The effect of density and mutual interference by a Predator: A laboratory study of predation by the Nudibranch Coryphella rufibranchialis on the hydroid Tubularia larynx

The nudibranch Coryphella rufibranchialis (JOHNSTON) feeds on a variety of hydroids, including Tubularia larynx Ellis & Solander. Experiments in which density of prey and predators were altered showed that more prey were eaten as prey density increased. However, more prey were consumed at low predator densities, presumably because of mutual interference among nudibranchs at the higher predator densities. The number of prey consumed per nudibranch was maximal with low predator densities and a ratio of 25–50 polyps per predator. Coryphella seems to show an opportunistic feeding strategy involving solitary predators rapidly depleting hydroid colonies and moving on to new colonies.     

The influence of prey behaviour on prey selection of the carnivorous plant Utricularia vulgaris

The mechanisms underlying differential prey selection of two microcrustaceans by the common bladderwort (Utricularia vulgaris) were studied in the laboratory. Functional response experiments with single prey showed that Utricularia had a higher attack rate coefficient and a longer handling time coefficient with the cladoceran Polyphemus pediculus than with the cyclopoid copepod Eucyclops serrulatus. Observation of predation rate, defined as number of prey eaten per unit time, from direct behavioural observation on single prey species, showed a higher predation rate with Polyphemus than on Eucyclops, at low prey densities. The opposite pattern was found at high prey density. When the two prey were presented simultaneously to the predator, Eucyclops was preferred over Polyphemus. Results from the situation with two prey and some of the results from the direct behavioural observations support field data on the diet of Utricularia, which shows that cyclopoid copepods are selected more frequently than Polyphemus.     

Effects of satiation on feeding and swimming behaviour of planktivores

Hunger affects the feeding and swimming behaviour in fish. After 36 h of food deprivation, the feeding and swimming behaviour of Pseudorasbora parva (Cyprinidae) was studied under different prey densities (0.5, 1, 2, 5, 10 and 25 of Daphnia pulex per liter). The initial feeding rates showed marked variations in relation to prey availability. Under high prey densities, the initial feeding rate of fish was higher and subsequently decreased faster, when compared to those feeding under low prey densities. At higher prey densities, two factors were involved: that of higher prey encounter rates and also the attainment of food satiation at a faster rate. Across all prey densities, the feeding rates of fish reached a plateau after satiation. The swimming speed of fish was found to be negatively related to the prey density and a significant change in swimming speed was noted as being directly related to the level of satiation. It was found that the increasing satiation level greatly influenced the handling time and reactive volume of predator, which finally caused reduced feeding rates.     

The swimming and feeding behavior of Mesocyclops

The swimming and feeding behaviors of Mesocyclops are described from a review of the literature and personal observations. Mesocyclops exhibits considerable behavioral flexibility in response to environmental stimuli. Mesocyclops edax exhibits an increase in horizontal looping behavior at high prey densities, and performs a tight vertical looping behavior in response to the loss of captured prey. Ingestion rates by Mesocyclops are a complex function of prey density, morphology, and behavior in addition to prey size. Vertebrate predators induce a rapid escape response in Mesocyclops and may be responsible at least in part for their extensive diel vertical migrations. The complex behavioral patterns of Mesocyclops suggest that its distribution and abundance in nature will be distinctly nonrandom and influenced as much by its own behavioral responses as by other external physical factors such as water circulation patterns.     

