Showing papers in "Oikos in 2008"

A consistent metric for nestedness analysis in ecological systems: reconciling concept and measurement

Abstract: Nestedness has been widely reported for both metacommunities and networks of interacting species. Even though the concept of this ecological pattern has been well-defined, there are several metrics by which it can be quantified. We noted that current metrics do not correctly quantify two major properties of nestedness: (1) whether marginal totals (i.e. fills) differ among columns and/or among rows, and (2) whether the presences (1’s) in less-filled columns and rows coincide, respectively, with those found in the more-filled columns and rows. We propose a new metric directly based on these properties and compare its behavior with that of the most used metrics, using a set of model matrices ranging from highly-nested to alternative structures in which no nestedness should be detected. We also used an empirical dataset to explore possible biases generated by the metrics as well as to evaluate correlations between metrics. We found that nestedness has been quantified by metrics that inappropriately detect this pattern, even for matrices in which there is no nestedness. In addition, the most used metrics are prone to type I statistical errors while our new metric has better statistical properties and consistently rejects a nested pattern for different types of random matrices. The analysis of the empirical data showed that two nestedness metrics, matrix temperature and the discrepancy measure, tend to overestimate the degrees of nestedness in metacommunities. We emphasize and discuss some implications of these biases for the theoretical understanding of the processes shaping species interaction networks and metacommunity structure.

Energy metabolism and animal personality

Abstract: In this paper we show how animal personality could explain some of the large inter-individual variation in resting metabolic rate (MR) and explore methodological and functional linkages between personality and energetics. Personality will introduce variability in resting MR measures because individuals consistently differ in their stress response, exploration or activity levels, all of which influence MR measurements made with respirometry and the doubly-labelled water technique. Physiologists try to exclude these behavioural influences from resting MR measurements, but animal personality research indicates that these attempts are unlikely to be successful. For example, because reactive animals "freeze" when submitted to a stress, their MR could be classified as "resting" because of immobility when in fact they are highly stressed with an elevated MR. More importantly, recent research demonstrating that behavioural responses to novel and highly artificial stimuli are correlated with both behaviour and fitness under more natural circumstances calls into question the wisdom of excluding these behavioural influences on MR measurements. The reason that intra-specific variation in resting MR are so weakly correlated with daily energy expenditure (DEE) and fitness, may be that the latter two measures fully incorporate personality while the former partially excludes its influence. Because activity, exploration, boldness and aggressiveness are energetically costly, personality and metabolism should be correlated and physiological constraints may underlie behavioural syndromes. We show how physiological ecologists can better examine behavioural linkages between personality and metabolism, as required to better understand the physiological correlates of personality and the evolutionary consequences of metabolic variability.

Distribution of specialist and generalist species along spatial gradients of habitat disturbance and fragmentation

Abstract: In this paper, we tested whether the spatial distribution of a given species in more or less fragmented and disturbed landscapes depends on the species habitat specialization. We studied 891 spatial replicates from the French Breeding Bird Survey (FBBS) monitored at least two years during 2001 �2005, and two independent landscape databases measuring respectively landscape fragmentation and recent landscape disturbance on each FBBS replicate. We used a continuous habitat specialization index for the 105 most common bird species monitored by the FBBS. We further modelled the spatial variation in abundance of each species according to fragmentation and disturbance across FBBS replicates, accounting for habitat differences and spatial trends. We then tested whether more or less specialized species responded to landscape fragmentation and disturbance. We found that the more specialist a species, the more negative its spatial response to landscape fragmentation and disturbance. Although there was a very high variation around these tendencies indicating that there are many other drivers of species distribution, our results suggest that measuring specialization may be helpful in predicting which species are likely to thrive in human degraded landscapes. We also emphasize the need to consider both positive and negative species responses when assessing consequences of habitat change in communities.

Search and navigation in dynamic environments – from individual behaviors to population distributions

Abstract: Animal movement receives widespread attention within ecology and behavior. However, much research is restricted within isolated sub-disciplines focusing on single phenomena such as navigation (e.g. homing behavior), search strategies (e.g. Levy flights) or theoretical considerations of optimal population dispersion (e.g. ideal free distribution). To help synthesize existing research, we outline a unifying conceptual framework that integrates individual-level behaviors and population-level spatial distributions with respect to spatio-temporal resource dynamics. We distinguish among (1) non-oriented movements based on diffusion and kinesis in response to proximate stimuli, (2) oriented movements utilizing perceptual cues of distant targets, and (3) memory mechanisms that assume prior knowledge of a target's location. Species' use of these mechanisms depends on life-history traits and resource dynamics, which together shape population-level patterns. Resources with little spatial variability should facilitate sedentary ranges, whereas resources with predictable seasonal variation in spatial distributions should generate migratory patterns. A third pattern, 'nomadism', should emerge when resource distributions are unpredictable in both space and time. We summarize recent advances in analyses of animal trajectories and outline three major components on which future studies should focus: (1) integration across alternative movement mechanisms involving links between state variables and specific mechanisms, (2) consideration of dynamics in resource landscapes or environments that include resource gradients in predictability, variability, scale, and abundance, and finally (3) quantitative methods to distinguish among population distributions. We suggest that combining techniques such as evolutionary programming and pattern oriented modeling will help to build strong links between underlying movement mechanisms and broad-scale population distributions.

