestal, Superficie del Fragmento y Conectividad con la Riqueza y Abundancia de Aves Neotropicales de Sotobosque     

Contribution of Urban Expansion and a Changing Climate to Decline of a Butterfly Fauna

Butterfly populations are naturally patchy and undergo extinctions and recolonizations. Analyses based on more than 2 decades of data on California's Central Valley butterfly fauna show a net loss in species richness through time. We analyzed 22 years of phenological and faunistic data for butterflies to investigate patterns of species richness over time. We then used 18–22 years of data on changes in regional land use and 37 years of seasonal climate data to develop an explanatory model. The model related the effects of changes in land‐use patterns, from working landscapes (farm and ranchland) to urban and suburban landscapes, and of a changing climate on butterfly species richness. Additionally, we investigated local trends in land use and climate. A decline in the area of farmland and ranchland, an increase in minimum temperatures during the summer and maximum temperatures in the fall negatively affected net species richness, whereas increased minimum temperatures in the spring and greater precipitation in the previous summer positively affected species richness. According to the model, there was a threshold between 30% and 40% working‐landscape area below which further loss of working‐landscape area had a proportionally greater effect on butterfly richness. Some of the isolated effects of a warming climate acted in opposition to affect butterfly richness. Three of the 4 climate variables that most affected richness showed systematic trends (spring and summer mean minimum and fall mean maximum temperatures). Higher spring minimum temperatures were associated with greater species richness, whereas higher summer temperatures in the previous year and lower rainfall were linked to lower richness. Patterns of land use contributed to declines in species richness (although the pattern was not linear), but the net effect of a changing climate on butterfly richness was more difficult to discern. Contribución de la Expansión Urbana y un Clima Cambiante a la Declinación de la Fauna de Mariposas     

Negative effects of nitrogen deposition on Swiss butterflies

Nitrogen (N) deposition from agriculture and combustion of fossil fuels is a major threat to plant diversity, but its effects on organisms at higher trophic levels are unclear. We investigated how N deposition may affect species richness and abundance (number of individuals per species) in butterflies. We reviewed the peer-reviewed literature on variables used to explain spatial variation in butterfly species richness and found that vegetation variables appeared to be as important as climate and habitat variables in explaining butterfly species richness. It thus seemed likely that increased N deposition could indirectly affect butterfly communities via its influence on plant communities. To test this prediction, we analyzed data from the Swiss biodiversity monitoring program for vascular plants and butterflies in 383 study sites of 1 km² that are evenly distributed throughout Switzerland. The area has a modeled N deposition gradient of 2–44 kg N ha⁻¹ year⁻¹. We used traditional linear models and structural equation models to infer the drivers of the spatial variation in butterfly species richness across Switzerland. High N deposition was consistently linked to low butterfly diversity, suggesting a net loss of butterfly diversity through increased N deposition. We hypothesize that at low elevations, N deposition may contribute to a reduction in butterfly species richness via microclimatic cooling due to increased plant biomass. At higher elevations, negative effects of N deposition on butterfly species richness may also be mediated by reduced plant species richness. In most butterfly species, abundance was negatively related to N deposition, but the strongest negative effects were found for species of conservation concern. We conclude that in addition to factors such as intensified agriculture, habitat fragmentation, and climate change, N deposition is likely to play a key role in negatively affecting butterfly diversity and abundance.     

Community ecology of invasions: direct and indirect effects of multiple invasive species on aquatic communities

With many ecosystems now supporting multiple nonnative species from different trophic levels, it can be challenging to disentangle the net effects of invaders within a community context. Here, we combined wetland surveys with a mesocosm experiment to examine the individual and combined effects of nonnative fish predators and nonnative bullfrogs on aquatic communities. Among 139 wetlands, nonnative fish (bass, sunfish, and mosquitofish) negatively influenced the probability of occupancy of Pacific treefrogs (Pseudacris regilla), but neither invader correlated strongly with occupancy by California newts (Taricha torosa), western toads (Anaxyrus boreas), or California red-legged frogs (Rana draytonii). In mesocosms, mosquitofish dramatically reduced the abundance of zooplankton and palatable amphibian larvae (P. regilla and T. torosa), leading to increases in nutrient concentrations and phytoplankton (through loss of zooplankton), and rapid growth of unpalatable toad larvae (through competitive release). Bullfrog larvae reduced the growth of native anurans but had no effect on survival. Despite strong effects on natives, invaders did not negatively influence one another, and their combined effects were additive. Our results highlight how the net effects of multiple nonnative species depend on the trophic level of each invader, the form and magnitude of invader interactions, and the traits of native community members.     

Landscape connectivity strengthens local-regional richness relationships in successional plant communities

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.     

