Vulnerability of newly settled plaice (Pleuronectes platessa L.) to predation: effects of habitat structure and predator functional response

Plaice (Pleuronectes platessa) nursery grounds on the Swedish west coast have been subject to increasing cover of annual green macroalgae during recent years, with growth of algae starting at the time of plaice settlement in April to May. A laboratory experiment was performed to investigate how the vulnerability to predation of metamorphosing plaice was affected by the presence of filamentous algae. Predation by shrimps (Crangon crangon) on settling plaice larvae was higher on sand than among algae, whereas predation by crabs (Carcinus maenas) was unaffected by habitat type, suggesting a lower overall mortality of plaice in the vegetated habitat. When predators and prey were presented with a combination of the two habitats, predation by shrimps was as high as that in the sand treatment alone, whereas predation by crabs was lower than that in the two treatments with one habitat. Based on these results, an additional experiment was performed, investigating the functional response of shrimps to six densities of juvenile plaice in a sand habitat with alternative prey present. The proportional mortality of juvenile plaice (12-16 mm total length (TL)) was density-dependent and was best described by a type III (sigmoid) functional response of the predatory shrimps. The results suggested that the combined predation pressure from shrimps and crabs was lower among algae than on sand, but settling plaice and predatory shrimps chose the sand habitat. Plaice densities in the sigmoid part of the obtained functional response curve represented normal to high field densities of plaice on the Swedish west coast, suggesting that shrimp predation could have a stabilising effect on plaice recruitment. The formation of macroalgae mats could therefore lead to a concentration of plaice juveniles in the remaining sand habitat and increased mortality through density-dependent predation by shrimps.     

An optimal foraging and migration model for juvenile plaice

Summary A state-dependent model has been used to predict daily and tidal patterns of migration, feeding and inactivity in juvenile plaice (Pleuronectes platessa L.) in their intertidal and shallow subtidal nursery areas. Vertical position, quantity of energy reserves and fullness of the gut characterized the state of individual fishes. If feeding is visually mediated, the model predicts that, by night, plaice should move to areas of low predation risk and become inactive, whereas by day, plaice should migrate to feed in areas of high prey encounter rate. Vertical zones adopted by day and night and, hence, patterns of migration, should depend on the distributions of predators and prey. When prey are more abundant in the intertidal zone, plaice should move onshore to feed as the tide rises and when prey are more abundant offshore, plaice should move offshore to feed by day. If predators are equally abundant in all zones, the fish should behave as if no predators were present, having no effective refuge. An increase in the abundance of predators with depth results in the restriction of plaice activities to shallower vertical zones, depending on the magnitude of the predation threat. Zones adopted thus depend on the trade-off between energy intake and predation risk. Concordance between predicted behaviour and observed patterns is evident in contrasting habitats. Migration and feeding in the Wadden Sea, where prey are more abundant on intertidal flats, is dominated by the tidal component, whereas on impoverished exposed beaches of the west coast of Scotland, the diurnal component is dominant. Tidally related behaviour persists in the latter environment, not predicted by the model and may be a consequence of using endogenous rhythms to approach optimal behavioural patterns.     

Effects of predation and habitat structure on the population dynamics of house mice in large outdoor enclosures

This paper examines the effect of different levels of protection from predation on feral house mice. Mice were contained in eight 50×50 m outdoor enclosures. Enclosures allowed access to a suite of freeliving vertebrate predators from the surrounding area, including feral foxes, feral cats and Australian raptors. A 10–15% cover of small, felled cypress pine trees was added in strips to low grassland to increase habitat complexity. Mice were not protected from predation when compared with low grassland pens, possibly because predators were able to focus their hunting activity in the strips. However, when felled trees were covered with wire netting, hence providing higher quality refuge, mouse populations achieved higher densities than in low grassland pens. A predator exclusion treatment was used to confirm the refuge effect was due to a reduction in the impact of predation. Survival rates under the different treatments were generally consistent with population level responses, with mice having lower survival in low grassland pens than in high refuge pens. This is the first study with mammals that confirms the importance of refuges from predators for prey populations.     

