Estimating predation rates in experimental systems: scale-dependent effects of aggregative behaviour

We studied the effect of arena size on estimates of prey mortality rate in predation experiments. In a laboratory experiment involving two of the dominating benthic species in the northern Baltic Sea, the predacious isopod Saduria entomon , and its main prey, the amphipod Monoporeia affinis , the prey mortality rate increased with container radius. The densities of both predator and prey close to the wall increased with container size. We hypothesised that this scale-dependent coaggregation of predator and prey caused the mortality rates to increase with arena size. We tested the hypothesis with the help of a simple model, by calculating the expected number of prey eaten in containers of different size from experimental data on the distributions of predator and prey within the arenas. A significant relationship between expected and observed numbers eaten supported our hypothesis. As the aggregative response was most pronounced in large arenas, this leads to the counterintuitive conclusion that large containers produced more biased estimates of morality rates than small containers.
To further study the effects of coaggregation we explored a general simulation model where both predator and prey preferred the habitat close to the arena wall. The model predicted a humpshaped relationship between encounter rate and arena size. This suggests that when predators and prey show a scale-dependent tendency to aggregate along arena walls, the most accurate estimates of predation rates may be obtained with very small or very large arenas.     

Multiple density dependence in two sub-populations of the amphipod Monoporeia affinis: a potential for alternative equilibria

Possible mechanisms for differences in population densities and dynamics were investigated in the amphipod Monoporeia affinis at two deep sites in the northern Bothnian Sea. The two sites were sampled yearly for 10 years. Average sizes, growth and mortality of the different age-classes were estimated from the cohort structure of the two populations. Laboratory experiments also investigated the ability of the common predatory isopod Saduria entomon to cause densitydependent (DD) mortality of the prey M. affinis. At site A, 43 m depth, the average density of M. affinis was twice as high as at site B, 81 m depth. The fluctuations in density were asynchronous between the two sites. Recruitment and subadult sizes of Monoporeia affinis were density dependent at both sites. The main functional difference between the two populations seemed to be the DD mortality of the 1 + cohort that occurred only at the low-density site B. A corresponding DD mortality was found in the predation experiments at densities of 1 + m. affinis corresponding to those found at site B. The potential importance of the predator was also indicated by a significant negative correlation between the biomass of S. entomon and the rate of change in M. affinis density in the field. The similarities in the abiotic factors between the two sites suggested that differences in carrying capacity should be small. The results could be explained by the predation regulation hypothesis for the low-density population at site B, while at site A M. affinis seemed to be regulated by intra-specific competition and limited by predation. It is suggested that in this simple predator-prey system there is potential for the existence of alternative equilibria.     

Functional response of early stages of the cuttlefish Sepia officinalis preying on the mysid Mesopodopsis slabberi

The functional response of Sepia officinalis early stages, preying on mysids of the species Mesopodopsis slabberi was investigated. The effects of five prey densities (12.5, 25, 37.5, 50 and 125 mysids l^-1) and two hatchling ages (1-day-old and 7-day-old) on consumption rate and the frequency of non-feeding animals were tested. Older animals were approximately 50% heavier than newly hatched ones. Hatchlings were individually assayed under 0.25 W m^-2 natural light, 37.8 psu in salinity and 19°C. The effect of prey density on consumption rate was highly significant and no effect of age was detected within the age range tested. Maximal values recorded for consumption rate were about 0.45 mysids h^-1. The frequency of non-feeding individuals was strongly reduced at saturating prey densities. The functional response curve showed an interval of prey densities for which density-dependent prey mortality is probable.     

Gastric evacuation of horse mackerel. II. The effects of different prey types on the evacuation model

Gastric evacuation experiments were performed on horse mackerel Trachurus trachurus. Discrete meals of different sizes of krill Meganyctiphanes norvegicus, mysids Neomysis integer , brown shrimp Crangon crangon , and herring Clupea harengus were fed to test the effect of prey type on the gastric evacuation. A general evacuation model without meal size as a variable was fitted to the data on dry weights by means of non-linear regression. With the exception of mysid evacuation the estimated parameters for curvature, predator weight effect, and temperature effect of the general evacuation model were very similar for all other food types and also confirmed the results obtained with smelt Osmerus eperlanus. In contrast the evacuation of mysids showed a strong exponential curvature, rather the effect of prey size than of prey type. The prey specific evacuation constants of the model revealed a linear relationship to the energy content. A general consumption model for field estimates is presented that can be used for prey types that have not yet been tested in the experiments, provided the energy content is known.     

Diet specialization in a fluctuating population of Saduria entomon: a consequence of resource or forager densities?

