Patterns and drivers of fish trophic trajectories over time

Fishing has wide-ranging impacts on marine ecosystems. One of the most pervasive signs of intensive fishing is "fishing down the food web", with landings increasingly dominated by smaller species from lower trophic levels. Decreases in the trophic level of landings are assumed to reflect those in fish communities, because size-selective mortality causes decreases in the relative abundance of larger species and in mean body size within species. However, existing analyses of fishing impacts on the trophic level of fish communities have focused on the role of changes in species composition rather than size composition. This will provide a biased assessment of the magnitude of fishing impacts, because fishes feed at different trophic levels as they grow. Here, we combine body size versus trophic level relationships for North Sea fishes (trophic level assessed using nitrogen stable-isotope analysis) with species–size–abundance data from two time-series of trawl-survey data (whole North Sea 1982–2000, central and northern North Sea 1925–1996) to predict long-term trends in the trophic structure of the North Sea fish community. Analyses of the 1982–2000 time-series showed that there was a slow but progressive decline in the trophic level of the demersal community, while there was no trend in the trophic level of the combined pelagic and demersal community. Analyses of the longer time-series suggested that there was no trend in the trophic level of the demersal community. We related temporal changes in trophic level to temporal changes in the slopes of normalised biomass size-spectra (which theoretically represent the trophic structure of the community), mean log2 body mass and mean log2 maximum body mass. While the size-based metrics of community structure showed long-term trends that were consistent with the effects of increased fishery exploitation, these trends were only correlated with trophic level for the demersal community. Our analysis suggests that the effects of fishing on the trophic structure of fish communities can be much more complex than previously assumed. This is a consequence of sampled communities not reflecting all the pathways of energy transfer in a marine ecosystem and of the absence of historical data on temporal and spatial changes in the trophic level of individuals. For the North Sea fish community, changes in size structure due to the differential effects of fishing on species and populations with different life histories are a stronger and more universal indicator of fishing effects than changes in mean trophic level.

Assessing the role of non-native species and artificial water bodies on the trophic and functional niche of Mediterranean freshwater fish communities

To study the trophic structure of fish community in Minjiang Estuary, we measured the carbon and nitrogen stable isotope ratios (δ13C and δ15N) of common fish species sampled in winter (January), spring (May), summer (August) and autumn (November) of 2015 by trawling, as well as seven quantitative metrics (ratios range of δ15N, NR; ratios range of δ13C, CR; total area, TA; standard ellipse area, SEAc; average distance of centroid, CD; mean nearest neighbor distance, MNND; standard deviation of nearest neighbor distance, SDNND). The results showed significant differences of δ13C values among all the seasons except between summer and autumn. There were significant differences of δ15N values between spring and other three seasons but no significant difference among summer, autumn, and winter. According to the results of the seven quantitative metrics, trophic structure of common fish community remarkably varied among the four seasons, as TA, SEAc, CR, NR, CD and MNND increased in succession from summer, autumn, winter to spring. Our results indicated that seasonal differences in trophic structure of common fish community were related to the fish spawning migration and the diversity of food source.

