A trophic framework for animal origins

Metazoans emerged in a microbial world and play a unique role in the biosphere as the only complex multicellular eukaryotes capable of phagocytosis. While the bodyplan and feeding mode of the last common metazoan ancestor remain unresolved, the earliest multicellular stem‐metazoans likely subsisted on picoplankton (planktonic microbes 0.2–2 μm in diameter) and dissolved organic matter (DOM), similarly to modern sponges. Once multicellular stem‐metazoans emerged, they conceivably modulated both the local availability of picoplankton, which they preferentially removed from the water column for feeding, and detrital particles 2–100 μm in diameter, which they expelled and deposited into the benthos as waste products. By influencing the availability of these heterotrophic food sources, the earliest multicellular stem‐metazoans would have acted as ecosystem engineers, helping create the ecological conditions under which other metazoans, namely detritivores and non‐sponge suspension feeders incapable of subsisting on picoplankton and DOM, could emerge and diversify. This early style of metazoan feeding, specifically the phagocytosis of small eukaryotic prey, could have also encouraged the evolution of larger, even multicellular, eukaryotic forms less prone to metazoan consumption. Therefore, the first multicellular stem‐metazoans, through their feeding, arguably helped bridge the strictly microbial food webs of the Proterozoic Eon (2.5–0.541 billion years ago) to the more macroscopic, metazoan‐sustaining food webs of the Phanerozoic Eon (0.541–0 billion years ago).     

The effect of dispersal between patches on the stability of large trophic food webs

Theoretical Ecology - Using computer simulations for the population dynamics of systems with many species, we investigate the stability of food webs distributed over several patches that are...     

Warming, plant phenology and the spatial dimension of trophic mismatch for large herbivores

Temporal advancement of resource availability by warming in seasonal environments can reduce reproductive success of vertebrates if their own reproductive phenology does not also advance with warming. Indirect evidence from large-scale analyses suggests, however, that migratory vertebrates might compensate for this by tracking phenological variation across landscapes. Results from our two-year warming experiment combined with seven years of observations of plant phenology and offspring production by caribou (Rangifer tarandus) in Greenland, however, contradict evidence from large-scale analyses. At spatial scales relevant to the foraging horizon of individual herbivores, spatial variability in plant phenology was reduced--not increased--by both experimental and observed warming. Concurrently, offspring production by female caribou declined with reductions in spatial variability in plant phenology. By highlighting the spatial dimension of trophic mismatch, these results reveal heretofore unexpected adverse consequences of climatic warming for herbivore population ecology.     

A cyanobacterial bloom prevents fish trophic cascades

1. We experimentally compared the impacts of visually feeding zooplanktivorous fish and filter‐feeding omnivorous fish in shallow tropical Dakar Bango reservoir, Senegal. We provoked a cyanobacterial Anabaena bloom under mesotrophic to eutrophic N‐limited conditions in 18 enclosures assigned to six Nile tilapia life‐stage treatments, at typical biomasses: fishless control (C), zooplanktivorous fry (Z), omnivorous juveniles (O), herbivorous fingerlings (H) and two combinations (OZ, OH). 2. All fish grew well, but as prevalent inedible phytoplankton dampened fish effects, community‐level trophic cascades did not occur. Planktivore types acted independently and affected differentially the biomasses of total zooplankton, cyclopoids, nauplii, cladocerans, invertebrate carnivores, large herbivores, colonial cyanobacteria and Chlorophyta. They neither influenced the total biomass of phytoplankton, nor most water chemistry characteristics. Responses were apparently not fish‐biomass related. The bloom collapsed synchronously in all enclosures, coinciding with enrichment ending, with a return to clear water within 12 days. 3. Our results support the hypothesis that excess nutrients and prevalent inedible cyanobacteria inhibit the cascading effects of natural biomass levels of both visually feeding zooplanktivores and filter‐feeding omnivores. In N‐limited meso‐eutrophic shallow tropical lakes with predominantly small herbivorous zooplankton, neither the type nor the biomass of planktivorous fish present seems likely to prevent the transient outburst of cyanobacterial blooms. Such fragile ecosystems may thus not sustain a trophic state suitable for drinking water production, unless human impacts are restricted. The generality of restoration approaches based on ecological engineering should be further explored.     