Intraspecific competition has been shown to favor diet specialization among individuals. However, the question whether the competition takes the form of interference or exploitative in driving diet specialization has never been investigated. We investigated individual diet specialization in the isopod Saduria entomon, in relation to forager and resource biomasses in a system that exhibits predator–prey fluctuations in density. We found that individual diet specialization was only affected by the biomass of their preferred prey (Monoporeia affinis) and not by Saduria biomass; diet specialization was higher when Monoporeia biomass was low compared to when there were high Monoporeia biomass. Population diet breadth increased at low Monoporeia biomass whereas individual diet breadths were marginally affected by Monoporeia biomass. Overall, this led to the increase in diet specialization at low Monoporeia biomass. This study shows that predator–prey dynamics might influence diet specialization in the predator and that resource biomass, not forager biomass might be important for individual diet specialization.     

Effects of experimental conditions on the feeding rate of Mysis mixta (Crustacea,Mysidacea)

Predation by the Baltic Sea mysid Mysis mixta onnewly hatched Artemia sp. and culturedDaphnia magna was examined in the laboratory. Wedesigned experiments to study the effects onconsumption rate of different experimental conditions:light, container size, diel variation, duration ofadaptation period, starvation and the duration of afeeding experiment.Consumption differed significantly between mysids keptin bright light vs mysids in complete darkness, withabout twice the consumption in darkness. Acclimationto light conditions prior to the experiment did notinfluence this result. The duration of an experimenthas large effect on estimated consumption. Predationrates in 1-h experiments were up to 60% higher thanthose recorded in 4-, 12- and 24-h incubations.Consumption rates of starved mysids were significantlyhigher (27% on average) than that of unstarvedindividuals. No significant diel variation inconsumption was observed over an experimental periodat 36 hours. We found a tendency, however notstatistically significant, that the food consumptionincreased when the container size ws increased between1 and 8 l.     

Vulnerability of early life intervals of Coregonus hoyi to predation by a freshwater mysid,Mysis relicta

Synopsis The bloater, Coregonus hoyi, deposits its eggs on deep sediments (70–100 m in Lake Michigan), where its eggs and embryos can be exposed to epibenthic predators. We investigated the vulnerability of early life intervals of bloaters to predation by mysids, Mysis relicta, which are mostly epibenthic by day and planktonic at night. Bloaters were raised from spawn in the laboratory and presented to field-collected mysids in laboratory predation trials. Eggs were not ingested by the mysids. Embryonic bloaters were vulnerable to predation by mysids only during the interval between hatching and swim up, usually 1–24 h under laboratory conditions. The mysids required about a day of exposure to this novel prey before they were able to kill significant numbers of the bloater embryos by making successive attacks with their thoracic legs. In experiments with experienced (2 and 3 days with bloater embryos) mysids, a functional response between embryo density and mysid predation rates was apparent. Temperature and the presence of alternative prey (zooplankton) did not alter the ‘kill rate’ (about 2.5 embryos mysid^-1d^-1) of experienced mysids at high bloater densities (>4 bloaters/mysid). However, more embryos were partially, rather than completely, ingested at 4 versus 9° C and in the presence of zooplankton.     

Predation by the isopod Saduria entomon on the amphipods Monoporeia affinis and Pontoporeia femorata: experiments on prey vulnerability

Summary Predation by Saduria entomon on the depositfeeding amphipods Monoporeia (=Pontoporeia) affinis and Pontoporeia femorata was studied in laboratory experiments. Prey vulnerabilities were compared in singleprey treatments and in mixed-prey treatments, where the proportions of the two species were varied at one total density. In a pilot experiment, P. femorata was the more vulnerable prey, both in single-prey and mixed-prey treatments. In later experiments, the amphipod species were equally vulnerable in single-prey treatments, while in mixed-prey treatments Saduria preyed preferentially on M. affinis, even when this prey was rare, i.e. it did not switch. We suggest that the different result in the later experiments may have been due to a change in the search mode of the isopods.     

Individual variation in prey choice in a predator-prey community

One predator-two prey community models are studied with an emphasis on individual variation in predator behavior. The predator behaves according to a well-known prey choice model. The behavioral model predicts that predators should always attack the primary prey (more profitable prey of the two), but only attack the alternative prey (less profitable prey of the two) when the density of the primary prey is below a threshold density. The predator that accepts the alternative prey does not discriminate between the primary and alternative prey (all-or-nothing preference for the alternative prey). However, empirical studies do not result in clear all-or-nothing responses. Previous models examined the relaxation of the all-or-nothing response by assuming partial preference (e.g., predators preferentially forage on the primary prey even when they also attack the alternative prey). In this study, I consider individual variation in two predator traits (prey density perception and handling time) as the sources of the variation in the threshold density, which can make empirical data appear deviated from the expectation. I examine how community models with partial preference and individual variation differ in their dynamics and show that the differences can be substantial. For example, the dynamics of a model based on individual variation can be more stable (e.g., stable in a wider parameter region) than that of a model based on partial preference. As the general statistical property (Jensen’s inequality) is a main factor that causes the differences, the results of the study have general implications to the interpretation of models based on average per-capita rates.     