Rainforest conversion to agricultural plantations such as rubber and oil palm in Indonesia is increasing in the last decades. In this thesis, I investigated effects of rainforest conversion to rubber and oil palm plantations in Jambi, Sumatra, Indonesia. The study area serves as model region for investigating consequences of land-use changes on biodiversity, the channeling of energy through soil food webs and changes in trophic niches of one dominant group of soil mesofauna, Collembola. Collembola are among the most abundant arthropods inhabiting the belowground system sensitively responding to changes in vegetation and soil conditions. In Chapter 2, consequences of the conversion of rainforest into plantations on the channelling of energy through soil food webs was investigated by using neutral lipid fatty acid analysis from six dominant soil fauna taxa. Rainforest conversion changed the biomarker FA composition of soil fauna at the community level. Conversion of rainforest into oil palm plantations enhanced the plant energy channel and reduced the bacterial energy channel. The changes in energy distribution within soil food webs may have significant implications for the functioning of tropical ecosystems and their response to environmental changes. In Chapter 3, I investigated effects of rainforest conversion on the density, species richness, functional and community composition of soil and litter Collembola. Land-use change negatively affected Collembola communities in the litter layer, but the impact was less pronounced in the soil layer. Water content and pH were identified as important factors driving Collembola community composition. Several pantropical genera of Collembola (i.e., Isotomiella, Pseudosinella and Folsomides) dominated across land-use systems, reflecting their high environmental adaptability or efficient dispersal calling for studies on their ecology and genetic diversity. The decline in species richness and density of litter-dwelling Collembola with the conversion of rainforest into plantation systems calls for management practices mitigating negative effects of the deterioration of the litter layer in rubber plantations, but even more in oil palm plantations. Specific traits, such as larger size, patterned or intensive pigmentation and number of ocelli may explain the adaptation of Collembola species to the environmental changes in particular to changes in soil water content and pH. In Chapter 4, the effect of rainforest conversion on the trophic niche structure of Collembola was investigated using stable isotope analysis. Across Collembola species, ∆13C values (calibrated to litter) were highest in rainforest suggesting more pronounced processing of litter resources by microorganisms and consumption of these microorganisms by Collembola in this system. Lower ∆13C values, but high ∆13C variation in Collembola in oil palm plantations indicated that Collembola shifted towards herbivory and used more variable resources in this system. Small range in ∆15N values in Collembola species in monoculture plantations in comparison to rainforest indicated that conversion of rainforest into plantations is associated with simplification in the trophic structure of Collembola communities. Across the studied ecosystems, atmobiotic species occupied the lowest, whereas euedaphic Collembola species occupied the highest trophic position, resembling patterns in temperate forests. A dominant species of the euedaphic life form and of high trophic position (Pseudosinella sp.1) was found in high abundance across rainforest and plantations suggesting that this species is best adapted to the studied land-use changes. Euedaphic species generally appear to be better adapted to cope with seasonal fluctuations than other life forms. Additionally, in Chapter 5, I investigated seasonal fluctuations of Collembola density and community composition in rainforest and plantation systems. Collembola density in the litter layer, was at a maximum at the beginning of the wet season. Euedaphic and hemiedaphic species, living in the lower litter layers and soil, fluctuated less with season than epedaphic and atmobiotic species, living in upper litter layers. Differences in community composition among land-use systems were most pronounced at the beginning of the dry season (June). Collembola community composition changed with season in all land-use systems, with the differences being more pronounced in the litter than in the soil layer. In line with the results from Chapter 3, water content and pH were identified as important factors driving Collembola community composition across seasons. Overall, rainforest conversion to rubber and oil palm have negative impacts on soil and litter Collembola biodiversity, alter the trophic structure of Collembola communities and the channeling of energy through soil food webs. To reduce the negative impacts of rainforest conversion into plantation systems as revealed in this thesis, improved management of agricultural plantations is needed.

SummaryBody size determines rates of respiration and production, energy requirements, mortality rates, patterns of predation and vulnerability to mortality. Body size distributions are often used to describe structure and energy flux in communities and ecosystems.If clear relationships can be established between body size and trophic level in fishes, they may provide a basis for integrating community and ecosystem analyses based on size spectra, food webs and life histories.We investigated relationships between the body sizes (weight and length) of north‐east Atlantic fishes and their trophic level. The abundance of15N, as determined by stable isotope analysis, was used as an index of trophic level.Cross‐species and comparative analyses demonstrated that body size was unrelated or weakly related to trophic level. Thus allometric relationships between body size and trophic level could not be used to predict the trophic structure of fish communities.The results of the cross‐species analyses contrasted with patterns in the size and trophic structure of entire fish communities. When fish communities were divided into size classes, there were strong positive relationships between size class and trophic level. The slope suggested a mean predator : prey body mass ratio of 80 : 1.Our results suggest that body size does not provide a useful surrogate of trophic level for individual species, but that body size is an excellent predictor of trophic level within the community, providing an empirical basis for integrating community analyses based on models of trophic structure and body size distributions.