The role of cladocerans reflecting the trophic status of two large and shallow Estonian lakes

The role of pelagic cladoceran communities is discussed on the basis of a comparative study conducted in two Estonian lakes, the moderately eutrophic Lake Peipsi (N_tot 700, P_tot 40 μg l^?1 as average of ice-free period of 1997–2003) and in a strongly eutrophic Lake Võrtsjärv (N_tot 1600, P_tot 54 μg l^?1). The cladoceran community was found to reflect the differences in the trophic state of these lakes. In L. Peipsi, characteristic species of oligo-mesotrophic and eutrophic waters co-dominated (making up 20% or more of total zooplankton abundance or biomass), whereas in L. Võrtsjärv only species of eutrophic waters occurred. In L. Peipsi, the dominant cladocerans were Bosmina berolinensis and Daphnia galeata, while Chydorus sphaericus was the most abundant cladoceran in L. Võrtsjärv. The cladocerans of L. Peipsi (mean individual wet weight 25 μg) were significantly (threefold) larger than those of L. Võrtsjärv (8 μg). The mean wet biomass of cladocerans was higher and total cladoceran abundance was lower in L. Peipsi compared to L. Võrtsjärv (biomass varied from 0.133 to 1.570 g m^?3; mean value 0.800 g m^?3 in L. Peipsi and from 0.201 to 0.706 g m^?3, mean 0.400 g m^?3 in L. Võrtsjärv; the corresponding data for abundances were: 8,000–43,000 ind. m^?3, mean 30,000 ind. m^?3 for L. Peipsi, 50,000–100,000, mean 52,000 ind. m^?3 for L. Võrtsjärv). Based upon differences in body size, cladocerans were more effective transporters of energy in L. Peipsi than in L. Võrtsjärv. Cladocerans proved to be informative indicators of the trophic status and of the efficiency of the food web in studied lakes.     

A trophic diversity index for presence-absence food data

Summary An index to assess trophic diversity from presence-absence food data is proposed. The index is computed according to the expression , where the 's are the frequencies of occurrence of the various prey categories. The upper and lower limits of D are derived. A test of the reliability of D was carried out by comparing D and H (Shannon's information function) values obtained from a set of twenty-three food analyses from vertebrate animals. Results show that, although a significant correlation exists between D and H, only a small fraction of H-variation is explained by D-variation. D contains two kinds of information, one referred to species richness and another relative to the degree of between-samples heterogeneity. The former is shared in common with H and this presumably explains the fairly weak correlation found between both measures.     

A call for applying trophic structure in ecological restoration

Ecological restoration projects have traditionally focused on vegetation as both a means (seeding, planting, and substrate amendments) and ends (success based upon primary productivity and vegetation diversity). This vegetation‐centric approach to ecological restoration stems from an historic emphasis on esthetics and cost but provides a limited measure of total ecosystem functioning and overlooks alternative ways to achieve current and future restoration targets. We advocate a shift to planning beyond the plant community and toward the physical and biological components necessary to initiate autogenic recovery, then guiding this process through the timely introduction of top predators and environmental modifications such as soil amendments and physical structures for animal nesting and refugia.     

A cross-ecosystem comparison of the strength of trophic cascades

Although trophic cascades (indirect effects of predators on plants via herbivores) occur in a wide variety of food webs, the magnitudes of their effects are often quite variable. We compared the responses of herbivore and plant communities to predator manipulations in 102 field experiments in six different ecosystems: lentic (lake and pond), marine, and stream benthos, lentic and marine plankton, and terrestrial (grasslands and agricultural fields). Predator effects varied considerably among systems and were strongest in lentic and marine benthos and weakest in marine plankton and terrestrial food webs. Predator effects on herbivores were generally larger and more variable than on plants, suggesting that cascades often become attenuated at the plant–herbivore interface. Top‐down control of plant biomass was stronger in water than on land; however, the differences among the five aquatic food webs were as great as those between wet and dry systems.     

