Linking multi-level population dynamics: state, role, and population.

The dynamics of an ecological community can be described at different focal scales of the species, such as individual states or the population level. More detailed descriptions of ecological dynamics offer more information, but produce more complex models that are difficult to analyze. Adequately controlling the model complexity and the availability of multiple descriptions of the concerned dynamics maximizes our understanding of ecological dynamics. One of the central goals of ecological studies is to develop links between multiple descriptions of an ecological community. In this article, starting from a nonlinear state-level description of an ecological community (generalized McKendrick-von Foerster model), role-level and population-level descriptions (Lotka-Volterra model) are derived in a consistent manner. The role-level description covers a wider range of situations than the population-level description. However, using the established connections, it is demonstrated that the population-level description can be used to predict the equilibrium status of the role-level description. This approach connects state-, role-, and population-level dynamics consistently, and offers a justification for the multiple choices of model description.

Poor nutritional quality of primary producers and zooplankton driven by eutrophication is mitigated at upper trophic levels.

Eutrophication and rising water temperature in freshwaters may increase the total production of a lake while simultaneously reducing the nutritional quality of food web components. We evaluated how cyanobacteria blooms, driven by agricultural eutrophication (in eutrophic Lake Köyliöjärvi) or global warming (in mesotrophic Lake Pyhäjärvi), influence the biomass and structure of phytoplankton, zooplankton, and fish communities. In terms of the nutritional value of food web components, we evaluated changes in the ω-3 and ω-6 polyunsaturated fatty acids (PUFA) of phytoplankton and consumers at different trophic levels. Meanwhile, the lakes did not differ in their biomasses of phytoplankton, zooplankton, and fish communities, lake trophic status greatly influenced the community structures. The eutrophic lake, with agricultural eutrophication, had cyanobacteria bloom throughout the summer months whereas cyanobacteria were abundant only occasionally in the mesotrophic lake, mainly in early summer. Phytoplankton community differences at genus level resulted in higher arachidonic acid, eicosapentaenoic acid (EPA), and docosahexaenoic acid (DHA) content of seston in the mesotrophic than in the eutrophic lake. This was also reflected in the EPA and DHA content of herbivorous zooplankton (Daphnia and Bosmina) despite more efficient trophic retention of these biomolecules in a eutrophic lake than in the mesotrophic lake zooplankton. Planktivorous juvenile fish (perch and roach) in a eutrophic lake overcame the lower availability of DHA in their prey by more efficient trophic retention and biosynthesis from the precursors. However, the most efficient trophic retention of DHA was found with benthivorous perch which prey contained only a low amount of DHA. Long-term cyanobacterial blooming decreased the nutritional quality of piscivorous perch; however, the difference was much less than previously anticipated. Our result shows that long-term cyanobacteria blooming impacts the structure of plankton and fish communities and lowers the nutritional quality of seston and zooplankton, which, however, is mitigated at upper trophic levels.

Ontogenetic shift and feeding habits of the European hake (Merluccius merluccius L., 1758) in Central and Southern Tyrrhenian Sea (Western Mediterranean Sea): A comparison between past and present data.

The present paper aims to investigate the ecological role of Merluccius merluccius, Linnaeus, 1758, in southern and central Tyrrhenian Sea (GSA 10, Resolution GFCM/33/2009/2 General Fisheries Commission for the Mediterranean), analyzing ontogenetic diet shifts, geographical variations on prey composition, and feeding habits. A total of 734 hake specimens ranging in size between 6 cm and 73 cm (Total Length, TL) were collected in 2018. In order to evaluate ontogenetic shifts in prey composition, samples were divided into five size classes and for each class the quantitative feeding indices have been calculated. The statistical analysis, based on index of relative importance percentage (%IRI), resulted in three trophic groups. The most abundant prey found in the immature hake specimens (size class I) were the Euphausiids, Stylocheiron longicorne and Mysidacea, while for samples with a total length over 10.5 cm were crustaceans and fish. Engraulis encrasicolus was the most abundant fish prey identified, followed by Boops boops and Myctophids. The high presence of Euphausiids, Mysids, Myctophidae, and Sternoptychidae in classes I, II, II, and IV (6-23 cm) showed the relevant role of mesopelagic fauna in hake diets, with an essential organic matter and energy flow from the mesopelagic to the epipelagic environment. Additionally, decapod crustaceans were found in the stomach contents of hakes belonging to class V (with size over 36 cm TL), which is notable considering that our study area includes an important decapod crustacean fishing area.

Fang shape varies with ontogeny and sex in the venomous elapid snake Pseudonaja affinis.