Coral reef mesopredator trophodynamics in response to reef condition

A combination of dietary guild analysis and nitrogen (δ(15) N) and carbon (δ(13) C) stable-isotope analysis was used to assess the trophic structure of the fish community in Rhode Island and Block Island Sounds, an area off southern New England identified for offshore wind energy development. In the autumn of 2009, 2010 and 2011, stomach and tissue samples were taken from 20 fish and invertebrate species for analysis of diet composition and δ(15) N and δ(13) C signatures. The food chain in Rhode Island and Block Island Sounds comprises approximately four trophic levels within which the fish community is divided into distinct dietary guilds, including planktivores, benthivores, crustacivores and piscivores. Within these guilds, inter-species isotopic and dietary overlap is high, suggesting that resource partitioning or competitive interactions play a major role in structuring the fish community. Carbon isotopes indicate that most fishes are supported by pelagic phytoplankton, although there is evidence that benthic production also plays a role, particularly for obligate benthivores such as skates Leucoraja spp. This type of analysis is useful for developing an ecosystem-based approach to management, as it identifies species that act as direct links to basal resources as well as species groups that share trophic roles.

The rupture of the Fundão dam is considered the largest mining failure in history, which had a particularly detrimental impact on fish populations, as the mud from the ore tailings significantly altered the water quality and habitat of Doce River basin. This study aimed to assess the trophic structure of fish communities in areas impacted and not impacted by the dam rupture in the Doce River basin. To evaluate the food web structure, community-wide trophic niche, and trophic positions of fish, stable isotopes of carbon (δ13C) and nitrogen (δ15N) were utilized across ten sites (seven impacted and three control). In general, fish appeared to assimilate resources such as invertebrates, algae, and periphyton, although the importance of each resource varied among sites. The site closest to the dam rupture exhibited a more simplified trophic structure compared to the control sites and those nearer the river mouth. In this site, most fish species occupied a similar trophic position. Trophic niches also exhibited the greatest dissimilarity between the site closest to the dam failure and those farther away from it, with an expansion of trophic niche breadth observed with an increase in the distance from the dam rupture. Our study provided valuable insights into the trophic structure of fish communities within the Doce River basin, shedding light on the trophic ecology of the 59 fish species investigated. We also emphasize the importance of our study for future assessments of ore tailings dam failure disasters and evaluating the effectiveness of mitigation measures for Doce River basin recovery.

The trophic spectrum provides a useful method to investigate the trophic structure of fish communities. However, ontogenetic shifts in diet can cause variations in the trophic level with body size, thereby influencing the trophic structure of fish communities. In this study, we investigated the effect of ontogenetic dietary shifts on the trophic structure of fish communities in Haizhou Bay based on trophic spectra and trophic indicators calculated under different scenarios of functional group classification. The results showed that the size threshold of ontogenetic dietary shifts was a viable basis for functional group classification. The biomass of fishes at trophic levels 3.4–3.8 was lower when ontogenetic dietary shifts were considered, which can influence the intensity of top–down control and hinder the formulation of harvest strategies. Ontogenetic dietary shifts can also redistribute the biomass of fishes across trophic levels, thereby regulating the trophic structure of fish communities. Different responses of trophic indicators to ontogenetic dietary shifts were identified, with high trophic indicator (HTI) being the most appropriate indicator reflecting the effect of ontogenetic dietary shifts in the trophic structure. This study provides a feasible way to optimize the trophic spectrum for elucidating the trophic structure of fish communities. And we recommend that ontogenetic dietary shifts should be considered and valued in fishery management.