Feeding selectivity of a tropical Chaoborus population

SUMMARY. The Chaoborus population of Lake Lanao, Philippines, was sampled weekly over a 65-week period. Specimens representing all four instars, all times of the year, and two points in the daily migration cycle were dissected for determination of gut contents. Major items in the diet of Lanao Chaoborus include Bosmina, Diaphanosoma, and the copepodid and adult stages of cyclopoid and calanoid copepods. Items that are available but seldom eaten include nauplii and rotifiers except Keratella. Feeding rate averages 2.5% of body weight per day in instars 3–4. Predator size affects but does not fully explain prey selection. Electivity values are much higher for Bosmina and Diaphanosoma than for copepods, even though these food items overlap in size. Bosmina, which has the highest electivity value of any prey, is virtually identical in size to the calanoid nauplius, which has one of the lowest electivity values. This and other similar data suggest that prey of the same size differ greatly in palatability or vulnerability. There is a marked variation in feeding rate and food composition from day to night. The smallest Chaoborus feed more during the day than at night, but the opposite is true for Chaoborus of moderate to large size. Large Chaoborus switch from a daytime emphasis on copepod copepodids and adults to a nighttime emphasis on Cladocera. Diurnal variation between instars in food composition cannot be explained simply on the basis of the vertical distributions of predators and prey. An interaction of vertical distribution with prey density and predator selectivity does explain the overall Chaoborus feeding pattern, however. During the day, larger Chaoborus move deep into the water column where food is scarce. Their daytime feeding rate is lower due to low prey density at great depths. Low prey density is partially compensated by relaxation of preference. At night, upward migration of large Chaoborus into an area of high prey density permits a resumption of marked selectivity and high feeding rates. Small Chaoborus do not descend deep into the water column during the day, as their lower hunting efficiency apparently requires higher food density and use of visual cues to sustain adequate feeding rates. Prey density thus affects both the vertical distribution and feeding selectivity of the Chaoborus population.     

Prey to predator size ratio influences foraging efficiency of larval Aeshna juncea dragonflies

We investigated foraging behaviour of larval dragonflies Aeshna juncea in order to examine the significance of prey density and body size in predator-prey dynamics. A. juncea were offered separately three size-classes of Daphnia magna at low and high densities. The data were collected with direct observations of the foraging individuals. We found that large A. juncea larvae could better enhance their intake of prey biomass as prey size and prey density increased than their smaller conspecifics. However, increasing feeding efficiency of both larval instars was constrained by declining attack success and search rate with increasing prey size and density. With small D. magna, in contrast to large A. juncea, small A. juncea increased their searching efficiency as prey density increased keeping D. magna mortality rate at a constant level. In a predator-prey relationship this indicates stabilizing potential and feeding thresholds set by both prey density and prey-predator size ratio. Attack success dropped with prey size and density, but did not change in the course of the foraging bout. For both A. juncea sizes prey handling times increased as more medium and large prey were eaten. The slope of the increase became steeper with increasing prey-predator size ratio. These observations indicate that components of the predator-prey relationship vary with prey density, contrary to the basic assumptions of functional response equations. Moreover, the results suggest that the effects of prey density change during the ontogeny of predators and prey.     

Diel vertical migrations of invertebrate predators (Leptodora kindtii ,Thermocyclops taihokuensis, andMesocyclops sp .) in a shallow, eutrophic lake

We studied vertical distribution patterns of three invertebrate predators – Leptodora kindtii, Mesocyclopssp., and Thermocyclops taihokuensis– in a shallow eutrophic lake, Lake Suwa , Japan. From June to October in 2000 and 2001, we collected samples in the lake center in order to examine the vertical distribution patterns and the densities of the predators in the water column during the day (0900) and at night (2330). We also examined phototactic behavior of Leptodora in the laboratory. The three invertebrate predators showed clear migration patterns. Leptodora and Thermocyclops displayed a typical migration, avoiding the surface and maintaining a high abundance in deeper water during the day, and being distributed uniformly during the night. Mesocyclops, on the other hand, showed no clear vertical distribution pattern in the water column. However, Mesocyclops showed higher densities in the water column during the night than during the day. It suggests that they stayed just above the bottom during the day and migrated upward during the night. Leptodora also showed such a density difference between day and night. In the laboratory, Leptodora showed strong negative phototactic behavior. The observed density changes between day and night in Leptodora and Mesocyclops suggests the possible underestimation of their population density by usual sampling methods, and thus the impact of predation on populations of prey zooplankton species may also be underestimated in shallow water bodies.     

Prey selection by Chaoborus punctipennis under laboratory conditions

Equal numbers of Diaphanosoma leuchtenbergianum, Daphnia parvula, and Diaptomus pallidus were offered to individual, fourth-instar larvae of Chaoborus punctipennis. When the prey species were presented separately to the larvae, 100% of the Diaphanosoma, 67% of the Daphnia, and 57% of the Diaptomus were consumed. However, when the three species were offered to the predator concurrently, 90% of the Diaphanosoma, 10% of the Daphnia and < 1% of the Diaptomus were eaten. This strong selection for Diaphanosoma was consistent at three prey densities.     

