Prey Patches Research Articles

Basking shark breaching behaviour observations west of Shetland

This study reports observations of basking sharks (Cetorhinus maximus) sighted during an offshore geophysical survey conducted in July and August 2013, west of Shetland, UK. During the 38-day survey, trained and dedicated marine wildlife observers recorded 19 sightings of basking sharks (n = 22 individuals). Of these observations, 17 were of single sharks, with one observation of two sharks and one observation of three sharks. All surface sightings occurred in water depths between 129 and 199 m, predominantly prior to noon local time (79%), and were mostly of sharks 6–8 m in length, although a young (2 m) individual was also recorded. Breaching behaviour was observed on 21 occasions, by individuals or in small groups. Breaching has been proposed as a male-male competitive behaviour during courtship displays and female basking sharks may breach to signal their readiness for mating. Aggregations of basking sharks at frontal systems are well documented and linked to the occurrence of prey patches; however, these oceanographic features may also be of importance to courtship. The high number of sightings of sharks recorded during a relatively short time frame in addition to breaching behaviour and presence of young individuals, suggest that this area west of Shetland may be an important habitat for the basking shark.

Constrained by consistency? Repeatability of foraging behavior at multiple timescales for a generalist marine predator

Marine predators frequently exhibit consistency in foraging behaviors despite the dynamic nature of marine ecosystems, which has the potential for ecological and evolutionary implications depending on the timescale at which it persists. We examined behavioral consistency in movements and diving behavior of adult female California sea lions (Zalophus californianus), which are abundant, generalist central-place foragers inhabiting an ecosystem characterized by small- and broad-scale oceanographic variability. We used biologging devices to measure repeatability of behavior within a season and stable isotope analysis of whiskers to quantify behavior across a 2-year period associated with anomalous environmental conditions that affected prey availability. Sea lions were significantly repeatable in all variables across multiple timescales (Radj = 0.26–0.82), although repeatability estimates were generally higher for variables related to characteristics of individual dives (e.g., dive depth) than those that described dive bouts (e.g., bout duration) or spatial use (e.g., volume of 3D utilization distribution). These differences may result from the fact that diving behaviors vary with prey type, whereas spatial use and bout variables may reflect the foraging success within prey patches or movement among patches. There was variation in how predictable individual sea lions were in their diving behaviors, which was largely unrelated or negatively related to foraging site fidelity. The strength of behavioral consistency decreased with time yet persisted across the 2-year period, suggesting that while sea lions alter their behavior in response to environmental change, the behavioral flexibility of individuals may ultimately be constrained by consistency.

Lévy flight movements prevent extinctions and maximize population abundances in fragile Lotka–Volterra systems

Multiple-scale mobility is ubiquitous in nature and has become instrumental for understanding and modeling animal foraging behavior. However, the impact of individual movements on the long-term stability of populations remains largely unexplored. We analyze deterministic and stochastic Lotka-Volterra systems, where mobile predators consume scarce resources (prey) confined in patches. In fragile systems (that is, those unfavorable to species coexistence), the predator species has a maximized abundance and is resilient to degraded prey conditions when individual mobility is multiple scaled. Within the Lévy flight model, highly superdiffusive foragers rarely encounter prey patches and go extinct, whereas normally diffusing foragers tend to proliferate within patches, causing extinctions by overexploitation. Lévy flights of intermediate index allow a sustainable balance between patch exploitation and regeneration over wide ranges of demographic rates. Our analytical and simulated results can explain field observations and suggest that scale-free random movements are an important mechanism by which entire populations adapt to scarcity in fragmented ecosystems.