The present study tested the hypothesis that artificial habitats (pier and bridge) harbour different fish trophic guilds compared with natural habitats (mangrove roots) and that the trophic structure of fish communities on estuarine artificial habitats resembles adjacent coral reefs. High-definition cameras were used to survey the fish community associated with the different structures over a 6-month period. Benthos was also analysed following the point intercept method on the different habitats. In the estuary, fish abundance was up to threefold higher and species richness twofold higher on artificial structures compared with the natural habitat. Mangrove roots were mainly inhabited by juvenile carnivores, whereas the pier and bridge were mostly inhabited by sessile invertebrate feeders and roving herbivores. A less diverse benthic community was found on mangrove roots, mostly composed of mud and algae. In contrast, benthos at the bridge and pier was more diverse and dominated by sponges, octocorals and oysters. In addition, fish trophic structure from an adjacent coral reef area showed more than 60% similarity with the fish community on the artificial structures surveyed. The results of the present study indicate that artificial hard structures support unique fish communities compared with natural estuarine mangrove habitats.

Seasonal variation in aquatic food web structure at Mad Island Marsh, Matagorda Bay, Texas, was examined using dietary information obtained from the analysis of gut contents from large samples of fish and crustacean specimens. Unique aspects of this study include the use of large samples of consumer gut contents (n=6,452), long-term sampling (bimonthly surveys over 18 mo), and standard methods of data collection and analysis facilitating comparisons with other aquatic food webs. Dietary data were partitioned for analysis into warm (summer) and cold (winter) seasons. Most consumers fed low in the food web, with trophic levels ranging from about 2 to 3.5 during both summer and winter. Vegetative detritus was more important in macroconsumer diets than live algae and macrophytes. Low trophic levels of consumers reflected the important role of abundant detritivores (e.g., striped mulletMugil cephalus, Gulf menhadenBrevortia patronum, and macroinvertebrates) in linking detritus to top predators via short food chains, a finding consistent with many other estuarine food web studies. Despite changes in community composition and population size structure of certain species, most food web properties revealed comparatively little seasonal variation. The summer food web had more nodes (86), more links (562), a higher density of links as indicated by connectance (0.08), and a slightly higher predator: prey ratio (0.51) compared to the winter food web (75 nodes, 394 links, connectance = 0.07, predator: prey ratio = 0.47). Proportions of top (0.06–0.07), intermediate (0.75–0.76), and basal (0.19) species did not vary significantly between seasons, but mean trophic level was higher during summer. Addition of feeding links based on information from the literature increased connectance to 0.13 during both seasons; other web parameters had values similar to those obtained for our directly estimated food webs. Seasonal variation in food web structure was influenced by changes in community composition (e.g., influxes of postlarval estuarine-dependent marine fishes during winter), availability of resources (e.g., more submerged macrophytes and amphipods during summer), and size structure and ontogenetic diet shifts of dominant consumer taxa. Our findings suggest that some basic properties of estuarine food web are resilient to seasonal changes in population and community structures and food web architecture.

Mangroves are important ecosystems of tropical and subtropical shorelines. Anthropogenic activity decreases their habitat quality, affecting structural and functional trophic features. We hypothesized that higher levels of anthropogenic intervention generate diversity loss and modify the trophic structure of tropical mangrove fish communities. We compared the taxonomic and isotopic (δ13C/δ15N) composition, abundance, trophic position, and isotopic niche of fish communities from three mangrove systems with different anthropogenic intervention levels in the Colombian Pacific. Non-parametric statistical tests and a Bayesian approach were used to analyze data. A total of 1254 specimens belonging to 23 families, 25 genera, and 30 species were identified, presenting higher abundance (821) in moderate anthropogenic intervention level mangrove (Moderate-AIL), with high dominance of one species (Lilestolifera). The low anthropogenic intervention level mangrove (Low-AIL) was the second in abundance (291) but exhibited a greater number of species (23), while the high anthropogenic intervention level mangrove (High-AIL) presented the least abundance (142) and species number (17). The isotopic composition ​​reveals that Moderate and High-AIL mangroves presented enriched 13C and 15N compared to Low-AIL (~ 2 to 4 ‰). Mean trophic position (TP) of communities was slightly higher in the more intervened systems (~ 1 to 2 orders of magnitude), as well as in specific species (Centropomus spp.). Isotopic niche width (TA and SEAc) was greater in High-AIL (41.1 and 9.2), more than doubled compared to Moderate-AIL (33.0 and 4.1). In High-AIL isotopic niche width increased, indicating lesser availability of prey and basal resources. The results obtained in this study support the proposed hypothesis and, suggest that anthropogenic intervention modifies diversity and food webs dynamics, affecting the transfer of matter and energy from macrotidal tropical mangroves to coastal ecosystems. However, it is recommended to be careful concluding differences based exclusively on the anthropogenic intervention level, since it is widely documented that mangrove settings also influence the analyzed trophic parameters.

