Showing papers in "Ecology in 2004"

Toward a metabolic theory of ecology

Abstract: Metabolism provides a basis for using first principles of physics, chemistry, and biology to link the biology of individual organisms to the ecology of populations, communities, and ecosystems. Metabolic rate, the rate at which organisms take up, transform, and expend energy and materials, is the most fundamental biological rate. We have developed a quantitative theory for how metabolic rate varies with body size and temperature. Metabolic theory predicts how metabolic rate, by setting the rates of resource uptake from the environment and resource allocation to survival, growth, and reproduction, controls ecological processes at all levels of organization from individuals to the biosphere. Examples include: (1) life history attributes, including devel- opment rate, mortality rate, age at maturity, life span, and population growth rate; (2) population interactions, including carrying capacity, rates of competition and predation, and patterns of species diversity; and (3) ecosystem processes, including rates of biomass production and respiration and patterns of trophic dynamics. Data compiled from the ecological literature strongly support the theoretical predictions. Even- tually, metabolic theory may provide a conceptual foundation for much of ecology, just as genetic theory provides a foundation for much of evolutionary biology.

Nitrogen mineralization: challenges of a changing paradigm

Abstract: Until recently, the common view of the terrestrial nitrogen cycle had been driven by two core assumptions—plants use only inorganic N and they compete poorly against soil microbes for N. Thus, plants were thought to use N that microbes “left over,” allowing the N cycle to be divided cleanly into two pieces—the microbial decomposition side and the plant uptake and use side. These were linked by the process of net mineralization. Over the last decade, research has changed these views. N cycling is now seen as being driven by the depolymerization of N-containing polymers by microbial (including mycorrhizal) extracellular enzymes. This releases organic N-containing monomers that may be used by either plants or microbes. However, a complete new conceptual model of the soil N cycle needs to incorporate recent research on plant–microbe competition and microsite processes to explain the dynamics of N across the wide range of N availability found in terrestrial ecosystems. We discuss the evolution of thinking abou...

Plant functional markers capture ecosystem properties during secondary succession

Abstract: Although the structure and composition of plant communities is known to influence the functioning of ecosystems, there is as yet no agreement as to how these should be described from a functional perspective. We tested the biomass ratio hypothesis, which postulates that ecosystem properties should depend on species traits and on species contribution to the total biomass of the community, in a successional sere following vineyard abandonment in the Mediterranean region of France. Ecosystem-specific net primary productivity, litter decomposition rate, and total soil carbon and nitrogen varied significantly with field age, and correlated with community-aggregated (i.e., weighed according to the relative abundance of species) functional leaf traits. The three easily measurable traits tested, specific leaf area, leaf dry matter content, and nitrogen concentration, provide a simple means to scale up from organ to ecosystem functioning in complex plant communities. We propose that they be called ''functional markers,'' and be used to assess the impacts of community changes on ecosystem properties induced, in particular, by global change drivers.

Interpolating, extrapolating, and comparing incidence-based species accumulation curves

Abstract: A general binomial mixture model is proposed for the species accumulation function based on presence-absence (incidence) of species in a sample of quadrats or other sampling units. The model covers interpolation between zero and the observed number of samples, as well as extrapolation beyond the observed sample set. For interpolation (sample- based rarefaction), easily calculated, closed-form expressions for both expected richness and its confidence limits are developed (using the method of moments) that completely eliminate the need for resampling methods and permit direct statistical comparison of richness between sample sets. An incidence-based form of the Coleman (random-placement) model is developed and compared with the moment-based interpolation method. For ex- trapolation beyond the empirical sample set (and simultaneously, as an alternative method of interpolation), a likelihood-based estimator with a bootstrap confidence interval is de- scribed that relies on a sequential, AIC-guided algorithm to fit the mixture model parameters. Both the moment-based and likelihood-based estimators are illustrated with data sets for temperate birds and tropical seeds, ants, and trees. The moment-based estimator is confi- dently recommended for interpolation (sample-based rarefaction). For extrapolation, the likelihood-based estimator performs well for doubling or tripling the number of empirical samples, but it is not reliable for estimating the richness asymptote. The sensitivity of individual-based and sample-based rarefaction to spatial (or temporal) patchiness is dis- cussed.