Volume-concentrated searching by an aerial insectivore, the common swift, Apus apus

How predators search for prey is a cornerstone question in behavioural ecology, which has yet to be investigated for animals foraging in 3D airspace. Do insectivorous birds such as swifts (Apodidae), swallows and martins (Hirundinidae) use similar strategies to those performed by terrestrial predators in 2D, or do they rely on different spatial search strategies because of some properties of the aerial open space? We addressed this question in the common swift, one of the most aerial birds, using a novel 3D optical tracking method. The analysis of fine-scale flight tracks revealed how birds distribute their presence in 3D space while foraging near their breeding colony. Common swifts concentrated the time spent per volume unit by adopting a tortuous path, and, to a much lesser extent, by decreasing their movement speed. By independently observing the birds' posture on tracking images, we were able to identify the occurrence of putative prey captures along flight tracks. We show that swifts' presence was concentrated mainly in the vicinity of prey captures, unveiling a volume-concentrated search (VCS) strategy in this aerial insectivore. This is an extension in 3D of the area-concentrated search classically described in terrestrial 2D space. VCS can (but does not necessarily) take place in thermal updrafts, where small insects can be concentrated in patches. In contrast to terrestrial and aquatic predators that can easily slow down or stop their movement in profitable places, a different speed–cost relationship underlying aerial movement prevents swifts from stopping in prey patches and explains why these birds rely mainly on movement tortuosity to perform intensive search. Our study thus shows how some physical properties of the environment can modulate the way an animal concentrates its search in profitable places.

Respiratory behaviors in sympatric rorqual whales: the influence of prey depth and implications for temporal access to prey

Energetically costly lunge feeding at depth causes the respiratory patterns and feeding performance of rorqual whales (Family Balaenopteridae) to hinge in part upon prey patch depth. This contingency has the potential to precipitate differences in prey preference and habitat suitability for sympatric species and may be a factor in competitive interactions, but comparative respiration studies are a necessary first step in assessing this hypothesis. We concurrently sampled dive behavior in sympatric, euphausivorous humpback (Megaptera novaeangliae) and fin whales (Balaenoptera physalus), as well as prey depth distribution within a British Columbia fjord system over the course of 2 summers. Ventilation and dive patterns differed significantly between species, including differential respiratory response to increasing prey depth, despite their foraging upon a common prey resource. Thanks to longer dives and shorter surface recoveries, fin whales spent a greater proportion of their time on dives. This behavior, coupled with faster swim speeds during descent and ascent reported in previous studies, afford fin whales greater periods of time at the depth of their prey. These interspecific discrepancies in dive behavior determine the whales' relative temporal access to prey. Simulations based on our observations indicate that the fin whale's relative advantage in this fjord system increases with increasing prey depth when all other prey parameters are held constant. Simulation results emphasize the importance of swim speed in rorqual foraging strategy. Small differences in prey access per dive can have important implications over the course of a foraging season, which may precipitate differences in habitat suitability. Our findings, when coupled with the body of knowledge from tagging studies, highlight this link and point to its potential role in the habitat preferences of foraging whales.

Life cycle of Harmonia axyridis in central Europe

The development and reproduction, and maximum number of generations across seasons were ascertained for populations of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Central Europe. Number of generations was calculated using the thermal characteristics of development and summation of daily average air temperatures, in 2007–2016. This species can produce 2–3 generations per year. However, there is insufficient time for it to achieve its full reproductive potential. Delayed emigration from hibernacula (late April) shortened the period of seasonal activity by 1–6%. Breeding occurred from May to mid September. Oviposition continued into late summer, which left insufficient time for the late-hatching immatures to complete their development before the onset of winter. A high mobility, opportunistic use of prey patches and weak tendency to enter dormancy provide the potential for population increase by extending the period of development of immatures at the risk of failure if the cold period comes early.

Costs and benefits of straight versus tortuous migration paths for Prairie Rattlesnakes (Crotalus viridis viridis) in seminatural and human-dominated landscapes