CONSEQUENCES OF ALTERNATIVE FUNCTIONAL RESPONSE FORMULATIONS IN MODELS EXPLORING WHALE-FISHERY INTERACTIONS

We evaluated the utility of Ecosim for exploring interactions between cetacean predators, their prey, and fisheries. We formulated six Ecosim parameterizations, representing alternative hypotheses of feeding interactions (functional response) between cetaceans and their main fish prey, and examined differences in the predicted responses to simulated harvesting regimes for minke whales and their prey. Regardless of the type of function response formulated, intense fishing on the main fish prey of minke whales had a longer-lasting negative impact on minke whales than when minke whale biomass was removed directly by harvesting. Consumption rate, biomass, feeding time and mortality of minke whales were all sensitive to the type of functional response specified. Inclusion of "handling time" limited minke whales consumption at high prey densities and predicted higher consumption at low prey densities; features characteristic of a type II functional response. Predicted decline and recovery rates of minke whales were slower than when consumption rates were not limited. Addition of "foraging time" adjustments resulted in more conservative estimates of decline and recovery. However, when "other mortality" was linked to time spent foraging, exposure to higher mortality at low prey densities, and reduced mortality at high prey densities resulted in dramatic differences in predicted biomass trajectory. Sensitivity to the "other mortality" assumption is important for cetaceans whose predation mortality is only a small proportion of total mortality. Differences in the feeding and biomass dynamics were also observed when prey availability to predators was represented by changes in prey vulnerability, confirming earlier reports that Ecosim predictions are sensitive to this parameter.     

Predation by Neomysis mercedis: effects of temperature, Daphnia magna size and prey density on ingestion rate and size selectivity

1. Neomysis mercedis predation rates on Daphnia magna were determined under laboratory conditions. There were generally no consistent differences between the number of Daphnia ingested at 10 and 14°C. 2. At each temperature, the number of prey consumed increased with mysid size and decreased with Daphnia size. 3. For small prey the relationship between ingestion rate and prey density represented a Type II functional response. However, for larger prey there was no significant relationship between density of prey and consumption by mysids. 4. The pattern of size-selective predation by Neomysis was studied to test the optimal foraging hypothesis. For prey populations with mixed size classes, the smallest size of prey was consumed most frequently but intermediate size prey provided the greatest biomass. These observations are contrary to our predictions based on calculations of profitability of different sizes of prey.     

Are digestive characteristics important contributors to the profitability of prey?

Summary In the field, Spinachia fed on four types of prey; copepods, isopods, mysids and amphipods. As fish size increased, mysids gradually succeeded amphipods as the most important food type in the diet. Prey dimensions and morphometry of the fish's mouth most accurately predicted capture efficiency for amphipods, whereas for mysids capture efficiency was determined by the prey's escape response and the fish's fast-start capability. Responses to model prey revealed the ability of fish to differentiate among contrasting prey characteristics, resulting in the adoption of appropriate predatory tactics. Amphipods were associated with a shorter gut evacuation time than mysids, although approximately equal proportions of energy were absorbed from each. Similar rations of mysids and amphipods were required to satiate fish. The lower energy content per unit dry mass of amphipods was off set by their lower water content. From pre-digestive behaviour, we predicted that mysids were more profitable than amphipods, and this was reflected in the fish's choice. Conversely, incorporating the net rate of energy uptake by the gut led us to predict that amphipods were more profitable. Although physiological constraints clearly influence the net rate of energy uptake, it appears that dietary preferences are based on pre-digestive predatory behaviour and hence on time minimisation.     