Spatially heterogeneous refugia and predation risk in intertidal salt marshes

The effect of habitat structure on interactions between predators and prey may vary spatially. In estuarine salt marshes, heterogeneity in refuge quality derives from spatial variation in vegetation structure and in tidal inundation. We investigated whether predation by blue crabs on periwinkle snails was influenced by distance from the seaward edge of the salt marsh and by characteristics of the primary habitat structure, smooth cordgrass ( Spartina alterniflora ). Spartina may provide refuge for snails and interfere with foraging by crabs. Furthermore, predation risk should decline with distance from the seaward edge because landward regions require more travel time for crabs during tidal inundation. We investigated these processes using a comparative survey of snails and habitat traits, an experiment that assessed the crab population and measured predation risk, and a size-structured model that estimated encounter rates. Taken together, these approaches indicated that predation risk for snails was lower where Spartina was present and was lower in a landward direction. Furthermore, Spartina architecture and distance from the seaward edge interacted. The strength of the predation gradient between seaward and landward regions of the marsh was greater where Spartina was tall or dense. These predation gradients emerge because vegetation and distance inland decrease encounter rates between crabs and snails. This study suggests that habitat modification, a process not uncommon in salt marshes, may have consequences for interactions among intertidal fauna.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

Vertical structure and predator refuge mediate oyster reef development and community dynamics

In addition to their economic value, oysters supply a number of critical ecosystem services such as controlling eutrophication, recycling nutrients, and providing habitat for many invertebrates and fishes. Thus, the maintenance and restoration of oyster reefs is vital for properly functioning estuarine ecosystems. We hypothesized that spatially heterogeneous and refuge-rich reefs are superior habitats for oysters and associated invertebrates and fishes. If this is shown to be the case, then vertical structure and refuge should be included in the design of newly constructed oyster reefs and conserved in existing reefs. To test our hypothesis, models were built as concrete blocks into which shells were placed horizontally, vertically, and widely spaced, and vertically and closely spaced. Respectively, the units represent horizontal orientation with no refuge (HNR), vertical orientation with no refuge (VNR), and vertical orientation with refuge (VWR). Flume studies of velocity distributions over each model type showed distinctive differences that may contribute to differential settlement on the models in the absence of predators or sediment. A subsequent settlement study in the same flume showed significantly higher settlement densities to horizontal shells than to vertical shells regardless of refuge (HNR>VNR=VWR). This result indicates the primary importance of settlement surface orientation (horizontal vs. vertical) over predation refuge when predators are absent. In contrast, two field recruitment studies indicated that the importance of orientation relative to predation refuge might depend on sedimentation rates. When sedimentation was low, the number of oysters and barnacles (live, dead, and total) was significantly higher on no-refuge models than on refuge models (HNR=VNR>VWR). However, percent mortality was lower on the refuge models than on the no refuge models (HNR=VNR>VWR). This leads to a conclusion that although fewer oysters may settle on refuge shells, those that do have a higher rate of survival. In a year when sedimentation was high, recruitment and survival of oysters, barnacles, and bryozoans was higher on vertical models regardless of refuge. In aggregate, our results indicate that both vertical orientation and predation refuge are critical to the early development of the oyster reef community and should be included in restoration designs.     

Variation in plant stem density and its effects on foraging success of juvenile bluegill sunfish

Synopsis Juvenile bluegill sunfish, Lepomis macrochirus, are known to use beds of aquatic vegetation as a refuge from predators. This study examines the effects of increasing plant stem density on juvenile bluegill foraging. Three stem densities (100, 250 and 500 stems m⁻²), varying in their refuge potential for bluegills from predators, were tested. Results demonstrate that stem densities chosen as a refuge from predation (i.e. 500 stems m⁻²) significantly reduced bluegill foraging success and increased time required to capture prey. Therefore, juvenile bluegills seeking safety in vegetation may be faced with a trade-off between foraging success and effective refuge from predation when choosing among plant stem densities.     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Feeding and Refuge Use by Small Fish in the Presence of Cyanobacteria Blooms