An individual’s migration path shape should affect its fitness, because patchily distributed features (e.g., prey) are encountered more often on straight than tortuous paths. We hypothesized that Prairie Rattlesnakes (Crotalus viridis viridis (Rafinesque, 1818)) with straighter migration paths should have better body condition, because they encounter prey patches more frequently, and higher migration mortality, because they also encounter predators and hazardous human land uses more frequently, than individuals with tortuous paths. If true, then a straighter path should be favoured when the benefit (resource acquisition) outweighs the cost (mortality risk). Humans pose a significant mortality risk for migrants; thus, the cost of straight-line movement should increase relative to the benefit in more human-dominated landscapes, favouring more tortuous movements. We tested these hypotheses using data on the body condition, mortality, and migration movements of 25 female Prairie Rattlesnakes in one human-dominated and one seminatural landscape. As hypothesized, we found better body condition and higher migration mortality for snakes with straighter migration paths, and that snakes followed more tortuous paths in the human-dominated landscape. Although selection for tortuous movements may reduce rates of migration mortality in human-dominated landscapes, this may ultimately contribute to population declines if poorer body condition reduces overwinter survival or reproductive success.

Taking movement data to new depths: Inferring prey availability and patch profitability from seabird foraging behavior.

Detailed information acquired using tracking technology has the potential to provide accurate pictures of the types of movements and behaviors performed by animals. To date, such data have not been widely exploited to provide inferred information about the foraging habitat. We collected data using multiple sensors (GPS, time depth recorders, and accelerometers) from two species of diving seabirds, razorbills (Alca torda, N = 5, from Fair Isle, UK) and common guillemots (Uria aalge, N = 2 from Fair Isle and N = 2 from Colonsay, UK). We used a clustering algorithm to identify pursuit and catching events and the time spent pursuing and catching underwater, which we then used as indicators for inferring prey encounters throughout the water column and responses to changes in prey availability of the areas visited at two levels: individual dives and groups of dives. For each individual dive (N = 661 for guillemots, 6214 for razorbills), we modeled the number of pursuit and catching events, in relation to dive depth, duration, and type of dive performed (benthic vs. pelagic). For groups of dives (N = 58 for guillemots, 156 for razorbills), we modeled the total time spent pursuing and catching in relation to time spent underwater. Razorbills performed only pelagic dives, most likely exploiting prey available at shallow depths as indicated by the vertical distribution of pursuit and catching events. In contrast, guillemots were more flexible in their behavior, switching between benthic and pelagic dives. Capture attempt rates indicated that they were exploiting deep prey aggregations. The study highlights how novel analysis of movement data can give new insights into how animals exploit food patches, offering a unique opportunity to comprehend the behavioral ecology behind different movement patterns and understand how animals might respond to changes in prey distributions.

How does hunger affect convergence on prey patches in a social forager?

AbstractInternal state, in this case hunger, is known to influence both the organisation of animal groups and the social foraging interactions that occur within them. In this study, we investigated the effects of hunger upon the time taken to locate and converge upon hidden simulated prey patches in a socially foraging fish, the threespine stickleback (Gasterosteus aculeatus). We predicted that groups of food‐deprived fish would find and recruit to prey patches faster than recently fed groups, reasoning that they might search more rapidly and be more attentive to inadvertent social information produced by other foragers. Instead we saw no difference between the two groups in the time taken to find the patches and found that in fact, once prey patches had been discovered, it was the recently fed fish that converged on them most rapidly. This finding is likely due to the fact that recently fed fish tend to organise themselves into fewer but larger subgroups, which arrived at the food patch together. Hunger has a significant impact upon the social organisation of the fish shoals, and it appears that this has a stronger effect upon the rate at which they converged upon the food patches than does internal state itself.

Seabird distribution patterns observed with fishing vessel\u2019s radar reveal previously undescribed sub-meso-scale clusters

Seabirds are known to concentrate on prey patches or at predators aggregations standing for potential feeding opportunities. They may search for prey using olfaction or by detecting visually feeding con-specifics and sub-surface predators, or even boats. Thus, they might form a foraging network. We hypothesized that conditionally to the existence of a foraging network, the visual detection ability of seabirds should have a bearing on their medium-scale distribution at sea. Using a fishing-boat radar to catch the instantaneous distribution of seabirds groups within 30 km around the vessel, we conducted a spatial clustering of the seabird-echoes. We found 7,657 clusters (i.e. aggregations of echoes), lasting less than 15 minutes and measuring 9.2 km in maximum length (median). Distances between seabirds groups within clusters showed little variation (median: 2.1 km; CV: 0.5), while area varied largely (median: 21.9 km2; CV: 0.8). Given existing data on seabirds’ reaction distances to boats or other marine predators, we suggest that these structures may represent active foraging sequences of seabirds spreading themselves in space such as to possibly cue on each others. These seabird clusters were not previously described and are size compatible with the existence of a foraging network.