How does plant richness affect pollinator richness and temporal stability of flower visits

Abstract: Pollinators play a key role in the reproduction of most plant species, and pollinator and plant diversity are often related. We studied an experimental gradient of plant species richness for a better understanding of plantpollinator community interactions and their temporal variability, because in non-experimental field surveys plant richness is often confounded with gradients in management, soil fertility, and community composition. We observed pollinator species richness and frequency of visits six times in 73 plots over two years, and used advanced statistical analysis to account for the high number of zeroes that often occur in count data of rare species. The frequency of pollinator visits increased linearly with both the blossom cover and the number of flowering plant species, which was closely related to the total number of plant species, whereas the number of pollinator species followed a saturation curve. The presence of particularly attractive plant species was only important for the frequency of flower visits, but not to the richness of pollinators. Plant species richness, blossom cover, and the presence of attractive plant species enhanced the temporal stability in the frequency of pollinator visits. In conclusion, grasslands with high plant diversity enhance and stabilize frequent and diverse flower visitations, which should sustain effective pollination and plant reproduction.

Historical bias in biodiversity inventories affects the observed environmental niche of the species

Abstract: It is well known that biodiversity data from historical inventories presents important geographic and taxonomic biases. Due to this, current knowledge on the distribution of most species could be incomplete and biased. We assess how the biases in historical biodiversity data might affect the description of the environmental niche of the species, using exhaustive data on the distribution of dung beetles in Madrid as a case study. We describe the historical process of survey and compare such historical data with the results of an exhaustive survey, identifying the environmental biases in the historical surveys during different periods, and assessing the completeness of the environmental niche of the species provided by historical data through time. Events like the Spanish Civil War affect the tempo and spread of surveys, but the exhaustive work since 1970 provides a good, though incomplete, coverage of the region by 1998. In spite of this, the biases in historical data result in a limited knowledge about the niche of an important number of species. Although nearly a half of the species had the 100% of their niche covered by data in 1998, roughly a third had less than 75%, nearly a fourth less than 50%, and 18 species had to be excluded from the analyses due to the lack of data. Our results point out that data from non-standardized inventories often provide an incomplete description of the environmental responses of most species. Due to this, we highlight that currently predictive models of species distributions present some limitations, since the results of models based in partial information about the environmental niche of the species will be compromised. Therefore, the biases in the available data must be evaluated before constructing predictive maps of species distributions, and taken into account when drawing conclusions or conservation strategies from these maps.

Year‐to‐year variation in the topology of a plant–pollinator interaction network

Abstract: Vigorous discussion of the degree of specialization in pollination interactions, combined with advances in the analysis of complex networks, has revitalized the study of entire plantpollinator communities. Noticeably rare, however, are attempts to quantify temporal variation in the structure of plantpollinator networks, and to determine whether the status of species as specialists or generalists is stable. Here we show that network structure varied through time in a montane meadow community from southern California, USA, in that pollinator species did not form the same links with plant species across years. Furthermore, composition of the generalized core group of species in the network varied among summers, as did the identity of those species involved in relationships that appeared to be reciprocally specialized within any one summer. These differences appear to be related to severe drought conditions experienced in the second summer of the 3 year study. In contrast to this variation, the pollinator community remained similarly highly nested in all three summers, even though species were packed into the nested matrix differently from year to year. These results suggest that plantpollinator networks vary in detail through time, while retaining some basic topological properties. This dynamic aspect of community-scale interactions has implications for both ecological and evolutionary inferences about pollination mutualisms.

Agricultural landscapes with organic crops support higher pollinator diversity

Abstract: Pollinators are traditionally thought to perceive non-flowering crop fields as hostile landscape matrix. In this study, we show that landscapes composed of higher proportions of organic crop fields support more bee species at greater abundances in fallow strips. An increase in organic cropping in the surrounding landscape from 5% to 20% enhanced bee species richness in fallow strips by 50%, density of solitary bees by 60% and bumble bee density by 150%. Bee species richness and bumble bee density responded strongest to organic cropping in landscape sectors with 500 m radius, solitary bee density in landscape sectors with 250 m radius. The most likely source of these results is that crop and noncrop habitats are strongly connected via bee foraging at the landscape scale. It seems likely that bees depending on nesting sites in fallow strips benefited from the more abundant flower resources provided by broadleaved weeds in organic crop fields. We conclude that the incorporation of organic crop fields into conventionally managed agricultural landscapes can provide food resources needed to sustain greater pollinator species richness in noncrop habitats.