We studied the effect of cyanobacteria on foraging and refuge use in small fish. We measured pike larval feeding in the presence of cyanobacteria by counting leftover prey. Our results showed that feeding by pike larvae on zooplankton prey decreased significantly in the presence of non-toxic cyanobacteria. The behaviour can be due to lowered vision caused by turbidity or clogging of the gills. Further, we tested whether the three-spined stickleback use toxic cyanobacteria as a refuge against predators in a choice experiment. The choice experiment was performed in a Y-maze fluviarum, where the fish could select between two different environments. Our results support the refuge use hypothesis because the three-spined stickleback clearly preferred toxic cyanobacteria to the chemical predator signal. To conclude, cyanobacteria decrease feeding rates in fish larvae, but may function as important refuge for e.g. sticklebacks, during predation pressure in pelagic algal blooms.     

Plaice nurseries: effects on recruitment

The function of coastal nursery areas is discussed in relation to the variability in North Sea plaice recruitment. Since 60% of the recruitment of juveniles originates from the Wadden Sea, special attention is paid to this area. The concentration of juveniles in a restricted area seems to evoke only adverse effects: an increased risk of food limitation and hence reduced growth, and an increased vulnerability to predation. Despite these expectations, growth of most of the plaice in the Wadden Sea has always been optimal within the wide range of year-class strength observed and depends only on ambient water temperature. The same situation is indicated for some British bays, where the growth of 0-group plaice is far lower than in the Wadden Sea, because of lower temperatures. Mortality through predation seems to be relatively low in the Wadden Sea and restricted to only a short period, because of the absence of almost all potential predators. In the more open British bays higher mortalities are found, probably due to the presence of a number of predatory fish species. The low variability in the recruitment of plaice might be the combined result of optimal growth with the absence of between-year fluctuations in predator abundance in the Wadden Sea. As a result, observed mortality only depends on prey abundance and is therefore density-dependent both within one year and especially between years, reducing variations in recruitment. This suggestion is supported by the situation in more open British bays: here too, growth is maximal, but the abundance of predators shows larger fluctuations between years and, as a result, greater fluctuations in mortality are observed.     

Habitat overlap of enemies: temporal patterns and the role of spatial complexity

Environmental heterogeneity can promote coexistence of conflicting species by providing spatial or temporal refuges from strong interactions (e.g., intraguild predation, competition). However, in many systems, refuge availability and effectiveness may change through time and space because of variability in habitat use by either species. Here I consider how the intensity of intraguild predation risk varies from day to night for aquatic insects that use both vegetated and open water habitats. Large (1,265 l) and small (42 l) mesocosms were used to test the hypothesis that Buenoa would choose an open-water habitat that minimized predation by the ambush predator Notonecta during the day, but that at night Buenoa would safely use both vegetated and open water. Regardless of container size, Notonecta remained in vegetated water during the day and exploited both habitats at night, despite exhibiting greatest instantaneous predation rates in open water during the day. In contrast, Buenoa maintained an even distribution throughout the mesocosms during day and night, even though habitat-specific predation risks were fivefold lower in open waters than in vegetation during the day and habitat-specific predation risk would have been reduced threefold by fully exploiting open waters. Thus, temporal heterogeneity was both beneficial and detrimental to Buenoa; darkness of night reduced predation, but spatial refuges also disappeared. Together, these patterns suggest that while environmental heterogeneity can dampen intense biotic interactions, enemies do not select habitats solely on the basis of conflict avoidance. Instead, it appears that habitat-specific variation in other biotic (e.g., visual predators) or physical factors (e.g., UV radiation) may also mediate species interactions by influencing habitat selection.     

