Persistence in food webs—I Lotka-Volterra food chains

Persistence-extinction in simple food chains modelled by Lotka-Volterra dynamics is governed by a single parameter which depends upon the interspecific interaction coefficients, the intraspecific interaction coefficients, and the length of the food chain. In persistent systems with nonzero carrying capacity, two new features predominate. Trophic level influence factors relate persistence on different trophic levels and determine, in conjunction with the persistence parameter, the magnitude of persistence. Equilibrium component ordering, which results in persistent systems, mandates once again that systems need to be studied on the complete ecosystem level; static field measurements reflect species location in the food chain, the total length of the food chain and assume characteristics according to these factors.     

Has the invasive round goby caused new links in Baltic food webs?

The Ponto-Caspian round goby (Neogobius melanostomus, Pallas 1814) most probably was established in the Gulf of Gdańsk, Baltic Sea, in the late 1980’s and has since become one of the dominant species in the region. In this study we assess the role of round gobies as prey for two important fish species in the Gulf of Gdańsk, cod (Gadus morhua) and perch (Perca fluviatilis). We compared their present diet with stomach analyses from the area prior the round goby establishment, as well as with diet analysis from Baltic regions where round gobies are absent. There were large differences in the diet between cods from the Gulf of Gdańsk 2003–2006 compared to cods in earlier studies (1977–1981) from the Southern Baltic Sea. There were also large differences in cod and perch diets from areas with and without round goby. Presently, round goby constitutes the most important prey for medium sized cods in Gulf of Gdańsk, and perch from the same area almost exclusively feed on gobiids. Stomach analysis, trophic level estimates, and stable isotope analyses all indicated that cod and perch in Gulf of Gdańsk after the round goby establishment belonged to a similar trophic level. Beside round goby, no mussel feeding fish contributed much to the diet of cod or at all to the diet of perch. Thus, it is likely that round gobies constitute a new energetic pathway from mussels to top predators. However, due to the short time elapsed after round goby establishment, we can only speculate on the species future impacts on Baltic food webs.     

Part–whole relations between food webs and the validity of local food-web descriptions

This work analyzes the relationship between large food webs describing potential feeding relations between species and smaller sub-webs thereof describing relations actually realized in local communities of various sizes. Special attention is given to the relationships between patterns of phylogenetic correlations encountered in large webs and sub-webs. Based on the current theory of food-web topology as implemented in the matching model, it is shown that food webs are scale invariant in the following sense: given a large web described by the model, a smaller, randomly sampled sub-web thereof is described by the model as well. A stochastic analysis of model steady states reveals that such a change in scale goes along with a re-normalization of model parameters. Explicit formulae for the re-normalized parameters are derived. Thus, the topology of food webs at all scales follows the same patterns, and these can be revealed by data and models referring to the local scale alone. As a by-product of the theory, a fast algorithm is derived which yields sample food webs from the exact steady state of the matching model for a high-dimensional trophic niche space in finite time.     

Predicting abundance–body size relationships in functional and taxonomic subsets of food webs

Abundance–body size relationships are widely observed macroecological patterns in complete food webs and in taxonomically or functionally defined subsets of those webs. Observed abundance–body size relationships have frequently been compared with predictions based on the energetic equivalence hypothesis and, more recently, with predictions based on energy availability to different body size classes. Here, we consider the ways in which working with taxonomically or functionally defined subsets of food webs affected the relationship between the predicted and observed scaling of biomass and body mass in sediment dwelling benthic invertebrate communities at three sites in the North Sea. At each site, the energy available to body size classes in the “whole” community (community defined as all animals of 0.03125–32.0 g shell-free wet weight) and in three subsets was predicted from estimates of trophic level based on nitrogen stable isotope analysis. The observed and predicted scalings of biomass and body size were not significantly different for the whole community, and reflected an increase in energy availability with body size. However, the results for subsets showed that energy availability could increase or decrease with body size, and that individuals in the subsets were likely to be competing with individuals outside the subsets for energy. We conclude that the study of abundance–body mass relationships in functionally or taxonomically defined subsets of food webs is unlikely to provide an adequate test of the energetic equivalence hypothesis or other relationships between energy availability and scaling. To consistently and reliably interpret the results of these tests, it is necessary to know about energy availability as a function of body size both within and outside the subset considered.     

