Impacts of fish adaptive migration on the dynamic stability and ecological functioning of the food web in the Beibu Gulf, China

Summary Food webs and trophic dynamics of coastal systems have been the focus of intense research throughout the world, as they prove to be critical in understanding ecosystem processes and functions. However, very few studies have undertaken a quantitative comparison of entire food webs from a key consumer perspective across a broad geographical area, limiting relevant comparisons among systems with distinct biotic and abiotic components. We investigate the structure and functioning of food webs in four tidal ecosystems of international importance for migratory shorebirds along the East Atlantic Flyway: Tejo estuary in Portugal, Sidi Moussa in Morocco, Banc d'Arguin in Mauritania and Bijagós archipelago in Guinea‐Bissau. Basal food sources, shorebirds and their prey (benthic invertebrates) were sampled in all areas, and Bayesian stable isotope mixing models and community‐wide metrics were used in a comparative analysis among areas. Significant differences among study areas were found in the structure of food webs, as well as in the relative importance of basal resource pools supporting each food web. Overall, the food web of Banc d'Arguin was characterized by lower trophic diversity and higher functional redundancy than the other sites. This result might be explained by the low number of trophic pathways of organic matter transfer in this seagrass‐dominated system which, as a fossil estuary, lacks inputs from both freshwater and nutrient‐rich offshore oceanic waters. Structure of shorebird communities was consistent with the main organizational patterns found for each food web, highlighting the less diverse character of the community of Banc d'Arguin. At Banc d'Arguin and Bijagós archipelago, which displayed the smallest and largest isotopic niche widths in bird assemblage, respectively, mean niche overlap among species was low, suggesting high interspecific partitioning in resource use. Tropical systems typically offer comparatively lower harvestable prey biomass for shorebirds and might thus strengthen interspecific competition, leading to low niche overlap among species. Our study reveals relevant differences in the structure of food webs and shorebird communities in coastal areas along an avian flyway. While differences in trophic redundancy of food webs point to distinct levels of ecosystem resilience, contrasts in the organization of shorebird communities highlight the plasticity in the foraging behaviour of species inhabiting areas with distinct environmental conditions.

Seasonality in temperate ecosystems shapes species phenology, influencing interactions and food web structure. Variations in species richness and biomass affect trophic interaction strength, a crucial factor for community stability, which can be assessed through energy fluxes – an essential indicator of ecosystem function. Yet, we still have limited understanding on how energy fluxes and food web structure vary overtime and how the seasonal dynamics influence stability. We examined how the structure, energy fluxes and stability of a highly resolved multitrophic lake food web change across seasons. Two estimates representing different facets of stability were considered: resilience and reactivity. We found a strong seasonal pattern in food web structure, with higher complexity observed in summer and autumn. In addition, we found that more complex food webs are more reactive to perturbations, and consequently, less stable than simpler ones over short time‐scale. This was influenced by a lower proportion of strong energy fluxes. Resilience was unaffected by seasonality, food web structure and energy fluxes, highlighting the need to account for multifaceted nature of stability. The seasonal variability of stability suggests that food webs may vary temporally in their vulnerability to environmental perturbations, which has important implications for ecosystem management.

Trophic interactions are central to ecosystem functioning, but the link between food web structure and ecosystem functioning remains obscure. Regularities (i.e. consistent patterns) in food web structure suggest the possibility of regularities in ecosystem functioning, which might be used to relate structure to function. We introduce a novel, genetic algorithm approach to simulate food webs with maximized throughput (a proxy for ecosystem functioning) and compare the structure of these simulated food webs to real empirical food webs using common metrics of food web structure. We repeat this analysis using robustness to secondary extinctions (a proxy for ecosystem resilience) instead of throughput to determine the relative contributions of ecosystem functioning and ecosystem resilience to food web structure. Simulated food webs that maximized robustness were similar to real food webs when connectance (i.e. levels of interaction across the food web) was high, but this result did not extend to food webs with low connectance. Simulated food webs that maximized throughput or a combination of throughput and robustness were not similar to any real food webs. Simulated maximum-throughput food webs differed markedly from maximum-robustness food webs, which suggests that maximizing different ecological functions can generate distinct food web structures. Based on our results, food web structure would appear to have a stronger relationship with ecosystem resilience than with ecosystem throughput. Our genetic algorithm approach is general and is well suited to large, realistically complex food webs. Genetic algorithms can incorporate constraints on structure and can generate outputs that can be compared directly to empirical data. Our method can be used to explore a range of maximization or minimization hypotheses, providing new perspectives on the links between structure and function in ecological systems.