Effect of prey density on behaviour and development of the predatory mosquito,Toxorhynchites towadensis

The effect of prey density on feeding behaviour, killing behaviour, and development of the predatory mosquito,Toxorhynchites towadensis, was investigated in the laboratory. The number of prey consumed per larva increased toward an upper asymptote as prey density increased. Prey consumption curves during fourth instar were concave at low prey densities but convex at high prey densities. This phenomenon was not observed during other instars. Killing without consuming any part of prey occurred at prey densities of 20 per container and over. The number of prey killed but not consumed increased linearly with the number of unconsumed prey in the container. Prey acquisition behaviour was not affected by prey densities during the prepupal period. Developmental time from first instar to adult emergence decreased with increasing prey densities, but remained constant at densities of 10 per container and over. Adult size increased with increasing prey densities but there was no effect at prey densities of 20 and over.     

Foraging activity of two species of predatory leeches exposed to active and sedentary prey

Summary The foraging activity of the freshwater leeches Nephelopsis obscura and Erpobdella punctata during acclimation to active or sedentary prey, starvation and refeeding was examined by filming over a 24 h cycle. For both species, the time of activity of individual leeches and the total number of leeches active were significantly correlated. Activity in the dark was significantly higher than in the light with the activity of N. obscura in the dark 16.2 times higher in terms of time and 6.7 times higher in terms of numbers of individuals moving than E. punctata. Large N. obscura were significantly more active than medium or small individuals and encountered more sedentary prey. With lower levels of activity E. punctata encounters and feeds on more active prey, and in lotic ecosystems its lower exposure to fish predation and drift is, at least partially, responsible for its numerical abundance over N. obscura. There were no significant differences in the activity of N. obscura or E. punctata fed on active or sedentary prey during either subsequent starvation of 1 to 10 d, or after refeeding, suggesting that unchanged foraging activity provides the optimun net energy gain during periods of change in prey type and availability.     

Preference and consumption of Macrolophus pygmaeus preying on mixed instar assemblages of Myzus persicae

This study examines the effects of changes in the prey frequency and abundance on prey selection among the four instars of Myzus persicae by the predator Macrolophus pygmaeus under laboratory conditions. The central hypothesis was that M. pygmaeus will become more selective as prey density increases. It was also observed that M. pygmaeus can occasionally abandon a prey item that had already been killed (non-consumptive prey mortality). It was assumed that the frequency of this behavior would increase with the prey size and prey density. For these purposes prey selection was evaluated by simultaneously presenting all instars of M. persicae to the predator in equal proportions and at increasing densities. M. pygmaeus showed a higher predation rate and a higher preference for smaller prey instars at all prey densities. However, if the predation rate by the predator is expressed in terms of biomass consumed, then biomass gain was higher when feeding on the larger instars of M. persicae. The prey selectivity was indicated by the total prey mortality (consumptive plus non-consumptive prey mortality) as well as by the non-consumptive prey mortality, was associated with relatively high prey densities, depending on the prey instar. Therefore, we argued that the predatory impact of M. pygmaeus on the various instars of the aphid depends not only on prey traits but also on their relative abundance in a patch. Observed decreases in biomass gain from larger prey were likely the result of high prey availability at densities before saturation, which might have caused confusion in the predator’s prey selection.     

An invasive crab alters interaction webs in a marine community

Over the last decade, the non-native, filter-feeding crab Petrolisthes armatus invaded oyster reefs of the South Atlantic Bight at densities of thousands m⁻². Mesocosm and field experiments demonstrated that P. armatus at ∼10–75% of mean summer densities: (1) suppressed growth of small oysters, biomass of benthic microalgae, and recruitment of native mud crabs, (2) enhanced oyster, mussel, and total bivalve recruitment, macroalgal cover, and survivorship of predatory oyster drills, but (3) did not affect native taxonomic richness. Laboratory feeding assays, field tethering experiments, and population changes in field and mesocosm experiments suggest that P. armatus is a preferred prey for native mud crabs and other consumers, thus relieving predation on native species and enhancing recruitment or survival of bivalves and oyster drills. In contrast, the invasive crab can consume crustacean larvae and via this feeding may suppress recruitment of native mud crabs. Our findings should be conservative given the low densities of P. armatus seeded into experimental plots and our inability to run longer-term experiments due to controls rapidly being colonized by non-native crabs recruiting from the plankton. Invasive crabs commonly impact native communities via predation, but community impacts of this invasive crab may be as much due to its role as a preferred prey of native consumers as to its predation on native prey. Given that oysters are foundation species for shallow reefs in the South Atlantic Bight, the long-term effects of this invasion could be considerable.     

