The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Three-dimensional barricading of a predatory trap reduces predation and enhances prey capture

Animal structures come at material, energetic, time, and expression costs. Some orb-web spiders add three-dimensional barrier structures to their webs, but many do not. Predator protection is considered to be the principal benefit of adding these structures. Accordingly, it remains paradoxical why some orb-web spiders might construct the barriers while others do not. Here, we experimentally determined whether the barrier structure added to the horizontal orb web of the spider Cyrtophora moluccensis deters predators at the cost of reducing the amount of prey captured in the field. We conducted experiments by day and night to assess whether the effects vary with the time of day. We found that the three-dimensional barriers not only offered protection from predatory wasps by day but also enhanced the amount of prey captured by day and night. Moreover, the barrier structure appears particularly useful at catching moths, the largest and most energetically profitable prey that it encounters. We, therefore, concluded that reducing the energetic and time costs associated with producing and depositing extra silk threads is the principal reason why barrier structures are used intermittently among orb-web spiders.     

An improved automatic recording apparatus for determining the filtration rate of Mytilus edulis as a function of size and algal concentration

An automatic recording apparatus for measuring the filtration rate in suspension-feeding bivalves is described. The concentration of algae in the experimental medium is kept constant throughout each experiment by addition of Phaeodactylum tricornutum from a chemostat. Within the range of body size 5.7 to 283 mg (W=dry weight of tissues), the filtration rate (F=ml min^-1) at 15°C in Mytilus edulis L. follows the allometric equation F=0.85 W ^0.72. Within the concentrations 0.18 to 0.70 mg algal dry weight l^-1, the filtration rate in mussels of 132 mg dry flesh weight ranges from 33.1 to 41.0 ml min^-1. At 0.18 mg algal dry weight l^-1 the mussels filter continuously for 20 h, with a high constant rate that presumably represents the water transport capacity under optimal laboratory conditions.     

Clam predation by whelks (Busycon spp.): Experimental tests of the importance of prey size,prey density,and seagrass cover

In 57 l-m² samples within a meadow of Halodule wrightii in Bogue Sound, North Carolina, USA, densities of the clams Mercenaria mercenaria and Chione cancellata were positively associated with seagrass cover. Where seagrass was experimentally removed, marked individuals of both clam species exhibited high rates of mortality in fine sand sediments during two successive experiments spanning 13 months. In the unaltered (control) seagrass meadow, M. mercenaria density remained constant over 13 months and C. cancellata density declined at a slower rate than in the unvegetated plots. Seagrass provides these clams with a refuge from whelk (Busycon carica, B. contrarium, and B. canaliculatum) predation, the major cause of mortality and population decline in experimentally unvegetated plots. In 2 factorial field experiments in unvegetated substratum in which densities of M. mercenaria and C. cancellata were varied independently, first over 5 levels (0 X, 1/2X, 1 X, 2 X, 4 X) and subsequently over 4 levels (0 X, 1/4 X, 1 X, 4 X), there was no repeatable intra- or interspecific effect of density on percent survival, or on the rate of any mortality type. Whelk predation fell preferentially on larger size classes of both species, whereas factors which contribute to clam disappearance usually acted more intensely on smaller sizes. Experimental exclusion of large predators by caging demonstrated that even in unvegetated substratum survivorship of both clam species was high in the absence of whelks and other predators. Individuals of C. cancellata live closer to the sediment surface than those of M. mercenaria, which may explain why seagrass does not serve as effectively to protect them from whelk predation. The mechanism of whelk inhibition may depend upon sediment binding by the H. wrightii root mat, which produces a demonstrable decrease in the physical penetrability of surface sediments.     

Cercopagis pengoi and Mysis spp. alter their feeding rate and prey selection under predation risk of herring (Clupea harengus membras)

The anti-predator behaviour of Baltic crustacean planktivores was studied in feeding experiments under predation pressure of herring. The experiments were conducted with pelagic mysids: Mysis mixta and Mysis relicta, and with Cercopagis pengoi, a non-indigenous cladoceran, which invaded the Baltic Sea in 1992. Zooplankton was offered as prey. Two kinds of experiments were performed in the absence and presence of chemical predator cues: (1) two-prey experiments with prey, which have poor or good escape responses and all three planktivores and (2) natural prey experiments with mysids in natural zooplankton assemblages. The results showed that all three species reacted to the chemical cue of herring by decreasing their feeding rate and altering prey selection. C. pengoi selected easily captured prey (rotifers) in two-prey experiments under predation risk while selection for any prey was evident in mysids in natural prey experiments only in the absence of predator cues. This indicates that planktivores have different anti-predator strategies, which are modified by their own prey capture abilities. C. pengoi was a very efficient predator on small prey with size-specific prey consumption rate 5 to 18 times the rate of mysids. Results show that the studied planktivores are capable of adjusting their feeding behaviour to decrease their conspicuousness in order to increase survival under predation risk. Further, results support the view that C. pengoi has adapted well to the Baltic ecosystem, sharing food niche with pelagic mysids and most probably having a strong influence on the whole pelagic food web.     