Exploring the role of large predators in marsh food webs: evidence for a behaviorally-mediated trophic cascade

The influence of large predators on lower trophic levels in oligotrophic, structurally complex, and frequently disturbed aquatic environments is generally thought to be limited. We looked for effects of large predators in two semi-permanent, spikerush-dominated marshes by excluding large fish (>12 mm body depth) and similarly sized herpetofauna from 1 m² cages (exclosures) for 2 weeks. The exclosures allowed for colonization by intermediate (in size and trophic position) consumers, such as small fish, shrimp, and crayfish. Exclosures were compared to control cages that allowed large fish to move freely in and out. At the end of the experiment, intermediate-consumer densities were higher in exclosures than in controls at both sites. Decapod crustaceans, especially the riverine grass shrimp (Palaemonetes paludosus), accounted for the majority of the response. Effects of large fish on shrimp were generally consistent across sites, but per capita effects were sensitive to estimates of predator density. Densities of intermediate consumers in our exclosures were similar to marsh densities, while the open controls had lower densities. This suggests that these animals avoided our experimental controls because they were risky relative to the surrounding environment, while the exclosures were neither avoided nor preferred. Although illuminating about the dynamics of open-cage experiments, this finding does not influence the main results of the study. Small primary consumers (mostly small snails, amphipods, and midges) living on floating periphyton mats and in flocculent detritus (“floc”) were less abundant in the exclosures, indicative of a trophic cascade. Periphyton mat characteristics (i.e., biomass, chlorophyll a, TP) were not clearly or consistently affected by the exclosure, but TP in the floc was lower in exclosures. The collective cascading effects of large predators were consistent at both sites despite differences in drought frequency, stem density, and productivity.     

Comparison of the trophic niches between two planktivorous fishes in two large lakes using stable isotope analysis

This paper studied the sources of food and the trophic levels of silver and bighead carps using stable isotopes (δ¹³C and δ¹⁵N) analysis in Lake Qiandao and Lake Taihu. The δ¹³C values of POM and phytoplankton indicated that Particulate Organic Matter (POM) is of terrestrial origin in LQ and phytoplankton origin in LT. The different signature of δ¹⁵N of phytoplankton and POM also showed the different extent of anthropogenic impacts on the two lakes, with higher impacts in LT than in LQ. In LQ the trophic levels of silver and bighead carps were significantly different (2.48 ± 0.19 and 2.66 ± 0.19, respectively), while in LT the difference was not significant (2.15 ± 1.54 and 2.36 ± 0.38, respectively). Using a mixing model, we found that silver and bighead carps consumed a major proportion on phytoplankton and/or POM in both lakes with higher contribution of phytoplankton in more eutrophic lake. Silver and bighead carps had clear differentiation in food resources with silver carp more on POM and bighead more on zooplankton in deep and mesotrophic lakes. Contrary to this, both carps fed primarily on phytoplankton (and POM) and may have more niche overlaps in shallow and eutrophic lakes.     

海州湾鱼类群落平均营养级和大型鱼类指数的变化特征

为探究近年来海州湾鱼类群落的变化特征,本研究根据2011年及2013-2017年春季(5月)和秋季(9-10月)在海州湾及其邻近海域进行的渔业资源底拖网调查,分析了海州湾鱼类群落平均营养级(MTL)和大型鱼类指数(LFI),对海州湾鱼类群落结构特征进行研究.结果表明:海州湾的优势鱼种主要有大泷六线鱼、方氏云鳚、尖海龙、小黄鱼、长蛇鲻等,且优势鱼种季节性变化明显.海州湾鱼类群落的平均营养级存在明显的年际和季节变化,总体上秋季的MTL高于春季,而且秋季MTL变化具有滞后性.LFI计算结果表明,近年来海州湾及其邻近海域大个体鱼类资源量有所减少,鱼类群落结构呈现出明显的小型化趋势.     