Pollen limitation of plant reproduction: Ecological and evolutionary causes and consequences

Abstract: Determining whether seed production is pollen limited has been an area of intensive empirical study over the last two decades. Yet current evidence does not allow satisfactory assessment of the causes or consequences of pollen limitation. Here, we critically evaluate existing theory and issues concerning pollen limitation. Our main conclusion is that a change in approach is needed to determine whether pollen limitation reflects random fluctuations around a pollen–resource equilibrium, an adaptation to stochastic pollination environments, or a chronic syndrome caused by an environmental perturbation. We formalize and extend D. Haig and M. Westoby's conceptual model, and illustrate its use in guiding research on the evolutionary consequences of pollen limitation, i.e., whether plants evolve or have evolved to ameliorate pollen limitation. This synthesis also reveals that we are only beginning to understand when and how pollen limitation at the plant level translates into effects on plant population dynamics...

Scaling of c:n:p stoichiometry in forests worldwide: implications of terrestrial redfield‐type ratios

Abstract: Inspired by the importance of globally well-constrained carbon:nitrogen: phosphorus (C:N:P) ratios in planktonic biomass to the understanding of nutrient cycles and biotic feedbacks in marine ecosystems, we looked for analogous patterns in forest ecosystems worldwide. We used data from the literature to examine the stoichiometry of C, N, and P in forest foliage and litter on both global and biome levels. Additionally, we examined the scaling of nutrient investments with biomass and production both globally and within biomes to determine if and when these ratios respond to macroscale ecosystem properties (such as nutrient availability). We found that, while global forest C:N:P ratios in both foliage and litter were more variable than those of marine particulate matter, biome level (temperate broadleaf, temperate coniferous, and tropical) ratios were as constrained as marine ratios and statistically distinct from one another. While we were more interested in the relative constancy of the C:N:P ratios than their numerical value we did note, as have others, that the atomic ratios calculated for foliage (1212:28:1) and litter (3007:45:1) reflect the increased proportion of C-rich structural material characteristic of terrestrial vegetation. Carbon : nutrient ratios in litter were consistently higher than in comparable foliar data sets, suggesting that resorption of nutrients is a globally important mechanism, particularly for P. Litter C:N ratios were globally constant despite biome-level differences in foliar C:N; we speculate that this strong coupling may be caused by the significant contribution of immobile cell wall bound proteins to the total foliar N pool. Most ratios scaled isometrically across the range of biomass stocks and production in all biomes sug- gesting that ratios arise directly from physiological constraints and are insensitive to factors leading to shifts in biomass and production. There were, however, important exceptions to this pattern: nutrient investment in broadleaf forest litter and coniferous forest foliage increased disproportionately relative to C with increasing biomass and production sug- gesting a systematic influence of macroscopic factors on ratios.

Dissecting the spatial structure of ecological data at multiple scales

Abstract: Spatial structures may not only result from ecological interactions, they may also play an essential functional role in organizing the interactions. Modeling spatial patterns at multiple spatial and temporal scales is thus a crucial step to understand the functioning of ecological communities. PCNM (principal coordinates of neighbor matrices) analysis achieves a spectral decomposition of the spatial relationships among the sampling sites, creating variables that correspond to all the spatial scales that can be perceived in a given data set. The analysis then finds the scales to which a data table of interest responds. The significant PCNM variables can be directly interpreted in terms of spatial scales, or included in a procedure of variation decomposition with respect to spatial and environmental components. This paper presents four applications of PCNM analysis to ecological data representing combinations of: transect or surface data, regular or irregular sampling schemes, univariate or multivariate data. The data sets include Amazonian ferns, tropical marine zooplankton, chlorophyll in a marine lagoon, and oribatid mites in a peat bog. In each case, new ecological knowledge was obtained through PCNM analysis.

Climate change uncouples trophic interactions in an aquatic ecosystem

Abstract: The largest uncertainty in forecasting the effects of climate change on eco- systems is in understanding how it will affect the nature of interactions among species. Climate change may have unexpected consequences because different species show unique responses to changes in environmental temperatures. Here we show that increasingly warmer springs since 1962 have disrupted the trophic linkages between phytoplankton and zoo- plankton in a large temperate lake because of differing sensitivity to vernal warming. The timing of thermal stratification and the spring diatom bloom have advanced by more than 20 days during this time period. A long-term decline in Daphnia populations, the keystone herbivore, is associated with an expanding temporal mismatch with the spring diatom bloom and may have severe consequences for resource flow to upper trophic levels.

