Interdependence of plankton spatial distribution and plancton biomass temporal oscillations: mathematical simulation

The dynamics of aquatic biological communities in a patchy environment is of great interest in respect to interrelations between phenomena at various spatial and time scales. To study the complex plankton dynamics in relation to variations of such a biologically essential parameter as the fish predation rate, we use a simple reaction-diffusion model of trophic interactions between phytoplankton, zooplankton, and fish. We suggest that plankton is distributed between two habitats one of which is fish-free due to hydrological inhomogeneity, while the other is fish-populated. We show that temporal variations in the fish predation rate do not violate the strong correspondence between the character of spatial distribution of plankton and changes of plankton biomass in time: regular temporal oscillations of plankton biomass correspond to large-scale plankton patches, while chaotic oscillations correspond to small-scale plankton patterns. As in the case of the constant fish predation rate, the chaotic plankton dynamics is characterized by coexistence of the chaotic attractor and limit cycle.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

Quantifying the interplay between environmental and social effects on aggregated-fish dynamics

Demonstrating and quantifying the respective roles of social interactions and external stimuli governing fish dynamics is key to understanding fish spatial distribution. If seminal studies have contributed to our understanding of fish spatial organization in schools, little experimental information is available on fish in their natural environment, where aggregations often occur in the presence of spatial heterogeneities. Here, we applied novel modeling approaches coupled to accurate acoustic tracking for studying the dynamics of a group of gregarious fish in a heterogeneous environment. To this purpose, we acoustically tracked with submeter resolution the positions of twelve small pelagic fish (Selar crumenophthalmus) in the presence of an anchored floating object, constituting a point of attraction for several fish species. We constructed a field-based model for aggregated-fish dynamics, deriving effective interactions for both social and external stimuli from experiments. We tuned the model parameters that best fit the experimental data and quantified the importance of social interactions in the aggregation, providing an explanation for the spatial structure of fish aggregations found around floating objects. Our results can be generalized to other gregarious species and contexts as long as it is possible to observe the fine-scale movements of a subset of individuals.     

Biotic invasions can alter nutritional composition of zooplankton communities

Ecologists and ecosystem managers often base their understanding of trophic dynamics on consumer and resource biomass. However, the factors that alter the relative nutritional value of resources are often poorly understood, despite their potential to decouple trophic interactions. Recent population declines in pelagic fishes of the upper San Francisco Estuary were not accompanied by an equivalent decrease in zooplankton biomass, which are the main resource for the fish and their larvae. It was hypothesized that changes in zooplankton nutritional conditions following the establishment of invasive species caused food‐quality related limitations for these higher‐order consumers. Using stable isotopes, elemental stoichiometry and fatty acid analyses for all dominant invasive and native zooplankton taxa and seston, we characterized the plankton community structure in the estuary and demonstrated taxon‐specific differences in their nutritional value. We then quantified the temporal dynamics in meso‐zooplankton proportions of docosahexaenoic acid (DHA), eicosapentaenoic acid (EPA), polyunsaturated fatty acids (PUFA), and ratio of n3:n6 fatty acids. We found temporal increase in the community‐level DHA, n3 to n6 fatty acid ratio, decrease in the community‐level EPA and PUFA in the brackish water region, but no change in the bulk PUFA proportions in the freshwater region of the estuary. These changes were caused mainly by declines of native cladocerans that are rich in EPA and by an increase in the dominance of invasive taxa with high DHA concentrations, similar to that of native taxa. Although we showed temporal shifts in individual fatty acid classes, the proportion of the essential fatty acids remained relatively high, suggesting that nutritional prey availability for fish remained unchanged with the shift in species composition. We argue that the nutritional content of resource communities should be considered when analyzing the long‐term trophic dynamics and designing effective management and restoration strategies.     

Towards resolving the paradox of enrichment: the impact of zooplankton vertical migrations on plankton systems stability

Eutrophication, often resulting from human activity, is a serious threat to aquatic communities. Theoretical analysis of this phenomenon, based on conceptual mathematical models, leads to controversial predictions known as Rosenzweig's paradox of enrichment. At the same time, field observations demonstrate that real plankton communities exhibit various mechanisms of self-regulation which can buffer negative effects of enrichment. In this paper, we study potential effects of zooplankton vertical migration on stability of plankton systems functioning. We consider an intrinsically unstable plankton model, which is characterized by an unlimited phytoplankton multiplication and population oscillations of increasing amplitude, and investigate whether vertical migrations of zooplankton can stabilize such a system at low plankton densities. By means of developing two different models accounting for different ecological situations, e.g. deep waters and shallow waters, we show that vertical migrations of zooplankton can result in stabilization of eutrophic plankton systems. Thus, we show that this mechanism, rarely taken into account in models of plankton dynamics, may be important for resolving the paradox of enrichment in plankton communities.     

