Effects of an invasive crayfish on trophic relationships in north‐temperate lake food webs

1. The introduction of invasive species is one of the main threats to global biodiversity, ecosystem structure and ecosystem processes. In freshwaters, invasive crayfish alter macroinvertebrate community structure and destroy macrophyte beds. There is limited knowledge on how such invasive species‐driven changes affect consumers at higher trophic levels. 2. In this study, we explore how the invasive rusty crayfish Orconectes rusticus, a benthic omnivore, affects benthic macroinvertebrates, as well as the broader consequences for ecosystem‐level trophic flows in terms of fish benthivory and trophic position (TP). We expected crayfish to decrease abundance of benthic macroinvertebrates, making most fish species less reliant on benthic resources. We expected crayfish specialists (e.g. Lepomis sp. and Micropterus sp.) to increase their benthic dependence. 3. In 10 northern Wisconsin lakes, we measured rusty crayfish relative abundance (catch per unit effort, CPUE), macroinvertebrate abundance, and C and N stable isotope ratios of 11 littoral fish species. We used stable isotope data and mixing models to characterise the trophic pathways supporting each fish species, and related trophic structure to crayfish relative abundance, fish body size and abiotic predictors using hierarchical Bayesian models. 4. Benthic invertebrate abundance was negatively correlated with rusty crayfish relative abundance. Fish benthivory increased with crayfish CPUE for all 11 fish species; posterior probabilities of a positive effect were >95%. TP also increased slightly with crayfish CPUE for some species, particularly smallmouth bass, largemouth bass, rock bass and Johnny darter. Moreover, both fish body size and lake abiotic variables explained variation in TP, while their effects on benthivory were small. 5. Rusty crayfish abundance explained relatively little of the overall variation in fish benthivory and TP. Although rusty crayfish appear to have strong effects on abundances of benthic macroinvertebrates, energy flow pathways and trophic niches of lentic fishes were not strongly influenced by invasive rusty crayfish.     

Microbial and classical food webs: A visit to a hypertrophic lake

Abstract: Plankton community structure and fluxes of carbon for bacteria (production and bacterivory) were investigated in the urban, hypertrophic Lake Rodó (Uruguay) using a short time interval for sampling (5–15 d) during one year. The lake sustains a high phytoplankton biomass (up to 335 μg l⁻¹ chlorophyll a ) always dominated by the filamentous cyanobacteria Planktothrix agardhii . The zooplankton community was numerically dominated by rotifers and ciliates; cladocerans were rare during most of the year. The rotifer abundance was very high (up to 10⁵ individual l⁻¹), the bacterivorous Anuraeopsis fissa being the most abundant species. Predation rates of heterotrophic nanoflagellates (HNF) on bacteria (range: 31–130 bacteria HNF⁻¹ h⁻¹) were higher than those reported in the literature for field studies. A carbon budget showed that HNF can consume on average 91 and 76% of the bacterial carbon production in summer and winter, respectively. Bacterial turnover times are the lowest reported until now from field conditions (5 to 42 h). Consequently, bacterial carbon production was extremely high (72 to 1071 μg C l⁻¹ d⁻¹). Bacterial production was positively correlated to bacterial abundance but the relationship was significantly improved by the inclusion of temperature (82% variability explained). My results support the general trend for increased bacterial production with increasing trophic status, and suggest a lower energy transfer efficiency to higher trophic levels in hypertrophic lakes due to the many trophic interactions involved.     

Top-down control of phytoplankton: the role of time scale, lake depth and trophic state

SUMMARY 1. One of the most controversial issues in biomanipulation research relates to the conditions required for top-down control to cascade down from piscivorous fish to phytoplankton. Numerous experiments have demonstrated that Phytoplankton biomass Top-Down Control (PTDC) occurs under the following conditions: (i) in short-term experiments, (ii) shallow lakes with macrophytes, and (iii) deep lakes of slightly eutrophic or mesotrophic state. Other experiments indicate that PTDC is unlikely in (iv) eutrophic or hypertrophic deep lakes unless severe light limitation occurs, and (v) all lakes characterised by extreme nutrient limitation (oligo to ultraoligotrophic lakes).
2. Key factors responsible for PTDC under conditions (i) to (iii) are time scales preventing the development of slow-growing inedible phytoplankton (i), shallow depth allowing macrophytes to become dominant primary producers (ii), and biomanipulation-induced reduction of phosphorus (P) availability for phytoplankton (iii).
3. Under conditions (iv) and (v), biomanipulation-induced reduction of P-availability might also occur but is insufficient to alter the epilimnetic P-content enough to initiate effective bottom-up control (P-limitation) of phytoplankton. In these cases, P-loading is much too high (iv) or P-content in the lake much too low (v) to initiate or enhance P-limitation of phytoplankton by a biomanipulation-induced reduction of P-availability. However, PTDC may exceptionally result under condition (iv) if high mixing depth and/or light attenuation cause severe light limitation of phytoplankton.
4. Recognition of the five different conditions reconciles previous seemingly contradictory results from biomanipulation experiments and provides a sound basis for successful application of biomanipulation as a tool for water management.     