Nonpoint source pollution entering rivers will pollute water quality, degrading the health of aquatic ecosystems. However, owing to the lack of quantitative research on the effects of nonpoint source pollution on the structure of aquatic food webs, there is a lack of quantitative basis for river management. Nonpoint source pollution is not only difficult to control effectively, but also the success rate of water ecological restoration projects is low. With the increasing proportion of nonpoint source pollution in water environmental problems, it is urgent to quantitatively assess and predict the impact of nonpoint source pollution on the structure of food webs. Therefore, this thesis presents a method for quantitatively assessing and predicting the impact of nonpoint source pollution on the structure of food webs through using fuzzy clustering to screen the typical points of the impact of nonpoint source pollution, then using canonical correspondence analysis (CCA) and partial least squares regression analysis to comprehensively filtrate the driving factors affect food web that results in nonpoint source pollution, and then determining the impact of each driving factor on the structure of food webs. Finally, the change trend of food web structure is predicted. The results show that (1) the driving factors that the nonpoint source pollution that affects the food web structure is NH3‐N and chemical oxygen demand (COD). The increase in NH3‐N and COD promotes the growth of phytoplankton, causing the change of the primary productivity of the ecosystem, and ultimately changes the entire food web structure; (2) NH3‐N and COD affect the stability, maturity, connectivity and complexity of the aquatic food web structure. The increase of NH3‐N increases the connectivity and maturity of the food web structure but reduces complexity and stability; the increase of COD increases the connection of the food web structure, while reducing the other three indicators; (3) in some areas with good water quality, aquatic species diversity is high, the relationship of interspecies dietary is complex, food web structure level index is high and the structure of food web is stable. The food web structure in the rainy season will be better than that in the dry season. In some areas with severe water pollution and poor food web structure, the ability of the food web to resist external interference is weak. The food web structure in the rainy season will be worse than that in the dry season owing to rainfall into the river. The methods and conclusions in this treatise can provide a reliable and quantitative scientific basis for river ecosystem management and ecosystem restoration and can improve the success rate of ecological restoration projects.