Evolution of body size in food webs: does the energetic equivalence rule hold?

The energetic equivalence rule (EER), which is derived from empirical observations linking population density and body size and from the allometric law linking metabolism and body size, predicts that the amount of energy used by the various species should be independent of body size. Here, we examine this hypothesis using a model that allows entire food webs to emerge from coevolution of interacting species. Body size influences both individual metabolism and interactions among species in the model. Overall, population density does decrease with body size roughly following a power law whose exponent is variable. We discuss this variability in the light of empirical data sets. The emerging relationship between the flux of resources exploited by the various species and their body size follows a decreasing power law, thus contradicting the EER. Our model emphasizes the importance of considering the influence of body size on species interactions in attempting to explain large-scale patterns related to body size.     

Patterns in benthic food webs: a role for omnivorous crayfish?

1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.     

Bt-maize in neotropical arthropod food webs: community-stress or lack thereof?

Although the agricultural use of genetically engineered (GE) plants has been employed extensively, its adoption is still controversial and its impact on arthropods has rarely been scrutinized at the community level. If this technology is aimed at a drastic reduction of a key community component, significant community-level impact is expected and needs to be assessed. Thus, food web analysis was used to assess the short-term impact of genetically modified maize plants, Zea mays L. (Poaceae), expressing insecticidal proteins of the bacterium Bacillus thuringiensis Berliner (i.e., Bt maize), on the associated arthropod assemblage in a neotropical scenario. Arthropods associated with winter and summer cultivations with Bt and non-Bt (isoline) maize were thus sampled using sweep nets and whole-plant collections throughout the plant phenological cycle. The collected information was used to build a plant-consumer-predator trivariate network based on data of individual arthropod body mass, numerical abundance, and biomass abundance using food web analysis. Eighty-five arthropod species were sampled, and whereas cultivation season significantly affected arthropod species richness and abundance, only marginal differences existed between maize hybrids (Bt vs. non-Bt). The recognized food webs also indicated significant differences between seasons. In contrast, Bt-maize-hosted food webs were similar to those of the non-Bt isoline indicating no significant impact on arthropod food webs. Nonetheless, Bt maize did not provide significant control of the target pest species, the fall armyworm, nor did it lead to higher crop yield, raising questions about its current usefulness in the region.     

Diversity–productivity–stability relationship in freshwater ecosystems: Whole-systemic view of all trophic levels

The idea of the diversity–productivity relationship in ecology was originated by Charles Darwin more than 100years ago and was highlighted by McNaughton in 1993. There have been extensive studies on this relationship, mostly conducted in grasslands. Many studies assert that a high level of biodiversity leads to high primary productivity, in accord with Darwin. However, these studies have mostly been done in grasslands (primary producer) and do not take into account effects from other trophic levels. The possibility remains that the positive diversity–production relationship does not hold in other types of ecological systems and that multitrophic effects influence and modify the relationship. In the present study, the diversity–productivity relationship is examined for freshwater foodwebs, including whole trophic compartments, by using indices of diversity and productivity more suitable for a whole systemic study on ecological systems. It seems that there are two trends in the relationship: productivity increases with the increase of diversity in ecosystems when diversity is small, and decreases with diversity in ecosystems when diversity is large. The first trend may be due to nutrient-limiting characteristics and the second trend may be due to photosynthetically active radiation–limiting characteristics. Also, both higher trophic and whole-systemic productivities are found to increase relative to primary productivity with the increase of diversity. Stability indices introduced previously, combined with the present results, reveal a positive correlation between relative productivity and stability in freshwater foodwebs, including all trophic compartments.     