Trophic cascades in a temperate seagrass community

Abstract: We assessed the relative importance of bottom-up and top-down processes in structuring an eelgrass community in Sweden, a system impacted both by eutrophication and overfishing. Using artificial seagrass as substrate, we manipulated nutrient levels and predator abundance in a full-factorial cage-experiment. The results revealed a seagrass community dominated by strong top-down processes controlling the aggregate biomass of mesograzers and macroalgae. In the absence of predators the large amphipod Gammarus locusta became very abundant resulting in a leaf community with low biomass of algae and smaller mobile fauna. One enclosed gobid fish predator reduced the abundance of adult G. locusta by > 90%, causing a three to six times increase in the biomass of algae, smaller mesograzers and meiofauna. Numerous small predators in uncaged habitats reduced the biomass of G. locusta and other mesograzers by > 95% in comparison to the fish treatment, further increasing the biomass of epiphytic algae and meiofauna. Although water column nutrient enrichment caused a temporal bloom of the filamentous macroalgae Ulva spp., no significant nutrient-effects were found on the algal community at the end of the experiment. The only lasting nutrient-effect was a significant increase in the biomass of G. locusta, but only in the absence of ambient predators. These results demonstrate that mesograzers can respond to enhanced food supply, increase their biomass and control the algal growth when predation rates are low. However, in the assessed system, high predation rates appear to make mesograzers functionally extinct, causing a community-wide trophic cascade that promotes the growth of ephemeral algae. This top-down effect could penetrate down, despite a complex food-web because the interaction strength in the community was strongly skewed towards two functionally dominant algal and grazer species that were vulnerable to consumption. These results indicate that overexploitation of gadoid fish may be linked to increased macroalgal blooms and loss of eelgrass in the area through a trophic cascade affecting the abundance of mesograzers.

Personality, habitat use, and their consequences for survival in North American red squirrels Tamiasciurus hudsonicus

Abstract: Personality affects many aspects of an individual's behaviour, life history and fitness, and has been shown to be moderately heritable in wild populations. Correlations between personality and risk-taking that lead to life history tradeoffs could act to maintain variation in personality within a population, but this has not yet been tested. In this study, we used females from a marked population of North American red squirrels in Kluane, Yukon, to determine whether personality predicts risk-taking in the wild, and whether these risk-taking behaviours result in life history tradeoffs. We measured personality in open field and mirror image stimulation tests and extracted two traits, activity and aggressiveness, using principal component analysis and mixed model techniques. Using trapping records for individuals from February to September 2005, we obtained three measures of risk-taking: the number of trapping events, the number of different trapping locations, and the maximum distance between the home territory and a trapping event. We used GLMs to determine whether the activity and aggressiveness of individuals are related to these risk-taking behaviours, and found that active squirrels were trapped significantly more frequently and at a greater number of locations. There was also a significant interaction between activity and aggressiveness to affect the maximum capture distance. To determine if there are fitness tradeoffs associated with these risk-taking behaviours, we examined female bequeathal behaviour and survival. Bequeathing a territory increases offspring probability of overwinter survival, and we found that an increasing number of trapping locations was associated with an increasing tendency to bequeath. Active females were less likely to survive until the following spring. Risk-taking is therefore predicted by personality in this population, and they affect both survival and territorial bequeathal. These fitness tradeoffs may therefore lead to the maintenance of variation in personality.

Any way the wind blows ‐ frequent wind dispersal drives species sorting in ephemeral aquatic communities

Abstract: Despite an upsurge of interest in spatial interactions between communities and in the impact of dispersal on ecological and evolutionary processes, dispersal patterns and dynamics in natural metacommunities remain poorly understood. Although passive aerial dispersal of freshwater invertebrates is generally accepted, the frequency and relative importance of wind as a vector is still subject of considerable debate. We assessed the importance of wind dispersal in an invertebrate metacommunity in a cluster of 36 temporary rock pools on an isolated mountaintop in South Africa. Wind dispersal was quantified every four days using nine windsocks (about 1.5 m above rock base), placed in the field during one month. Distance to the nearest pool varied from 2 up to 16 m. Wind direction and speed were monitored for the entire period. About 850 propagules (mostly resting eggs) of 17 taxa were captured. The presence of water in the pools (level of exposure of the dormant propagule bank) and the dominant wind direction were the key factors affecting the yield. Wind speed was much less important. Our results suggest that wind dispersal of propagules from temporary aquatic systems is more frequent than previously thought. This may stabilise the metacommunity by mediating gene flow among populations and facilitating rapid (re)colonisation of patches. On the other hand, wind erosion of the dormant propagule bank may lead to egg bank depletion and local extinction. The measured frequent wind dispersal most likely fuels strong species sorting processes ultimately shaping the structure of the local communities as observed in an earlier study. To elucidate the link between local dispersal rates and their contribution to long range dispersal is a major challenge for future research on aerial dispersal of aquatic invertebrates.