Global climate change is driving species' distributions towards the poles and mountain tops during both non-breeding and breeding seasons, leading to changes in the composition of natural communities. However, the degree of season differences in climate-driven community shifts has not been thoroughly investigated at large spatial scales. We compared the rates of change in the community composition during both winter (non-breeding season) and summer (breeding) and their relation to temperature changes. Based on continental-scale data from Europe and North America, we examined changes in bird community composition using the community temperature index (CTI) approach and compared the changes with observed regional temperature changes during 1980-2016. CTI increased faster in winter than in summer. This seasonal discrepancy is probably because individuals are less site-faithful in winter, and can more readily shift their wintering sites in response to weather in comparison to the breeding season. Regional long-term changes in community composition were positively associated with regional temperature changes during both seasons, but the pattern was only significant during summer due to high annual variability in winter communities. Annual changes in community composition were positively associated with the annual temperature changes during both seasons. Our results were broadly consistent across continents, suggesting some climate-driven restructuring in both European and North American avian communities. Because community composition has changed much faster during the winter than during the breeding season, it is important to increase our knowledge about climate-driven impacts during the less-studied non-breeding season.

Fish communities associated with coral reefs worldwide are threatened by habitat degradation and overexploitation. We assessed coral reefs, mangrove fringes, and seagrass meadows on the Caribbean coast of Panama to explore the influences of their proximity to one another, habitat cover, and environmental characteristics in sustaining biomass, species richness and trophic structure of fish communities in a degraded tropical ecosystem. We found 94% of all fish across all habitat types were of small body size (≤10 cm), with communities dominated by fishes that usually live in habitats of low complexity, such as Pomacentridae (damselfishes) and Gobiidae (gobies). Total fish biomass was very low, with the trend of small fishes from low trophic levels over-represented, and top predators under-represented, relative to coral reefs elsewhere in the Caribbean. For example, herbivorous fishes comprised 27% of total fish biomass in Panama relative to 10% in the wider Caribbean, and the small parrotfish Scarus iseri comprised 72% of the parrotfish biomass. We found evidence that non-coral biogenic habitats support reef-associated fish communities. In particular, the abundance of sponges on a given reef and proximity of mangroves were found to be important positive correlates of reef fish species richness, biomass, abundance and trophic structure. Our study indicates that a diverse fish community can persist on degraded coral reefs, and that the availability and arrangement within the seascape of other habitat-forming organisms, including sponges and mangroves, is critical to the maintenance of functional processes in such ecosystems.

Understand how environmental factors correlate with the trophic structure of fish communities is a fundamental step to allow designing models for the functioning of stream ecosystems and planning conservation actions. Thus, this study aims to characterize the trophic structure of fish communities of streams of the Contas River basin and evaluate the relationships between local environmental factors (stream structure and limnological variables) and trophic structure. We sample nine streams of the Upper Contas sub-basin and nine of the Gongogi sub-basin using electric fishing. We analyzed the diet of 24 species, arranged in six trophic guilds, according to stomach contents: detritivores, omnivores, insectivores, invertivores, piscivores, and algivores. The results show high proportions of detritivores (46.62%), omnivores (28.55%), and insectivores (11.8%). Invertivores were correlated to colder streams, and with little proportion of sand in the substrate. Algivores were positively related to temperature and proportion of sand in the substrate. The relationship between other guilds and the environmental factors was not significant, probably due to limitations in the number of streams in the analysis. The sub-basins did not differ regarding the trophic structure, corroborating the perception that the trophic structure is more influenced by local factors than by broader scale factors.