Zooplankton biomass variation in relation to climatic conditions in the Barents Sea

Over the last two decades, there have been large changes in the zooplankton biomass in the Barents Sea. These biomass variations are mainly attributed to predation pressure and environmental factors (e.g. advective transport). When stock size of capelin (Mallotus villosus), a major planktivorous fish in the Barents Sea ecosystem, was quite low as in 1986 and 1994, the zooplankton biomass showed marked increase. However, the increase in the zooplankton biomass occurred in different water masses during 1986 and 1994. In 1986, a climatically cold year, the plankton biomass was highest in the Arctic waters of the northeastern Barents Sea. This is probably due to the increase in larger Arctic amphipod species, such as Themisto libellula. In 1994, a climatically warm year, the zooplankton biomass was high in the Atlantic waters of the southwestern Barents Sea. The large increase in zooplankton biomass in the Atlantic waters in 1994 was presumably due to the higher inflow of advected organisms, e.g. Calanus spp., as well as high temperatures, which may lead to high growth rates of zooplankton. Throughout the studied region, the plankton biomass in the "cold year" of 1986 was generally much lower than in the "warm year" of 1994.     

State of the Ichthyoplankton Community along the Crimean Peninsula in August 2011

Data on species composition, spatial distribution of ichthyoplankton, and feeding of fish larvae in August 2011 are presented for the shelf and the open waters along the Crimean Peninsula from Kerch Strait to Karkinit Bay. In the ichthyoplankton, fish eggs and larvae represented 19 species from 16 families. The average egg abundance was 111 egg/m² and larvae abundance was 22 ind./m². The elimination ratio of the European anchovy Engraulis encrasicolus, which dominates in the plankton from the final stages of development to the prelarvae in 2011, was similar to that observed in 1957. The high number of larvae of the older age group in the western halistatic zone is explained by the peculiarities of the hydrological regime. Decrease in the pressure of predatory jelly macroplankton and an fodder zooplankton (zooplankton abundance) contribute to the survival of the fish larvae.     

Rising temperature and marine plankton community dynamics: Is warming bad?

Global warming is a major threat to the natural environment worldwide with potential adverse impact on plankton community. This will ultimately lead to a change in the dynamics of aquatic food webs. In this study we used seasonally forced multi-species version of the classic Rosenzweig–MacArthur predator–prey model to understand the role and stochastic influence of increasing temperature on marine plankton. First, stable coexistence of four phytoplankton and three zooplankton species was created in a system and then the level of temperature changed to achieve our research goal. We found that the stable coexistence of phytoplankton and zooplankton was related to periodic shifts in species biomass, variation in inter-specific competition and niche configuration. Warming significantly reduced total plankton biomass and changed turnover time of a species, with gradual warming breaking the stable coexistence of phytoplankton and zooplankton. In addition, we found that warming make specialist species more vulnerable than generalist species. After adding noise, a significant variation was observed in plankton biomass and amplification of noise was higher for phytoplankton compared to zooplankton. These results suggest that stochastic or unpredictable nature of temperature fluctuations may create a window of opportunity for the emergence of new species. Overall, warming would induce a shift in plankton dynamics and thereby exert pressure on plankton dependent communities such as fish in the long run.     

Evidence for Changes in Estuarine Zooplankton Fostered by Increased Climate Variance

Estuaries are among the most valuable aquatic systems in terms of their services to human welfare. They offer an ideal framework to assess multiscale processes linking climate and food web dynamics through the hydrological cycle. Resolving food web responses to climate change is fundamental to resilience management of these threatened ecosystems under global change scenarios. Here, we examined the temporal variability of the plankton food web in the Mondego Estuary, central Iberian Peninsula, over the period 2003 to 2012. The results pointed out a cascading effect from climate to plankton communities that follow a non-stationary behavior shaped by the climate variance envelope. Concurrent changes in hydrographic processes at the regional, that is, upwelling intensity, and local, that is, estuarine hydrology, scales were driven by climatic forcing promoted by the North Atlantic Oscillation; the influence of which permeated the physical environment in the estuary affecting both autotrophic and heterotrophic communities. The most conspicuous change arose around 2008 and consisted of an obvious decrease in freshwater taxa along with a noticeable increase in marine organisms, mainly driven by gelatinous zooplankton. The observed increase in small-sized cosmopolitan copepods, that is, Clausocalanus arcuicornis, Oithona plumifera, thermophilic species, that is, Penilia avirostris, and gelatinous zooplankton suggests a structural change in the Mondego plankton community. These results provide empirical support to the expectation that expanding climate variance changes plankton structure and functioning, likely fostering trophic interactions in pelagic food webs.     