Balancing multiple mutualists: asymmetric interactions among plants, arbuscular mycorrhizal fungi, and fungal endophytes

Abstract: Most organisms engage in beneficial interactions with other species; however, little is known regarding how individuals balance the competing demands of multiple mutualisms. Here we examine three-way interactions among a widespread grass, Schedonorus phoenix, a protective fungal endophyte aboveground, Neotyphodium coenophialum, and nutritional symbionts (arbuscular mycorrhizal fungi) belowground. In a greenhouse experiment, we manipulated the presence/ absence of both fungi and applied a fertilizer treatment to individual plants. Endophyte presence in host plants strongly reduced mycorrhizal colonization of roots. Additionally, for plants with the endophyte, the density of endophyte hyphae was negatively correlated with mycorrhizal colonization, suggesting a novel role for endophyte abundance in the interaction between the symbionts. Endophyte presence increased plant biomass, and there was a positive correlation between endophyte hyphal density and plant biomass. The effects of mutualists were asymmetric: mycorrhizal fungi treatments had no significant impact on the endophyte and negligible effects on plant biomass. Fertilization affected all

A neutral-niche theory of nestedness in mutualistic networks

Abstract: Recently, there has been a vigorous interest in community ecology about the structure of mutualistic networks and its importance for species persistence and coevolution. However, the mechanisms shaping mutualistic networks have been rarely explored. Here we extend for the first time the neutral theory of biodiversity to a multi trophic system. We focus on nestedness, a distinctive pattern of mutualistic community assembly showing two characteristics, namely, asymmetrical specialization (specialists interacting with generalists) and a generalist core (generalists interacting with generalists). We investigate the importance of relative species abundance (RSA) for the nested assembly of plant–animal mutualistic networks. Our results show that neutral mutualistic communities give rise to networks considerably more nested than real communities. RSA explains 60–70% of nested patterns in two real communities studied here, while 30–40% of nestedness is still unexplained. The nested pattern in real communities is better explained when we introduce interaction-specific species traits such as forbidden links and intensity of dependence (relative importance of fruits for the diet of a frugivore) in our analysis. The fact that neutral mutualistic communities exhibit a perfectly nested structure and do not show a random or compartmentalized structure, underlines the importance of RSA in the assembly of mutualistic networks.

Size does matter: variation in herbivory between and within plants and the plant vigor hypothesis

Abstract: The plant stress and plant vigor hypotheses (PVH) are two of the most widely recognized hypothesis invoked to explain differential distribution of insect herbivores among their host plants. In both cases, the emphasis is on bottomup processes (i.e. host-plant quality), but a recent meta-analytical review of the literature has shown that the plant stress hypothesis might have limited support among insect herbivores. In this study, we conducted a meta-analysis of the effects of plant vigor on insect herbivore abundance and survivorship by reviewing 71 published articles that explicitly tested the PVH and enabled 161 independent comparisons. Z-transform was used as the metric to standardize the results of all independent comparisons. Our quantitative results have shown that Hymenoptera (sawflies) was the most abundant group in the reviewed studies, representing 28.1% of the independent comparisons, followed by Diptera (25.1%) and Homoptera (22.6%). Amongst all the guilds studied, gall-formers were the most representative group (68.0%), whereas leaf-miners and stem-borers were underrepresented (less than 4.0% of the available comparisons). Insect herbivores were significantly more abundant on more vigorous plants (E 0.6432, CI0.75580.7280), but no significant effect was detected on herbivore survivorship. When herbivores were categorized into feeding guilds, effects of plant vigor on herbivore abundance were stronger for sap-suckers, leaf-miners and gall-formers. Our results have shown a strong herbivore preference for more vigorous plants, although our results do not support a preferenceperformance linkage.

Differences in aggression, activity and boldness between native and introduced populations of an invasive crayfish.

Abstract: Aggressiveness, along with foraging voracity and boldness, are key behavioral mechanisms underlying the competitive displacement and invasion success of exotic species However, do aggressiveness, voracity and boldness of the invader depend on the presence of an ecologically similar native competitor in the invaded community? We conducted four behavioral assays to compare aggression, foraging voracity, threat response and boldness to forage under predation risk of multiple populations of exotic signal crayfish Pacifastacus leniusculus across its native and invaded range with and without a native congener, the Shasta crayfish P fortis We predicted that signal crayfish from the invaded range and sympatric with a native congener (IRS) should be more aggressive to outcompete a close competitor than populations from the native range (NR) or invaded range and allopatric to a native congener (IRA) Furthermore, we predicted that IRS populations of signal crayfish should be more voracious, but less bold to forage under predation risk since native predators and prey likely possess appropriate behavioral responses to the invader Contrary to our predictions, results indicated that IRA signal crayfish were more aggressive towards conspecifics and more voracious and active foragers, yet also bolder to forage under predation risk in comparison to NR and IRS populations, which did not differ in behavior Higher aggression/voracity/boldness was positively correlated with prey consumption rates, and hence potential impacts on prey We suggest that the positive correlations between aggression/voracity/boldness are the result of an overall aggression syndrome Results of stream surveys indicated that IRA streams have significantly lower prey biomass than in IRS streams, which may drive invading signal crayfish to be more aggressive/voracious/bold to acquire resources to establish a population

Does Liebig's law of the minimum scale up from species to communities?

