Community Size Spectra Research Articles

Systematic deviations from linear size spectra of lake fish communities are correlated with predator–prey interactions and lake‐use intensity

Size structure of organisms at logarithmic scale (i.e. size spectrum) can often be described by a linear function with a negative slope; however, substantial deviations from linearity have often been found in natural systems. Theoretical studies suggest that greater nonlinearity in community size spectrum is associated with high predator–prey size ratios but low predator–prey abundance ratios; however, empirical evaluation of the effects of predator–prey interactions on nonlinear structures remains scarce. Here, we aim to empirically explore the pattern of the size‐specific residuals (i.e. deviations from the linear regression between the logarithmic fish abundance and the logarithmic mean fish size) by using size spectra of fish communities in 74 German lakes. We found that nonlinearity was strong in lakes with high predator–prey abundance ratios but at low predator–prey size ratios. More specifically, our results suggest that only large predators, even if occurring in low abundances, can control the density of prey fishes in a broad range of size classes in a community and thus promote linearity in the size spectrum. In turn, the lack of large predator fishes may cause high abundances of fish in intermediate size classes, resulting in nonlinear size spectra in these lakes. Moreover, these lakes were characterized by a more intense human use including high fishing pressure and high total phosphorus concentrations, which have negative impacts on the abundance of large, predatory fish. Our findings indicate that nonlinear size spectra may reflect dynamical processes potentially caused by predator–prey interactions. This opens a new perspective in the research on size spectrum, and can be relevant to further quantify the efficiency of energy transfer in aquatic food webs.

A size-structured matrix model to simulate dynamics of marine community size spectrum.

Several types of size-based models have been developed to model the size spectra of marine communities, in which abundance scales strongly with body size, using continuous differential equations. In this study, we develop a size-structured matrix model, which can be seen as a discretization of the Mckendrick-von Foerster equation, to simulate the dynamics of marine communities. The developed model uses a series of simple body size power functions to describe the basic processes of predator–prey interactions, reproduction, metabolism, and non-predation mortality based on the principle of mass balance. Linear size spectra with slopes of approximately –1 are successfully reproduced by this model. Several examples of numerical simulations are provided to demonstrate the model’s behavior. Size spectra with cut-offs and/or waves are also found for certain parameter values. The matrix model is flexible and can be freely manipulated by users to answer different questions and is executed over a shorter numerical calculation running time than continuous models.

Temporal and spatial variability of zooplankton on the Faroe shelf in spring 1997–2016

Abstract Zooplankton availability during spring and summer determines the growth and survival of first-feeding fish larvae, and thus impacts the recruitment to both fish prey species and commercial fish stocks. On the Faroe shelf, however, the relative importance of oceanic versus neritic zooplankton species has hitherto not been well understood. In this study, spatio-temporal variability in zooplankton community structure and size spectra on the Faroe shelf is investigated using observations from late April during the period 1997–2016. The main objective was to explore which environmental variables influence the zooplankton community structure in early spring. The zooplankton community in the permanently well mixed central shelf inside the tidal front consists of a mixture of neritic, cosmopolitan and oceanic species. In this region, redundancy analyses showed that chlorophyll concentration had a positive effect on abundance of neritic copepods and meroplankton as well as all zooplankton Calanus finmarchicus , is however observed off-shore from the tidal front, especially on the western side of the Faroe Plateau. A shift in C. finmarchicus phenology occurred around 2007, resulting in earlier reproduction of this species, and this variability could not be explained by the employed regional environmental parameters. Our results indicate that the Faroe shelf biological production is more dependent on the local primary production and neritic zooplankton species than on the large oceanic C. finmarchicus stock.