Chaos and order in plankton dynamics. Complex behavior of a simple model

The role of the diffusive interaction between fish-populated and fish-free habitats in a patchy environment in plankton pattern formation is studied by means of a minimal reaction-diffusion model of the nutrient-plankton-fish food chain. It is shown that such interaction can give rise to spatio-temporal plankton patterns. The fractal dimension of the patterns is shown dependent on the fish predation rate. The spatially averaged plankton dynamics depending on both fish predation rate and distance between fish-populated habitats can exhibit chaotic and regular behavior. The chaotic plankton dynamics is characteristic of a wide parameter range.     

Nutrients and toxin producing phytoplankton control algal blooms — a spatio-temporal study in a noisy environment

A phytoplankton-zooplankton prey-predator model has been investigated for temporal, spatial and spatio-temporal dissipative pattern formation in a deterministic and noisy environment, respectively. The overall carrying capacity for the phytoplankton population depends on the nutrient level. The role of nutrient concentrations and toxin producing phytoplankton for controlling the algal blooms has been discussed. The local analysis yields a number of stationary and/or oscillatory regimes and their combinations. Correspondingly interesting is the spatio-temporal behaviour, modelled by stochastic reaction-diffusion equations. The present study also reveals the fact that the rate of toxin production by toxin producing phytoplankton (TPP) plays an important role for controlling oscillations in the plankton system. We also observe that different mortality functions of zooplankton due to TPP have significant influence in controlling oscillations, coexistence, survival or extinction of the zoo-plankton population. External noise can enhance the survival and spread of zooplankton that would go extinct in the deterministic system due to a high rate of toxin production.     

Water level and fish-mediated cascading effects on the microbial community in eutrophic warm shallow lakes: a mesocosm experiment

Information on the effects of water level changes on microbial planktonic communities in lakes is limited but vital for understanding ecosystem dynamics in Mediterranean lakes subjected to major intra- and inter-annual variations in water level. We performed an in situ mesocosm experiment in an eutrophic Turkish lake at two different depths crossed with presence/absence of fish in order to explore the effects of water level variations and the role of top-down regulation at contrasting depths. Strong effects of fish were found on zooplankton, weakening through the food chain to ciliates, HNF and bacterioplankton, whereas the effect of water level variations was overall modest. Presence of fish resulted in lower biomass of zooplankton and higher biomasses of phytoplankton, ciliates and total plankton. The cascading effects of fish were strongest in the shallow mesocosms as evidenced by a lower zooplankton contribution to total plankton biomass and lower zooplankton:ciliate and HNF:bacteria biomass ratios. Our results suggest that a lowering of the water level in warm shallow lakes will enhance the contribution of bacteria, HNF and ciliates to the plankton biomass, likely due to increased density of submerged macrophytes (less phytoplankton); this effect will, however, be less pronounced in the presence of fish.     

Time delay as a key factor of model plankton dynamics

Studies of the mechanisms underlying complex dynamics of ecological systems at various spatial and time scales bring increasing awareness that complexity is an intrinsic feature of ecological functioning. This paper is to investigate the role of such an ecologically significant parameter as the time delay due to maturation processes in the complex plankton dynamics. We show that the time lag T1, associated with the zooplankton maturation period can lead to essential changes in the plankton dynamics. Particularly, we show that the coexistence of limit cycle and chaotic attractor we have recently found to be typical of the system at T1 = 0 [A.B. Medvinsky, I.A. Tikhonova, R.R. Aliev, B.-L. Li, Z.-S. Lin, H. Malchow, Patchy environment as a factor of complex plankton dynamics, Phys. Rev. E 64 (2001) 021915] is replaced by pure chaotic plankton dynamics as T1 becomes more than a critical value. The results obtained imply that chaos is a rather common phenomenon in the plankton functioning.     