Abstract: Liebig's law of the minimum, which states that only one element limits the growth of organisms at any given time, is widely used in ecology. This principle is routinely applied to organisms, populations and communities, but can it really be applied indistinguishably across these different scales? Here we show, by prediction of a resource ratio conceptual model and with an experimental test carried out in microcosms with bacteria that, unlike single species, communities are likely to adjust their stoichiometry to that of their resources. This adjustment results from competitive exclusion and coexistence mechanisms, and is sensitive to the overall diversity of species in the community. It guaranties co-limitation, i.e. simultaneous limitation by multiple resources, at the community scale and optimal use of resources and maximization of community biomass for wide ranges of resource ratios. These results question the applicability of the Liebig's law of the minimum at the community level, and the relevance of ecosystem models relying on this principle.

Oak seedling survival and growth along resource gradients in Mediterranean forests: implications for regeneration in current and future environmental scenarios

Abstract: Understanding seedling performance across resource gradients is crucial for defining the regeneration niche of plant species under current environmental conditions and for predicting potential changes under a global change scenario. A 2-year field experiment was conducted to determine how seedling survival and growth of two evergreen and two deciduous Quercus species vary along gradients of light and soil properties in two Mediterranean forests with contrasting soils and climatic conditions. Half the seedlings were subjected to an irrigation treatment during the first year to quantify the effects on performance of an alteration in the summer drought intensity. Linear and non-linear models were parameterized and compared to identify major resources controlling seedling performance. We found both site-specific and general patterns of regeneration. Strong site-specificity was found in the identity of the best predictors of seedling survival: survival decreased linearly with increasing light (i.e. increasing desiccation risk) in the drier site, whereas it decreased logistically with increasing spring soil water content (i.e. increasing waterlogging risk) in the wetter site. We found strong empirical support for multiple resource limitation at the drier site, the response to light being modulated by the availability of soil resources (water and P). Evidence for regeneration niche partitioning among Quercus species was only found at the wetter site. However, at both sites Quercus species shared the same response to summer drought alleviation through water addition: increased first-year survival but not final survival (i.e. after two years). This suggests that extremely dry summers (i.e. the second summer in the experiment) can cancel out the positive effects of previous wetter summers. Therefore, an increase in the intensity and frequency of summer drought with climate change might cause a double negative impact on Quercus regeneration, due to a general reduction in survival probability and the annulment of the positive effects of (infrequent) ‘wet’ years. Overall, results presented in this study are a major step towards the development of a mechanistic model of Mediterranean forest dynamics that incorporates the idiosyncrasies and generalities of tree regeneration in these systems, and that allow simulation and prediction of the ecological consequences of resource level alterations due to global change.

Intraspecific competition drives multiple species resource polymorphism in fish communities

Abstract: It has been hypothesized that inter-specific competition will reduce species niche utilization and drive morphological evolution in character displacement. In the absence of a competitor, intra-specific competition may favor an expansion of the species niche and drive morphological evolution in character release. Despite of this theoretical framework, we sometimes find potential competitor species using the same niche range without any partitioning in niche. We used a database on test fishing in Sweden to evaluate the factors (inter- and intraspecific competition, predation, and abiotic factors) that could influence habitat choice of two competitor species. The pattern from the database shows that the occurrence of perch and roach occupying both littoral and pelagic habitats of lakes in Sweden is a general phenomenon. Furthermore, the results from the database suggest that this pattern is due to intra-specific competition rather than inter-specific competition or predation. In a field study, we estimated the morphological variation in perch and roach and found that, individuals of both species caught in the littoral zone were more deeper bodied compared to individuals caught in the pelagic zone. Pelagic perch fed more on zooplankton compared to littoral perch, independent of size, whereas the littoral perch had more macroinvertebrates and fish in their diet. Pelagic roach fed more on zooplankton compared to littoral roach, whereas littoral individuals fed more on plant material. Furthermore, we sampled littoral and pelagic fish from another lake to evaluate the generality of our first results and found the same habitat associated morphology in both perch and roach. The results show a consistent multi-species morphological separation in the littoral and pelagic habitats. This study suggests that intra-specific competition is possibly more important than inter-specific competition for the morphological pattern in the perch-roach system.