Interference by wood ant influences size selection and retrieval rate of prey by Formica fusca

Summary Dominance relations between the territorial wood ant Formica polyctena and the submissive F. fusca generate testable predictions on the mechanisms of their coexistence. Here I tested the influence of interference competition by the dominant F. polyctena on the foraging of F. fusca. In the presence of F. polyctena, the activity and the retrieval rate of items of F. fusca decreased significantly. When F. fusca were given a choice between small chironomids and flies a hundred times heavier they selected nearly always flies in the absence of wood ants; when disturbed F. fusca took proportionately more chironomids. In nature, irrespective of distance from the wood-ant mound the size distribution of potentially available food items was the same. F. fusca collected smaller items close to the wood-ant mound, where the density of interfering wood ants was high, than far from it. Also, F. polyctena carried larger protein items from the outskirts of its territory than from the center. The items of F. fusca were on average smaller than those of F. polyctena although the item-size overlap was substantial. When encountering a F. polyctena, those F. fusca workers carrying a fly always lost their booty to the dominant but always managed to bring the chironomids to the nest. F. fusca and F. polyctena were equally efficient in detecting single chironomids placed on the surface of ground, but the presence of either species decreased the discovery rate of the other. This implies mutual exploitation competition between the species for the locally most abundant protein resource. F. polyctena found single flies faster than F. fusca. In an earlier study I showed that close to the wood-ant mound the colony size and production of sexual offspring of F. fusca were reduced. I suggest that the suppressed colony success of F. fusca is attributable to the diminished size and decreased retrieval rate of prey items close to the woodant mound abounding with interfering wood-ant foragers.     

Effects of surrounding land use and water depth on seagrass dynamics relative to a catastrophic algal bloom

Seagrasses are the foundation of many coastal ecosystems and are in global decline because of anthropogenic impacts. For the Indian River Lagoon (Florida, U.S.A.), we developed competing multistate statistical models to quantify how environmental factors (surrounding land use, water depth, and time [year]) influenced the variability of seagrass state dynamics from 2003 to 2014 while accounting for time‐specific detection probabilities that quantified our ability to determine seagrass state at particular locations and times. We classified seagrass states (presence or absence) at 764 points with geographic information system maps for years when seagrass maps were available and with aerial photographs when seagrass maps were not available. We used 4 categories (all conservation, mostly conservation, mostly urban, urban) to describe surrounding land use within sections of lagoonal waters, usually demarcated by land features that constricted these waters. The best models predicted that surrounding land use, depth, and year would affect transition and detection probabilities. Sections of the lagoon bordered by urban areas had the least stable seagrass beds and lowest detection probabilities, especially after a catastrophic seagrass die‐off linked to an algal bloom. Sections of the lagoon bordered by conservation lands had the most stable seagrass beds, which supports watershed conservation efforts. Our results show that a multistate approach can empirically estimate state‐transition probabilities as functions of environmental factors while accounting for state‐dependent differences in seagrass detection probabilities as part of the overall statistical inference procedure.     

The filtration rate of Mytilus edulis and its dependence on algal concentration,measured by a continuous automatic recording apparatus

A new apparatus for long-term, continuous automatic measurements of filtration rates in suspension-feeding organisms is described. As the concentration of algae in the experimental medium is diminished by the filter-feeding activity of the experimental animals, algal suspension is automatically added, thus keeping the algal concentration constant. In this way, accurate determinations of filtration rates in relation to particle concentration are made possible. For determination of filtration rates in the common mussel Mytilus edulis L., individuals of different body size (shell length 8.5 to 56.5 mm) were used. Within the range of 10x10⁶ to 40x10⁶ cells of Dunaliella marina/l, mussels of the same body size filter-out approximately the same amount of algae at high or low concentrations. A low algal concentration is counterbalanced by a corresponding higher filtration rate. Within the range of body size (W=dry weight of tissues) and algal concentrations used, the filtration rate (F) follows the general allometric equation F=a·W ^b, where a and b are constants at specific experimental conditions. At a temperature of 12 °C, the values obtained for a are 2410 at a concentration of 20x10⁶, and 1313 at a concentration of 40x10⁶ Dunaliella cells/l; correspondingly, the filtration rates of a mussel of 1 g dry-tissue weight are 2410 ml/h and 1313 ml/h. b, the slope of the regression line (0.73 to 0.74), is independent of algal concentration. However, examination of all known measurements reveals that, most probably, the general allometric equation is an oversimplification; in large individuals there is a more pronounced decrease in filtration rate. The relationship between filtration rate, body size of mussels, and algal concentrations used is discussed.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung — Abwässer in Küstennähe.     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Propulsion efficiency and cost of transport for copepods: a hydromechanical model of crustacean swimming

