Predator size interacts with habitat structure to determine the allometric scaling of the functional response

While both predator body size and prey refuge provided by habitat structure have been established as major factors influencing the functional response (per capita consumption rate as a function of prey density), potential interactions between these factors have rarely been explored. Using a crab predator (Panopeus herbstii) – mussel prey (Brachidontes exustus) system, we examined the allometric scaling of the functional response in oyster (Crassostrea virginica) reef habitat, where crevices within oyster clusters provide mussels refuge from predation. A field survey of mussel distribution showed that mussels attach closer to the cluster periphery at high mussel density, indicating the potential for saturation of the refuge. In functional response experiments, the consumption rate of large crabs was depressed at low prey density relative to small crabs, while at high prey density the reverse was true. Specifically, the attack rate coefficient and handling time both decreased non‐linearly with crab size. An additional manipulation revealed that at low prey densities, the ability of large crabs to maneuver their claws and bodies to extract mussels from crevices was inhibited relative to small crabs by the structured habitat, reducing their attack rate. At high prey densities, crevices were saturated, forcing mussels to the edge of clusters where crabs were only limited by handling time. Our study illuminates a potentially general mechanism where the quality of the prey refuge provided by habitat structure is dependent on the relative size of the predator. Thus anthropogenic influences that alter the natural crab size distribution or degrade reef habitat structure could threaten the long‐term stability of the crab –mussel interaction in reefs.     

Processes regulating early post-settlement habitat use in a subtidal assemblage of brachyuran decapods

In highly mobile animals post-settlement dispersion of juveniles can strongly influence the observed patterns of abundance and distribution. To explore the relative importance of factors regulating the use of habitat by crabs we performed a multi-species manipulative experiment in a subtidal environment of the central Chilean coast. First, demographic patterns were established by performing a year-round crab survey in three discrete and well-known subtidal crab habitats: (1) algal turf, (2) cobbles and (3) shell hash. Second, habitat preferences were experimentally evaluated using concrete trays that were filled with different substrate types that simulate natural habitats. Settlement and recruitment rates were estimated from experimental trays that were left in the field and surveyed after 2 weeks (complete experiment was repeated 7 times throughout 1 year). Third, mortality, due to predation, was assessed by covering 50% of the trays with a 4-mm mesh-size screen that excluded large predators (i.e., fishes, shrimps). Fourth, habitat colonization rates were evaluated by quantifying the arrival, into open trays, of large juveniles (secondary dispersal). The most abundant species in this system (Paraxhantus barbiger, Cancer setosus, Taliepus dentatus and Pilumnoides perlatus) displayed clear habitat preferences at the time of settlement, evidenced by differences in density of recruits among habitats. Recruitment regulation by predation seemed to explain the observed patterns in only one case. For most species, however, evidence of ontogenetic change in the use of habitat, through active habitat redistribution by large juveniles, was detected. Thus, secondary dispersal among habitats seems to outweigh the influence of megalopae's habitat selection and post-settlement mortality as responsible for the observed demographic patterns.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Universal Optimization of Flight Initiation Distance and Habitat-Driven Variation in Escape Tactics in a Namibian Lizard Assemblage

Some aspects of escape predicted by theoretical models are intended to apply universally. For example, flight initiation distance (distance between an approaching predator and prey when escape begins) is predicted from predation risk and the costs of escaping. Escape tactics and refuge selection are not currently predicted by theoretical models, but are expected to vary with structural features of the habitat. One way of studying such variation is to compare aspects of antipredatory behavior among sympatric species that differ in habitat or microhabitat use. In an assemblage of lizards in northwestern Namibia, we conducted experiments to test predictions of escape theory for three risk factors in representatives of three families and observed escape tactics in additional species. As predicted by escape theory, flight initiation distance increased with directness of a predator's approach and predator speed in Agama planiceps, Mabuya acutilabris, and Rhotropus boultoni, and with distance from refuge in M. acutilabris. As predicted by theory, the probability of entering refuge increased with risk in R. boultoni. All available data indicate that flight initiation distance and refuge entry by lizards conform to theoretical predictions. Escape tactics varied greatly as a function of habitat type: (1) arboreal species fled up and around trees and sometimes entered tree holes; (2) saxicolous species used rock crevices as refuges, but differed in tactics prior to entering refuges; and (3) terrestrial species fled into bushes or other vegetation, often to the far sides of them. Some M. acutilabris entered small animal burrows or buried themselves in sand beneath bushes. Escape tactics varied even among congeners in Mabuya, highlighting the important effect of habitat structure on them. Although habitat partitioning has traditionally been viewed as favoring species coexistence, an interesting by‐product appears to be structuring of escape tactics in lizard communities.     