北部湾秋季底层鱼类多样性和优势种数量的变动趋势

Natural ecosystems comprise an innumerable amount of different organisms. These organisms are not separated, they interact and depend on each other. Today’s ecosystems are facing an enormous decline in biodiversity due to human impacts with thus far unknown consequences. One key objective of ecological research is to understand the mechanisms generating and maintaining this incredible amount of diversity. However, comprehensive analyses of natural ecosystems are impeded by their complexity and diversity. Food webs, therefore, provide an excellent tool to analyze the complexity of ecosystems. They depict the system‘s diversity and species interactions in a condensed form. Furthermore, food-web structure can help to predict the interaction strengths between species and the energy pathways through the system. In my thesis, I use food web structure to analyze structural properties which separate food webs from other network types and furthermore I investigate generalities and differences of food-web structure across different ecosystems. 
\nOne of the most important ecosystems is the soil ecosystem, as it provides the base for aboveground productivity. However, detailed soil food webs are scarce. In chapter 2, I assembled the complex food webs of 48 forest soil communities and analyzed if soil food webs differ in their topological parameters from those of other ecosystems. I found that soil food webs are characterized by a higher number of omnivorous and cannibalistic species. Moreover, they comprise more trophic chains and intraguild-predation motifs than food webs from other ecosystems. Finally, soil food webs showed high average and maximum trophic levels. These differences in network structure to other ecosystem types may be a result of ecosystem-specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food-web level. Despite these differences, soil food webs showed the same scaling of their properties with connectance and size. In a second analysis of land-use effects, I found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. This study has unravelled systematic structures of soil food-webs, extending our mechanistic understanding how their environmental characteristics determine patterns at the community level. Additionally, I have shown that the general scaling laws also apply for soil food webs.
\nIn addition to purely topological properties, I analyzed another important aspect of food webs. The distributions of body masses and degrees across species are key determinants of food-web structure and dynamics. In chapter 3, I analyzed body masses of species and their systematic distributions across food-web structure. In particular, allometric degree distributions combine both aspects in the relationship between degrees and body masses. They are of critical importance for the stability of complex ecological networks. I used an entirely novel global body-mass database including food-web structures of four different ecosystem types to analyze body-mass distributions, cumulative degree distributions, and allometric degree distributions regarding differences among ecosystem types. My results demonstrate some general patterns across ecosystems: the body masses are either roughly log-normally (terrestrial and stream ecosystems) or multimodally (lake and marine ecosystems) distributed, and most networks exhibit exponential cumulative degree distributions except stream networks that most often possess uniform degree distributions. Additionally, with increasing species body masses we found significant decreases in vulnerability in 70% of the food webs and significant increases in generality in 80% of the food webs. Overall, these analyses document striking generalities in the body-mass and degree structure across ecosystem types as well as surprising exceptions (uniform degree distributions in stream ecosystems). This suggests general constraints of body masses on the link structure of natural food webs irrespective of ecosystem characteristics. 
\nWhile I revealed general patterns of food-web topology in chapter 2 and 3, I investigated the drivers of these general patterns in chapter 4. Therefore, I analyzed the influence of different external factors on community (beta diversity) and food-web structure. Two main theoretical bodies explain β-diversity, the niche theory and neutral theory. However, neutral theory predicts only distributions for trophically identical species, whereas influences of local niches or neutral effects on food-web structure as a crucial part of the multitrophic structure of ecosystems are not taken into account. In chapter 4, I therefore analyzed the effects of spatial distance and environmental dissimilarity on the species dissimilarity (beta diversity) and food web dissimilarity (structural dissimilarity) of multitrophic forest communities. I showed that the mechanisms proposed by neutral theory can adequately predict the beta diversity of multitrophic species communities. Furthermore, food-web structure was robust and affected neither by spatial distance (random dispersal, neutral theory) nor by environmental filtering (niche theory). I additionally analyzed model food webs (random and niche topology) and compared their dissimilarities to empirical food webs. The highest dissimilarity was reached by random food webs whereas niche model food webs were in between and the lowest distances were expressed by empirical food webs. Further, random food webs displayed the highest mean trophic level (115), while niche model food webs showed lower (5) and empirical food webs the lowest (4) mean trophic level values. Hence, food-web structure appears to be energetically optimized with local species adapted to energetic niches within the food web while species identity within these niches remains random. This suggests that different species could be adapted to the same energetic niches and, while following random drift, still assemble into similar food web structures.
\nAltogether, the results of this thesis demonstrate the practicality of food-web structure in unravelling generalities across different ecosystems. Furthermore, food-web structure explains species distributions across the environment and provides additional important information on the ecosystem. 
\nThe observed generalities indicate constraints on food-web structure. The allometric degree distributions demonsrate such constraints on food-web structure by distributing the links in dependence of the species body masses. Finally, my results from chapter 4 indicate that, additionally to global topological constraints, local communities have to meet certain energetic constraints to explain the similarity found across food webs.