Remarkable size-spectra stability in a marine system undergoing massive invasion

The Mediterranean Sea is an invasion hotspot, with non-indigenous species suspected to be a major driver behind community changes. We used size spectra, a reliable index of food web structure, to examine how the influx of Red Sea fishes into the Mediterranean Sea has impacted the indigenous species community. This is the first attempt to use changes in the size spectra to reveal the effect of biological invasions. We used data from trawl catches along Israel's shoreline spanning 20 years to estimate changes in the community size spectra of both indigenous and non-indigenous species. We found that the relative biomass of non-indigenous species increased over the 20 years, especially for small and large species, leading to a convergence with the indigenous species size spectra. Hence, the biomass of indigenous and non-indigenous species has become identical for all size classes, suggesting similar energetic constraints and sensitivities to fishing. However, over this time period the size spectrum of indigenous species has remained remarkably constant. This suggests that the wide-scale invasion of non-indigenous species into the Mediterranean may have had little impact on the community structure of indigenous species.

Effects of lower trophic level biomass and water temperature on fish communities: A modelling study

Physical and biogeochemical changes of the oceans have complex influences on fish communities. Variations of resource and temperature affect metabolic rates at the individual level, biomass fluxes at the species level, and trophic structure as well as diversity at the community level. We use a Dynamic Energy Budget-, trait-based model of the consumers’ community size-spectrum to assess the effects of lower trophic level biomass and water temperature on communities at steady state. First, we look at the stressors separately in idealized simulations, varying one while the second remains constant. A multi-domain response is observed. Linked to the number of trophic levels sustained in the consumers’ community, the regimes highlighted present similar properties when lower trophic level biomass is increased or temperature decreased. These trophic-length domains correspond to different efficiencies of the transfer of biomass from small to large individuals. They are characterized by different sensitivities of fish communities to environmental changes. Moreover, differences in the scaling of individuals’ metabolism and prey assimilation with temperature lead to a shrinking of fish communities with warming. In a second step, we look at the impact of simultaneous variations of stressors along a mean latitudinal gradient of lower trophic level biomass and temperature. The model explains known observed features of global marine ecosystems such as the fact that larger species compose fish communities when latitude increases. The structure, diversity and metabolic properties of fish communities obtained with the model at different latitudes are interpreted in light of the different trophic-length domains characterized in the idealized experiments. From the equator to the poles, the structure of consumers’ communities is predicted to be heterogeneous, with variable sensitivities to environmental changes.

Acoustically derived fish size spectra within a lake and the statistical power to detect environmental change

Fisheries acoustic surveys are increasingly being used to monitor the abundance of fish stocks, yet their adoption as a tool to monitor changes in community size spectra has not been well explored. In this study, we use a series of historical acoustic surveys of the pelagic zones of three arms of Lake Opeongo to determine if acoustically derived size-spectra indicators (slope and height) can be effectively measured and used as a monitoring tool. Acoustic size-spectra indicators were successfully measured for every survey and resembled the same indicators found in netting surveys. From 2005 to 2009, the slope of the size spectra became shallower, likely due to a decrease in abundance in schooling prey fish. Estimates of sources of survey variation, including fish size estimates and interbasin differences, were low (<10% coefficients of variation), suggesting that monitoring programs would detect annual changes in the size-spectra slopes ranging from 2% to 15% within 10 years. Size-spectra heights did not change very much over the time series of surveys. Using lake trout (Salvelinus namaycush) fishery data from Lake Opeongo, we estimate that sources of natural variation in size spectra at the population level could be much higher and potentially require longer monitoring periods. We noted from our study that standardized surveys and data analyses are critical if acoustically derived size spectra are to be adopted as a monitoring tool. We also suggest that size spectra be calculated through echo-integration methods rather than echo-counting methods because, in this study, echo counting led to lower estimates of abundance and size-spectra indicators, which are likely underestimates of the true fish community’s abundance.

