Multiple resource limitation: nonequilibrium coexistence of species in a competition model using a synthesizing unit

During the last two decades, the simple view of resource limitation by a single resource has been changed due to the realization that co-limitation by multiple resources is often an important determinant of species growth. Hence, the multiple resource limitation hypothesis needs to be taken into account, when communities of species competing for resources are considered. We present a multiple species–multiple resource competition model which is based on the concept of synthesizing unit to formulate the growth rates of species competing for interactive essential resources. Using this model, we demonstrate that a more mechanistic explanation of interactive effects of co-limitation may lead to the known complex dynamics including nonequilibrium states as oscillations and chaos. We compare our findings with earlier investigations on biological mechanisms that can predict the outcome of multispecies competition. Moreover, we show that this model yields a periodic state where more species than limiting complementary resources can coexist (supersaturation) in a homogeneous environment. We identify two novel mechanisms, how such a state can emerge: a transcritical bifurcation of a limit cycle and a transition from a heteroclinic cycle. Furthermore, we demonstrate the robustness of the phenomenon of supersaturation when the environmental conditions are varied.     

Seasonal variation in exploitative competition between honeybees and bumblebees

Honeybees (Apis mellifera) and bumblebees (Bombus spp.) often undergo exploitative competition for shared floral resources, which can alter their foraging behaviour and flower choice, even causing competitive exclusion. This may be strongest in summer, when foraging conditions are most challenging for bees, compared to other times of the year. However, the seasonal dynamics of competition between these major pollinator groups are not well understood. Here, we investigate whether the strength of exploitative competition for nectar between honeybees and bumblebees varies seasonally, and whether competitive pressure is greatest in summer months. We carried out experimental bee exclusion trials from May to late September, using experimental patches of lavender, variety Grosso, in full bloom. In each trial, we compared the numbers of honeybees (HB) foraging on patches from which bumblebees had been manually excluded (bumblebee excluded, BBE) versus control (CON) patches, HB(BBE-CON). This measure of exploitative competition varied significantly with season. As expected, mean HB(BBE-CON) was significantly greater in summer trials than in spring or autumn trials. This was despite high nectar standing crop volumes in BBE patch flowers in spring and autumn trials. Mean HB(BBE-CON) was not different between spring and autumn trials. Our results show that nectar competition between honeybees and bumblebees varies seasonally and is stronger in summer than spring or autumn, adding to current understanding of the seasonality of resource demand and competition between bee species. This information may also help to inform conservation programs aiming to increase floral resources for bees by showing when these resources are most needed.     

Nutrient limitation along a productivity gradient in wet meadows

Conservation management in meadows often focuses on reducing soil fertility and consequently community productivity as to promote and sustain species-rich vegetations. The productivity level to which nutrients are limiting growth is, however, unclear, as well as the relationship between productivity and the type of nutrient limitation. We carried out a fertilization experiment with N, P and K in six annually mown meadows along an aerial phytomass gradient (200–650 g m^–2). All meadows were found to be growth-limited by nutrients. Low-productive meadows were N-limited, or N+P co-limited, whereas our higher productive meadows were co-limited by a combination of N, P and/or K. The results from our experiments were compared with the results from 45 other fertilization experiments with N, P and K in grasslands and wetlands (aerial phytomass range 50–1500 g m^–2). Our results were consistent in nitrogen being the most frequent (co)-limiting nutrient, and regarding the equal frequence of occurrence of P (co)-limitation and K (co)-limitation (both in ca. 25–30% of all sites). Co-limitation occurred more often in our sites than in the other experiments. There was no clear relationship between aerial phytomass and type of nutrient limitation, except that K (co)-limitation only occurred at sites with phytomass above 200 g m^–2, and P (co)-limitation below 600 g m^–2. A comparison of productivity and nutrient concentrations in aerial phytomass among two years indicated that the type of nutrient limitation is not a static site characteristic but may vary with dynamic environmental conditions such as soil wetness; drought seems to enhance N-availability which may induce P- and K-limitation.     