Metabolism Predicts Ecological Response to Warming

Food webs show the architecture of trophic relationships, revealing the biodiversity and species interactions in an ecosystem. Understanding which factors modulate the structure of food webs offers us the ability to predict how they will change when influential factors are altered. To date, most of the research about food webs has focused on species interactions whereas the influences of surrounding environments have been overlooked. Here, using network analysis, we identified how the structure of aquatic food webs varied across a range of geophysical conditions within a whole stream system. Within a headwater basin in the Cascade Mountain Range, Oregon, USA, macroinvertebrate and vertebrate composition was investigated at 18 sites. Predator–prey interactions were compiled based on existing literature and dietary analysis. Several structural network metrics were calculated for each food web. We show that the structure of food webs was predictable based on geophysical features at both local (i.e., slope) and broader (i.e., basin size) spatial extents. Increased omnivory, greater connectance, shorter path lengths, and ultimately greater complexity and resilience existed downstream compared to upstream in the stream network. Surprisingly, the variation in food web structure was not associated with geographic proximity. Structural metric values and abundance of omnivory suggest high levels of stability for these food webs. There is a predictable variation in the structure of food webs across the network that is influenced by both longitudinal position within streams and patchy discontinuities in habitat. Hence, findings illustrate that the slightly differing perspectives from the River Continuum Concept, Discontinuity Patch Dynamics, and Process Domains can be integrated and unified using food web networks. Our analyses extend ecologists’ understanding of the stability of food webs and are a vital step toward predicting how webs and communities may respond to both natural disturbances and current global environmental change.

Surface water connectivity controls fish food web structure and complexity across local- and meta-food webs in Arctic Coastal Plain lakes

Food web structure and energy flux dynamics, but not taxonomic richness, influence microbial ecosystem functions in a Sphagnum-dominated peatland

Structure of and carbon flux through soil food webs of temperate grassland as affected by land use management

Soil biogeochemical cycles are regulated by soil food webs. However, variation of soil food web structure and functioning across key environmental gradients remains unknown, hampering generalisations of any suggested links between fauna and biogeochemistry. Here, we used two complementary approaches to quantify soil animal food web variation across forest types, from southern taiga to rainforests. First, we applied the energy flux approach to explore patterns of energy distribution across micro-, meso- and macrofauna. We showed that tropical soil food webs have consistently higher energy flux, proportionally higher predation rates (31 vs 18-27% of the total energy flux) and relied more on the plant energy channel (21 vs 10%), but less on the bacterial (5 vs 9-18%) and litter energy channels (14 vs 18-32%), than temperate soil food webs. Second, we compiled a large database (>8000 records) of stable isotope composition of soil animals to see how detritivory and microbivory in soil animal communities change with environmental temperature and litter quality. Despite little effect of temperature, shift in 15N concentrations suggested that in most cases low litter quality (high %C and low %N) result in a switch from feeding directly on litter to feeding on microorganisms. Thus, soil animals change their functional role from competitors to consumers of microbes. Our studies show how the functioning of soil animal food webs changes across biomes with different climate and litter quality and summarise functional roles animals play in different biomes.

Seasonal shifts in hydrology are known to alter the abundance and diversity of basal production resources and habitats and hence strongly influence the structure and function of river ecosystems. However, equivalent knowledge of natural lake ecosystems in floodplain regions is lacking. Here, we used stable isotope ratios of carbon and nitrogen to assess available primary production sources and consumer taxa during the dry and wet seasons in a large floodplain lake connected to the Yangtze River. Fish species showed distinct δ13C values between two hydrological periods but only small changes in δ15N values. Most of the fish species had higher estimated trophic levels in the dry season, likely indicating greater carnivory. Results of Bayesian mixing models revealed that benthic algae and benthic organic matter (BOM), combined with C3 vegetation, were the principal food sources supporting the biomass of most fish species during the low-water period, whereas benthic algae and seston were the most important carbon sources during the flood period. Overall, these findings demonstrate that seasonal hydrological changes, such as water-level fluctuations, can affect the trophic structure and ecosystem functioning of floodplain lake food webs in the subtropical zone.