The effects of seasonal processes on size spectrum dynamics

The recent advent of dynamic size spectrum models has allowed the analysis of life processes in marine ecosystems to be carried out without the high complexity arising from interspecies interactions within dense food webs. In this paper, we use “mizer”, a size spectrum modelling framework, to investigate the consequences of including the seasonal processes of plankton blooms and batch spawning in the model dynamics. A multispecies size spectrum model is constructed using 12 common North Sea fish species, with growth, predation, and mortality explicitly modelled, before simulating both seasonal plankton blooms and batch spawning of fish (using empirical data on the spawning patterns of each species). The effect of seasonality on the community size spectrum is investigated; it is found that with seasonal processes included, the species spectra are more varied over time, while the aggregated community spectrum remains fairly similar. Growth of seasonally spawning mature individuals drops significantly during peak reproduction, although lifetime growth curves follow nonseasonal ones closely. On analysing properties of the community spectrum under different fishing scenarios, seasonality generally causes more varied spectrum slopes and lower yields. Under seasonal conditions, increasing fishing effort also results in greater temporal variability of fisheries yields due to truncation of the community spectrum towards smaller sizes. Further work is needed to evaluate robustness of management strategies in the context of a wider range of seasonal processes and behavioural strategies, as well as longer term environmental variability and change.

Implementing a multispecies size-spectrum model in a data-poor ecosystem

Multispecies ecological models have been used for predicting the effects of fishing activity and evaluating the performance of management strategies. Size-spectrum models are one type of physiologically-structured ecological model that provide a feasible approach to describing fish communities in terms of individual dietary variation and ontogenetic niche shift. Despite the potential of ecological models in improving our understanding of ecosystems, their application is usually limited for data-poor fisheries. As a first step in implementing ecosystem-based fisheries management (EBFM), this study built a size-spectrum model for the fish community in the Haizhou Bay, China. We describe data collection procedures and model parameterization to facilitate the implementation of such size-spectrum models for future studies of data-poor ecosystems. The effects of fishing on the ecosystem were exemplified with a range of fishing effort and were monitored with a set of ecological indicators. Total community biomass, biodiversity index, W-statistic, LFI (Large fish index), MeanW (mean body weight) and Slope (slope of community size spectra) showed a strong non-linear pattern in response to fishing pressure, and largest fishing effort did not generate the most drastic responses in certain scenarios. We emphasize the value and feasibility of developing size-spectrum models to capture ecological dynamics and suggest limitations as well as potential for model improvement. This study aims to promote a wide use of this type of model in support of EBFM.

Acoustic measures of lake community size spectra

Acoustic methods were tested as an alternative to catch data to measure size spectra of freshwater fish communities in Lac du Bonnet (LdB), Manitoba, Canada. Eleven daytime hydroacoustic surveys conducted in 2011 and 2012 enabled length-frequency spectra of fish communities to be calculated within LdB. Fish echoes tracked and converted to length (>5 cm) formed size spectra with significant linear regressions in all cases (p values < 0.05, R2 values from 0.87 to 0.99). Fish abundance (spectral height) was greater in 2011 (mean ± SD = 0.19 ± 0.15) than in 2012 (–0.19 ± 0.20), decreasing through the summer sampling season. Both abundance and size composition (spectral slope) of fish communities differed between three basins of varying bathymetry, with steepest slopes and lowest heights associated with the shallowest waters. We conclude that acoustically derived size spectra provide an efficient means to detect and monitor fish community dynamics over varied spatial and temporal scales in aquatic ecosystems where fish are not highly aggregated and hence single acoustic targets predominate.

Implications of scaled δ15N fractionation for community predator-prey body mass ratio estimates in size-structured food webs.