The nutrient-load hypothesis: patterns of resource limitation and community structure driven by competition for nutrients and light

Resource competition theory predicts that the outcome of competition for two nutrients depends on the ratio at which these nutrients are supplied. Yet there is considerable debate whether nutrient ratios or absolute nutrient loads determine the species composition of phytoplankton and plant communities. Here we extend the classical resource competition model for two nutrients by including light as additional resource. Our results suggest the nutrient-load hypothesis, which predicts that nutrient ratios determine the species composition in oligotrophic environments, whereas nutrient loads are decisive in eutrophic environments. The underlying mechanism is that nutrient enrichment shifts the species interactions from competition for nutrients to competition for light, which favors the dominance of superior light competitors overshadowing all other species. Intermediate nutrient loads can generate high biodiversity through a fine-grained patchwork of two-species and three-species coexistence equilibria. Depending on the species traits, however, competition for nutrients and light may also produce multiple alternative stable states, suppressing the predictability of the species composition. The nutrient-load hypothesis offers a solution for several discrepancies between classical resource competition theory and field observations, explains why eutrophication often leads to diversity loss, and provides a simple conceptual framework for patterns of biodiversity and community structure observed in nature.     

The influence of 3-years of warming and N-deposition on ecosystem dynamics is small compared to past land use in subalpine meadows

Background and aims

Drivers of ecosystem dynamics that are under human influence range from local, land-management decisions to global processes such as warming temperatures and N deposition. The goal of this study was to understand how multiple, potentially interacting factors influence net primary production, N mineralization, and water and soil CO₂ fluxes.

Methods

Here I report on a three-year experiment that manipulated air temperature using ITEX passive warming cones and N deposition in a mountain meadow ecosystems that were historically grazed or protected from grazing.

Results

The strongest and most consistent effect was due to the legacy of grazing, with previously grazed sites having lower primary production, lower soil respiration rates, lower soil moisture, and lower soil C and N stocks than historically ungrazed sites. Warming increased soil respiration, but the effect was transient, and decreased over the 3-year study. Nitrogen addition increased primary production in the second and third year of the experiment but had no significant effect on soil respiration. The effect of historical grazing on primary production was approximately double the effect of N addition. Temperature and N deposition rarely interacted except for increasing N availability during the warm, wet growing season of 2004.

Conclusions

These findings indicate that the legacies of land use, with their influence on plant community composition and hydrologic processes, are locally more important than short-term step changes in temperature and nutrient availability.     

Influence of local and landscape factors on bumblebees in semi-natural meadows: a multiple-scale study in a forested landscape

Understanding the effects of local and landscape factors on bumblebees is relevant for the conservation of this group of pollinators. Bumblebees have been well-studied in agricultural landscapes of Western Europe, Asia and North America, but few studies have been developed on bumblebees in forest-dominated landscapes of Eastern Europe. We developed this study in 22 semi-natural meadows located in a patchy forested landscape of Estonia. We investigated the influence of habitat characteristics and landscape factors (calculated at four spatial scales: 250, 500, 1,000 and 2,000 m radius) on the total species richness and abundance of bumblebees. Correlation analysis, partial least squares (PLS) and stepwise forward-selection multiple regression analysis were applied in this study. The presence of a high diversity of flowering plants in semi-natural meadows may benefit the abundance of bumblebees. At the local level, patch area and shape seem to have positive and negative influences, respectively, on bumblebee species richness. At the landscape level, human settlements with the presence of gardens may favour bumblebee richness and abundance. Also, bumblebee species may increase with a high presence of meadows in the landscape, and may decrease with high percentages of forest and young forest. Overall, forested landscapes with a strong presence of edges and a diverse matrix may support a higher species richness and abundance of bumblebees. Both local and landscape factors should be considered when designing conservation strategies and agri-environmental measures.     