Extracting more out of relocation data: building movement models as mixtures of random walks

Abstract: We present a framework for fitting multiple random walks to animal move- ment paths consisting of ordered sets of step lengths and turning angles. Each step and turn is assigned to one of a number of random walks, each characteristic of a different behavioral state. Behavioral state assignments may be inferred purely from movement data or may include the habitat type in which the animals are located. Switching between different behavioral states may be modeled explicitly using a state transition matrix estimated directly from data, or switching probabilities may take into account the proximity of animals to landscape features. Model fitting is undertaken within a Bayesian framework using the WinBUGS software. These methods allow for identification of different movement states using several properties of observed paths and lead naturally to the formulation of movement models. Analysis of relocation data from elk released in east-central Ontario, Canada, suggests a biphasic movement behavior: elk are either in an ''encamped'' state in which step lengths are small and turning angles are high, or in an ''exploratory'' state, in which daily step lengths are several kilometers and turning angles are small. Animals encamp in open habitat (agricultural fields and opened forest), but the exploratory state is not associated with any particular habitat type.

Measuring plant interactions: a new comparative index

Abstract: We propose an index to measure the relative interaction intensity in plants (RII) with strong mathematical and statistical properties which overcome problems shown by other frequently used indices. RII has defined limits [−1, +1]; is symmetrical around zero, with identical absolute values for competition and facilitation; is linear; and does not have discontinuities in its range. It is therefore safe to use in statistical and mathematical operations. RII distribution is approximately normal, with means equal to the true population index and a sampling variance that can be derived. Its strong statistical properties make RII proper for use in parametric meta-analyses. It can be applied to any kind of interaction (from competitive exclusion to symbiosis) and in commonly published ranges of interaction intensity it offers the most consistent results. Because RII uses basic arithmetical operators, it can be scaled up and used to measure multispecific interactions at the community level.

Maternal age as a determinant of larval growth and survival in a marine fish, sebastes melanops

Abstract: Relative body size has long been recognized as a factor influencing repro- ductive success in fishes, but maternal age has only recently been considered. We monitored growth and starvation resistance in larvae from 20 female black rockfish (Sebastes melan- ops), ranging in age from five to 17 years. Larvae from the oldest females in our experiments had growth rates more than three times as fast and survived starvation more than twice as long as larvae from the youngest females. Female age was a far better predictor of larval performance than female size. The apparent underlying mechanism is a greater provisioning of larvae with energy-rich triacylglycerol (TAG) lipids as female age increases. The volume of the oil globule (composed primarily of TAG) present in larvae at parturition increases with maternal age and is correlated with subsequent growth and survival. These results suggest that progeny from older females can survive under a broader range of environmental conditions compared to progeny from younger females. Age truncation commonly induced by fisheries may, therefore, have severe consequences for long-term sustainability of fish populations.

Adaptive variation in the vulnerability of woody plants to xylem cavitation

Abstract: The ability of plants to supply water to their leaves is intimately associated with survival. Water supply to leaves depends on maintaining an intact water column in the xylem from the roots to shoots. Because this hydraulic pathway is under tension, it is vulnerable to breakage through the induction of air emboli (cavitation). Although the phys- iological benefit of resistance to water-stress-induced xylem cavitation for desiccation tol- erance is clear, there is considerable interspecific variation within and across climates. To understand the adaptive significance of this variation and the potential trade-off with water transport, we compiled a database of 167 species from 50 seed plant families and examined relationships among resistance to xylem cavitation, water transport capacity (as determined by the specific conductivity of xylem ( KS)), and climate. Relationships were evaluated using standard cross-species correlations ( r). Because inferences about the adaptive significance of these correlations can be biased by the potential similarity of closely related species, we also analyzed our data using phylogenetically independent contrast correlations (PIC) calculated over a range of alternate seed plant phylogenies. Resistance to cavitation, ex- pressed as the xylem tension at which 50% of hydraulic conductivity was lost (C50), ranged from 20.18 to 29.9 MPa for angiosperms and from 21.5 to 214.1 MPa for conifers. Conifers were most resistant to cavitation, with mean C50 80% more negative than angio- sperms. In contrast, KS was 270% higher in angiosperms than conifers. Across all species, cavitation resistance increased with decreasing mean annual precipitation. However, sig- nificant phylogenetically independent contrast correlations between C50 and annual precip- itation were found within the evergreen angiosperms and conifers but not in the deciduous angiosperms. Thus, the adaptive significance of increased resistance to cavitation as a mechanism of drought tolerance may be of primary importance in evergreen angiosperms and conifers. In contrast, analysis of independent contrasts indicated that KS increased with decreasing rainfall in deciduous angiosperms, whereas there was no association between KS and water availability for evergreen angiosperms and conifers. These results suggest that the evolution of increased KS may be a critical adaptation to water limitation in de- ciduous angiosperms. Although there was a significant cross-species correlation between C50 and KS, this relationship was not supported by the independent contrast correlation, suggesting that the evolutionary basis for a trade-off between cavitation resistance and water transport capacity is weak.