Introduction of new fish species to lakes that contain fish is known to disrupt food webs, but few studies have quantified the effects of introduced fish on flow of organic matter and mechanisms governing food web structure in historically fishless lakes. Contrasting flow of organic matter and food web structure in macroinvertebrate communities in lakes with and without fish provides an opportunity to characterise resiliency of macroinvertebrate food web function when there are changes in community composition and reductions in biomass. Organic matter flow and food web structure were characterised in six lakes with and six lakes without fish at high elevation in the southern Rocky Mountains of Colorado, U.S.A. Carbon (δ13C) and nitrogen (δ15N) were used to determine the proportionate contribution of organic matter from primary organic matter sources (periphyton, phytoplankton, terrestrial plants) to consumers including herbivores and higher trophic levels. Contributions of organic matter sources to macroinvertebrates were combined with estimates of littoral macroinvertebrate production in quantifying the contribution of each primary source to total invertebrate production. Although introduced fish altered community composition, body size, and biomass, they did not alter contributions of the organic matter sources to production of macroinvertebrates. The dominant resources contributing to macroinvertebrate production were periphyton and phytoplankton, which combined contributed to 80% of macroinvertebrate production. Despite occupying the highest trophic position (difference in δ15N relative to primary sources), fish did not alter trophic structure of littoral macroinvertebrate communities in maximum macroinvertebrate trophic position and community trophic position. Community trophic position weighted by production for macroinvertebrates was related to macroinvertebrate production per unit area but was independent of lake area. In lakes with fish, maximum macroinvertebrate trophic position was similar to that of lakes without fish because large predators (predaceous beetles and dragonflies) were replaced by small predators (tanypod chironomids). The present study indicates that macroinvertebrate food webs can exhibit resilience to introduced or invasive fish even when there are changes in community composition and biomass. This suggests that changes in structure (e.g. biomass, community composition) associated with introduced or invasive species do not necessarily equate to changes in food web function (e.g. flow of organic matter, maximum trophic position). Further, the present study suggests that although maximum trophic position has been shown to increase with lake size for lakes containing fish, the pattern is unlikely to extend to lakes without fish. The relationship between lake size and maximum trophic position for lakes containing fish may occur because fish have expandable niches associated with high motility and indeterminate growth, in contrast to macroinvertebrates.

Summary1. We studied the effect of substratum movement on the communities of adjacent mountain and spring tributaries of the Ivishak River in arctic Alaska (69°1′N, 147°43′W). We expected the mountain stream to have significant bed movement during summer because of storm flows and the spring stream to have negligible bed movement because of constant discharge.2. We predicted that the mountain stream would be inhabited only by taxa able to cope with frequent bed movement. Therefore, we anticipated that the mountain stream would have lower macroinvertebrate species richness and biomass and a food web with fewer trophic levels and lower connectance than the spring stream.3. Substrata marked in situ indicated that 57–66% of the bed moved during summer in the mountain stream and 4–20% moved in the spring stream.4. Macroinvertebrate taxon richness was greater in the spring (25 taxa) than in the mountain stream (20 taxa). Mean macroinvertebrate biomass was also greater in the spring (4617 mg dry mass m−2) than in the mountain stream (635 mg dry mass m−2). Predators contributed 25% to this biomass in the spring stream, but only 7% in the mountain stream.5. Bryophyte biomass was >1000 times greater in the spring stream (88.4 g ash‐free dry mass m−2) than the mountain stream (0.08 g ash‐free dry mass m−2). We attributed this to differences in substratum stability between streams. The difference in extent of bryophyte cover between streams probably explains the high macroinvertebrate biomass in the spring stream.6. Mean food‐web connectance was similar between streams, ranging from 0.18 in the spring stream to 0.20 in the mountain stream. Mean food chain length was 3.04 in the spring stream and 1.83 in the mountain stream. Dolly Varden char (Salvelinus malma) was the top predator in the mountain stream and the American dipper (Cinclus mexicanus) was the top predator in the spring stream. The difference in mean food chain length between streams was due largely to the presence of C. mexicanus at the spring stream.7. Structural differences between the food webs of the spring and mountain streams were relatively minor. The difference in the proportion of macroinvertebrate biomass contributing to different trophic levels was major, however, indicating significant differences in the volume of material and energy flow between food‐web nodes (i.e. food web function).