Habitat context influences predator interference interactions and the strength of resource partitioning

Despite increasing evidence that habitat structure can shape predator–prey interactions, few studies have examined the impact of habitat context on interactions among multiple predators and the consequences for combined foraging rates. We investigated the individual and combined effects of stone crabs (Menippe mercenaria) and knobbed whelks (Busycon carica) when foraging on two common bivalves, the hard clam (Mercenaria mercenaria) and the ribbed mussel (Geukensia demissa) in oyster reef and sand flat habitats. Because these species co-occur across these and other estuarine habitats of varying physical complexity, this system is ideal for examining how habitat context influences foraging rates and the generality of predator interactions. Consistent with results from previous studies, consumption rates of each predator in isolation from the other were higher in the sand flat than in the more structurally complex oyster reef habitat. However, consumption by the two predators when combined surprisingly did not differ between the two habitats. This counterintuitive result probably stems from the influence of habitat structure on predator–predator interactions. In the sand-flat habitat, whelks significantly reduced their consumption of their less preferred prey when crabs were present. However, the structurally more complex oyster reef habitat appeared to reduce interference interactions among predators, such that consumption rates when the predators co-occurred did not differ from predation rates when alone. In addition, both habitat context and predator–predator interactions increased resource partitioning by strengthening predator dietary selectivity. Thus, an understanding of how habitat characteristics such as physical complexity influence interactions among predators may be critical to predicting the effects of modifying predator populations on their shared prey.     

Recruitment of shore crabsCarcinus maenas on tidal flats: Mussel clumps as an important refuge for juveniles

During the late summer and early fall, juvenile shore crabs (Carcinus maenas L.) occurred in high abundances in mussel clumps scattered on tidal flats of the Wadden Sea. Abundances were much lower on bare tidal flats without mussel clumps and decreased substantially from July to November, whereas numbers in mussel clumps remained high. Large crabs left the tidal flats in early fall, whereas juveniles undertook tidal migrations only in the late fall. In March very few shore crabs were found in the intertidal area. The size of juvenile shore crabs living between mussels did not increase significantly during fall. On the bare tidal flats surrounding the mussels, a size increase was observed. Mussel beds and mussel clumps serve as a spatial refuge for the early benthic phases of juvenile shore crabs. Between mussels they can hide effectively from their epibenthic predators. Juvenile shore crabs do not leave the intertidal area and the mussel habitats before their major predators have left the area. Mussel clumps scattered over the tidal flats may be a critical refuge for juvenile shore crabs settling on tidal flats. Intensified efforts in mussel culturing in the European Wadden Sea during recent decades may have caused an increased abundance of mussel clumps on tidal flats, thus enhancing habitat availability for some mussel-clump inhabitants.     

The relative importance of habitat-specific settlement, predation and juvenile dispersal for distribution and abundance of young juvenile shore crabs Carcinus maenas L.