Trophic interactions involving mysid shrimps (Mysidacea) in the near-bottom habitat in the Baltic Sea

In a six month mesocosm tank experiment, hypotheses were tested concerning the role of benthopelagic mysid shrimps (Mysidacea) in the near-bottom food web of the Bothnian Sea, in the northern Baltic Sea. The first hypothesis tested was that the mysids interact, through predation, with benthic deposit-feeding Monoporeia affinis amphipods. A second hypothesis tested was that the sediment type is important for the overwintering success of the mysids. Changes in abundance and mass were recorded for M. affinis and mysids when separate and when coexisting, in two sediment types differing in organic content (food level); soft muddy clay (rich) and fine sand (poor). Despite the fact that newborn M. affinis offspring, a plausible target for predation by mysids, were present in substantial numbers in the tanks, no consistent evidence for any interaction between these taxa was found. The biomass of mysids was slightly higher in the muddy clay than in the sand tanks, and the mechanism behind this substrate effect is discussed. A third hypothesis, that the mysids interact with near-bottom zooplankton, was investigated. The tanks were continually supplied with in situ near-bottom sea-water containing a semi-natural assemblage of near-bottom plankton. As a result of mysid predation, tanks with mysids had lower abundance and biomass of cyclopoid copepods than tanks without mysids. Thus, the major interaction found was predation on near-bottom zooplankton by mysids and it is suggested that this interaction could potentially be an important food link, especially during periods with low food availability in the pelagic system.     

Role of predation in short-term population fluctuations of some birds and mammals in Fennoscandia

Summary We tested the hypothesis that synchronous fluctuations in small game species in boreal Fennoscandia are caused by varying predation pressure. The main prey of predators are the cyclically superabundant voles. Small game species (alternative prey) are rare compared to voles. The following 4 predictions were checked: (1) Predators should shift their diet from main prey to alternative prey as main prey decline. — This was confirmed using data on red fox (Vulpes vulpes L.) diet.; (2) The mortality rate of alternative prey should be inversely correlated to the abundance of main prey. — This was true for mountain hare (Lepus timidus L.) mortality rates and the rate of nest predation on black grouse (Tetrao tetrix L.).; (3) The total consumption of prey by all the predators should at least equal the critical losses in alternative prey during a decline year. — A tentative estimate of predator consumption amounted to 10 times the losses in grouse and hare.; and (4) The absence of synchrony between the species in the boreonemoral region should be associated with a more diverse diet of predators. — This was the case for red fox diets throughout Sweden. Although all 4 predictions were confirmed, we could not necessarily exclude other hypotheses involving changes in quality or quantity of plant food.     

Assessment of the interaction between the white shrimp, Palaemon longirostris, and the exotic oriental shrimp, Palaemon macrodactylus, in a European estuary (SW Spain)

Results of field surveys, stomach content analysis, and laboratory measurements of oxygen consumption at different salinities in the exotic shrimp Palaemon macrodactylus and its native counterpart P. longirostris from the Guadalquivir estuary (SW Spain) were used to assess interaction between both species. After its first record in this estuary in 1998, P. macrodactylus has shown a gradual increase in density and its estuarine population has been clearly dominated by juveniles. Feeding habits of both shrimps indicated a strong trophic overlap between them, with mysids as main prey. Their salinity-related and spatial distribution patterns show that maximal inter-specific overlap between the two populations occurs in the inner, less saline part of the estuary. Specific oxygen consumption rates under different salinities and water oxygen concentrations suggest a more efficient metabolism and a higher tolerance to hypoxic conditions in brackish waters by P. macrodactylus than by P. longirostris. Such inter-specific physiological differences could have made it possible for the exotic P. macrodactylus to invade an estuarine stretch that, prior to its arrival, was infra-utilized by native species.     

Spatial-temporal patterns in intra-annual gray whale foraging: Characterizing interactions between predators and prey in Clayquot Sound, British Columbia, Canada

In Clayquot Sound, British Columbia, gray whales (Eschrichtius robustus) forage primarily on mysids (Family Mysideae) and also on crab larvae (Family Porcellanidae) that are constrained to specific habitat, which relate to bathymetric depths. In this paper we characterize the interactions of gray whales and their prey by analyzing fine scale spatial‐temporal patterns in foraging gray whale distribution within a season. Kernel density estimators are applied to two seasons (1998 and 2002) of high‐resolution data on foraging by gray whales. By partitioning data from each foraging season into several time periods (12 in 1998 and 11 in 2002), using a temporal autocorrelation function, and generating kernel density estimated surfaces for each time period, it is possible to identify discrete areas of increasing and declining foraging effort. Our results indicate that gray whales forage on mysids throughout a season and opportunistically forage on crab larvae. The episodic crab larvae feeding may reduce, but not eliminate, pressure to mysid populations enabling mysids to reassemble swarms and continue to support gray whale foraging in the latter part of the season. Results suggest that when managing marine environments, gray whale populations require multiple and connected habitats for summer foraging.     