Variation in the phenology and abundance of flowering by native and exotic plants in subalpine meadows

The timing and abundance of flower production is important to the reproductive success of angiosperms as well as pollinators and floral and seed herbivores. Exotic plants often compete with native plants for space and limiting resources, potentially altering community floral dynamics. We used observations and a biomass-removal experiment to explore the effects of an invasive exotic flowering plant, Linaria vulgaris, on community and individual species flowering phenology and abundance in subalpine meadows in Colorado, USA. Invasion by L. vulgaris was associated with a shift in both the timing and abundance of community flowering. Invaded plant communities exhibited depressed flowering by 67% early in the season relative to uninvaded communities, but invaded sites produced 7.6 times more flowers than uninvaded sites once L. vulgaris began flowering. This increase in flowers at the end of the season was driven primarily by prolific flowering of L. vulgaris. We also found lower richness and evenness of resident flowering species in invaded plots during the period of L. vulgaris flowering. At the species level, a common native species (Potentilla pulcherrima) produced 71% fewer flowers in invaded relative to uninvaded plots, and the species had reduced duration of flowering in invaded relative to uninvaded sites. This result suggests that L. vulgaris does not simply alter the flowering of subordinate species but also the flowering of an individual common species in the plant community. We then used observational data to explore the relationship between L. vulgaris density and resident floral production but found only partial evidence that higher densities of L. vulgaris were associated with stronger effects on resident floral production. Taken together, results suggest that a dominant invasive plant can affect community and individual-species flowering.     

The effects of snowpack properties and plant strategies on litter decomposition during winter in subalpine meadows

Background and aims

Approximately 50 % of belowground organic carbon is present in the northern permafrost region and due to changes in climate there are concerns that this carbon will be rapidly released to the atmosphere. The release of carbon in arctic soils is thought to be intimately linked to the N cycle through the N cycle’s influence on microbial activity. The majority of new N input into arctic systems occurs through N₂-fixation; therefore, N₂-fixation may be the key driver of greenhouse gases from these ecosystems.

Methods

At Alexandra Fjord lowland, Ellesmere Island, Canada concurrent measurements of N₂-fixation, N mineralization and nitrification rates, dissolved organic soil N (DON) and C, inorganic soil N and surface greenhouse gas fluxes (CO₂, N₂O and CH₄) were taken in two ecosystem types (Wet Sedge Meadow and Dryas Heath) over the 2009 growing season (June-August). Using Structural Equation Modelling we evaluated the hypothesis that CO₂, CH₄ and N₂O flux are linked to N₂-fixation via the N cycle.

Results

The soil N cycle was linked to CO₂ flux in the Dryas Heath ecosystem via DON concentrations, but there was no link between the soil N cycle and CO₂ flux in the Wet Sedge Meadow. Methane flux was also not linked to the soil N cycle, nor surface soil temperature or moisture in either ecosystem. The soil N cycle was closely linked to N₂O emissions but via nitrification in the Wet Sedge Meadow and inorganic N in the Dryas Heath, indicating the important role of nitrification in net N₂O flux from arctic ecosystems.

Conclusions

Our results should be interpreted with caution given the high variability in both the rates of the N cycling processes and greenhouse gas flux found in both ecosystems over the growing season. However, while N₂-fixation and other N cycling processes may play a more limited role in instantaneous CO₂ emissions, these processes clearly play an important role in controlling N₂O emissions.     

Pollen and resource limitation in a gynodioecious species

Differences between plant sex morphs in pollen or resource availability may affect their relative fitness and thereby the sex ratio of dimorphic species. In gynodioecious species, in which hermaphroditic and female plants coexist, a variety of factors (e.g., hermaphrodite self-fertility or rarity or pollinator discrimination against females) might be expected to lead to stronger pollen limitation in females than in hermaphrodites. On the other hand, females have been found to be superior compared to hermaphrodites in low-nutrient conditions. The effects of supplemental hand-pollination and resource addition on the reproductive output of the self-fertile gynodioecious perennial Geranium sylvaticum (Geraniaceae) were tested for several populations that differ in their female frequency (4.4-23.0%). Both pollen and resource availability limited fruit set and the number of seeds produced per plant; however, seed set (i.e., the number of seeds produced per fruit) was limited only by resources. Because pollen limitation in females did not correlate with female frequency, our results suggest that pollen limitation in females does not depend on the frequency of the pollen-producing hermaphrodites. Furthermore, because pollen and resource availability limited reproductive output of both sex morphs, these factors may not contribute significantly to maintenance and evolution of gynodioecy in G. sylvaticum.     