Nitrogen stable isotope ratios (δ(15) N) may be used to estimate community-level relationships between trophic level (TL) and body size in size-structured food webs and hence the mean predator to prey body mass ratio (PPMR). In turn, PPMR is used to estimate mean food chain length, trophic transfer efficiency and rates of change in abundance with body mass (usually reported as slopes of size spectra) and to calibrate and validate food web models. When estimating TL, researchers had assumed that fractionation of δ(15) N (Δδ(15) N) did not change with TL. However, a recent meta-analysis indicated that this assumption was not as well supported by data as the assumption that Δδ(15) N scales negatively with the δ(15) N of prey. We collated existing fish community δ(15) N-body size data for the Northeast Atlantic and tropical Western Arabian Sea with new data from the Northeast Pacific. These data were used to estimate TL-body mass relationships and PPMR under constant and scaled Δδ(15) N assumptions, and to assess how the scaled Δδ(15) N assumption affects our understanding of the structure of these food webs. Adoption of the scaled Δδ(15) N approach markedly reduces the previously reported differences in TL at body mass among fish communities from different regions. With scaled Δδ(15) N, TL-body mass relationships became more positive and PPMR fell. Results implied that realized prey size in these size-structured fish communities are less variable than previously assumed and food chains potentially longer. The adoption of generic PPMR estimates for calibration and validation of size-based fish community models is better supported than hitherto assumed, but predicted slopes of community size spectra are more sensitive to a given change or error in realized PPMR when PPMR is small.

Shift in a large river fish assemblage: body-size and trophic structure dynamics.

As the intensity and speed of environmental change increase at both local and global scales it is imperative that we gain a better understanding of the ecological implications of community shifts. While there has been substantial progress toward understanding the drivers and subsequent responses of community change (e.g. lake trophic state), the ecological impacts of food web changes are far less understood. We analyzed Wabash River fish assemblage data collected from 1974-2008, to evaluate temporal variation in body-size structure and functional group composition. Two parameters derived from annual community size-spectra were our major response variables: (1) the regression slope is an index of ecological efficiency and predator-prey biomass ratios, and (2) spectral elevation (regression midpoint height) is a proxy for food web capacity. We detected a large assemblage shift, over at least a seven year period, defined by dramatic changes in abundance (measured as catch-per-unit-effort) of the dominant functional feeding groups among two time periods; from an assemblage dominated by planktivore-omnivores to benthic invertivores. There was a concurrent increase in ecological efficiency (slopes increased over time) following the shift associated with an increase in large-bodied low trophic level fish. Food web capacity remained relatively stable with no clear temporal trends. Thus, increased ecological efficiency occurred simultaneous to a compensatory response that shifted biomass among functional feeding groups.

Interactions between temperature and nutrients across levels of ecological organization.

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits--growth, respiration, body size, and elemental content--that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population-level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature-nutrient interactions influence processes at the whole-ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature-nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.

Mizer: an R package for multispecies, trait-based and community size spectrum ecological modelling.

Size spectrum ecological models are representations of a community of individuals which grow and change trophic level. A key emergent feature of these models is the size spectrum; the total abundance of all individuals that scales negatively with size. The models we focus on are designed to capture fish community dynamics useful for assessing the community impacts of fishing.We present mizer, an R package for implementing dynamic size spectrum ecological models of an entire aquatic community subject to fishing. Multiple fishing gears can be defined and fishing mortality can change through time making it possible to simulate a range of exploitation strategies and management options.mizer implements three versions of the size spectrum modelling framework: the community model, where individuals are only characterized by their size; the trait-based model, where individuals are further characterized by their asymptotic size; and the multispecies model where additional trait differences are resolved.A range of plot, community indicator and summary methods are available to inspect the results of the simulations.

Scaling body size fluctuations

The size of an organism matters for its metabolic, growth, mortality, and other vital rates. Scale-free community size spectra (i.e., size distributions regardless of species) are routinely observed in natural ecosystems and are the product of intra- and interspecies regulation of the relative abundance of organisms of different sizes. Intra- and interspecies distributions of body sizes are thus major determinants of ecosystems' structure and function. We show experimentally that single-species mass distributions of unicellular eukaryotes covering different phyla exhibit both characteristic sizes and universal features over more than four orders of magnitude in mass. Remarkably, we find that the mean size of a species is sufficient to characterize its size distribution fully and that the latter has a universal form across all species. We show that an analytical physiological model accounts for the observed universality, which can be synthesized in a log-normal form for the intraspecies size distributions. We also propose how ecological and physiological processes should interact to produce scale-invariant community size spectra and discuss the implications of our results on allometric scaling laws involving body mass.