Boldness predicts an individual's position along an exploration-exploitation foraging trade-off.

Individuals do not have complete information about the environment and therefore they face a trade‐off between gathering information (exploration) and gathering resources (exploitation). Studies have shown individual differences in components of this trade‐off but how stable these strategies are in a population and the intrinsic drivers of these differences is not well understood.Top marine predators are expected to experience a particularly strong trade‐off as many species have large foraging ranges and their prey often have a patchy distribution. This environment leads these species to exhibit pronounced exploration and exploitation phases but differences between individuals are poorly resolved. Personality differences are known to be important in foraging behaviour but also in the trade‐off between exploration and exploitation. Here we test whether personality predicts an individual exploration–exploitation strategy using wide ranging wandering albatrosses (Diomedea exulans) as a model system.Using GPS tracking data from 276 wandering albatrosses, we extract foraging parameters indicative of exploration (searching) and exploitation (foraging) and show that foraging effort, time in patch and size of patch are strongly correlated, demonstrating these are indicative of an exploration–exploitation (EE) strategy. Furthermore, we show these are consistent within individuals and appear stable in the population, with no reproductive advantage.The searching and foraging behaviour of bolder birds placed them towards the exploration end of the trade‐off, whereas shy birds showed greater exploitation. This result provides a mechanism through which individual foraging strategies may emerge. Age and sex affected components of the trade‐off, but not the trade‐off itself, suggesting these factors may drive behavioural compensation to maintain resource acquisition and this was supported by the evidence that there were no fitness consequence of any EE trait nor the trade‐off itself.These results demonstrate a clear trade‐off between information gathering and exploitation of prey patches, and reveals for the first time that boldness may drive these differences. This provides a mechanism through which widely reported links between personality and foraging may emerge.

Dynamic oceanography determines fine scale foraging behavior of Masked Boobies in the Gulf of Mexico.

During breeding, foraging marine birds are under biological, geographic, and temporal constraints. These contraints require foraging birds to efficiently process environmental cues derived from physical habitat features that occur at nested spatial scales. Mesoscale oceanography in particular may change rapidly within and between breeding seasons, and findings from well-studied systems that relate oceanography to seabird foraging may transfer poorly to regions with substantially different oceanographic conditions. Our objective was to examine foraging behavior of a pan-tropical seabird, the Masked Booby (Sula dactylatra), in the understudied Caribbean province, a moderately productive region driven by highly dynamic currents and fronts. We tracked 135 individuals with GPS units during May 2013, November 2013, and December 2014 at a regionally important breeding colony in the southern Gulf of Mexico. We measured foraging behavior using characteristics of foraging trips and used area restricted search as a proxy for foraging events. Among individual attributes, nest stage contributed to differences in foraging behavior whereas sex did not. Birds searched for prey at nested hierarchical scales ranging from 200 m—35 km. Large-scale coastal and shelf-slope fronts shifted position between sampling periods and overlapped geographically with overall foraging locations. At small scales (at the prey patch level), the specific relationship between environmental variables and foraging behavior was highly variable among individuals but general patterns emerged. Sea surface height anomaly and velocity of water were the strongest predictors of area restricted search behavior in random forest models, a finding that is consistent with the characterization of the Gulf of Mexico as an energetic system strongly influenced by currents and eddies. Our data may be combined with tracking efforts in the Caribbean province and across tropical regions to advance understanding of seabird sensing of the environment and serve as a baseline for anthropogenic based threats such as development, pollution, and commercial fisheries.