Seasonal dynamics of denitrification activity in two water meadows

Seasonal variation in denitrification activity was measured in twoflooded water meadows, one on peaty and one on sandy soil, over a three-yearperiod. Measurements were taken during flooded and drained periods, usingthe acetylene-blockage technique, and the rates were compared to massbalance estimates of nitrate removal in the percolating water.Denitrification activity was higher in sandy soil than in peaty soil. Higherwater infiltration rate and thereby higher nitrate load was considered to bethe cause of the higher denitrification in the sandy soil. Floodingsignificantly increased denitrification, and the rates were higher in autumnand winter than in spring. This was considered to be a result of highernitrogen concentration in inflowing stream water during winter. Annualdenitrification was estimated to 430–460 kg N ha^-1yr^-1 in the sandy soil meadow, and 220 kg N ha^-1yr^-1 in the peaty soil meadow. In the sandy soil there was alarge discrepancy between nitrate removal rates and denitrification rates,which can be explained by nitrification of ammonium released from the soil.In the peaty soil nitrate disappearance and denitrification correspondedfairly well.     

旅游对北京东灵山亚高山草甸物种多样性影响的初步研究

在1965、1983、1994和2000年4个时期调查数据的基础上,对北京东灵山区亚高山草甸物种多样性变化和生活型谱进行了对比研究,结果显示,近年来东灵山亚高山草甸的物种多样性呈下降趋势,尤其是1983～1994年的下降速度明显加快,利用重要值作为权重参数对不同时期的东灵山亚高山草甸生活型谱进行了比较,所得到的结果反映了20世纪80年代以来,伴随旅游发展而迅速增加的马匹散牧已经对亚高山草甸的群落外貌造成了严重的不良影响,群落高度降低,物种组成改变及植被盖度下降等,说明过度放牧是东灵山亚高山草甸退化的重要影响因素,根据东灵山风景区亚高山草甸目前存在的问题,我们建议当地有关部门科学地确定东灵山风景区旅游容量,首先要严格限制风景区马匹的散牧,关闭风景区内的跑马场和牧马帐蓬,并通过加强宣传、管理和景区的基本建设,充分发挥生态系统的服务功能,实现在保护生态环境的前提下实现资源的持续利用。     

Gamete evolution and sperm numbers: sperm competition versus sperm limitation

Both gamete competition and gamete limitation can generate anisogamy from ancestral isogamy, and both sperm competition (SC) and sperm limitation (SL) can increase sperm numbers. Here, we compare the marginal benefits due to these two components at any given population level of sperm production using the risk and intensity models in sperm economics. We show quite generally for the intensity model (where N males compete for each set of eggs) that however severe the degree of SL, if there is at least one competitor for fertilization (N − 1 ≥ 1), the marginal gains through SC exceed those for SL, provided that the relationship between the probability of fertilization (F) and increasing sperm numbers (x) is a concave function. In the risk model, as fertility F increases from 0 to 1.0, the threshold SC risk (the probability q that two males compete for fertilization) for SC to be the dominant force drops from 1.0 to 0. The gamete competition and gamete limitation theories for the evolution of anisogamy rely on very similar considerations: our results imply that gamete limitation could dominate only if ancestral reproduction took place in highly isolated, small spawning groups.     

Information use and resource competition: an integrative framework

Organisms may reduce uncertainty regarding how best to exploit their environment by collecting information about resource distribution. We develop a model to demonstrate how competition can facilitate or constrain an individual''s ability to use information when acquiring resources. As resource distribution underpins both selection on information use and the strength and nature of competition between individuals, we demonstrate interdependencies between the two that should be common in nature. Individuals in our model can search for resources either personally or by using social information. We explore selection on social information use across a comprehensive range of ecological conditions, generalizing the producer–scrounger framework to a wide diversity of taxa and resources. We show that resource ecology—defined by scarcity, depletion rate and monopolizability—determines patterns of individual differences in social information use. These differences suggest coevolutionary processes linking dominance systems and social information use, with implications for the evolutionary demography of populations.     