Investigation of Fishing and Climate Effects on the Community Size Spectra of Eastern Bering Sea Fish

AbstractThe eastern Bering Sea (EBS) is a highly productive subarctic marine ecosystem that is exposed to considerable climate variability and is noted for conservative management of its fishery resources. The community size spectrum (CSS; relationship between animal abundance and size) of fish captured in EBS shelf bottom trawl surveys was examined for evidence of change over time and fishing and climate effects. The slope (indicative of fish size) and height (indicative of ecosystem productivity, or essentially fish abundance) of the CSS can change due to changes in fishing intensity and climate variability. Linear trends were not observed in EBS groundfish size or abundance during 1982–2006. The abundance of large‐sized fish increased in 2001–2006 relative to 1982–2000. Observed changes in CSS height partially corroborated evidence for decreased productivity of the main fishery target species since the late 1990s, which may have been due to changes in relative species composition. In addition, abundance and size composition of nontarget fish decreased during 1982–2006, possibly related to changes in water temperature. In contrast, however, the size and productivity of fish that were primarily bycatch species increased during 1982–2006. Unlike in other ecosystems, changes in CSS slopes and heights for the EBS were not related to exploitation rates. Changes in the abundance of fish in areas that were normally occupied by cold bottom water (northwest inner and middle shelf) may have been related to the effects of temperature on fish distribution.

Warming alters the size spectrum and shifts the distribution of biomass in freshwater ecosystems

Organism size is one of the key determinants of community structure, and its relationship with abundance can describe how biomass is partitioned among the biota within an ecosystem. An outdoor freshwater mesocosm experiment was used to determine how warming of∼4 °C would affect the size, biomass and taxonomic structure of planktonic communities. Warming increased the steepness of the community size spectrum by increasing the prevalence of small organisms, primarily within the phytoplankton assemblage and it also reduced the mean and maximum size of phytoplankton by approximately one order of magnitude. The observed shifts in phytoplankton size structure were reflected in changes in phytoplankton community composition, though zooplankton taxonomic composition was unaffected by warming. Furthermore, warming reduced community biomass and total phytoplankton biomass, although zooplankton biomass was unaffected. This resulted in an increase in the zooplankton to phytoplankton biomass ratio in the warmed mesocosms, which could be explained by faster turnover within the phytoplankton assemblages. Overall, warming shifted the distribution of phytoplankton size towards smaller individuals with rapid turnover and low standing biomass, resulting in a reorganization of the biomass structure of the food webs. These results indicate future environmental warming may have profound effects on the structure and functioning of aquatic communities and ecosystems.

Scaling in Ecosystems and the Linkage of Macroecological Laws

Scaling provides an elegant framework for understanding power-law behavior and deducing relationships between critical exponents. We demonstrate that scaling theory can be generalized to develop a framework for the analysis of diverse empirical macroecological relationships traditionally treated as independent. Our mathematical arguments predict links between the species-area relationship, the relative species abundance and community size spectra in excellent accord with empirical data.