Within‐patch oviposition site shifts by spider mites in response to prior predation risks decrease predator patch exploitation

AbstractIn egg‐laying animals with no post‐oviposition parental care, between‐ or within‐patch oviposition site selection can determine offspring survival. However, despite the accumulation of evidence supporting the substantial impact predators have on oviposition site selection, few studies have examined whether oviposition site shift within patches (“micro‐oviposition shift”) reduces predation risk to offspring. The benefits of prey micro‐oviposition shift are underestimated in environments where predators cannot disperse from prey patches. In this study, we examined micro‐oviposition shift by the herbivorous mite Tetranychus kanzawai in response to the predatory mite, Neoseiulus womersleyi, by testing its effects on predator patch exploitation in situations where predatory mites were free to disperse from prey patches. Adult T. kanzawai females construct three‐dimensional webs on leaf surfaces and usually lay eggs under the webs; however, females that have experienced predation risks, shift oviposition sites onto the webs even in the absence of current predation risks. We compared the predation of eggs on webs deposited by predator‐experienced females with those on leaf surfaces. Predatory mites left prey patches with more eggs unpredated when higher proportions of prey eggs were located on webs, and egg survival on webs was much higher than that on leaf surfaces. These results indicate that a micro‐oviposition shift by predator‐experienced T. kanzawai protects offspring from predation, suggesting adaptive learning and subsociality in this species. Conversely, fecundity and longevity of predator‐experienced T. kanzawai females were not reduced compared to those of predator‐naïve females; we could not detect any costs associated with the learned micro‐oviposition shift. Moreover, the previously experienced predation risks did not promote between‐patch dispersal of T. kanzawai females against subsequently encountered predators. Based on these results, the relationships of between‐patch oviposition site selection and micro‐oviposition shift are discussed.

Costs and benefits of interlarval cannibalism in Harmonia axyridis

AbstractCannibalism, which is rather common in ladybirds, has been usually studied at the individual level: benefits of cannibalism for cannibals were estimated. Our study was conducted at the group level: we evaluated the overall effect of interlarval cannibalism on a group of Harmonia axyridis larvae of the fourth instar deprived of food, including both cannibals and their victims. Experiments showed that the probability of pupation in larvae which were kept individually was significantly higher than in larvae kept in groups of five, other conditions being the same. The proportion of samples in which at least one of five individuals pupated among the larvae kept individually was also higher than among those kept in groups suggesting that the eventual benefit of cannibalism was outweighed by the negative impact of aggressive interlarval interactions. The mean and minimum survival time in samples where none of five larvae pupated were longer when larvae were kept individually than when larvae were kept in groups. However, the maximum survival time (the survival time of the last larva in a sample) increased when larvae were kept in groups, which was the only one benefit of cannibalism found in our study. Under natural conditions, the possible adaptive value of this effect is that in the absence of natural prey, longer larval survival time proportionally increases the possibility of finding a new prey patch thereby ensuring survival of the population.

Finding patches in a heterogeneous aquatic environment: pH‐taxis by the dispersal stage of choanoflagellates

AbstractMicrobial eukaryotes that feed on bacteria are critical links in aquatic food webs. We used the choanoflagellate Salpingoeca rosetta, whose life history includes fast‐ and slow‐swimming unicellular forms and multicellular colonies, to study performance of these different forms at finding and accumulating in patches of prey. Video microscopy of behavior in microfluidic experiments showed that only unicellular fast swimmers exhibited taxis into patches of attractant. Of the chemical cues tested, only pH affected behavior: fast swimmers moved toward patches of water that had pH in the range of 6–7 from water with higher or lower pH. Since bacteria can lower the pH of the water in their vicinity, we suggest that seeking regions with a moderately more acidic pH than average seawater helps the fast‐swimming dispersal stage of choanoflagellates find particles or surfaces rich in prey.

How Elephant Seals (Mirounga leonina) Adjust Their Fine Scale Horizontal Movement and Diving Behaviour in Relation to Prey Encounter Rate.