Young juveniles of many motile benthic species are concentrated in structurally complex habitats, but the proximate causes of this distribution are usually not clear. In the present study, I assessed three potentially important processes affecting distribution and abundance of early benthic stages in the shore crab (Carcinus maenas): (1) selection of habitat by megalopae (postlarvae); (2) habitat-specific predation; and (3) post-settlement movements by juveniles. These processes were assessed concurrently over 3-9 days at two spatial scales: at the scale of square meters using cage techniques within nursery areas, and at the scale of hectares using isolated populations of juvenile shore crabs in small nursery areas as mesocosms. The results were compared to habitat-specific distribution in the field.Shore crab megalopae and first instar juveniles (settlers) were distributed non-randomly among micro-habitats in the assessed nursery areas, with great densities in both mussel beds, eelgrass and filamentous algal patches (on average 114-232 settlers m⁻²), and significantly smaller densities on open sand habitats at all times (on average 4 settlers m⁻²). The same habitat-specific settlement pattern was found in cages where predators were excluded, suggesting that active habitat selection at settlement was responsible for the initial distribution. Older juveniles (second to ninth instar crabs) were also sparse on sand, but in contrast to settlers, were concentrated in mussel beds, which showed significantly greater densities than eelgrass and algal habitats. The cage experiment demonstrated a dynamic distribution of juvenile crabs. Young juveniles constantly migrated over open sand habitats (20 m or further) and colonized the experimental plots in a habitat-specific pattern that reflected the distribution in the field. This pattern was also found for very small crabs colonizing predator-exclusion cages, suggesting that selection of habitat by migrating juveniles caused the ontogenetic change in habitat use. Although post-settlement movements were great within nursery areas, juvenile dispersal at a regional scale appeared to be small, and the recruitment of juvenile shore crabs to the shallow bays occurred mainly through pelagic megalopae.Conservative estimates at the scale of whole nursery areas, based on migration trap data and field samples, indicated great mortality of settlers and early benthic stages of shore crabs. Results from the cage experiment suggest that predation by crabs and shrimp were responsible for the high settlement mortality. Both enclosed cannibalistic juvenile crabs and local predators on uncaged habitat plots caused significant losses of settlers in all habitats (on average 22% and 64% 3 day⁻¹, respectively). The effect of predators was highly variable between trials, but differed little between habitat types, and predation had no detectable proximate effect on juvenile distribution, despite the great losses. Small settlement densities on sand habitats in combination with a refuge at low prey numbers, and an aggregation of cannibalistic juvenile crabs in nursery habitats appear to decrease the effect of habitat-specific predation rates on the distribution of juvenile shore crabs. This study demonstrates that active habitat selection at settlement followed by a dynamic redistribution of young juveniles can be the proximate processes responsible for habitat-specific distribution of epibenthic juveniles, and indicate that predation represents a major evolutionary process reinforcing this behavior.     

Guppy populations differ in cannibalistic degree and adaptation to structural environments

There is considerable variation in cannibalism between different species and also between individuals of different species, although relatively little is known about what creates this variation. We investigated the degree of cannibalism in guppy (Poecilia reticulata) populations originating from high and low predation environments in Trinidad, and also how cannibalism was affected by the presence of refuges. Females from two populations were allowed to feed on juveniles from two populations in aquaria trials. The cannibalism was size-dependent and varied depending on both juvenile and female origin. Low predation females were more efficient cannibals and low predation juveniles were better at avoiding cannibalism compared to high predation guppies when no refuges were present. The high predation females were superior cannibals and the high predation juveniles were better at escaping cannibalism than the low predation guppies when refuges were present. We discuss whether the differences in cannibalism and response to refuge addition relate to predation-induced habitat shifts and differences in the guppies’ natural environment.     

Can protective attributes of artificial refuges offset predation risk in lizards?