Glacial cycling, large mammal community composition, and trophic adaptations in the Western Cape, South Africa

Some of the earliest evidence for modern human behavior has been recovered from the Western Cape Province, South Africa. Archaeological and paleontological sites in the Western Cape are typically described as “glacial” or “interglacial” in aspect based on the numbers of grazers found in the faunal assemblage, as glacial periods are often thought to have been characterized by spreading C₄ grasslands that replaced endemic C₃ shrubland vegetation found in the Western Cape today. Here, we test the hypothesis that glacial and interglacial time periods were associated with a predictable change in large mammal trophic adaptations by analyzing the proportions of grazing larger mammals from 118 levels of 15 Western Cape fossil assemblages sampling marine isotope stage (MIS) 6 to the present time to determine whether there is a change in composition in these communities that might reflect a shift in ecology and habitat. Our results indicate that trophic proportions did not significantly change over time in the Western Cape as a whole, and thus the hypothesis for habitat changes affecting the subsistence ecology of modern humans during the development of modern behavior is not supported. However, our results show that the southwestern subregion of the Western Cape was characterized by the presence of more grazing species through time than the western subregion. Thus, if ecological and population isolation during glacial periods were integral to catalyzing the development of modern behaviors in the Western Cape region of South Africa, then a complex model including the development of possible mosaic habitats is needed.     

A bioenergetic framework for the temperature dependence of trophic interactions

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature‐dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature‐dependent processes that are common to all consumer–resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability.     

Monitoring of trophic chains

In the paper simple trophic chains of the type resource-producer-primary consumer are considered. For an analysis of the dynamic state monitoring of this system, the concept of observability of mathematical systems theory is proposed. Using a linearization method of non-linear observation systems, biologically interpretable sufficient conditions are obtained to guarantee local observability. The latter means that based on the dynamic observation of a single component, the state of the whole system can be uniquely recovered, at least near a positive equilibrium.     

Diet composition,feeding niche partitioning and trophic organisation of large pelagic predatory fishes in the eastern Arabian Sea

Information on the ecology and feeding behaviour of the large oceanic predatory fishes is crucial for the ecosystem approaches to fisheries management models. Co-existing large pelagic predators in the open oceans may avoid competition for the limited forage by resource partitioning on spatial, temporal or trophic levels. To test this, we studied the prey species composition, diet overlap, trophic level, and trophic organisation of 12 large predatory fishes co-existing in the eastern Arabian Sea. Stomach contents of 1,518 specimens caught by exploratory longline operations in the Indian Exclusive Economic Zone during the years 2006–2009 were analysed. Finfishes were dominant prey of all species except blue marlin (Makaira nigricans) and yellowfin tuna (Thunnus albacares), which fed mainly on cephalopods, and long-snouted lancetfish (Alepisaurus ferox) and pelagic stingray (Pteroplatytrygon violacea), which fed mainly on crustaceans. Common dolphinfish (Coryphaena hippurus) and yellowfin tuna fed on a wider variety of prey than the other species, while the diets of lancetfish and black marlin (Istiompax indica) were narrowest. Pelagic stingray and great barracuda (Sphyraena barracuda) fed on species occupying epipelagic waters, whereas the contribution of mesopelagic prey was higher in the diets of swordfish (Xiphias gladius) and pelagic thresher (Alopias pelagicus). Trophic levels of these fishes ranged from 4.13 to 4.37. Diet overlap index revealed that some of the large pelagic predatory fishes share common prey species. Cluster analysis of the diets revealed four distinct trophic guilds namely ‘flyingfish feeders’ (common dolphinfish and great barracuda); ‘mesopelagic predators’ (pelagic thresher and swordfish); ‘crab feeders’ (lancetfish, pelagic stingray and silky shark) and ‘squid feeders’ (yellowfin tuna, Indo-Pacific sailfish (Istiophorus platypterus), skipjack tuna (Katsuwonus pelamis), black marlin and blue marlin). Large predatory fishes of the eastern Arabian Sea target different prey types, and limit their vertical extent and time of feeding to avoid competing for prey.     