A predator's preferred prey often changes over the course of its life as it grows from an inexperienced juvenile through to a sexually mature adult. For species with highly specialised feeding strategies, this may require its anatomy to change over the course of its life. The dugite (Pseudonaja affinis, Günther 1872) is a venomous snake from Australia that displays such a diet shift, with juveniles feeding on small reptiles, while adults mainly target mammals. We examined the morphology of fangs across both sexes and throughout ontogeny using geometric morphometrics and cross-sectional sharpness measurements of key functional regions on these teeth. This highlighted key differences in shape that likely relate to the varied properties of their adult and juvenile diet. We found that juveniles display a more robust and blunter fang, which likely relates to feeding on scaly lizard prey, whereas adults have slender fangs with sharper tips, which reflects their diet of softer mammalian prey. There were also differences between males and females, with male snakes having significantly more slender fangs than females, which might be an indication of niche partitioning between the sexes. Using snout-vent length as a proxy for age, we found that the ontogenetic shift in fang shape occurs when P. affinis is around 60 cm long, corresponding with previous studies that found this size to be the moment where these snakes switch from their juvenile to adult diet.

Developmental Change in Predators Drives Different Community Configurations.

AbstractTheoreticians who first observed alternative stable states in simple ecological models warned of grave implications for unexpected and irreversible collapses of natural systems (i.e., regime shifts). Recent ecosystem-level shifts engendering considerable economic losses have validated this concern, positioning bistability at the vanguard of coupled human-environment systems management. While the perturbations that induce regime shifts are known, the ecological forces that uphold alternative stable states are often unresolved or complex and system specific. Thus, the search continues for general mechanisms that can produce alternative stable states under realistic conditions. Integrating model predictions with long-term zooplankton community experiments, we show that the core feature of ontogenetic development, food-dependent maturation, enables a single community to reach different configurations within the same constant environment. In one configuration, predators regulate prey to foster coexistence, while in the other, prey counterintuitively exclude their predators via maturation-limiting competition. The concordance of these findings with the unique outcome and underlying mechanism of a general model provides empirical evidence that developmental change, a fundamental property of life, can support bistability of natural systems.

Trade-Offs with Growth Limit Host Range in Complex Life-Cycle Helminths.

AbstractParasitic worms with complex life cycles have several developmental stages, with each stage creating opportunities to infect additional host species. Using a data set for 973 species of trophically transmitted acanthocephalans, cestodes, and nematodes, we confirmed that worms with longer life cycles (i.e., more successive hosts) infect a greater diversity of host species and taxa (after controlling for study effort). Generalism at the stage level was highest for middle life stages, the second and third intermediate hosts of long life cycles. By simulating life cycles in real food webs, we found that middle stages had more potential host species to infect, suggesting that opportunity constrains generalism. However, parasites usually infected fewer host species than expected from simulated cycles, suggesting that generalism has costs. There was no trade-off in generalism from one stage to the next, but worms spent less time growing and developing in stages where they infected more taxonomically diverse hosts. Our results demonstrate that life-cycle complexity favors high generalism and that host use across life stages is determined by both ecological opportunity and life-history trade-offs.

Functional traits explain trophic allometries of cephalopods.

Individual body size strongly influences the trophic role of marine organisms and the structure and function of marine ecosystems. Quantifying trophic position-individual body size relationships (trophic allometries) underpins the development of size-structured ecosystem models to predict abundance and the transfer of energy through ecosystems. Trophic allometries are well studied for fishes but remain relatively unexplored for cephalopods. Cephalopods are important components of coastal, oceanic and deep-sea ecosystems, and they play a key role in the transfer of biomass from low trophic positions to higher predators. It is therefore important to resolve cephalopod trophic allometries to accurately represent them within size-structured ecosystem models. We assessed the trophic positions of cephalopods in an oceanic pelagic (0-500 m) community (sampled by trawling in a cold-core eddy in the western Tasman Sea), comprising 22 species from 12 families, using bulk tissue stable isotope analysis and amino acid compound-specific stable isotope analysis. We assessed whether ontogenetic trophic position shifts were evident at the species-level and tested for the best predictor of community-level trophic allometry among body size, taxonomy and functional grouping (informed by fin and mantle morphology). Individuals in this cephalopod community spanned two trophic positions and fell into three functional groups on an activity level gradient: low, medium and high. The relationship between trophic position and ontogeny varied among species, with the most marked differences evident between species from different functional groups. Activity-level-based functional group and individual body size are best explained by cephalopod trophic positions (marginal R2  = 0.43). Our results suggest that the morphological traits used to infer activity level, such as fin-to-mantle length ratio, fin musculature and mantle musculature are strong predictors of cephalopod trophic allometries. Contrary to established theory, not all cephalopods are voracious predators. Low activity level cephalopods have a distinct feeding mode, with low trophic positions and little-to-no ontogenetic increases. Given the important role of cephalopods in marine ecosystems, distinct feeding modes could have important consequences for energy pathways and ecosystem structure and function. These findings will facilitate trait-based and other model estimates of cephalopod abundance in the changing global ocean.