In the absence of direct measurement, costs of locomotion to small swimming Crustacea (<10 mm) have been derived exclusively through application of the fluid dynamic theory. Results indicate very low swimming costs, and contradict experimental data on larger Crustacea (15 to 100 mm) that suggest a three-fold increase in metabolic rate with increasing swimming speed. This paper introduces a swimming model that analyzes the hydrodynamic forces acting on a crustacean swimming at non-steady velocity. The model treats separately the hydrodynamic forces acting on the body and the swimming appendages, approximating the simultaneous solution of equations quantifying the drag and added-mass forces on each by stepwise integration. Input to the model is a time-series of instantaneous swimming-appendage velocities. The model output predicts a corresponding time-series of body velocities as well as the mechanical energy required to move the swimming appendages, dissipated kinetic energy, and metabolic cost of swimming. Swimming of the calanoid copepod Pleuromamma xiphias (Calanoida) was analyzed by extrapolating model parameters from data available in the literature. The model predictions agree well with empirical observations reported for larger crustaceans, in that swimming for copepods is relatively costly. The ratio of active to standard metabolism for P. xiphias was >3. Net cost of transport was intermediate to the values found experimentally for fish and larger crustaceans. This was a consequence of the predicted mechanical efficiency (34%) of the copepod's paddle propulsion, and of increased parasitic resistance resulting from non-steady velocity swimming.     

The use of space by animals as a function of accessibility and preference

Heterogeneous usage of space by individual animals or animal populations is partly due to their preference for particular resources that are, themselves, heterogeneously distributed. When all points in the environment are equally accessible, a direct relationship between usage and preference can be assumed. However, when accessibility is restricted, spatial variations in usage can no longer be attributed entirely to preference. In such cases, it is necessary to control for the effects of accessibility on observed usage before conclusions about preference can be drawn. In this paper, I develop a modelling framework that treats the use of space by animals as a joint function of preference and accessibility. I specify a null version of the framework that assumes no preference and propose that its output can be used to control for the effect of accessibility on the observed, spatial distribution of usage. I briefly discuss how the framework can subsequently be used to provide insights about the animals’ preference for different resources and types of movement, and to predict usage in areas where no usage data exist. I explore the properties of the methodology using data from a population of simulated animals and present the first results of its application to a sub-set of the British population of grey seals (Halichoerus grypus).     

Behavioral games involving a clever prey avoiding a clever predator: An individual-based model of dusky dolphins and killer whales

Faced with an intermittent but potent threat, animals exhibit behavior that allows them to balance foraging needs and avoid predators and over time, these behaviors can become hard-wired adaptations with both species trying to maximize their own fitness. In systems where both predator and prey share similar sensory modalities and cognitive abilities, such as with marine mammals, the dynamic nature of predator-prey interactions is poorly understood. The costs and benefits of these anti-predator adaptations need to be evaluated and quantified based on the dynamic engagement of predator and prey. Many theoretic models have addressed the complexity of predator-prey relationships, but few have translated into testable mechanistic models. In this study, we developed a spatially-explicit, geo-referenced, individual-based model of a prototypical adult dusky dolphin off Kaikoura, New Zealand facing a more powerful, yet infrequent predator, the killer whale. We were interested in two primary objectives, (1) to capture the varying behavioral game between a clever prey and clever predator based on our current understanding of the Kaikoura system, (2) to compare evolutionary costs vs. benefits (foraging time and number of predator encounters) for an adult non-maternal dusky dolphin at various levels of killer whale-avoidance behaviors and no avoidance rules. We conducted Monte Carlo simulations to address model performance and parametric uncertainty. Mantel tests revealed an 88% correlation (426 × 426 distance matrix, km²) between observed field sightings of dusky dolphins with model generated sightings for non-maternal adult dusky dolphin groups. Simulation results indicated that dusky dolphins incur a 2.7% loss in feeding time by evolving the anti-predator behavior of moving to and from the feeding grounds. Further, each evolutionary strategy we explored resulted in dolphins incurring an additional loss of foraging time. At low killer whale densities (appearing less than once every 3 days), each evolutionary strategy simulated converged towards the evolutionary cost of foraging, that is, the loss in foraging time approached the 2.7% loss experienced by evolving near shore-offshore movement behavior. However, the highest level of killer whale presence resulted in 38% decreases in foraging time. The biological significance of these losses potentially incurred by a dusky dolphin is dependent on various factors from dolphin group foraging behavior and individual energy needs to dolphin prey availability and behavior.     