Edge effects on insect–plant food webs: assessing the influence of geographical orientation and microclimatic conditions

1. Geographical orientation affects the magnitude of microclimatic edge effects, mainly by altering the exposure to solar radiation. There is considerable evidence available concerning the influence of microclimatic edge effects on organisms at the population level, although little is known about effects at the community level, and variations with edge geographical orientation have barely been addressed. The present study investigated the influence of microclimatic conditions at the forest interior and edges with different geographical orientation on plant–leafminer–parasitoid food webs in the fragmented Chaco Serrano forest located in Central Argentina. 2. Microclimatic variables were measured in three locations: interior, north, and south-facing edges of six forest fragments. In each location, plant resource availability was assessed and mined leaves were collected to investigate plant–leafminer–parasitoid communities by rearing leafminer and parasitoid adults. 3. The results obtained revealed changes in the specific composition of plant, leafminer and parasitoid communities among locations, although they failed to detect location influences on most variables linked to abundance (except for leafminers), richness or food web structure. Leafminer abundance was negatively related to temperature, whereas their richness was related, also negatively, to moisture. Instead, parasitoids were favoured by higher temperatures and light levels, stronger winds, and lower humidity. 4. The results of the present study indicate that the effects of microclimatic conditions may depend on the trophic level considered. Moreover, the findings suggest that possible trade-offs between direct and indirect effects (such as plant quality, floral resources, etc.) of microclimatic conditions could influence plant and insect responses to location.     

Environmentally-friendly and organic management practices enable complementary diversification of plant–bumblebee food webs

Plant and pollinator diversity have declined concurrently in Europe in the last half century. We studied plant–bumblebee food webs to understand the effects of two agri-environmental schemes (AES, organic farming and environmentally-friendly management practice) vs. conventional farming as control group, landscape structure (heterogeneous vs. homogeneous landscapes) and seasonality (June, July, and August) interactions using Estonian AES monitoring data. In the summer of 2014, we observed foraging bumblebees (20 species) on 64 farms that varied in agricultural management and landscape structure, yielding a total of 2303 flower visits on 76 plant species. We found that both management practice and landscape structure influenced the generality (redundancy in the use of flower resources) of food webs. In homogeneous landscapes, environmentally-friendly management practices, including restrictions on the application of glyphosates, enhancement of bumblebee habitats, such as permanent grassland field margins, the allocation of a minimum of 15% of arable land (including rotational grasslands) to legumes, contributed to a higher number of visited plant species (generality) in July, whereas organic farming did so in August. Therefore, both environmentally-friendly and organic management practices are needed to support plant–bumblebee food webs in agricultural landscapes. Food web generality and diversity (Shannon index) are affected by a significant interaction between landscape structure and seasonality: food web diversity varied in homogeneous landscapes between the three different survey months, whereas food webs were more diverse in heterogeneous landscapes. We did not find any significant interaction effect of management, landscape structure and seasonality on linkage density and vulnerability. A full list of the most visited plant species by bumblebees based on species-specific flower visitation was also assembled. In homogeneous landscapes, resource limitation is an issue for bumblebees in certain time periods. For supporting bumblebees in the agricultural landscapes, avoiding resource limitation is important and this can be secured with a combination of AES management practices.     

Do fall additions of salmon carcasses benefit food webs in experimental streams?

Research showing that salmon carcasses support the productivity and biodiversity of aquatic and riparian ecosystems has been conducted over a variety of spatial and temporal scales. In some studies, carcasses were manipulated in a single pulse or loading rate or manipulations occurred during summer and early fall, rather than simulating the natural dynamic of an extended spawning period, a gradient of loading rates, or testing carcass effects in late fall-early winter when some salmon stocks in the US Pacific Northwest spawn. To address these discrepancies, we manipulated salmon carcass biomass in 16 experimental channels located in the sunlit floodplain of the Cedar River, WA, USA between mid-September and mid-December, 2006. Total carcass loads ranged from 0?C4.0 kg/m² (0, 0.001, 0.01, 0.1, 0.5, 1.0, 2.0 and 4.0 kg/m², n = 2 per treatment) and were added to mimic the temporal dynamic of an extended spawning period. We found little evidence that carcasses influenced primary producer biomass or fish growth; however, nutrients and some primary consumer populations increased with loading rate. These effects varied through time, however. We hypothesize that the variable effects of carcasses were a result of ambient abiotic condition, such as light, temperature and disturbance that constrained trophic response. There was some evidence to suggest peak responses for primary producers and consumers occurred at a loading rate of ~1.0?C2.0 kg/m², which was similar to other experimental studies conducted during summer.     