Plant water stress and its consequences for herbivorous insects: a new synthesis

Abstract: Traditionally, herbivorous insects are thought to exhibit enhanced performance and outbreak dynamics on water-stressed host plants due to induced changes in plant physiology. Recent experimental studies, however, provide mixed support for this historical view. To test the plant-stress hypothesis (PSH), we employed two methods (the traditional vote-counting approach and meta-analysis) to assess published studies that investigated insect responses to experimentally induced water-deficit in plants. For insects, we examined how water deficit affects survivorship, fecundity, density, relative growth rate, and oviposition preference. Responses were analyzed by major feeding guild (sap-feeding insects and chewing insects) and for the subguilds of sap-feeders (phloem, mesophyll, and xylem feeders) and chewing insects (free-living chewers, borers, leaf miners, and gall-formers). Both vote counting and meta-analysis found strong negative effects of water stress on the performance of sap-feeding insects at large and...

Historical landscape connectivity affects present plant species diversity

Abstract: Transformation of landscapes is considered to be one of the main drivers behind species loss, regionally and globally. Theory and empirical studies suggest that landscape structure influences speci ...

Changes in turbulent mixing shift competition for light between phytoplankton species

Abstract: The intriguing impact of physical mixing processes on species interactions has always fascinated ecologists. Here, we exploit recent advances in plankton models to develop competition theory that predicts how changes in turbulent mixing affect competition for light between buoyant and sinking phytoplankton species. We compared the model predictions with a lake experiment, in which the turbulence structure of the entire lake was manipulated using artificial mixing. Vertical eddy diffusivities were calculated from the measured temperature microstructure in the lake. Changes in turbulent mixing of the lake caused a dramatic shift in phytoplankton species composition, consistent with the predictions of the competition model. The buoyant and potentially toxic cyanobacterium Microcystis dominated at low turbulent diffusivity, whereas sinking diatoms and green algae dominated at high turbulent diffusivity. These findings warn that changes in the turbulence structure of natural waters, for instance driven by climate change, may induce major shifts in the species composition of phytoplankton communities.

Plant functional traits in relation to fire in crown-fire ecosystems

Abstract: Disturbance is a dominant factor in many ecosystems, and the disturbance regime is likely to change over the next decades in response to land-use changes and global warming. We assume that predictions of vegetation dynamics can be made on the basis of a set of life-history traits that characterize the response of a species to disturbance. For crown-fire ecosystems, the main plant traits related to postfire persistence are the ability to resprout (persistence of individuals) and the ability to retain a persistent seed bank (persistence of populations). In this context, we asked (1) to what extent do different life- history traits co-occur with the ability to resprout and/or the ability to retain a persistent seed bank among differing ecosystems and (2) to what extent do combinations of fire- related traits (fire syndromes) change in a fire regime gradient? We explored these questions by reviewing the literature and analyzing databases compiled from different crown-fire ecosystems (mainly eastern Australia, California, and the Mediterranean basin). The review suggests that the pattern of correlation between the two basic postfire persistent traits and other plant traits varies between continents and ecosystems. From these results we predict, for instance, that not all resprouters respond in a similar way everywhere because the associated plant traits of resprouter species vary in different places. Thus, attempts to generalize predictions on the basis of the resprouting capacity may have limited power at a global scale. An example is presented for Australian heathlands. Considering the com- bination of persistence at individual (resprouting) and at population (seed bank) level, the predictive power at local scale was significantly increased.