The tempo-spatial variations of phytoplankton diversities and their correlation with trophic state levels in a large eutrophic Chinese lake

Lake Chaohu is one of the most eutrophic lakes in China. Research on this lake's seasonal and spatial variations in phytoplankton diversity is needed to understand the distribution of eutrophication, as well as to find appropriate comprehensive biodiversity indices to assess the eutrophication status of the lake. The present study indicated that the Margalef index of all samples was as low as 0.799 ± 0.543 in summer (August 2011) and as high as 1.467 ± 0.653 in winter (February 2012). The Margalef index of the river samples had a high mean value and substantial variation compared with the lake samples. The Peilou index of the lake samples was higher than that of the river samples in summer and autumn (November 2011) but lower than that of the river samples in winter. In spring (May 2012), the Peilou index of the western samples was lower than that of the eastern samples. The spatial distribution of the Shannon–Wiener index was more similar to that of the Peilou index in autumn and winter, while in spring and summer, the spatial distribution was affected by both species richness and evenness. High eutrophication levels occurred in the western lake in spring and summer, whereas high levels occurred in the eastern lake, especially in the middle of the lake, in autumn and winter. The total trophic state index (TSI) in all samples exhibited a significant negative correlation with the Margalef (r = −0.726) and Peilou (r = −0.530) indices but a significant positive correlation with the Shannon–Wiener (r = 0.654) index. The partial correlation analysis results implied that these phytoplankton biodiversity indices could serve as synthetic ecological indicators to assess the eutrophication condition of Lake Chaohu.     

A novel trophic relationship between cassid gastropods and mysticete whale carcasses

Large cassid gastropods found in close association with Oligocene mysticete whale skeletons in a deep water setting indicate a novel trophic relationship. Bones of early mysticetes are preserved in close association with a recurring assemblage of bivalve taxa that are common components of fossil and recent hydrothermal vent and cold methane seep localities. Living members of these bivalve groups harbor sulfide-oxidizing autotrophic bacteria and comparisons with recent whale-falls indicate that they utilized bone-oil seepage. But unlike recent whale-falls, this fossil invertebrate assemblage is volumetrically dominated by large individuals of the cassid gastropod Liracassis apta that are preserved with their apertures adhering to the bone surfaces. Comparisons with recent cassoidean taxa and their ecological settings suggests that Liracassis apta belongs to the subfamily Oocorythinae, not Cassininae, and was an opportunistic scavenger in deep water, low oxygen environments. Oocorythine gastropods radiated rapidly on the northeastern Pacific margin during the Late Eocene and Oligocene to become the most commonly recorded gastropod in shelf and slope settings.     

Four decades of functional community change reveals gradual trends and low interlinkage across trophic groups in a large marine ecosystem

The rate at which biological diversity is altered on both land and in the sea, makes temporal community development a critical and fundamental part of understanding global change. With advancements in trait‐based approaches, the focus on the impact of temporal change has shifted towards its potential effects on the functioning of the ecosystems. Our mechanistic understanding of and ability to predict community change is still impeded by the lack of knowledge in long‐term functional dynamics that span several trophic levels. To address this, we assessed species richness and multiple dimensions of functional diversity and dynamics of two interacting key organism groups in the marine food web: fish and zoobenthos. We utilized unique time series‐data spanning four decades, from three environmentally distinct coastal areas in the Baltic Sea, and assembled trait information on six traits per organism group covering aspects of feeding, living habit, reproduction and life history. We identified gradual long‐term trends, rather than abrupt changes in functional diversity (trait richness, evenness, dispersion) trait turnover, and overall multi‐trait community composition. The linkage between fish and zoobenthic functional community change, in terms of correlation in long‐term trends, was weak, with timing of changes being area and trophic group specific. Developments of fish and zoobenthos traits, particularly size (increase in small size for both groups) and feeding habits (e.g. increase in generalist feeding for fish and scavenging or predation for zoobenthos), suggest changes in trophic pathways. We summarize our findings by highlighting three key aspects for understanding functional change across trophic groups: (a) decoupling of species from trait richness, (b) decoupling of richness from density and (c) determining of turnover and multi‐trait dynamics. We therefore argue for quantifying change in multiple functional measures to help assessments of biodiversity change move beyond taxonomy and single trophic groups.