Artificial refuges are often used to supplement habitat in areas where natural shelters have been degraded or removed. Although artificial refuges are intended to support particular species, they may be equally attractive and accessible to others, including predators. We explored the influence of snake predation risk and shelter attributes on the overnight use of different artificial refuges (timber, tiles, and iron) using the predator‐prey relationship between Boulenger's skink, Morethia boulengeri and the curl snake, Suta suta. We collected adult M. boulengeri from two bioregions in south‐eastern Australia: the Riverina, where the two species co‐occur, and the South Western Slopes, where S. suta does not occur. Two adult S. suta were collected for use as chemical donors. We conducted four experiments on overnight refuge choice to determine: (i) predator‐scent avoidance, (ii) artificial refuge preferences, (iii) a trade‐off between a preferred refuge and predator‐avoidance, and (iv) the effect of gap height on refuge preference. We found that skinks avoided predator‐scented refuges in favour of identical, but unscented refuges. Skinks preferred timber refuges, and most skinks maintained this preference when predator‐scent was added. However, when gap height was manipulated, skinks shifted to the refuge with the smallest gap. Skinks displayed complex regional variation in behaviour; skinks from both bioregions avoided predator‐scent, but in the trade‐off experiment, skinks from the South Western Slopes were less likely to avoid predator‐scented timber refuges than those from the Riverina. Our findings suggest that protective refuge attributes, such as small gap height, can offset the risk implied by predator‐scent within a refuge. This study highlights the need to consider predator‐prey interactions when designing and using artificial refuges for habitat restoration or biological monitoring.     

The role of geologic grazing refuges in structuring Mediterranean grassland plant communities

We tested the hypothesis that small rock-enclosed geologic refuges have an important role in maintaining grazing-sensitive species in grassland with a long history of intense grazing. The study was carried out in Mediterranean grassland in a basalt landscape in northern Israel. Community composition was compared in 63 sites between samples of 1 m² quadrats in two microhabitats: (1) rock-enclosed, presumed “refuges” and (2) rock-adjacent, “near refuges” but just outside them, accessible to cattle grazing. Median refuge area was 2.75 m², median rock height and diameter around refuges were 1.10 m and 1.50 m. Median height of residual dry herbage was 1.41 m in refuges, compared to 0.38 m outside, indicating the difference in grazing intensity. Species richness at three scales (quadrat, site, all sites) was significantly greater in the near-refuge than in the refuge habitat. In the latter, many annuals were excluded by dominance of tall perennials. Twelve species (of 103) had significantly higher cover in refuges, including tall perennial grasses, tall annuals, climbers, and a shrub. A total of 53 species with a strong significant negative response to refuges were mostly small and medium height annuals. The intermediate group of 38 species with weak or non-significant responses to refuges included, among others, dominant tall grasses that were abundant both in refuges and just outside them. The latter, as well as most refuge-positive species had shown a positive response to protection in exclosures. The results support the hypothesis that small rock-enclosed habitats—more so than artificial exclosures—are effective grazing refuges for rare, grazing-susceptible species. The contribution of refuges to species richness at the landscape scale is much greater than their proportion of the area. Dispersion from refuges maintains small populations of rare species near refuges and can initiate expansion into the landscape when grazing pressure is lowered.     

Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

Lack of strong refuges allows top-down control of soybean aphid by generalist natural enemies

Refuges have been shown to be important mediators of predator–prey interactions, and in particular, have been proposed as a potential mechanism allowing herbivore populations to reach outbreak levels. However, very little research on the role of refuges has been conducted in systems dominated by generalist predators. We investigated the existence of refuges from predation for the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae) at multiple scales. This species invaded North America and in spite of previous studies demonstrating strong suppression by generalist natural enemies, its populations periodically cause significant economic losses. Using naturally occurring populations of soybean aphid and its natural enemies, we tested for the presence of A. glycines spatial and dynamic refuges at the within-field, single plant, and within-plant scale. At the within-field level, we found only weak and transient spatial patterns in aphid populations suggesting the lack of spatial refuges at this scale. Similarly, at the plant level we found no individual colonies that escaped predation and aphid suppression was 9- to 28-fold greater in comparison with caged controls regardless of initial aphid density. When high aphid populations were exposed to predation they were rapidly reduced to levels close to the average field density and showed reduced per capita growth rates, indicating an absence of dilution of predation risk at increased aphid density. Finally, we found a significant shift in the distribution of aphids to the lower portions of the plant in the presence of generalist predators, suggesting a partial refuge from predation at the within-plant scale. Overall, we found the naturally occurring community of generalist predators to exert strong top-down suppression of soybean aphid populations at multiple scales, and no evidence that the presence of refuges at the scales studied can lead to outbreak populations. The partial refuge from predation at the within-plant scale revealed in our study may have important consequences for the within-season population dynamics of A. glycines, since it may be associated with low plant quality tradeoffs, and therefore warrants further research.     