The uplift of soil nutrients by plants: biogeochemical consequences across scales

Abstract: Although the bulk of plant biomass contains relatively light, atmospherically derived elements (C, H, O, N, and S), 5-10% of biomass is composed of heavier elements from soil minerals, such as Ca, Mg, K, and P. Plant uptake and cycling transport these heavier elements to the soil surface, resulting in shallower vertical distributions for strongly cycled elements than for other elements. In this paper, we evaluate the biogeochemical consequences of this process at different spatial and temporal scales based on chronose- quence studies and soil database analyses. In the bare coastal dunes of Argentina, the vertical distributions of exchangeable K 1 (strongly cycled) and Na 1 (more weakly cycled) were similar initially but diverged 15 years after pine afforestation, with K distributions becoming significantly concentrated in the surface and Na distributions becoming deeper. To evaluate the effects of plant stoichiometry on micronutrient distributions, chronose- quences of paired native grasslands (low Mn cycling) and eucalypt plantations (high Mn cycling) in the pampas of Argentina were also used. Within 50 years, eucalypts dramatically redistributed Mn pools toward the soil surface, reducing total pools by half at medium depths (20-60 cm) and increasing concentrations by up to an order of magnitude at the surface. Globally, we used generalized contrasts among exchangeable K, Na, and Mg in 7661 soil profiles to estimate the global magnitude of K uplift due to plant activity. Based on this calculation, the exchangeable K pool in the top 20 cm of soils without plant uplift would be 4-6 3 10 15 g smaller globally, one-third to one-half smaller than its current size. Vegetation change alters the vertical distribution and bioavailability of mineral elements. Understanding how the stoichiometry of plant cycling affects soil nutrient distributions will help refine predictions of the biogeochemical consequences of current vegetation change.

The intermediate disturbance hypothesis: patch dynamics and mechanisms of species coexistence

Abstract: The intermediate disturbance hypothesis (IDH) has been used for several decades as an explanation for the coexistence of species in ecological communities. It is intuitively simple, but deceptively so. We show, via discussion and examples, that the IDH is not one mechanism of coexistence, but rather summarizes a set of similar phenomena that can arise from the action of several different coexistence mechanisms. These underlying mechanisms are defined by the various ways in which species differ in their response to disturbance-induced spatial and temporal variability in resources and environmental conditions. As an example, the original specification of the IDH required patchy disturbances for coexistence. However, because the underlying mechanisms of coexistence can also operate at the within-patch scale, patchy disturbances are not a necessary requirement for coexistence under intermediate-disturbance regimes. These conclusions are illustrated through the analysis of three models: a spatial within-patch model, a spatial between-patch model, and a purely temporal model. All three generate similar patterns of coexistence under intermediate disturbance, yet underlying that coexistence lie at least two quite-distinct mechanisms of species coexistence: the storage effect and relative nonlinearity. The results from our analyses suggest that, as a promoter of species coexistence, the IDH is both broader in scope and richer in detail than has previously been recognized.

Fish invasion restructures stream and forest food webs by interrupting reciprocal prey subsidies

Abstract: Habitat alteration and biotic invasions are the two leading causes of global environmental change and biodiversity loss. Recent innovative experiments have shown that habitat disturbance can have drastic effects that cascade to adjacent ecosystems by altering the flow of resource subsidies from donor systems. Likewise, exotic species in- vasions could alter subsidies and affect distant food webs, but very few studies have tested this experimentally. Here we report evidence from a large-scale field experiment in northern Japan that invasion of nonnative rainbow trout (Oncorhynchus mykiss) interrupted reciprocal flows of invertebrate prey that drove stream and adjacent riparian forest food webs. Rainbow trout usurped terrestrial prey that fell into the stream, causing native Dolly Varden charr (Salvelinus malma) to shift their foraging to insects that graze algae from the stream bottom. This indirectly increased algal biomass, but also decreased biomass of adult aquatic insects emerging from the stream to the forest. In turn, this led to a 65% reduction in the density of riparian-specialist spiders in the forest. Thus, species invasions can interrupt flows of resources between interconnected ecosystems and have effects that propagate across their boundaries, effects that may be difficult to anticipate without in-depth understanding of food web relationships.