Junk-food in marine ecosystems

Abstract: The abundance and availability of food are critical determininants of reproductive success and population dynamics of marine top predators. However, recent work has indicated that the quality of the food may also be critically important for some marine predators. The ‘junkfood hypothesis’ was originally suggested as a potential explanation for a dramatic population decline of Stellers sea lions Eumetopias jubatus in the Gulf of Alaska. According to the hypothesis, a dietary switch to prey of low energy content led to detrimental effects on the population of sea lions. A number of observations indicate that the hypothesis is relevant for several population parameters. Recent work on piscivorous seabirds has provided substantial evidence indicating the relevance of this hypothesis in food webs in e.g. the North Pacific, the North Sea and the Baltic Sea. The emergence of ‘junk-food’ in these systems may be coupled to large scale changes in climatological and oceanographic forcing, although predation, fishing and competition provide additional plausible hypotheses. It may be possible to predict which kinds of animals will be particularly sensitive to food quality; these seem to be species with limited ability to carry food loads, with energetically-expensive foraging behaviour, and with digestive anatomy evolved to minimize mass at the cost of digestive efficiency. This review suggests that the junk-food hypothesis is a highly relevant factor in relation to sustaining ecosystem resilience, and is an important consideration in ecosystem management. Sustaining healthy populations of marine top-predators requires an understanding of the role of food quality, in addition to food abundance and availability.

Food-web connectance and predator interference dampen the paradox of enrichment

Abstract: Classic consumer-resource models with hyperbolic functional responses predict that enrichment increases the average biomasses of the species, but eventually leads to species' extinction due to accelerated oscillations ("paradox of enrichment"). However, empirical studies have stressed the complexity of natural food webs and the dominance of sigmoid or predator-interference functional responses, which may dampen population oscillations due to enrichment. Using analytical and numerical methods, we study enrichment effects on simple consumer-resource pairs and complex food webs with hyperbolic Holling type II (hereafter: type II), sigmoid Holling type III (hereafter: type III) and Beddington-De Angelis predator-interference functional responses (hereafter: BDA). Consumer-resource systems with a type III or BDA functional response are highly robust against accelerated oscillations due to enrichment, and the "paradox of enrichment" is resolved under certain parameter combinations. Subsequently, we simulated complex food webs with empirically-corroborated body-size structures of consumers that are ten times larger than their average resource. Our analyses demonstrate positive connectance-stability relationships with BDA or type III functional responses. Moreover, increasing connectance of these food webs also increases the robustness against enrichment in models with a BDA functional response. These results suggest that the well-known destabilising effects of connectance and enrichment found in classic models with type II functional responses may be inverted into stabilising effects in more realistic food-web models that are based on empirically-corroborated body-size structures and BDA or type III functional responses.

Does scale-dependent feedback explain spatial complexity in salt-marsh ecosystems?

Abstract: Complexity theory highlights scale-dependent feedback mechanisms as an explanation for regular spatial patterning in ecosystems. To what extent scale-dependent feedback clarifies spatial structure in more complex, non-regular systems remains unexplored so far. We report on a scale-dependent feedback process generating patchy landscapes at the interface of intertidal flats and salt marshes. Here, vegetation was characterized by Spartina anglica tussocks, surrounded by erosion gullies. To demonstrate the presence of a scale-dependent feedback, we determined if vegetation induced habitat modification resulted in local facilitation and large scale-inhibition of plant growth. Field surveys revealed that larger tussocks have deeper gullies, suggesting that gully erosion is caused by increased water flow around tussocks. This was confirmed by flume experiments, showing that feedback effects vary with current velocity and water depth. Transplantation of small Spartina units inside and just outside present tussocks revealed that the growth of Spartina transplants compared to transplant growth on bare sediment was higher within the raised Spartina tussocks, but lower in the gully just outside Spartina tussocks, providing clear evidence of scale-dependent feedback. Our results emphasize that scale-dependent feedback is a more general explanation for spatial complexity in ecosystems than previously considered.

Can a seed bank maintain the genetic variation in the above ground plant population

Abstract: There are indications that a persistent seed bank can protect small and isolated plant populations from local extinction. Genetic mechanisms contributing to this phenomenon are the increase of local effective population size - and hence the decrease of genetic drift - through a reservoir of persistent seeds, and the accumulation of intergenerational genetic diversity in the seed bank. To find evidence for these mechanisms, we conducted two formal meta-analyses. First, we analyzed 42 published habitat fragmentation studies and investigated whether the degree of genetic differentiation between fragmented plant populations was mediated by seed longevity. Second, we reviewed 13 published studies reporting the genetic diversity of both the seed bank and the above ground plants, aiming at comparing genetic diversity contained in the seed bank with the above ground vegetation. We conclude that a persistent seed bank may indeed mitigate the consequences of habitat fragmentation and protect a species from genetic drift and population genetic differentiation. We found no evidence, however, of high levels of genetic diversity accumulating in the soil seed bank. If genetic differences are present between the standing crop and the seed bank, they are very likely the result of local selection acting either directly or indirectly as a filter on the alleles present in the seed bank. We finally suggest that 1) the role of the seed bank should not be neglected in habitat fragmentation studies and 2) it is not very fruitful to continue comparing seed bank genetic diversity with above ground plant genetic diversity, unless this is performed under different selection regimes.