The role of alien fish (the centrarchid Micropterus salmoides) in lake food webs highlighted by stable isotope analysis

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Copepoda – Cladocera – Tunicata: The role of three major mesozooplankton groups in pelagic food webs

Copepods, cladocerans and tunicates form major groups of herbivorous mesozooplankton. The former two are found in fresh and marine waters, while the latter are restricted to marine systems. In the present review, we compile existing ecophysiological knowledge about between-group differences in metabolic and reproductive rates, feeding selectivity and elemental composition. From this, we derive predictions about their impact on the lower trophic levels (phytoplankton and microbial food web) and predictions about their prevalence under different ecological conditions (e.g. nutrient richness, Si : N ratio, phytoplankton size structure and top-down control).     

A stable carbon and nitrogen isotope study of lake food webs in Canada's Boreal Plain

1.  Stable carbon and nitrogen isotope and fish stomach content analyses were used to investigate food webs in five relatively undisturbed lakes on the Boreal Plain of Canada. Stable isotope analysis was also used to determine the importance of external and internal carbon sources.
2.  Overlap in the carbon and nitrogen signatures of primary producers made it difficult to determine unambiguously the feeding habits of many invertebrates. However, isotope analysis suggested that external carbon inputs were detectable in the aquatic food chains of the one lake with a short water residence time («1 year). In the other four lakes, with water residence times ≥1 year, autochthonous carbon was the only detectable carbon source in the food webs.
3.  Food webs in these lakes spanned a range of four to five trophic levels. Both invertebrates and fish appeared to eat a variety of food, often feeding at more than one trophic level.
4.  With the exception of one lake (SPH20), top predators in these lakes, northern pike ( Esox lucius ) and fathead minnows ( Pimephales promelas ), occupied similar trophic positions despite large differences in body size and trophic morphology. In SPH20, where there were two additional fish species, pike occupied a higher trophic position. However, all the top predators in each lake appeared to be omnivores and generalists.
5.  The prevalence of omnivory and the apparent generalist feeding habits of fish in these lakes suggest that organisms are flexible in their feeding habits and that these food webs will be resilient to disturbance.     

Polar bears make little use of terrestrial food webs: evidence from stable-carbon isotope analysis

Summary The mean stable-carbon isotope ratios (¹³C) for polar bear (Ursus maritimus) tissues (bone collagen –15.7, muscle –17.7, fat –24.7) were close to those of the same tissues from ringed seals (Phoca hispida) (–16.2, –18.1, and –26.1, respectively), which feed exclusively from the marine food chain. The ¹³C values for 4 species of fruits to which polar bears have access when on land in summer ranged from –27.8 to –26.2, typical of terrestrial plants in the Arctic. An animal's ¹³C signature reflects closely the ¹³C signature of it's food. Accordingly, the amount of food that polar bears consume from terrestrial food webs appears negligible, even though some bears spend 1/3 or more of each year on land during the seasons of greatest primary productivity.     

The dynamics of spatially coupled food webs

The dynamics of ecological systems include a bewildering number of biotic interactions that unfold over a vast range of spatial scales. Here, employing simple and general empirical arguments concerning the nature of movement, trophic position and behaviour we outline a general theory concerning the role of space and food web structure on food web stability. We argue that consumers link food webs in space and that this spatial structure combined with relatively rapid behavioural responses by consumers can strongly influence the dynamics of food webs. Employing simple spatially implicit food web models, we show that large mobile consumers are inordinately important in determining the stability, or lack of it, in ecosystems. More specifically, this theory suggests that mobile higher order organisms are potent stabilizers when embedded in a variable, and expansive spatial structure. However, when space is compressed and higher order consumers strongly couple local habitats then mobile consumers can have an inordinate destabilizing effect. Preliminary empirical arguments show consistency with this general theory.     