Seasonal strengths of the abiotic and biotic drivers of a zooplankton community

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Long-term dynamics of Asplanchna priodonta in Lake Windsborn with comments on the diet

We evaluate trophic interactions between the rotifer Asplanchna priodonta and the plankton community of Lake Windsborn (Eifel District, Germany). The long-term population dynamics of this species and of the two most abundant zooplankton species, Keratella cochlearis and Bosmina longirostris, were analyzed in relation to hydrochemical and climatic variates. The maximum abundance of A. priodonta was limited to the summer months with temperature being the only significant predictor of abundance. Intrinsic rate of growth and abundance of the three species were independent from each other suggesting the absence of trophic interactions. In Lake Windsborn, A. priodonta was a grazer rather than a predator. Its diet included high proportions of colonial algae and cyanobacteria which cannot be ingested by the other abundant zooplankton species, whereas zooplanktonic food were rarely found. Thus, our study supports the hypothesis that A. priodonta is an opportunistic feeder whose trophic position entirely depends on the structure of the plankton community.     

Long-period endogenous oscillations in fish population size: Mathematical modeling

We present a mathematical model of an aquatic community, where the size-and-age structure of hydrobiont populations is taken into account and the corresponding trophic interactions between zooplankton, peaceful fish, and predatory fish are described. We show that interactions between separate components of the aquatic community can give rise to long-period oscillations in fish population size. The period of these oscillations is on the order of decades. With this model we also show that an increase in the zooplankton growth rate may entail a sequence of bifurcations in the fish population dynamics: steady states → regular oscillations → quasicycles → dynamic chaos.     

Predators, resources, and trophic chains in the regulation of plankton population and biomass in oligothrophic lakes

The relative strength of "top-down" versus "bottom-up" control of plankton community structure and biomass in two small oligotrophic lakes (with and without fish), located near the Polar circle (Russia), has been investigated for two years, 1996 and 1997. The comparative analyses of zooplankton biomass and species abundance showed strong negative effect of fish, stickeback (Pungitius pungitius L.), on the zooplankton community species, size structure and biomass of particular prey species but no effect on the biomass of the whole trophic level. An intensive predation in Verkhneye lake has lead to: 1) sixfold decline in biomass of large cladoceran Holopedium gibberum comparing to the lake lacking predator, 2) shift in the size mode in zooplankton community and the replacement of the typical large grazers by small species--Bosmina longirostris and rotifers. Their abundance and biomass even increased, demonstrating the stimulating effect of fish on the "inefficient" and unprofitable prey organisms. The analysis of contributions of different factors into the cladoceran's birth rate changes was applied to demonstrate the relative impact of predators and resources on zooplankton abundance. An occasional introduction of the stickleback to Vodoprovodnoye lake (the reference lake in 1996) in summer 1997 lead to drastic canges in this ecosystem: devastating decrease of zooplankton biomass and complete elimination of five previously dominant grazer species. The abundance of edible phytoplankton was slightly higher in the lake with fish in 1996 and considerably higher in the lake where fish has appeared in 1997 showing the prevailing "top-down" control of phytoplankton in oligotrophic ecosystem. The reasons of trophic cascade appearance in oligotrophic lakes are also discussed.     

Phytoplankton blooms and fish recruitment rate: Effects of spatial distribution

We consider the spatio-temporal dynamics of a spatially-structured generalization of the phytoplankton-zooplankton-fish larvae model system proposed earlier (Biktashev et al., 2003, J. Plankton Res. 5, 21–33; James et al., 2003, Ecol. Model. 160, 77–90). In contrast to Pitchford and Brindley (2001, Bull. Math. Biol. 63, 527–546), who were concerned with small scale patchiness (i.e., 1–10m), on which the (stochastic) raptorial behaviour of individual larvae is important, we address here the much larger scale ‘patchy’ problems (i.e., 10–100 km), on which both larvae and plankton may be regarded as passive tracers of the fluid motion, dispersed and mixed by the turbulent diffusion processes. In particular, we study the dependence of the fish recruitment on carrying capacities of the plankton subsystem and on spatio-temporal evolution of that subsystem with respect to the larvae hatching site(s). It is shown that the main features found both in the nonstructured and age-structured spatially uniform models are observed in the spatially structured case, but that spatial effects can significantly modify the overall quantitative outcome. Spatial patterns in the metamorphosed fish distribution are a consequence of quasi-local interaction of larvae with plankton, in which the dispersion of larvae by large scale turbulent eddies plays little part due to the relatively short timescale of the larvae development. As a result, in a strong phyto/zooplankton subsystem, with fast reproduction rate and large carrying capacity of phytoplankton and high conversion ratio of zooplankton, recruitment success depends only on the localization and timing of the hatching with respect to the plankton patches. In a weak phyto/zooplankton system, with slow reproduction rate and small carrying capacity of phytoplankton and low conversion ratio of zooplankton, the larvae may significantly influence the evolution of the plankton patches, which may lead to nontrivial cooperative effects between different patches of larvae. However, in this case, recruitment is very low.