Egg limitation in host-parasitoid dynamics: an individual-based perspective

Theoretical models of parasitoid-host dynamics predict that egg limitation in parasitoids destabilizes community dynamics. However, although egg limitation is experienced by individual parasitoids with variable success of encountering hosts, such details were neglected in previous models. This study developed an individual-based parasitoid-host model that explicitly incorporates egg limitation and host encounters of individual parasitoids. The model indicates that the combination of egg limitation and variation in the success of encountering hosts stabilizes parasitoid-host dynamics. The stabilizing mechanism emerges from Jensen’s inequality because egg limitation makes the number of offspring inherently concave down in the number of encountered hosts. Reasons for the inconsistent predictions of the effect of egg limitation between the current model and previous models are discussed.     

The allelopathic effects of Festuca paniculata depend on competition in subalpine grasslands

Allelopathy is recognized as an important process in plant–plant interactions, but how it affects plant communities growing in competitive conditions has not been assessed. This article investigates whether the allelopathic effect of Festuca paniculata is modified by competition between target plants in subalpine grasslands. We hypothesized that plants growing in mixed stands will be more affected by allelochemicals than the same species in monoculture. At Lautaret pass (Northern French Alps), a pot experiment was designed. We used leachates from donor pots (Treatments: 1. Bare soil, 2. F. paniculata clipped, and 3. F. paniculata unclipped) to water target pots (Treatments: 1. Control (soil only), 2. Dactylis glomerata, 3. Agrostis capillaris, and 4. D. glomerata and A. capillaris). Target plants were cultivated during one growing season. The effects of leachates from donor pots and interspecific competition in target pots were evaluated by measuring the final biomass of plants. Soil fertility was controlled in all target pots by measuring NO₃ ^?, NH₄ ⁺, N, and C % of the soil. Effect of target treatment under bare soil : Both D. glomerata and A. capillaris grew better in monocultures than in mixture. Effect of donor treatment on monocultures : Under bare soil, D. glomerata grew better than under F. paniculata leachates. By contrast, A. capillaris did not respond to donor pot treatment. Effect of donor treatment on mixtures: However, when both species were cultivated together under F. paniculata leachates, the biomass of D. glomerata was similar to that in monoculture under bare soil. Differences in sensitivity to allelopathy reversed the impact of interspecific competition: A. capillaris facilitated D. glomerata under allelopathy, which made allelopathy of F. paniculata on D. glomerata inefficient. The complexity of overlapping mechanisms of plant–plant interactions are highlighted by this semi-natural experiment. In subalpine grasslands, allelopathy not only limits the growth of neighboring plants, but it may also modify community assembly by affecting other plant–plant interactions such as competition. This study contributes to explore the way allelopathy interacts with other plant–plant interactions in natural systems.     

Effects of weather on foraging-flights of bumblebees (Hymenoptera, Apidae) in a subalpine/alpine area

The foraging activity of bumblebees in a subalpine/alpine area in northern Sweden was related to weather factors (temperature, light, wind and rain) by counting the number of specimens visiting a square (5 × 5 m) with a high density of forage plants. The combined effect of light and temperature was the most important factor regulating the flight activity. The condition of the arctic light-climate allows bumblebees to fly even at the less light hours of the day. No clear differences according to flight activity were observed between the subalpine/alpine species Bombus balteatus Dahlbom, B. lapponicus (Fabricius) and the species B. jonellus (Kirby), B. pascuorum (Scopoli), B. pratorum (L.), observed in the subalpine habitats. A comparison between the flight activity of queens, workers and males shows that queens are able to fly at the lowest temperatures followed by males and workers at slightly higher temperatures. The relevance of the overlap of the body sizes between the casts is discussed.     

Evidence for competition between honeybees and bumblebees; effects on bumblebee worker size

Numerous studies suggest that honeybees may compete with native pollinators where introduced as non-native insects. Here we examine evidence for competition between honeybees and four bumblebee species in Scotland, a region that may be within the natural range of honeybees, but where domestication greatly increases the honeybee population. We examined mean thorax widths (a reliable measure of body size) of workers of Bombus pascuorum, B. lucorum, B. lapidarius and B. terrestris at sites with and without honeybees. Workers of all four species were significantly smaller in areas with honeybees. We suggest that reduced worker size is likely to have implications for bumblebee colony success. These results imply that, for conservation purposes, some restrictions should be considered with regard to placing honeybee hives in or near areas where populations of rare bumblebee species persist.