Toward a predictive theory of risk effects: hypotheses for prey attributes and compensatory mortality

Risk effects, or the costs of antipredator behavior, can comprise a large proportion of the total effect of predators on their prey. While empirical studies are accumulating to demonstrate the importance of risk effects, there is no general theory that predicts the relative importance of risk effects and direct predation. Working toward this general theory, it has been shown that functional traits of predators (e.g., hunting modes) help to predict the importance of risk effects for ecosystem function. Here, I note that attributes of the predator, the prey, and the environment are all important in determining the strength of antipredator responses, and I develop hypotheses for the ways that prey functional traits might influence the magnitude of risk effects. In particular, I consider the following attributes of prey: group size and dilution of direct predation risk, the degree of foraging specialization, body mass, and the degree to which direct predation is additive vs. compensatory. Strong tests of these hypotheses will require continued development of methods to identify and quantify the fitness costs of antipredator responses in wild populations.     

The development of a multi-species algal ecodynamic model for urban surface water systems and its application

An ecodynamic model that can simulate four phytoplankton species has been developed to deal with the unique characteristics of urban river systems which has manmade river profile, flow controlled by gates, severe eutrophication status, and fragile aquatic ecosystem. The ecodynamic model was developed referencing two typical models: the water quality simulation model WASP and ecological model CAEDYM. The model can simulate 11 state variables: dissolved oxygen, carbonaceous biochemical oxygen demand, ammonia nitrogen, nitrate nitrogen, organic nitrogen, inorganic phosphorus, organic phosphorus and four phytoplankton species with zooplankton as a boundary condition. The ecodynamic model was applied to Sihai section of the Beijing urban river system, where serious algal blooms broke out in recent years. The dominant phytoplankton species are Cyanophyta, Chlorophyta, Bacillariophyta, and Cryptophyta. Site-specific data on geometry, meteorology, pollution sources, and existing ecosystem parameters were collected and used for model calibration and verification The model results mimic observed trends of water quality and phytoplankton species succession and can be used for forecasting algal blooms as well as assessment of river management measures.     

Inhibition of photosynthesis by ultraviolet radiation as a function of dose and dosage rate: Results for a marine diatom

The effects of ultraviolet radiation on phytoplankton are usually described as a function of dose (J m^–2, weighted appropriately). Experiments conducted in 1988 and 1989 on a marine diatom,Thalassiosira pseudonana (Clone 3H), demonstrate that during lightlimited photosynthesis in visible radiation, the inhibition of photosynthesis by supplemental ultraviolet radiation (principally UV-B: 280 to 320 nm) is a function of irradiance (W m^–2) as well as of dose: for equal doses of UV-B, a relatively short exposure to high UV-B irradiance is more damaging to photosynthesis than a longer exposure to lower irradiance. In fact, photoinhibition by UV-B is well described as a monotonic, nonlinear function of irradiance for time scales of 0.5 to 4 h. A nitrate-limited culture was about nine times more sensitive to UV-B than was a nutrient-replete culture, but the kinetics of photoinhibition were similar. These results have some bearing on efforts to describe the effects of ultraviolet radiation on marine primary productivity. Action spectra of photoinhibition by UV can be constructed, but they should only be used to describe photoinhibition for specified time scales. Vertical profiles of relative photoinhibition must be interpreted cautiously because photoinhibition by UV-B is likely to be a function of incubation time and results must therefore be interpreted in the context of vertical mixing.     

The removal of trimethoprim and sulfamethoxazole by a high infiltration rate artificial composite soil treatment system

Sulfamethoxzole (SMX) and trimethoprim (TMP), two combined-using sulfonamide antibiotics, have gained increasing attention in the surface water, groundwater and the drinking water because of the ecological risk. The removal of TMP and SMX by artificial composite soil treatment system (ACST) with different infiltration rates was systematically investigated using K⁺, Na⁺, Ca²⁺, Mg²⁺ hydrogeochemical indexes. Batch experiments showed that the sorption onto the low-cost and commercially available clay ceramsites was effective for the removal of SMX and TMP from water. The column with more silty clay at high infiltration rate (1.394 m·d^–1) had removal rates of 80% to 90% for TMP and 60% to 70% for SMX. High SMX and TMP removal rates had a higher effluent concentration of K⁺, Ca²⁺ and Mg²⁺ and had a lower effluent Na⁺ concentration. Removal was strongly related to sorption. The results showed that the removal of SMX and TMP was related to hydrogeochemical processes. In this study, ACST is determined to be applicable to the drinking water plants.