Stable isotopes of carbon and nitrogen as indicators of diet and trophic structure of the fish community in a shallow hypereutrophic lake

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes were employed to elucidate energy flows and trophic interactions in Lake Apopka, a hypereutrophic lake in central Florida, U.S.A. Isotope compositions of lake biota ranged from −27·1 to −3·0‰ for δ¹³C, and from 3·7 to 13·9‰ for δ¹⁵N. The food web was based primarily on plankton production with diatoms, Microcystis and zooplankton dominating the diet of fish. Carbon isotope evidence showed that pico- and nano-phytoplankton were not a direct carbon source for fish, but were important to zooplankton. δ¹⁵N mass balance estimates indicated that planktivorous fish obtained 48–85% of their diets from zooplankton. The ∼3‰ range of δ¹⁵N in gizzard shad reflected increasing dependence on zooplankton as fish grew whereas the positive relationship between total length and δ¹⁵N of largemouth bass reflected increasing predation on larger planktivorous fish with growth. The broad ranges of δ¹³C (−25·9 to −9·5‰) and δ¹⁵N (5·8 to 14·4‰) of blue tilapia were indicators of diet diversity. Two presumed omnivores (brown bullhead and white catfish) and piscivores (black crappie, largemouth bass and Florida gar) were found to depend on planktivorous fish. However, stable isotope data revealed no trophic links between blue tilapia, an abundant fish in the near-shore area, and piscivores.     

The underestimated importance of belowground carbon input for forest soil animal food webs

The present study investigated the relative importance of leaf and root carbon input for soil invertebrates. Experimental plots were established at the Swiss Canopy Crane (SCC) site where the forest canopy was enriched with ¹³C depleted CO₂ at a target CO₂ concentration of c . 540 p.p.m. We exchanged litter between labelled and unlabelled areas resulting in four treatments: (i) leaf litter and roots labelled, (ii) only leaf litter labelled, (iii) only roots labelled and (iv) unlabelled controls. In plots with only ¹³C-labelled roots most of the soil invertebrates studied were significantly depleted in ¹³C, e.g. earthworms, chilopods, gastropods, diplurans, collembolans, mites and isopods, indicating that these taxa predominantly obtain their carbon from belowground input. In plots with only ¹³C-labelled leaf litter only three taxa, including, e.g. juvenile Glomeris spp. (Diplopoda), were significantly depleted in ¹³C suggesting that the majority of soil invertebrates obtain its carbon from roots. This is in stark contrast to the view that decomposer food webs are based on litter input from aboveground.     

植物入侵对食物网的影响及其途径

生态系统中的物质和能量主要沿食物链在食物网中流通,诸多研究表明入侵植物对生态系统功能的影响是通过改变当地原有的食物网结构而实现的,因此外来植物入侵对食物网的影响受到人们越来越多的关注.本文分析了植物入侵引起食物网变化的原因以及改变食物网的途径,探讨了土著食物网特点对群落可入侵性的影响,得出以下结论:(1)食物网的变化主要是由于入侵植物引起的消费者基础食物资源或者周围环境条件的变化造成的;(2)入侵植物通过3种途径影响食物网:一是入侵植物具有较好的可利用性,能够直接被土著草食者取食,通过上行效应按照原有的路径进入土著食物网;二是当入侵植物的可利用性较差时,入侵植物所固定的能量通过引入新的消费者或者转变流通路径形成新的食物网结构;三是入侵植物通过非营养作用造成食物网中各级消费者的种群密度和行为活动等发生变化,进而影响土著生物群落和食物网结构;(3)食物网的大小及其连通性可能影响外来植物的入侵成功率.在今后的研究中,为深入揭示生物入侵对食物网的影响,在短期控制实验的基础上应加强长时间尺度的动态研究,对于食物网变化的定量描述对象也需要进一步加以明确.此外,食物网对入侵植物生态修复的响应以及食物网变化对入侵植物的反馈作用等也具有重要的研究价值.     