Influence of a dominant consumer species reverses at increased diversity

Theory and experiments indicate that changes in consumer diversity affect benthic community structure and ecosystem functioning. Although the effects of consumer diversity have been tested in the laboratory and the field, little is known about effects of consumer diversity in the subtidal zone, one of the largest marine habitats. We investigated the grazing effects of sea urchins on algal abundance and benthic community structure in a natural subtidal habitat of the Galápagos Islands. Three species of urchins (Eucidaris, Lytechinus, and Tripneustes) were manipulated in inclusion cages following a replacement design with three levels of species richness (one, two, and three species) with all possible two-species urchin combinations. Identity was the main factor accounting for changes in the percentage of substrate grazed and benthic community structure. Two out of the three two-species assemblages grazed more than expected, suggesting a richness effect, but analyses revealed that this increased grazing was due to a sampling effect of the largest and commercially valued urchin species, Tripneustes. Benthic community structure in treatments with Eucidaris, Lytechinus, and Tripneustes alone was significantly different at the end of the experiment, suggesting that resource use differentiation occurred. Communities in Tripneustes enclosures were characterized by abundant crustose coralline algae and grazed substrate, while those without it contained abundant green foliose algae (Ulva sp.). An unexpected emergent property of the system was that the most species-rich urchin assemblage underyielded, grazing less than any other assemblage with Tripneustes, effectively reversing its dominant influence observed in the two-species treatments. While further experiments are needed to discern the mechanisms of underyielding, it may be related to changing interspecific interactions as richness increases from two to three species or to density-dependent Tripneustes grazing. This study highlights the general importance of evaluating consumer richness effects across the entire range of species richness considered, as the performance of the most species-rich consumer assemblage could not be predicted by manipulations of intermediate levels of consumer species richness.     

Land‐Cover Change and Avian Diversity in the Conterminous United States

Abstract: Changes in land use and land cover have affected and will continue to affect biological diversity worldwide. Yet, understanding the spatially extensive effects of land‐cover change has been challenging because data that are consistent over space and time are lacking. We used the U.S. National Land Cover Dataset Land Cover Change Retrofit Product and North American Breeding Bird Survey data to examine land‐cover change and its associations with diversity of birds with principally terrestrial life cycles (landbirds) in the conterminous United States. We used mixed‐effects models and model selection to rank associations by ecoregion. Land cover in 3.22% of the area considered in our analyses changed from 1992 to 2001, and changes in species richness and abundance of birds were strongly associated with land‐cover changes. Changes in species richness and abundance were primarily associated with changes in nondominant types of land cover, yet in many ecoregions different types of land cover were associated with species richness than were associated with abundance. Conversion of natural land cover to anthropogenic land cover was more strongly associated with changes in bird species richness and abundance than persistence of natural land cover in nearly all ecoregions and different covariates were most strongly associated with species richness than with abundance in 11 of 17 ecoregions. Loss of grassland and shrubland affected bird species richness and abundance in forested ecoregions. Loss of wetland was associated with bird abundance in forested ecoregions. Our findings highlight the value of understanding changes in nondominant land cover types and their association with bird diversity in the United States.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

Forest age influences oak insect herbivore community structure, richness, and density.

Plant succession is one of many factors that may affect the composition and structure of herbivorous insect communities. However, few studies have examined the effect of forest age on the diversity and abundance of insect communities. If forest age influences insect diversity, then the schedule of timber harvest rotation may have consequent effects on biodiversity. The insect herbivore community on Quercus alba (white oak) in the Missouri Ozarks was sampled in a chronoseries, from recently harvested (2 yr) to old-growth (approximately 313 yr) forests. A total of nine sites and 39 stands within those sites were sampled in May and August 2003. Unique communities of plants and insects were found in the oldest forests (122-313 yr). Density and species richness of herbivores were positively correlated with increasing forest age in August but not in May. August insect density was negatively correlated with heat load index; in addition, insect density and richness increased over the chronoseries, but not on the sunniest slopes. Forest structural diversity (number of size classes) was positively correlated with forest age, but woody plant species richness was not. In sum, richness, density, and community structure of white oak insect herbivores are influenced by variation in forest age, forest structure, relative abundance of plant species, and abiotic conditions. These results suggest that time between harvests of large, long-lived, tree species such as white oak should be longer than current practice in order to maintain insect community diversity.     

Climate correlates of 20 years of trophic changes in a high-elevation riparian system

The consequences of climate change for ecosystem structure and function remain largely unknown. Here, I examine the ability of climate variation to explain long-term changes in bird and plant populations, as well as trophic interactions in a high-elevation riparian system in central Arizona, USA, based on 20 years of study. Abundances of dominant deciduous trees have declined dramatically over the 20 years, correlated with a decline in overwinter snowfall. Snowfall can affect overwinter presence of elk, whose browsing can significantly impact deciduous tree abundance. Thus, climate may affect the plant community indirectly through effects on herbivores, but may also act directly by influencing water availability for plants. Seven species of birds were found to initiate earlier breeding associated with an increase in spring temperature across years. The advance in breeding time did not affect starvation of young or clutch size. Earlier breeding also did not increase the length of the breeding season for single-brooded species, but did for multi-brooded species. Yet, none of these phenology-related changes was associated with bird population trends. Climate had much larger consequences for these seven bird species by affecting trophic levels below (plants) and above (predators) the birds. In particular, the climate-related declines in deciduous vegetation led to decreased abundance of preferred bird habitat and increased nest predation rates. In addition, summer precipitation declined over time, and drier summers also were further associated with greater nest predation in all species. The net result was local extinction and severe population declines in some previously common bird species, whereas one species increased strongly in abundance, and two species did not show clear population changes. These data indicate that climate can alter ecosystem structure and function through complex pathways that include direct and indirect effects on abundances and interactions of multiple trophic components.