Towards a consensus for calculating dendrogram-based functional diversity indices

Abstract: The widely used FD index of functional diversity is based on the construction of a dendrogram. This index has been the subject of a strong debate concerning the choice of the distance and the clustering method to be used, since the method chosen may greatly affect the FD values obtained. Much of this debate has been centred around which method of dendrogram construction gives a faithful representation of species distribution in multidimensional functional trait space. From artificially generated datasets varying in species richness and correlations between traits, we test whether any single combination of clustering method(s) and distance consistently produces a dendrogram that most closely corresponds to the matrix of functional distances between pairs of species studied. We also test the ability of consensus trees, which incorporate features common to a range of different dendrograms, to summarize distance matrices. Our results show that no combination of clustering method(s) and distance constantly outperforms the others due to the complexity of interactions between correlations of traits, species richness, distance measures and clustering methods. Furthermore, the construction of a consensus tree from a range of dendrograms is often the best solution. Consequently, we recommend testing all combinations of distances and clustering methods (including consensus trees), then selecting the most reliable tree (with the lowest dissimilarity) to estimate FD value. Furthermore we suggest that any index that requires the construction of functional dendrograms potentially benefits from this new approach.

Reconciling neutral community models and environmental filtering: theory and an empirical test

Abstract: It is widely believed that the neutral theory of biodiversity cannot be used for parameter inference if the assumption of neutrality is not met. The goal of this work is to extend this neutral framework to quantify the intensity of recruitment limitation (limited dispersal plus environmental filtering) in natural species assemblages. We model several local communities as part of a larger metacommunity, and we assume that neutrality holds in each local community, but not in the metacommunity. The immigration rate m does not only reflect dispersal limitation into a given local community, but also the intensity of environmental filtering. We develop a novel statistical method to infer the immigration parameter m in each local community. Using simulated datasets, we show that m indeed depends on both dispersal limitation and on the intensity of environmental filtering. We then apply this method to a network of tropical tree plots in central Panama. Inferred recruitment rates m were positively correlated with the fraction of trees dispersed by mammals, and with annual rainfall, possibly due to a weaker environmental filtering as rainfall increases. Finally, m, as estimated from trees greater than 1 cm trunk diameter, were significantly larger than an estimation based on trees greater than 10 cm trunk diameter. This suggests a cumulative effect of environmental filtering upon trees throughout their ontogeny.

Trophic trickles and cascades in a complex food web: impacts of a keystone predator on stream community structure and ecosystem processes

Abstract: Chalk streams are among the most species-rich and productive of all temperate ecosystems Despite this, a few keystone species have the potential to exert disproportionately powerful effects on community structure and ecosystem processes Two of these are the bullhead Cottus gobio, a small benthic fish that is an extremely abundant, voracious predator, and the freshwater shrimp Gammaruspulex, which dominates the prey assemblage and is the principal detritivore Field experiments detected a bullhead–Gammarus–detritus trophic cascade, with detrital processing rates slowed dramatically in the presence of the predator In addition, survey data also revealed strong negative density-dependence between bullhead and brown trout, adding a further link in the cascade However, although bullhead also depressed the abundance of a dominant grazer, the snail Potamopyrgus antipodarum, there was no cascading effect upon algal production, suggesting that autochthonous inputs were not controlled by top–down effects This skewed effect of the predator upon autochthonous versus allochthonous basal resources stresses the need to consider both pathways of energy flux into the food web, whereas many previous studies have potentially overemphasized the importance of predator–herbivore–primary producer cascades The wider community food web contained 142 species and 1383 feeding links This complex network exhibited “small world” properties, such as high clustering (unlike many other food webs) and shortest path lengths between species were small (in common with many other food webs) In particular, each of the four members of the detrital cascade could be connected to any other species by three links or fewer Our data revealed that powerful cascading effects can be imbedded within even very complex ecological networks

Positive interactions among plant species for pollinator service: assessing the 'magnet species' concept with invasive species

Abstract: Plants with poorly attractive flowers or with little floral rewards may have inadequate pollinator service, which in turn reduces seed output. However, pollinator service of less attractive species could be enhanced when they are associated with species with highly attractive flowers (so called ‘magnet-species’). Although several studies have reported the magnet species effect, few of them have evaluated whether this positive interaction result in an enhancement of the seed output for the beneficiary species. Here, we compared pollinator visitation rates and seed output of the invasive annual species Carduus pycnocephalus when grow associated with shrubs of the invasive Lupinus arboreus and when grow alone, and hypothesized that L. arboreus acts as a magnet species for C. pycnocephalus. Results showed that C. pycnocephalus individuals associated with L. arboreus had higher pollinator visitation rates and higher seed output than individuals growing alone. The higher visitation rates of C. pycnocephalus associated to L. arboreus were maintained after accounting for flower density, which consistently supports our hypothesis on the magnet species effect of L. arboreus. Given that both species are invasives, the facilitated pollination and reproduction of C. pycnocephalus by L. arboreus could promote its naturalization in the community, suggesting a synergistic invasional process contributing to an ‘invasional meltdown’. The magnet effect of Lupinus on Carduus found in this study seems to be one the first examples of indirect facilitative interactions via increased pollination among invasive species.