Tracing biogeochemical subsidies from glacier runoff into Alaska's coastal marine food webs

Nearly half of the freshwater discharge into the Gulf of Alaska originates from landscapes draining glacier runoff, but the influence of the influx of riverine organic matter on the trophodynamics of coastal marine food webs is not well understood. We quantified the ecological impact of riverine organic matter subsidies to glacier‐marine habitats by developing a multi‐trophic level Bayesian three‐isotope mixing model. We utilized large gradients in stable (δ¹³C, δ¹⁵N, δ²H) and radiogenic (Δ¹⁴C) isotopes that trace riverine and marine organic matter sources as they are passed from lower to higher trophic levels in glacial‐marine habitats. We also compared isotope ratios between glacial‐marine and more oceanic habitats. Based on isotopic measurements of potential baseline sources, ambient water and tissues of marine consumers, estimates of the riverine organic matter source contribution to upper trophic‐level species including fish and seabirds ranged from 12% to 44%. Variability in resource use among similar taxa corresponded to variation in species distribution and life histories. For example, riverine organic matter assimilation by the glacier‐nesting seabirds Kittlitz's murrelet (Brachyramphus brevirostris) was greater than that of the forest‐nesting marbled murrelet (B. marmoratus). The particulate and dissolved organic carbon in glacial runoff and near surface coastal waters was aged (12100–1500 years BP ¹⁴C‐age) but dissolved inorganic carbon and biota in coastal waters were young (530 years BP ¹⁴C‐age to modern). Thus terrestrial‐derived subsidies in marine food webs were primarily composed of young organic matter sources released from glacier ecosystems and their surrounding watersheds. Stable isotope compositions also revealed a divergence in food web structure between glacial‐marine and oceanic sites. This work demonstrates linkages between terrestrial and marine ecosystems, and facilitates a greater understanding of how climate‐driven changes in freshwater runoff have the potential to alter food web dynamics within coastal marine ecosystems in Alaska.     

Algivory in food webs of three temperate Andean rivers

Food web analyses have been fundamental in understanding community organization and ecosystem functioning. To date, a number of studies demonstrate that stream food webs depend to a large extent on allochthonous detritus, but there are more recent studies that show a high degree of autochthony. Our food‐web study was carried out in three Andean rivers (Coilaco, Guampoe and Trancura) within the catchment area of Toltén River in southern Chile. Based on the analyses of 4251 invertebrate gut contents, we found that these Andean stream food webs are dominated by herbivores (range: 50–73% of all species) supported by a species‐rich algal (basal) component, and characterized by a low proportion of omnivores (range: 8–27% of all species) and predatory species (range: 10–24%). Significant differences in the number of feeding links of the herbivores Meridialis diguillina and Antarctoperla michaelseni and the omnivore Smicridea chilensis were found between seasons. The spring herbivore Aubertoperla sp. showed significant differences between rivers. S. chilensis fed on 50 different prey items as compared with the herbivores whose maximum number of links ranged between 37 and 40. Web sizes ranged between 93 and 131 species and the proportion of top species was distinctly lower than those of basal (up to 0.651 in Coilaco River) and intermediate species. Direct connectance (links per species²) values were low and similar among rivers (range: 0.051–0.074), whereas mean food chain length ranged between 2.23 and 2.90. The distributions of web property values from the Andean rivers differed from those previously published. In contrast to previous predictions, mean food chain length in these Andean streams displayed a scale‐invariant pattern across different web sizes, but it was significantly related to the proportion of intermediate species.     

持久性有机污染物在水生食物网中的传递行为

食物网是持久性有机污染物(POPs)在水生生态系统中传递的重要途径,了解其传递行为与机制是POPs生态暴露风险评价的科学基础。从4个方面展开了讨论和分析:(1)食物网主要特征(营养级和食物链长度)与POPs环境行为的关系;(2)POPs在底栖及底栖-浮游耦合食物网中的环境行为;(3)微食物网对POPs环境行为的作用;(4)食物网的变化对POPs环境行为的影响。主要结论如下:(1)已有研究对水生生物中POPs生物放大作用存在较大争议。一般营养级越高,POPs生物富集性越强,但由于各种生态和生理性质的影响,也存在例外情况。食物链长度与POPs生物富集性呈正相关。(2)POPs通过底栖食物网将沉积物中的POPs向上传递,底栖-浮游食物网的耦合提高了高营养级消费者的暴露风险,目前就POPs在底栖食物网中的生物放大性是否大于浮游食物网存在争议。(3)微生物具有较大的比表面积,是吸附POPs的重要载体。另,沉积物中的微生物通过分解有机质,将POPs释放到水柱中。微生物降解也是环境中POPs脱离环境的重要途径。(4)在内、外压力下,食物网结构和功能发生变化,使物质和能量的传递方向和效率发生改变,并与环境理化性质的变化互相耦合,影响POPs的环境行为。当前研究的重点多集中在POPs在浮游食物网,尤其是高营养级浮游食物网中的环境行为,对POPs在底栖及底栖-浮游耦合食物网和微食物网中环境行为的研究相对缺乏。有关POPs在食物网中环境行为的研究多集中在食物网的某个部分,时间尺度较短,缺乏对POPs环境行为动态变化的研究,未来需深入开展多尺度和多角度的POPs在食物网中环境行为的动态变化研究。新型POPs的生产和使用量不断增加,但有关其在食物网中环境行为的相关分析还较为匮乏,需加强研究。     