Effects of global environmental changes on parasitoid–host food webs and biological control

Global environmental changes threaten biodiversity and the interactions between species, and food-web approaches are being used increasingly to measure their community-wide impacts. Here we review how parasitoid–host food webs affect biological control, and how their structure responds to environmental change. We find that land-use intensification tends to produce webs with low complexity and uneven interaction strengths. Dispersal, spatial arrangement of habitats, the species pool and community differences across habitats have all been found to determine how webs respond to landscape structure, though clear effects of landscape complexity on web structure remain elusive. The invasibility of web structures and response of food webs to invasion have been the subject of theoretical and empirical work respectively, and nutrient enrichment has been widely studied in the food-web literature, potentially driving dynamic instability and altering biomass ratios of different trophic levels. Combined with food-web changes observed under climate change, these responses of food webs could signal changes to biological control, though there have been surprisingly few studies linking food-web structure to pest control, and these have produced mixed results. However, there is strong potential for food-web approaches to add value to biological control research, as parasitoid–host webs have been used to predict indirect effects among hosts that share enemies, to study non-target effects of biological control agents and to quantify the use of alternative prey resources by enemies. Future work is needed to link food-web interactions with evolutionary responses to the environment and predator–prey interactions, while incorporating recent advances in predator biodiversity research. This holistic understanding of agroecosystem responses and functioning, made possible by food-web approaches, may hold the key to better management of biological control in changing environments.     

Revisiting Beverton–Holt recruitment in the presence of variation in food availability

Understanding density-dependent changes in juvenile survival and growth rates is of great importance because these rates determine recovery rates for imperiled populations and/or sustainable harvest rates. Unfortunately, the mechanisms leading to density dependent survival and growth are among the least understood process in biology and fisheries. Previous work has shown that small fish may vary foraging times to achieve a target growth rate, resulting in the well-known Beverton–Holt recruitment function with variation in food availability affected the initial slope of the recruitment curve. We amend their derivation to show that incorporating fish growth under a variety of evolutionary strategies for balancing foraging time and predation risk still leads to recruitment approximately as expected under the Beverton–Holt recruitment model but that changing food availability affects both the initial slope and maximum recruitment level. We demonstrate that when food availability is known to vary over time, these models often result in a more parsimonious alternative than the standard Beverton–Holt function. Further, Beverton–Holt recruitment is expected when foraging times are adjusted to balance fitness gains from growth against mortality risk. Finally, linking recruitment success to food availability warns that species with high scope for density dependent survival (high compensation ratio or steepness) may be extremely sensitive to changes in available food densities. This work emphasizes the sensitivity of stock-recruitment parameters to food availability and strongly suggests a need to carefully monitor lower trophic levels to better understand and predict dramatic changes in juvenile recruitment and carrying capacity.     

Dynamics of planktonic food webs in three mining lakes across a pH gradient (pH 2–4)

Acidic mining lakes (ML) in Lusatia (Germany) are characterised by their geogenically determined chemistry. The present study describes the structure, main components and relationships within the food webs of three acidic mining lakes with different pH values (ML 111: pH 2.6; ML 117: pH 2.8; ML Felix: pH 3.6) in order to show their typical characteristics. The investigation covered the period 1995–1997. The number of species and the biomass are both low, but increase with increasing pH. Planktonic components in the most acidic ML 111 (pH 2.6–2.9) comprise bacteria, Ochromonas spp. and Chlamydomonas spp. and a few rotifers (E. worallii, C. hoodi). Heliozoans are the top-predators. In ML 117 (pH 2.8–3) Gymnodinium sp., ciliates, the rotifer B. sericus and the pioneer crustacean Chydorus sphaericus join the pelagial community. Heliozoans were not found in ML 117 or ML Felix (pH 3.4–3.8). ML Felix had the most taxa. The benthic food chain of all three lakes includes phytobenthic algae as producers, chironomids as primary consumers and corixids as top predators in the profundal. Corixids predate on small cladocerans inhabiting the pelagial in lakes with a pH above 2.8 such as ML Felix. They invade the pelagial and act as a connecting link between the benthic and the pelagic food chains, which are isolated in lakes with a lower pH. Occasionally primary producers and consumers were abundant in all three lakes. These organisms do not depend on the degree of acidity, but on the availability of essential ressources. Mass variations covered up any seasonal variation in the extremely acidic ML 111 (0.9 mm³ l^–1), while in the other two lakes seasonal patterns of biomass were found.     