Carbon sequestration in ecosystems: the role of stoichiometry

Abstract: The fate of carbon (C) in organisms, food webs, and ecosystems is to a major extent regulated by mass-balance principles and the availability of other key nutrient elements. In relative terms, nutrient limitation implies excess C, yet the fate of this C may be quite different in autotrophs and heterotrophs. For autotrophs nutrient limitation means less fixation of inorganic C or excretion of organic C, while for heterotrophs nutrient limitation means that more of ingested C will “go to waste” in the form of egestion or respiration. There is in general a mismatch between autotrophs and decomposers that have flexible but generally high C:element ratios, and consumers that have lower C:element ratios and tighter stoichiometric regulation. Thus, C-use efficiency in food webs may be governed by the element ratios in autotroph biomass and tend to increase when C:element ratios in food approach those of consumers. This tendency has a strong bearing on the sequestration of C in ecosystems, since more C will be di...

Mapping the fundamental niche: physiology, climate, and the distribution of a nocturnal lizard

Abstract: The fundamental niche can be viewed as the set of conditions and resources that allow a given organism to survive and reproduce in the absence of biotic interactions. Quantitative descriptions of the environmental variables with which organisms are asso- ciated are becoming common with the advent of geographic information systems (GIS). Although such descriptive approaches to the niche are useful for interpolating species distributions, they implicitly incorporate biotic interactions and therefore do not represent the fundamental niche. A mechanistic understanding of the fundamental niche, when com- bined with GIS data, can provide us with greater insight into the causes of distribution and abundance, a solid foundation for exploring the role of biotic interactions, and greater confidence in extrapolating to novel circumstances such as climate change and species introductions. We apply such a mechanistic approach to study the climatic component of the fundamental niche of a nocturnal lizard, Heteronotia binoei, across an entire continent. We combine physiological measurements of this species (thermal requirements for egg development, thermal preferences and tolerances, metabolic and evaporative water loss rates), and high-resolution climatic data for the Australian continent (air temperature, cloud cover, wind speed, humidity, and radiation), with biophysical models to calculate the cli- matic component of the fundamental niche of this lizard and map it onto the Australian landscape at high resolution. We also use this approach to predict the effects of a mild global warming on the degree-days in the soil for egg development and the potential for aboveground activity of the study organism.

Recognition and selection of settlement substrata determine post-settlement survival in corals

Abstract: Habitat recognition and selective settlement by dispersive propagules greatly increases the post-settlement survival chances of sessile organisms. To better understand the key role some species can play in the structure of highly complex coral reef ecosystems, we compare the role of two independent, but sequential, processes: settlement choice and post-settlement survival. This study describes the chemical and physical recognition and ranking of specific settlement substrata by coral larvae. Several species of crustose coralline algae (CCA) are known to induce coral settlement; however they also employ physical and biological anti-settlement defense strategies that vary greatly in effectiveness. We examine the interactions between settling larvae of two common reef building coral species (Acropora tenuis and A. millepora) and five species of CCA (Neogoniolithon fosliei, Porolithon onkodes, Hydrolithon reinboldii, Titanoderma prototypum, and Lithoporella melobesioides) that co-occur on reef crests and slopes of the Great Barrier Reef, Australia. Distinct settlement patterns were observed when coral larvae were provided with a choice of settlement substrata. Settlement on the most preferred substratum, the CCA species T. prototypum, was 15 times higher than on N. fosliei, the least preferred substratum. The rates of post-settlement survival of the corals also varied between CCA species in response to their anti-settlement strategies (shedding of surface cell layers, overgrowth, and potential chemical deterrents). Rates of larval settlement, post-settlement survival, and the sensitivity of larvae to chemical extracts of CCA were all positively correlated across the five species of CCA. Nonliving settlement substrata on coral reefs is sparse; consequently the fact that only a few CCA species (notably T. prototypum) facilitate coral recruitment, has important implications for structuring the reef ecosystem.