The second life of terrestrial and plastic carbon as nutritionally valuable food for aquatic consumers

Primary production is the basis for energy and biomolecule flow in food webs. Nutritional importance of terrestrial and plastic carbon via mixotrophic algae to upper trophic level is poorly studied. We explored this question by analysing the contribution of osmo- and phagomixotrophic species in boreal lakes and used ¹³C-labelled materials and compound-specific isotopes to determine biochemical fate of carbon backbone of leaves, lignin–hemicellulose and polystyrene at four-trophic level experiment. Microbes prepared similar amounts of amino acids from leaves and lignin, but four times more membrane lipids from lignin than leaves, and much less from polystyrene. Mixotrophic algae (Cryptomonas sp.) upgraded simple fatty acids to essential omega-3 and omega-6 polyunsaturated fatty acids. Labelled amino and fatty acids became integral parts of cell membranes of zooplankton (Daphnia magna) and fish (Danio rerio). These results show that terrestrial and plastic carbon can provide backbones for essential biomolecules of mixotrophic algae and consumers at higher trophic levels.     

On the different nature of top-down and bottom-up effects in pelagic food webs

SUMMARY 1. Each individual planktonic plant or animal is exposed to the hazards of starvation and risk of predation, and each planktonic population is under the control of resource limitation from the bottom up (growth and reproduction) and by predation from the top down (mortality). While the bottom-up and top-down impacts are traditionally conceived as compatible with each other, field population-density data on two coexisting Daphnia species suggest that the nature of the two impacts is different. Rates of change, such as the rate of individual body growth, rate of reproduction, and each species' population growth rate, are controlled from the bottom up. State variables, such as biomass, individual body size and population density, are controlled from the top down and are fixed at a specific level regardless of the rate at which they are produced.
2. According to the theory of functional responses, carnivorous and herbivorous predators react to prey density rather than to the rate at which prey are produced or reproduced. The predator's feeding rate (and thus the magnitude of its effect on prey density) should hence be regarded as a functional response to increasing resource concentration.
3. The disparity between the bottom-up and top-down effects is also apparent in individual decision making, where a choice must be made between accepting the hazards of hunger and the risks of predation (lost calories versus loss of life).
4. As long as top-down forces are effective, the disparity with bottom-up effects seems evident. In the absence of predation, however, all efforts of an individual become subordinate to the competition for resources. Biomass becomes limited from the bottom up as soon as the density of a superior competitor has increased to the carrying capacity of a given habitat. Such a shift in the importance of bottom-up control can be seen in zooplankton in habitats from which fish have been excluded.     

Effects of climate‐driven primary production change on marine food webs: implications for fisheries and conservation

Climate change is altering the rate and distribution of primary production in the world's oceans. Primary production is critical to maintaining biodiversity and supporting fishery catches, but predicting the response of populations to primary production change is complicated by predation and competition interactions. We simulated the effects of change in primary production on diverse marine ecosystems across a wide latitudinal range in Australia using the marine food web model Ecosim. We link models of primary production of lower trophic levels (phytoplankton and benthic producers) under climate change with Ecosim to predict changes in fishery catch, fishery value, biomass of animals of conservation interest, and indicators of community composition. Under a plausible climate change scenario, primary production will increase around Australia and generally this benefits fisheries catch and value and leads to increased biomass of threatened marine animals such as turtles and sharks. However, community composition is not strongly affected. Sensitivity analyses indicate overall positive linear responses of functional groups to primary production change. Responses are robust to the ecosystem type and the complexity of the model used. However, model formulations with more complex predation and competition interactions can reverse the expected responses for some species, resulting in catch declines for some fished species and localized declines of turtle and marine mammal populations under primary productivity increases. We conclude that climate‐driven primary production change needs to be considered by marine ecosystem managers and more specifically, that production increases can simultaneously benefit fisheries and conservation. Greater focus on incorporating predation and competition interactions into models will significantly improve the ability to identify species and industries most at risk from climate change.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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