Cladoceran assemblages, seasonal succession and the importance of a hypolimnetic refuge

SUMMARY. 1. We investigate the importance of a refuge from fish predation to the abundance, species composition and seasonal succession of zooplankton. Thirty lakes representing a range of depths were sampled twice in summer for physical/chemical parameters and zooplankton community structure.
2. We define the refuge from centrarchid predators to be that space between the thermocline and the zone of anoxia. As lakes vary in rate of oxygen depletion from the hypolimnion. the refuge size and lake depth are independent: refuge size decreases during the summer period.
3. Lake depth and refuge size independently explain variation among lakes in zooplankton species composition, but seasonal community change within lakes is best predicted by loss of refuge size.
4. Refuge size also explains the substantial variation in the relative dominance of the two major daphnid species. Lakes possessing a large refuge are dominated by D. pulicaria ; those with a small refuge are dominated by the smaller, D. galeata mendotae . We suggest that lakes of intermediate refuge size, which are characterized by high species diversity, represent a more equitable balance of predation and competition.     

Effect of habitat complexity on the predation of Buenoa fuscipennis (Heteroptera: Notonectidae) on mosquito immature stages and alternative prey

Notonectids are well‐known predators in aquatic habitats, where mosquito larvae, chironomids, and cladocerans constitute their main diet. Our purpose was to assess the effect of structural complexity on the predatory ability of Buenoa fuscipennis, a common predator in aquatic habitats of Buenos Aires city (Argentina). Buenoa fuscipennis showed type 2 functional responses in both the presence and absence of prey refuge and no differences in attack rate or handling time between refuge treatments. Regarding mosquito size classes, B. fuscipennis exhibited a significantly higher preference for 2^nd instar larvae and no predation on pupae. In the presence of mosquito larvae and alternative prey, B. fuscipennis preferred mosquitoes over chironomid larvae and adult cladocerans over mosquito larvae. No switching behavior was detected in our experiments. Habitat structure only slightly affected the predator´s consumption rates on mosquito larvae. Overall, preference for prey did not vary with the presence of refuge, except for the preference for mosquitoes over chironomid larvae, which was significantly decreased in the presence of refuge as a consequence of reduced predation on mosquito larvae. The results suggest that B. fuscipennis could efficiently control mosquitoes in structurally simple habitats where chironomids are the most abundant alternative prey but not in temporary pools where cladocerans are abundant.     

Size-specific predation on marine invertebrate larvae

Predation on planktonic larval stages is frequently a major source of mortality for the offspring of benthic marine invertebrates. Mortality rate likely varies with larval size and developmental stage, but few experiments have measured how these factors affect predation rates. I used experimental reductions in egg size to test how variation in larval size affects the likelihood of predation during planktonic development. Blastomeres of the sand dollar Dendraster excentricus were separated at the two-cell stage to produce half-sized zygotes. Larvae resulting from this manipulation were tested for their susceptibility to predation relative to whole-sized siblings at four ages. Individuals from each size class were simultaneously presented as prey items to five predators (crab zoeae, crab megalopae, chaetognaths, solitary tunicates, and postlarval fish) in the laboratory. Four predators consumed significantly more half-sized larvae than whole-sized larvae, but one predator type (postlarval fish) consumed more whole-sized larvae. Predators that consumed more half-sized larvae also preferentially consumed younger larvae. In contrast, postlarval fish showed no significant prey preference based on larval age. These results suggest that assumptions of constant mortality rates during development should be modified to account for the effects of larval size and age.     

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.     

Observation of epigeic predators and predation on artificial prey in a cereal field

Predation activity of different epigeic predators on artificially supplied prey was studied in a cereal field during several 24 h periods from spring to early summer 1993. Carabids were the most frequently observed predators. Among themPoecilus cupreus was the most voracious species during day-time, andAgonum muelleri during night-time. Predation rates were highest between 10.00 and 18.00 h and lowest between 02.00 and 10.00 h. Linear regressions showed significantly positive relations between temperature and predation rates, and between predation and activity density of epigeic predators. The latter relation was stronger when only activity density of predators which were observed to consume baits was included. Quadrat sampling and pitfall sampling showed that with the former only a small proportion of bait-consuming predators were caught whereas with the latter method this proportion was high.     