Time – size tradeoffs: a phylogenetic comparative study of flowering time, plant height and seed mass in a north‐temperate flora

Abstract: Parents face a timing problem as to when they should begin devoting resources from their own growth and survival to mating and offspring development. Seed mass and number, as well as maternal survival via plant size, are dependent on time for development. The time available in the favorable season will also affect the size of the developing juveniles and their survival through the unfavorable season. Flowering time may thus represent the outcome of such a time partitioning problem. We analyzed correlations between flowering onset time, seed mass, and plant height in a north-temperate flora, using both cross-species comparisons and phylogenetic comparative methods. Among perennial herbs, flowering onset time was negatively correlated with seed mass (i.e. plants with larger seeds started flowering earlier) while flowering onset time was positively correlated with plant height. Neither of these correlations was found among woody plants. Among annual plants, flowering onset time was positively correlated with seed mass. Cross-species and phylogenetically informed analyses largely agreed, except that flowering onset time was also positively correlated with plant height among annuals in the cross-species analysis. The different signs of the correlations between flowering onset time and seed mass (compar. gee regression coefficient=−7.8) and flowering onset time and plant height (compar. gee regression coefficient=+30.5) for perennial herbs, indicate that the duration of the growth season may underlie a tradeoff between maternal size and offspring size in perennial herbs, and we discuss how the partitioning of the season between parents and offspring may explain the association between early flowering and larger seed mass among these plants.

First insights into specificity of belowground tritrophic interactions

Abstract: Tritrophic interactions involving plants, herbivores and parasites have been only recently documented for belowground systems, where entomopathogenic nematodes can exploit root herbivore induced volatile compounds to locate their hosts. Little is known, however, about whether the specificity of such interactions rivals that of the remarkable interactions found in aboveground studies. Using a belowground six-arm olfactometer that allows recording of nematode attraction, specificity of nine economically important species of different trophic levels, including plants, root feeders and entomopathogenic nematodes, was tested. We found that belowground tritrophic interactions are variable at the level of plant volatiles that are induced, elicitation by herbivores, as well as behavior of nematodes. We argue that studies on specificity and variability of belowground responses should be included in plant defense theories and in efforts to exploit tritrophic interactions to improve biological control practices.

Interactive effects of predators and a pesticide on aquatic communities

Abstract: Chemical contaminants are ubiquitous in nature and a major goal of ecologists has been to understand and predict their impacts on natural communities. While direct toxic effects can be garnered from single-species laboratory studies, the full suite of possible effects can only be observed when organisms are embedded within a community. In this study, we manipulated the concentration of malathion (a broad-spectrum insecticide) to determine the impacts on aquatic communities containing phytoplankton, periphyton, and 27 species of animals (16 zooplankton, 5 snails, 3 tadpoles and 3 predatory insects). Using relatively low concentrations (0.13 to 0.46 mg l -1 ), we found important direct (and interactive) effects of predators and malathion on the food web as well as a number of apparent density- and trait-mediated indirect effects. Malathion initiated an indirect effect by decreasing zooplankton diversity and abundance, which propagated an increase in phytoplankton, a decrease in periphyton, and a subsequent decrease in the growth of leopard frog tadpoles. There also was an apparent trait-mediated indirect effect whereby increased amounts of the pesticide reduced predation rates on amphibians without affecting the survival of the primary amphibian predator (larval Anax dragonflies). In contrast, snail survival and growth was unaffected by the pesticide but there were strong, species-specific effects from their primary predator (adult Belostoma water bugs). This is one of few studies to examine the impacts of malathion on aquatic communities across a range of concentrations, despite the fact that it is currently the most commonly applied insecticide in the United States, it is applied around the world, and it can be legally directly sprayed over aquatic habitats to control the mosquitoes that carry malaria and West Nile virus. Our results suggest several mechanisms by which a wide variety of pesticides with similar modes of action might impact aquatic communities.

Linking dendritic network structures to population demogenetics: The downside of connectivity

Abstract: Spatial structures strongly influence ecological processes. Connectivity is known to positively influence metapopulation demography and genetics by increasing the rescue effect and thus favoring individual and gene flow between populations. This result has not been tested in the special case of dendritic networks, which encompass stream and cave ecosystem for instance. We propose a first approach using an individual based model to explore the population demography and genetics in various dendritic networks. To do so, we first generate a large number of different networks, and we analyze the relationship between their hydrographical characteristics and connectivity. We show that connectivity mean and variance of connectivity are strongly correlated in dendritic networks. Connectivity segregates two types of networks: Hortonian and non-Hortonian networks. We then simulate the population dynamics for a simple life cycle in each of the generated networks, and we analyze persistence time as well as populations structure at quasi-stationary state. Our main results show that connectivity in dendritic networks can promote local extinction and genetic isolation by distance at low dispersal and diminish the size of the metapopulation at high dispersal. We discuss these unexpected findings in the light of connectivity spatial distribution in dendritic networks in the case of our model.