Foraging behaviors and the interaction of alewife,Alosa pseudoharengus,and bloater,Coregonus hoyi

Synopsis Alewife, Alosa pseudoharengus, and bloater, Coregonus hoyi, are common planktivores in Lake Michigan. Both alewife and bloater use a variety of feeding modes. Alewives can filter, gulp and particulate feed; bloaters can only gulp and particulate feed. We examined handling time per prey and probability of capture for alewife and bloater particulate feeding on Mysis relicta. Using these estimates and available data for filtering alewives, cost curves were derived for alewife and bloater particulate feeding and for alewife using all three modes of feeding. Alewives filter small prey relative to their own body size and particulate feed on larger prey. Feeding mode appears to be dependent on prey size and density and shifts in feeding mode are apparently based on maximizing biomass eaten per time. The ability to filter confers a competitive advantage on alewife when small prey are abundant as they were in the mid 1960s in Lake Michigan. If the zooplankton are large, bloater young-of-year do not suffer this relative disadvantage. In fact, large bloaters can consume prey on the bottom not available to alewife. This shifting competitive balance may explain, in part, the observed dynamics of alewife and bloater.     

The effects of host plant on the coccinellid functional response: Is the conifer specialist Aphidecta obliterata (L.) (Coleoptera: Coccinellidae) better adapted to spruce than the generalist Adalia bipunctata (L.) (Coleoptera: Coccinellidae)?

Female Aphidecta obliterata consumed an average of 26.6 ± 5.8 Elatobium abietinum and males consumed an average of 19.9 ± 3.2 Elatobium abietinum, but there was no significant difference in the number consumed between the sexes. In Petri dish trials, the larval stages of A. obliterata and all stages of Adalia bipunctata appeared to exhibit a Type II response to prey density, although A. obliterata adults showed a more linear response to prey density. There was no significant difference between the functional responses of the 3rd instars of the two coccinellid species, but there was a significant difference between the responses shown by the 1st instars, with A. obliterata larvae consuming more than those of A. bipunctata, especially at low densities, suggesting that the two species respond differently to an increase in prey density.At low prey densities, adults and 4th instars of both species exhibited a similar response to an increase in prey density but at higher densities the 4th instars and adult stages of A. bipunctata showed higher attack rates when compared with the same stages of A. obliterata.Adult and 4th instar A. obliterata exhibited Type II functional responses on spruce sections. The 4th instar A. obliterata larva appeared to be a more effective predator than the adult of the species, and was more effective when compared with adult A. bipunctata at lower prey densities but A. bipunctata adults appeared to be a more effective predator at higher prey densities.The host plant affected the rate of consumption by adult A. obliterata as adults on Sitka spruce sections consumed significantly higher numbers of aphids than their counterparts on Norway spruce. This was most noticeable at densities above 16 aphids.The distributions of the two coccinellid species in the olfactometer were significantly affected by the presence of host plant material. Aphidecta obliterata adults were found in significantly higher numbers in the Sitka spruce chambers than the control chambers (those without plant material). Adalia bipunctata adults were found in significantly lower numbers in the Norway spruce chamber than the control chamber.Although A. bipunctata has a higher level of voracity than A. obliterata, the latter is more adapted to the spruce environment and the boom and bust population dynamics of E. abietinum.     

Functional responses of five cyprinid species to planktonic prey

Synopsis The functional responses of five species of cyprinids (Chalcalburnus chalcoides, Vimba vimba, Abramis brama, Rutilus rutilus, and Scardinius erythrophthalmus) feeding on four planktonic prey types were measured in the laboratory. Although no alternative prey types were present, the response curves were sigmoid in most cases, because attack rates were not independent of prey density. The findings are explained as being the overt expression of the fishes& foraging tactics. The chief way of maximizing food uptake, according to our interpretation, is accelerating attack rates with increasing prey density. The ability of prey to escape or relative prey size may interfere with this strategy. C. chalcoides, the only obligatory planktivore among the species studied, attacks at higher rates and responds most markedly to changes in prey density.     