Predation by sprat and herring on pelagic fish eggs in a plaice spawning area in the Irish Sea

Predation on planktonic fish eggs was examined by stomach content analysis of sprat and herring sampled in a plaice spawning area to the east of the Isle of Man in March 1993. Plankton samples were taken to examine prey selection. The clupeids showed a selection for the later developmental stages of plaice eggs. Plaice eggs had a refuge in size from predation by sprat <80 mm total length which selected smaller non-plaice eggs. However, herring and sprat >80 mm total length showed a strong selection for plaice eggs, presumably due to their large size.     

Predation rates on decapod crustaceans in latitudinally separated seagrass communities: a study of spatial and temporal variation using tethering techniques

Field experiments were conducted over a 5-yr period to test the relative risk of predation for decapod crustaceans in seagrass and adjacent sand habitats in New Jersey, Florida, and Bermuda. These studies showed that vegetation provides a significant but temporally variable refuge from predation in all geographic locations. Brachyuran crabs were preyed on most heavily in lower latitudes and hermit crabs were taken only in Florida. There was no observable relationship between predation rates on hermit crabs and presumed antipredatory architectures or ornamentation of their shells. Spatial and temporal variation in predation risk of the nature we observed should influence both life history characteristics and the behavioral ecology of prey species.     

Population dynamics of thrips prey and their mite predators in a refuge

Prey refuges are expected to affect population dynamics, but direct experimental tests of this hypothesis are scarce. Larvae of western flower thrips Frankliniella occidentalis use the web produced by spider mites as a refuge from predation by the predatory mite Neoseiulus cucumeris. Thrips incur a cost of using the refuge through reduced food quality within the web due to spider mite herbivory, resulting in a reduction of thrips developmental rate. These individual costs and benefits of refuge use were incorporated in a stage-structured predator-prey model developed for this system. The model predicted higher thrips numbers in presence than in absence of the refuge during the initial phase. A greenhouse experiment was carried out to test this prediction: the dynamics of thrips and their predators was followed on plants damaged by spider mites, either with or without web. Thrips densities in presence of predators were higher on plants with web than on unwebbed plants after 3 weeks. Experimental data fitted model predictions, indicating that individual-level measurements of refuge costs and benefits can be extrapolated to the level of interacting populations. Model-derived calculations of thrips population growth rate enable the estimation of the minimum predator density at which thrips benefit from using the web as a refuge. The model also predicted a minor effect of the refuge on the prey density at equilibrium, indicating that the effect of refuges on population dynamics hinges on the temporal scale considered.     

Diel variation in predator abundance, predation risk and prey distribution in shallow-water estuarine habitats

Predation by visual predators is often affected by light conditions and may therefore exhibit strong diel variation. The dominant predators on grass shrimp, Palaemonetes pugio, are finfish predators that are thought to locate their prey by visual cues. We examined the response of grass shrimp to diel variation in predation risk in the nearshore shallow waters of the Chesapeake Bay. We used diel shoreline seines to assess the relative abundance of predators. We assessed the relative risk of predation with shrimp tethered at refuge (30 cm) and nonrefuge (60 cm) depths. To measure grass shrimp response to predation risk, we used dipnets to monitor habitat use. Four predominantly visual predators dominated the shoreline seine catches, Fundulus heteroclitus, Micropogonias undulatus, Morone americana and Morone saxatilis. Total predator abundance had a diel component, with dramatic nighttime decreases in total abundance, whereas guild composition and relative abundance remained unchanged. Relative predation risk for tethered shrimp exhibited significant time by habitat interaction. During the day, depth negatively affected survivorship of tethered shrimp while at night overall survivorship increased and there was no effect of depth. Shrimp habitats use reflected diel predation risks. Abundances in the near shore were highest during the day with decreased abundances at night. Together, the seine and tethering data highlight the importance for a refuge (e.g., shallow water) from predation during the daytime and a relaxation of predation pressure at night.