Refuge function of marine algae complicates selection in an intertidal snail

Species with restricted gene flow often show trait-shifts from one type of environment to another. In those rock-dwelling marine gastropods that lack larval dispersal, size generally decreases in wave-exposed habitats reducing risk of dislodgement, while increases in less exposed habitats to resist crab-crushing. In Littorina fabalis, however, snails of moderately exposed shores are generally much larger (11–14 mm) than snails of sheltered shores (5–8 mm). Observations from the White Sea (where crabs are not present) indicate that in the absence of crabs snails are small (6–7 mm) in both habitats. We assumed that the optimal size for L. fabalis in the absence of crabs is less than 8 mm, and thus that increased size in moderately exposed habitats in areas with crabs might be a response to crab predation. In a crab-rich area (Sweden) we showed that crab predation is an important mortality factor for this snail species in both sheltered and moderately exposed habitats. In sheltered habitats, snails were relatively more protected from crab-predation when dwelling on their habitual substrate, fucoid algae, than if experimentally tethered to rocks below the algae. This showed that algae function as snail refuges. Snail dislodgement increased, however, with wave exposure but tethering snails in moderately exposed habitats showed that large snails survived equally well on rocks under the algae as in the canopy of the algae. Thus in sheltered habitats a small snail size is favored, probably due to life-history reasons, while increased risk of being dislodged from the algae refuges promotes a large size in moderately exposed habitats. This study shows an example of selection of a trait depends on complex interactions of different factors (life-history optimization, crab predation, wave induced dislodgement and algal refuges).     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Avoiding the flow: refuges expand the swimming potential of coral reef fishes

While many coral reef fishes utilise substratum refuges, the direct influence of water flow and swimming ability on such refuging patterns is yet to be established. This study examined the swimming ability and refuging behaviour of a labrid (Halichoeres margaritaceus) and a pomacentrid (Pomacentrus chrysurus) that inhabit high flow, wave-swept coral reef flats. Field observations of refuging patterns were combined with experimental evaluations in a flow tank using a replica of a substratum hole frequently used by these species. Under a range of flow speeds commonly found on the reef flat (0–60 cm s⁻¹), flow within the substratum refuge was reduced to speeds of 0–12 cm s⁻¹, representing a 75–100% flow reduction. Swimming ability of each species was then tested at 60 cm s⁻¹ with and without access to this flow refuge. Both species were able to maintain activity within the 60 cm s⁻¹ flow for considerably longer when provided with a refuge, with increases from approximately 39 min to 36 h for H. margaritaceus and 8 min to 88 h for P. chrysurus. Despite H. margaritaceus having the strongest swimming ability without access to a refuge, P. chrysurus was able to maintain swimming activity more than twice as long as H. margaritaceus when provided with a refuge. These increases in activity are probably due to energetic savings, with this type of refuge providing an estimated 95% energy saving over swimming directly into a unidirectional flow of 60 cm s⁻¹. These results highlight the major advantages provided by refuging behaviour and emphasise the importance of habitat refuges in shaping patterns of habitat use in reef fishes.     