Direct uptake of nitrogen by Pfiesteria piscicida and Pfiesteria shumwayae, and nitrogen nutritional preferences

The rates of uptake of a range of forms of nitrogenous nutrients were measured in cultures of Pfiesteria piscicida and Pfiesteria shumwayae maintained at varying physiological states. The measured rates of dissolved N uptake under some conditions approached the rates of N uptake that are achieved through phagotrophy. Rates of dissolved N uptake by P. piscicida contributed <10% of the cellular N of flagellated cells feeding on algae, but were equal to or greater than phagotrophic N acquisition in cells recently removed from fish cultures. Specific N uptake rates (V, h⁻¹) were higher for cells that were maintained on algal prey for long periods (months) than those that were grown with live fish. However, rates of N uptake on a cellular basis for cells grown on or recently removed from fish were comparable to those maintained on algal prey, likely reflecting differences in the sizes of cells of different physiological condition. Preferences for form of N generally followed a decreasing trend of amino acids > urea > NH₄⁺ > NO₃⁻. Nitrate consistently was not a preferred form of N. Although Pfiesteria spp. are often found in eutrophic environments, the relationship between Pfiesteria spp. and nutrient availability is likely to be primarily indirect, mediated through the production of various prey on which Pfiesteria spp. feed. These findings also confirm, however, that when dissolved N concentrations are elevated, they can contribute to the supplemental nutrition of these cells, and thus may provide a significant source of N to Pfiesteria spp. in nature.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Temporal clumping of prey and coexistence of unequal interferers: experiments on social forager groups of brown trout feeding on invertebrate drift

Environmental fluctuations have been proposed to enhance the coexistence of competing phenotypes. Evaluations are here presented on the effects of prey density and short‐term temporal clumping of prey availability on the relative foraging success of unequal interferers in social forager groups of juvenile brown trout Salmo trutta feeding on drifting invertebrate prey (frozen chironomids). Groups of three trout with established linear dominance hierarchies (dominant, intermediate and subordinate) were subjected to three different total numbers of prey, combined with three different levels of temporal clumping of prey arrival, resulting in nine treatment combinations. Higher total number of prey increased the consumption for all dominance ranks, while higher temporal clumping decreased the consumption for the dominant individuals and increased the consumption for the subordinate individuals. The proportion of prey eaten was smaller at high prey numbers. Similarly, there was a trend that increased temporal clumping also decreased the proportion of prey eaten. We conclude that density and temporal clumping of prey contribute to the coexistence of unequal interferers, and that there is a potential positive feedback between prey behaviour and phenotypic coexistence through decreased per capita predation risk for prey that drift synchronously in high densities.     

Feeding by the aquatic heteropteran,Diplonychus rusticum (Belostomatidae): an effect of prey density on meal size

The behaviour of Diplonychus rusticum feeding on chironomid larvae has been investigated under laboratory conditions. Changes in the percentage of material extracted from prey indicated that feeding for the first two minutes enabled the predator to obtain approximately 33% of the available food; feeding for 10 minutes resulted in only 60% extraction. Comparing the percentage of each prey consumed by D. rusticum exposed to various prey densities, it was apparent that predators were more wasteful and ate less of each prey as chironomid density increased. Because the rate of food intake declined as a greater proportion of each prey was extracted, predators exposed to high chironomid densities reduced the amount of each prey consumed thereby conforming to a simple optimal feeding model.     

The effect of water temperature on the functional response of the water stick insect Ranatra dispar (Heteroptera: Nepidae)

Functional response curves were constructed for adult Ranatra dispar feeding on four different densities of notonectid prey at 15, 20, 25 and 30°C. Values for the attack-rate and handling time were estimated from Roger's random predator equation. The most generally applicable response was the type II, with the mean number of prey eaten increasing with increase in temperature. The attack-rate was linearly related to temperature while handling time decreased exponentially with increase in temperature, although values changed very little between 20 and 30°C. It is suggested that changes in metabolic activity and related ‘hunger’ effects on various components of predatory behaviour account for the observed number of prey eaten up to 25°C, however, an increase in the level of prey activity at 30°C may influence the observed number of prey caught (and eaten) at this higher temperature.