Facilitative interactions among plants via shared pollinators

Abstract: Outcrossing in plants is influenced by the availability of pollinators and compatible mates, both of which may be modified by the population and community context in which plant–pollinator interactions occur. Although indirect interactions among plants through shared pollinators are often expected to be competitive, pollinator sharing may be beneficial when plant species jointly attract or maintain populations of pollinators. In this study, I tested the hypothesis that pollinator-sharing congeners facilitate reproduction in a focal taxon, Clarkia xantiana ssp. xantiana, and that positive interactions are most pronounced in small and sparse populations. Population surveys revealed that C. x. xantiana frequently coexists with pollinator-sharing congeners except at the periphery of its range. Populations varied extensively in size and density, with small populations more likely associated with pollinator-sharing congeners; conversely, populations occurring alone were more likely large. Flowering schedules in...

Fire regimes at the transition between mixedwood and coniferous boreal forest in northwestern quebec

Abstract: Fire history was reconstructed for an area of 15 000 km 2 located in the transition zone between the mixed and coniferous forests in Quebec's southern boreal forest. We used aerial photographs, archives, and dendroecological data (315 sites) to reconstruct a stand initiation map for the area. The cumulative distribution of burnt area in relation to time since fire suggests that the fire frequency has decreased drastically since the end of the Little Ice Age (about 1850) in the entire region. However, a large part of the area was burned between 1910 and 1920 during intensive colonization and when the climate was very conducive to fire. For the period 1920-1945, large fires have mainly been concentrated in the more populated southern area, while few fires have been observed in the virgin coniferous forest in the north. Despite slight differences between the south and the north, fire cycles or the average number of years since fire are not significantly different. Since 1945, there have been far more fires in the south, but the mean fire size was smaller than in the north. These results suggest that the transition between the mixed and coniferous forests observed in the southern boreal forest cannot be explained by a difference in fire frequency, at least during the last 300 years. As climatic factors and species potential distribution did not vary significantly from south to north, we suggest that the transition from mixedwood to coniferous forests is mainly controlled by fire size and severity. Smaller and less severe fires would favor species associated with the mixedwood forests as many need survivors to reinvade burnt areas. The abundance of deciduous species in mixedwood forests, together with the presence of more lakes that can act as firebreaks, may contribute to decreases in fire size and severity. The transition between the two vegetation zones could be related to the initial setting following the vegetation invasion of the area during the Holocene. In this context, the limit of vegetation zones in systems controlled by disturbance regimes such as fires may not have reached a balance with current climatic conditions. Historical legacies and strong positive feedback between disturbance regimes and com- position may filter and delay the responses to changes in climate.

How do different measures of functional diversity perform

Abstract: Biodiversity can influence ecosystem functioning through changes in the amount of resource use complementary among species. Functional diversity is a measure of biodiversity that aims to quantify resource use complementarity and thereby explain and predict ecosystem functioning. The primary goal of this article is to compare the explanatory power of four measures of functional diversity: species richness, functional group richness, functional attribute diversity, and FD. The secondary goal is to showcase the novel methods required for calculating functional attribute diversity and FD. We find that species richness and functional group richness explain the least variation in above- ground biomass production within and across grassland biodiversity manipulations at six European locations; functional attribute diversity and FD explain greater variation. Rea- sons for differences in explanatory power are discussed, such as the relatively greater amount of information and fewer assumptions included in functional attribute diversity and FD. We explore the opportunities and limitations of the particular methods we used to calculate functional attribute diversity and FD. These mainly concern how best to select the information used to calculate them.

Competition between native perennial and exotic annual grasses: implications for an historical invasion

Abstract: Though established populations of invasive species can exert substantial competitive effects on native populations, exotic propagules may require disturbances that decrease competitive interference by resident species in order to become established. We compared the relative competitiveness of native perennial and exotic annual grasses in a California coastal prairie grassland to test whether the introduction of exotic propagules to coastal grasslands in the 19th century was likely to have been sufficient to shift community composition from native perennial to exotic annual grasses. Under experimental field con- ditions, we compared the aboveground productivity of native species alone to native species competing with exotics, and exotic species alone to exotic species competing with natives. Over the course of the four-year experiment, native grasses became increasingly dominant in the mixed-assemblage plots containing natives and exotics. Although the competitive interactions in the first growing season favored the exotics, over time the native grasses significantly reduced the productivity of exotic grasses. The number of exotic seedlings emerging and the biomass of dicot seedlings removed during weeding were also significantly lower in plots containing natives as compared to plots that did not contain natives. We found evidence that the ability of established native perennial species to limit space available for exotic annual seeds to germinate and to limit the light available to exotic seedlings reduced exotic productivity and shifted competitive interactions in favor of the natives. If interactions between native perennial and exotic annual grasses follow a similar pattern in other coastal grassland habitats, then the introduction of exotic grass propagules alone without changes in land use or climate, or both, was likely insufficient to convert the region's grasslands.