Seasonal variation of gelatinous plankton consumption by fish in the South-western Atlantic Ocean: a question of strategy*

Gelatinous plankton is an important food resource for several species of fishes in the Southwestern Atlantic Ocean. Some fish depend heavily on these organisms and are specialized to feed on ctenophores, salps and medusae, while others only consume gelatinous plankton occasionally. We hypothesize that consumption of gelatinous plankton by fishes represents an alternative food resource when primary prey are not available during cold periods in the study area. To determine seasonal variations of gelatinous plankton consumption by fishes, data samples were grouped into cold and warm periods. A total of 64,567 stomachs belonging to 106 species of fish were analysed, of which 32,943 (51%) contained food items. Of those containing food items, 2719 (8.2%), from 38 fish species, contained gelatinous items. Fourteen species ingested gelatinous in warm period, nine in cold period, and 15 in both periods. The proportion of stomachs with gelatinous was significantly greater during the cold period. Ctenophores were the most predominant prey in both periods, followed by salps and medusae. Consumption of ctenophores, salps and medusae was unevenly distributed within the area during the different periods. Classification methods (group average sorting utilizing Bray–Curtis similarity measures based on log (X + 1) identified eight areas of consumption. SIMPER (similarity percentages) analyses revealed that nine fish species contributed most to gelatinous plankton consumption. The seasonal and spatial variation of gelatinous consumption by fish would be related to the availability of food in each period. Strategies of gelatinous consumption, including survival, feeding opportunities and prey specialization, are discussed.     

Cyclic voles, prey switching in red fox, and roe deer dynamics – a test of the alternative prey hypothesis

Medium-sized predators sometimes switch to alternative prey species as their main prey declines. Our objective of this study was to test the alternative prey hypothesis for a medium sized predator (red fox, Vulpes vulpes ), a small cyclically fluctuating main prey (microtine voles) and larger alternative prey (roe deer fawns, Capreolus capreolus ). We used long-term time series (28 years) on voles, red fox and roe deer from the Grimsö Wildlife Research Area (59°40'N, 15°25'E) in south-central Sweden to investigate interspecific relationships in the annual fluctuations in numbers of the studied species. Annual variation in number of roe deer fawns in autumn was significantly and positively related to vole density and significantly and negatively related to the number of fox litters in the previous year. In years of high vole density, predation on roe deer fawns was small, but in years of low vole density predation was more severe. The time lag between number of fox litters and predation on fawns was due to the time lag in functional response of red fox in relation to voles. This study demonstrates for the first time that the alternative prey hypothesis is applicable to the system red fox, voles and roe deer fawns.     

Effects of visual conditions and prey density on feeding kinetics of paralarvae of Octopus vulgaris from a laboratory spawning

Consumption rate and feeding success of newly hatched paralarvaeof the cephalopod Octopus vulgaris preying on Artemia larvaewere investigated in relation to visual conditions and preydensity. Each paralarva was tested individually using a small-scaleexperimental setup; consumption over one day was measured at20°C. A factorial experiment was designed to investigatethe effects on the consumption rate of two predictor variables:illumination/background (three levels: 7.5 Wm^–2 whitelight + white background, 7.5 Wm^–2 white light +blue background, darkness) and prey density (four levels: 2.35,4.70, 9.40 and 14.10 Artemia metanauplii ml^–1). Consumptionrate varied significantly between different conditions of illuminationand prey density. Light enhanced consumption rate, but differentbackgrounds yielded similar rates. The maximal consumption rateunder illumination was close to 16 Artemia paralarva^–1day^–1, and it was around 5 Artemia paralarva^–1 day^–1for assays in darkness. The predatory efficiency, measured asthe proportion of prey consumed, was significantly affectedby prey density, pointing to a type III functional response.The number of nonfeeding paralarvae was significantly higherin darkness and at low prey density. (Received 14 February 2006; accepted 23 December 2006)     

Evaluation of predation abilities of Blattisocius dolichus (Acari: Blattisociidae) on a plant-parasitic nematode, Radopholus similis (Tylenchida: Pratylenchidae)

Predation and predatory behavior of Blattisocius dolichus on Radopholus similis were tested both in experimental arenas and on potted plants. Predation occurred in all active stages of B. dolichus. Blattisocius dolichus preferred live R. similis when offered together with Caneorhabditis elegans and dead R. similis in a choice test. Consumption rate was affected by temperature, prey density and duration of starvation. Maximum consumption rates were observed at 25 °C, for both adult males and females after being starved for 96 and 72 h, respectively. Consumption rate increased with increasing prey density until satiation was reached, when the predator–prey ratio was 1:250 for both male and female predators. Anthurium andraeanum seedlings, artificially infested with R. similis (1,000 per pot), were used to evaluate the biological control efficiency of B. dolichus. The nematode density decreased by 66 % 10 days after a release of 500 mites per pot.