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The clearance rate of microzooplankton as the key element for describing estimated non-linear dilution plots demonstrated by a model

The objective of this study was to determine whether the clearance rate of grazers (as an individual response) was sensitive enough to describe non-linear plots estimated by dilution experiments for measuring the instant grazing rate of microzooplankton. The study was based on an initial analysis of a non-linear feeding pattern based on the food concentration dependence of clearance rate of microzooplankton. In contrast to the traditional assumption of a linear functional response, I assumed that the microzooplankton functional response was non-linear and that the dependence of the clearance rate can be sub-divided into four intervals of food concentration (Sections I–IV) as follows: in Section I clearance rate is zero; Section II is a transitional interval in which the clearance rate increases from zero to a maximum value; in Section III, the clearance rate is maximal and constant, and in Section IV, the clearance rate decreases from its maximum value due to saturated ingestion rate. A set of derived differential equations describes the phytoplankton growth rate in each section, leading to the possibility of comparing predicted non-linear dilution plots with observed non-linear dilution data, using only the specific solutions for Sections III and IV. One should evaluate the quality of fit provided by the non-linear and linear models, rather than uncritically accepting only the linear model for observed non-linear dilution data, using calculated expected non-linear and linear dilution plots as alternative hypotheses. It can be demonstrated that the non-linear model provided a better fit to estimated non-linear dilution data from the Red Sea, Rhode River Estuary (USA) and Kiel Fjord (Germany) than the standard linear model. Published dilution experiments which had a non-linear shape were also selected as illustrative examples to demonstrate the superior fit of the non-linear model.     

Assessment of patch quality by ladybirds: relative response to conspecific and heterospecific larval tracks a consequence of habitat similarity?

Summary. Aphid colonies can reach high levels of abundance but last for short periods of time. The larvae of aphidophagous ladybirds (Coleoptera: Coccinellidae) that feed on these colonies might therefore suffer from starvation, which favours the occurrence of cannibalism and intraguild predation. Thus, the assessment of patch quality becomes crucial and it has been shown that female ladybirds refrain from laying eggs in the presence of an oviposition deterring semiochemical deposited by their larvae. Adalia bipunctata (L.), Adalia decempunctata (L.) and Coccinella septempunctata L. are 3 sympatric species of ladybirds, which can co-occur in aphid colonies. As a consequence, their eggs and larvae are under threat, not only from cannibalism but also intraguild predation. Females should, therefore, also use the tracks deposited by heterospecific larvae to assess the quality of aphid colonies as oviposition sites. The expectation is that: 1- the strength of the reaction to each other’s larval tracks should be correlated with percentage habitat overlap and that 2- the reaction to conspecific larval tracks should be stronger than to heterospecific tracks. In order to test these hypotheses, females’ oviposition behaviour was analysed and a chemical analysis of the tracks of their larvae undertaken. The results show that oviposition behaviour is not related to habitat overlap. Both species of Adalia react to tracks of their own larvae and those of C. septempunctata, but A. decempunctata reacted more strongly than A. bipunctata. C. septempunctata reacted very slightly to its own tracks but not to those of either species of Adalia. The larval tracks are mainly composed of alkanes. Those of the two species of Adalia are qualitatively 100% similar and 60% so when the quantitative results are compared. They are, however, only 24–29% similar to those of Coccinella septempunctata.     

焦化厂多环芳烃污染场地的环境评价实证研究

通过实地调查某焦化厂污染场地污染现状,结果表明：该场地土壤和地下水以多环芳烃（PAHs）有机类污染为主,土壤中苯并（仅）芘和多环芳烃污染以1．5m以内表层土污染为主,随着深度的增加浓度逐渐降低,弱透水层表现出良好的阻隔作用。沿着地下水流动方向,地下水中多环芳烃浓度逐渐降低,经过约400m的迁移,由17296μg／L降至111μg/L;苯并（α）芘经过200m的迁移,由18．3μg／L降至0,地下水流动导致污染物在包气带横向扩散。1号孔、3号孔和4号孔所在地表层土壤中苯并（α）芘的单个污染物的致癌风险分别为0．0278、0．0209和1．496,远远超出可接受水平,因此上述孔位所在地附近表层土壤需重点治理。