Asymmetric specialization: a pervasive feature of plant-pollinator interactions

Abstract: Although specialization in species interactions has usually been equated to reciprocal specialization, asymmetric specialization (i.e., a specialist interacting with a generalist) is also likely. Recent studies have suggested that asymmetric specialization in species interactions could be more common than previously thought. We contrasted patterns of asymmetric specialization observed in 18 plant-pollinator interaction webs with pre- dictions based on null models. We found that asymmetric specialization is common in plant-pollinator interactions, and that its occurrence is more frequent than expected under a simple null model that assumed random interactions among species; furthermore, large assemblages with many pairs of interacting species tend to have more asymmetric inter- actions than smaller assemblages. A second null model, which incorporated a correlation between species frequency of interaction and degree of specialization observed in most data sets produced patterns that were generally closer to those present in the data. At least three kinds of explanations could account for the observed asymmetric specialization, including random interactions among individuals (rather than species), adaptive conse- quences of specialization, and artifacts, such as data aggregation and sampling biases. Future studies should be aimed at understanding the relative importance of each of these alternative explanations in generating asymmetric specialization in species interactions.

Habitat complexity disrupts predator–prey interactions but not the trophic cascade on oyster reefs

Abstract: Despite recognition of the significance of both food web interactions and habitat complexity in community dynamics, current ecological theory rarely couples these two processes Experimental manipulations of the abundance of the two predators in an oyster-reef trophic cascade, and the structural complexity provided by reefs of living oysters, demonstrated that enhanced habitat complexity weakened the strengths of trophic interactions The system of tri-trophic interactions included oyster toadfish (Opsanus tau) as the top predator that consumed the mud crab (Panopeus herbstii), which preys upon juvenile oysters (Crassostrea virginica) On reefs of low complexity, toadfish controlled mud crab abundances and indirectly determined the level of mortality of juvenile oysters The indirect effects of toadfish on oysters emerged through their influence on how intensely mud crabs preyed on oysters Augmentation of habitat complexity by substituting vertically oriented, living oysters for the flat shells of dead o

Age‐dependent tree‐ring growth responses to climate in larix decidua and pinus cembra

Abstract: Dendrochronology generally operates under the assumption that climate–growth relationships are age independent, once growth trends and/or disturbance pulses have been accounted for. However, several studies have demonstrated that tree physiology undergoes changes with age. This may cause growth-related climate signals to vary over time. Using chronology statistics and response functions, we tested the consistency of climate–growth responses in tree-ring series from Larix decidua and Pinus cembra trees of four age classes. Tree-ring statistics (mean sensitivity, standard deviation, correlation between trees, and first principal component) did not change significantly with age in P. cembra, whereas in L. decidua they appeared to be correlated with age classes. Response function analysis indicated that climate accounts for a high amount of variance in tree-ring widths in both species. The older the trees are, the higher the variance explained by climate, the significance of the models, and the percentage of ...

A predictive model of edge effects

Abstract: Edge effects are among the most extensively studied ecological phenomena, yet we lack a general, predictive framework to understand the patterns and variability observed We present a conceptual model, based on resource distribution, that predicts whether organismal abundances near edges are expected to increase, decrease, or remain unchanged for any species at any edge type Predictions are based on whether resources are found predominantly in one habitat (decreased abundance in preferred habitat, increase in non-preferred), divided between habitats (predicts an increase near both edges), spread equally among habitats (predicts a neutral edge response), or concentrated along the edge (increase) There are several implications of this model that can explain much of the variability reported in the edge literature For instance, our model predicts that a species may show positive, negative, and neutral responses, depending on the edge type encountered, which explains some intraspecific variability observed