Latitudinal variation in plankton size spectra in the Atlantic Ocean

Abstract The size distribution of the pelagic community has the potential to compare ecosystems with different species composition as well as to identify the main functional properties of the system. Plankton size spectra are an effective approach of summarising the size structure of the community and have the potential to indicate the transfer of energy up the trophic food web. However, data on which open ocean plankton biomass-size structure can be constructed are scarce. Here we present the latitudinal distribution in the Atlantic Ocean of normalised biomass-size spectra (NB-S), plankton biomass and abundance, as well as mean zooplankton size between 70°N and 50°S. Samples were collected from three Atlantic Meridional Transect (AMT) cruises during May/June 2003, September/October 2003 and April/June 2004, as well as from a Marine Productivity (MarProd) cruise farther north in the Irminger Sea during spring 2002. The biomass-size distribution covered a body size range from nano- to mesoplankton and was based on a depth range of 50–0 m on the AMT and 120–0 m on the Irminger Sea cruise. The distribution of normalised biomass versus size was linear on a double log plot at all of the 95 stations. The slopes of the NB-S spectra ranged from −0.93 to −1.26 and −1.12 to −1.46 on the AMT and Irminger Sea cruises, respectively. A “dome-shaped” pattern in the slopes of community size spectra was observed in the Atlantic, indicating a decrease in the trophic transfer efficiency of energy with increasing latitude and phytoplankton biomass. Mesozooplankton biomass, abundance, and mean size followed a distribution similar to ecosystem productivity.

Spatial distribution of zooplankton size spectra on the French continental shelf of the Bay of Biscay during spring 2000 and 2001

During two cruises in springtime (18 March to 13 April 2000 and 27 March to 4 June 2001), the whole French continental shelf of the Bay of Biscay was sampled to obtain an overview on the zooplankton community size structure. A laboratory optical plankton counter (OPC‐1L) was used to process plankton net tow samples and estimate abundance, biovolume, and general characteristics of size spectra. In a second step, biomass estimates were extrapolated from size by using a conversion factor. Both biomass and abundance estimates show spatial patterns with a clear coastal‐open sea gradient for both years. The coastal area was characterized by the highest biomasses and abundances per volume. A first analysis of the zooplankton community size spectra was made by using the slope of the normalized biomass size spectrum. Different spatial patterns of zooplankton size spectra were highlighted for spring 2000 and 2001. The highest slopes were found for the coastal zone, showing a large ratio of small organisms, although this was less marked in the springtime 2000. Stations characterized by high proportions of large organisms were located in majority in the north of the bay and at the shelf break. A second analysis using the size probability distributions of organisms revealed a nearly permanent nonlinearity of probability distributions. This implied the community structure was not in an equilibrium state during spring and this nonlinearity could be locally related to dominant species dynamics.

Seasonal change of community structure and size spectra of zooplankton in the Kaw River estuary (French Guiana)

The zooplankton community of the Kaw River estuary in French Guiana and its relationship to its physical and chemical environment are described. Sampling was carried out in November 1998, June 1999, and November 2001, corresponding to the end of the dry season in 1998 and 2001 and to the end of the rainy season in 1999. The Kaw is a small coastal river that drains a vast swamp and flows into the Atlantic Ocean, close to the equator and to the mouth of the Amazon. The hydrodynamic and hydrochemical functioning of the estuary, and the development of littoral mangroves, is strongly constrained by the transport of sediments from the Amazon. During the rainy season the zooplankton, originating from the highly oligotrophic upstream sectors of the Kaw River, was of a strictly freshwater type. It was highly dispersed within the flooding water and dominated (in terms of density) by highly diversified rotifer taxa and (in terms of biomass) by cladocerans. During the dry season the estuary, turbid but enriched by exports from adjacent mudflats and mangroves, was colonized by a large and abundant zooplankton community that was dominated by tintinnids, with copepods as their main associated taxa. Exploiting the rich autochthonous phytoplankton and allochthonous phytobenthos, the microzooplankton components were only constrained when strong hydrodynamic exchanges allowed mixing between the opportunistic estuarine community and coastal mesozooplankton (copepods, chaetognaths, bivalve veligers). Between 1998 and 2001, these two communities were partially isolated from each other due to the gradual arrival of a mud bank, causing the blockage of the estuary. This isolation resulted in the under-exploitation of the microzooplankton. The intensification of tidal currents (spring tide) that occurred during the subsequent stabilization phase of the mudflat induced a more balanced zooplankton community. While the “estuarine” zooplankton of the Kaw River estuary therefore relies on the relative isolation of its water mass, its contribution to the coastal ecosystem also implies the existence of strong tidal currents that temporarily break this isolation.