Selection of diurnal refuges by the nocturnal squirrelfish, <Emphasis Type="Italic">Holocentrus rufus</Emphasis>

We examined the diurnal refuges occupied by the nocturnal squirrelfish, Holocentrus rufus, to describe refuges and the behavior associated with their use and to determine which, if any, refuge characteristics were selected. We tagged 21 H. rufus on two sites on a fringing reef in Barbados, West Indies, identified the refuges they used (n = 57), measured ten characteristics of each refuge and the surrounding microhabitat, and monitored their refuge use for 4 weeks. To evaluate refuge selection, we measured the same characteristics on a comparable number of unused potential refuges (n = 67) on the same reefs and used classification tree models to determine which characteristics separated used from unused refuges. Each fish used 1–9 refuges, which did not overlap among individuals and were defended against intrusion by conspecifics and some heterospecifics. Fish with more than one refuge frequently moved among them. There was strong site fidelity with no immigration of untagged fish or emigration of tagged fish on either reef during the study period and no additional refuges being occupied over the 4-week period. Refuges were primarily holes, open at one or two ends, which varied in size, distance from the reef edge, entrance orientation, and vertical relief at the entrance. Holes used as refuges differed significantly from unused holes mainly in characteristics related to the vertical position of their entrance, but the classification tree models differed for the two sites. This study provides the first detailed information on characteristics of daytime refuges used by a nocturnally active reef fish and the first evidence of selectivity of refuges. It suggests that the abundance and characteristics of holes on reefs could influence the density of H. rufus on natural reefs.     

The shifting balance of littoral predator-prey interaction in regimes of hydrodynamic stress

Above lowshore levels of wave-beaten rocky shores, desiccation from tidal exposure and hydrodynamics stresses from wave action are thought to create refuges from predation, allowing concentrations of sedentary prey such as mussel beds. Underwater time-lapse photography on rocky shores in Southern California revealed that dense aggregations of spiny lobsters prey on mussels during nocturnal high tides. In contradiction of the refuge hypothesis, the densest aggregations occurred on midshore levels of the most wave-exposed site, a semi-protected site showed intermediate densities, and a protected site showed only sparse numbers of lobsters. On wave-beaten shores, the lobsters' high mobility and rapid prey handling allowed them to exploit intertidal prey in the brief period at extreme high tide, when both desiccation and hydrodynamic stresses were at a minimum. The spatial differences in lobster densities were, however, positively related to the recruitment rates of juvenile mussels, the preferred prey. A field experiment demonstrated that predation by lobsters within a mussel bed affects the age/size structure of the bed without changing primary percent coverage. Therefore, concentrations of adult prey on some wave-swept sites appear to result from elevated rates of prey recruitment that surpass rates of predation, rather than absolute refuges from predation.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Experimental manipulation reveals the importance of refuge habitat temperature selected by lizards

Refuges provide shelter from predators, and protection from exposure to the elements, as well as other fitness benefits to animals that use them. In ectotherms, thermal benefits may be a critical aspect of refuges. We investigated microhabitat characteristics of refuges selected by a heliothermic scincid lizard, Carlia rubrigularis, which uses rainforest edges as habitat. We approached lizards in the field, simulating a predator attack, and quantified the refuge type used, and effect of environmental temperatures (air temperature, substrate temperature and refuge substrate temperature) on the amount of time skinks remained in refuges after hiding (emergence time). In respone to our approach, lizards were most likely to flee into leaf litter, rather than into rocks or woody debris, and emergence time was dependent on refuge substrate temperature, and on refuge substrate temperature relative to substrate temperature outside the refuge. Lizards remained for longer periods in warmer refuges, and in refuges that were similar in temperature to outside. We examined lizard refuge choice in response to temperature and substrate type in large, semi‐natural outdoor enclosures. We experimentally manipulated refuge habitat temperature available to lizards, and offered them equal areas of leaf litter, woody debris and rocks. When refuge habitat temperature was unmanipulated, lizards (85%) preferred leaf litter, as they did in the field. However, when we experimentally manipulated the temperature of the leaf litter by shading, most skinks (75%) changed their preferred refuge habitat from leaf litter to woody debris or rocks. These results suggest that temperature is a critical determinant of refuge habitat choice for these diurnal ectotherms, both when fleeing from predators and when selecting daytime retreats.