The ecological potentials of Phytomyxea (“plasmodiophorids”) in aquatic food webs

The Phytomyxea (“plasmodiophorids”) including both Plasmodiophorida and Phagomyxida is a monophyletic group of Eukaryotes composed of obligate biotrophic parasites of green plants, brown algae, diatoms and stramenopiles commonly found in many freshwater, soil and marine environments. However, most research on Phytomyxea has been restricted to plant pathogenic species with agricultural importance, thereby missing the huge ecological potential of this enigmatic group of parasites. Members of the Phytomyxea can induce changes in biomass in their hosts (e.g. hypertrophies of the host tissue) under suitable environmental conditions. Upon infection they alter the metabolism of their hosts, consequently changing the metabolic status of their host. This results in an altered chemical composition of the host tissue, which impacts the diversity of species which feed on the tissues of the infected host and on the zoospores produced by the parasites. Furthermore, significant amounts of nutrients derived from the hosts, both primary producers (plants and algae) and primary consumers (litter decomposers and plant parasites [Oomycetes]), can enter the food web at different trophic levels in form of zoospores and resting spores. Large numbers of zoospores and resting spores are produced which can be eaten by secondary and tertiary consumers, such as grazing zooplankton and metazoan filter-feeders. Therefore, these microbes can act as energy-rich nutrient resources which may significantly alter the trophic relationships in fresh water, soil and marine habitats. Based on the presented data, Phytomyxea can significantly contribute to the complexity and energy transfer within food webs.     

Amino acid–dependent stability of the acyl linkage in aminoacyl-tRNA

Aminoacyl-tRNAs are the biologically active substrates for peptide bond formation in protein synthesis. The stability of the acyl linkage in each aminoacyl-tRNA, formed through an ester bond that connects the amino acid carboxyl group with the tRNA terminal 3′-OH group, is thus important. While the ester linkage is the same for all aminoacyl-tRNAs, the stability of each is not well characterized, thus limiting insight into the fundamental process of peptide bond formation. Here, we show, by analysis of the half-lives of 12 of the 22 natural aminoacyl-tRNAs used in peptide bond formation, that the stability of the acyl linkage is effectively determined only by the chemical nature of the amino acid side chain. Even the chirality of the side chain exhibits little influence. Proline confers the lowest stability to the linkage, while isoleucine and valine confer the highest, whereas the nucleotide sequence in the tRNA provides negligible contribution to the stability. We find that, among the variables tested, the protein translation factor EF-Tu is the only one that can protect a weak acyl linkage from hydrolysis. These results suggest that each amino acid plays an active role in determining its own stability in the acyl linkage to tRNA, but that EF-Tu overrides this individuality and protects the acyl linkage stability for protein synthesis on the ribosome.     

Are simple environmental indicators of food web dynamics reliable: Exploring the kittiwake–temperature relationship

This study examined the inter-relationships between kittiwake breeding success and sandeel abundance, Calanus copepods, chlorophyll and temperature. The validity of past proxies of sandeel prey (Calanus and temperature) suggested in the literature was also examined. Winter temperature was not found to be a reliable indicator of the abundance of sandeel or lower trophic levels, although the present study did support past evidence for a dependence of kittiwake breeding success on local sandeel abundance as, with the exception of two years, there was a linear relationship with log age-0 sandeel abundance. The abundance of sandeel was positively linked to the dates of predicted peak egg abundance in C. finmarchicus and C. helgolandicus. This supported earlier evidence that the match with prey timing is important to sandeel recruitment. Neither Calanus species was associated with temperature in the February to March period nor in the April–May period, which may explain the lack of a temperature effect that propagates through the trophic levels. Consequently, although kittiwake breeding success appears to show some sensitivity and specificity to changes in their prey, this responsiveness was not found to extend to the lowest trophic level (Calanus abundance) or to temperature. As such kittiwake breeding success was not a reliable indicator of climate-driven changes in the local food web.