Contrasting size and fate of juvenile crown-of-thorns starfish linked to ontogenetic diet shifts.

Population dynamics of organisms are shaped by the variation in phenotypic traits, often expressed even among individuals from the same cohort. For example, individual variation in the timing of ontogenetic shifts in diet and/or habitat greatly influences subsequent growth and survival of some organisms, with critical effects on population dynamics. Few studies of natural systems have, however, demonstrated that marked phenotypic variation in growth rates or body size among individuals within a modelled cohort is linked to dietary shifts and food availability. Population irruptions of the crown-of-thorns starfish are one of the foremost contributors to the global degradation of coral reefs, but causes of irruptions have been debated for decades. Here we demonstrate, based on extensive field sampling of juvenile starfish (n = 3532), that marked variation in body size among juvenile starfish is linked to an ontogenetic diet shift from coralline algae to coral. This transition in diet leads to exponential growth in juveniles and is essential for individuals to reach maturity. Because smaller individuals experience higher mortality and growth is stunted on an algal diet, the ontogenetic shift to corallivory enhances individual fitness and replenishment success. Our findings suggest that the availability of coral prey facilitates early ontogenetic diet shifts and may be fundamental in initiating population irruptions.

Life-history genomic regions explain differences in Atlantic salmon marine diet specialization.

Animals employ various foraging strategies along their ontogeny to acquire energy, and with varying degree of efficiencies, to support growth, maturation and subsequent reproduction events. Individuals that can efficiently acquire energy early are more likely to mature at an earlier age, as a result of faster energy gain which can fuel maturation and reproduction. We aimed to test the hypothesis that heritable resource acquisition variation that covaries with efficiency along the ontogeny would influence maturation timing of individuals. To test this hypothesis, we utilized Atlantic salmon as a model which exhibits a simple, hence trackable, genetic control of maturation age. We then monitored the variation in diet acquisition (quantified as stomach fullness and composition) of individuals with different ages, and linked it with genomic regions (haploblocks) that were previously identified to be associated with age-at-maturity. Consistent with the hypothesis, we demonstrated that one of the life-history genomic regions tested (six6) was indeed associated with age-dependent differences in stomach fullness. Prey composition was marginally linked to six6, and suggestively (but non-significantly) to vgll3 genomic regions. We further showed Atlantic salmon switched to the so-called 'feast and famine' strategy along the ontogeny, where older age groups exhibited heavier stomach content, but that came at the expense of running on empty more often. These results suggest genetic variation underlying resource utilization may explain the genetic basis of age structure in Atlantic salmon. Given that ontogenetic diet has a genetic component and the strong spatial diversity associated with these genomic regions, we predict populations with diverse maturation age will have diverse evolutionary responses to future changes in marine food web structures.

Size structuring and allometric scaling relationships in coral reef fishes.

Temperate marine fish communities are often size-structured, with predators consuming increasingly larger prey and feeding at higher trophic levels as they grow. Gape limitation and ontogenetic diet shifts are key mechanisms by which size structuring arises in these communities. Little is known, however, about size structuring in coral reef fishes. Here, we aimed to advance understanding of size structuring in coral reef food webs by examining the evidence for these mechanisms in two groups of reef predators. Given the diversity of feeding modes amongst coral reef fishes, we also compared gape size-body size allometric relationships across functional groups to determine whether they are reliable indicators of size structuring. We used gut content analysis and quantile regressions of predator size-prey size relationships to test for evidence of gape limitation and ontogenetic niche shifts in reef piscivores (n = 13 species) and benthic invertivores (n = 3 species). We then estimated gape size-body size allometric scaling coefficients for 21 different species from four functional groups, including herbivores/detritivores, which are not expected to be gape-limited. We found evidence of both mechanisms for size structuring in coral reef piscivores, with maximum prey size scaling positively with predator body size, and ontogenetic diet shifts including prey type and expansion of prey size. There was, however, little evidence of size structuring in benthic invertivores. Across species and functional groups, absolute and relative gape sizes were largest in piscivores as expected, but gape size-body size scaling relationships were not indicative of size structuring. Instead, relative gape sizes and mouth morphologies may be better indicators. Our results provide evidence that coral reef piscivores are size-structured and that gape limitation and ontogenetic niche shifts are the mechanisms from which this structure arises. Although gape allometry was not indicative of size structuring, it may have implications for ecosystem function: positively allometric gape size-body size scaling relationships in herbivores/detritivores suggests that loss of large-bodied individuals of these species will have a disproportionately negative impact on reef grazing pressure.