Understanding the diving behaviour of diving predators in relation to concomitant prey distribution could have major practical applications in conservation biology by allowing the assessment of how changes in fine scale prey distribution impact foraging efficiency and ultimately population dynamics. The southern elephant seal (Mirounga leonina, hereafter SES), the largest phocid, is a major predator of the southern ocean feeding on myctophids and cephalopods. Because of its large size it can carry bio-loggers with minimal disturbance. Moreover, it has great diving abilities and a wide foraging habitat. Thus, the SES is a well suited model species to study predator diving behaviour and the distribution of ecologically important prey species in the Southern Ocean. In this study, we examined how SESs adjust their diving behaviour and horizontal movements in response to fine scale prey encounter densities using high resolution accelerometers, magnetometers, pressure sensors and GPS loggers. When high prey encounter rates were encountered, animals responded by (1) diving and returning to the surface with steeper angles, reducing the duration of transit dive phases (thus improving dive efficiency), and (2) exhibiting more horizontally and vertically sinuous bottom phases. In these cases, the distance travelled horizontally at the surface was reduced. This behaviour is likely to counteract horizontal displacement from water currents, as they try to remain within favourable prey patches. The prey encounter rate at the bottom of dives decreased with increasing diving depth, suggesting a combined effect of decreased accessibility and prey density with increasing depth. Prey encounter rate also decreased when the bottom phases of dives were spread across larger vertical extents of the water column. This result suggests that the vertical aggregation of prey can regulate prey density, and as a consequence impact the foraging success of SESs. To our knowledge, this is one of only a handful of studies showing how the vertical distributions and structure of prey fields influence the prey encounter rates of a diving predator.

Chemical Ecology of Wave-Swept Shores: the Primacy of Contact Cues in Predation by Whelks.

Wave-swept shores are valuable for developing and testing key ecological principles. A synthesis of research is nonetheless missing a critical component: the chemosensory basis for behavioral interactions that determine population- and community-wide attributes. Chemical signaling environments on wave-swept shores, given their intense, turbulent mixing and complex topographies, would be difficult or impossible to simulate in a laboratory setting. For this reason, appropriately scaled field studies are needed to advance understanding of chemical stimuli and their biotic effects. Here, we performed a field investigation to establish the relative roles of dissolved and contact cues in predation by whelks (Acanthinucella spirata) on barnacles (Balanus glandula), their preferred prey. Experiments tested responses of whelks to seawater drawn above dense prey patches (10,240-12,180 barnacles m-2) and also over adjacent sand flats (no prey present). There was no evidence of waterborne stimuli associated with prey, even when sea states were nearly tranquil. Field trials also tested faux prey, which were constructed from cleaned barnacle shells and flavored gels. Prospective contact cues were presented to whelks at concentrations typical of epidermal tissue and cuticle in live, intact barnacles. These compounds were highly effective inducers of attack behavior and feeding. Selective enzyme degradations showed that the bioactive material was proteinaceous. Moreover, whelks did not distinguish faux barnacles with a single, purified glycoprotein (named "MULTIFUNCin") from live counterparts. Combined field results thus demonstrate the importance of contact cues, and indicate little, if any, effect of waterborne cues on predation by whelks under native conditions. Our findings underscore the need for appropriately scaled field experiments, and highlight surface chemistry as a critical factor that drives trophic interactions on rocky, wave-swept shores.

Biosonar detection range of mesopelagic patches by spinner dolphins in Hawaii

Spinner dolphins (Stenella longirostris) in the near-shore waters of the Hawaiian islands forage on the mesopelagic boundary community (mbc) of organisms consisting of myctophids, mid-water shrimp, and small squids. They forage at night in a coordinated fashion swimming parallel to shore hunting for patches of prey that they can encircle and herd into a tight three-dimensional patch. A profiler housing a broadband echo-ranger that projected dolphin-like biosonar signals was used to measure the target strength of the mbc. Echoes consisted of a number of highlights bunched together with target strength between -45 and -55 dB based on a dolphin’s integration window of 264 ms. Noise values collected by an autonomous acoustic recorder at midnight in the location where the profiler data were obtained were used to estimate the biosonar detection range of spinner dolphins for mesopelagic patches. The receiving directivity index and the width of the auditory filter for Tursiops trun catus were used to estimate the biosonar detection ranges of Stenella longirostris searching for mbc patches. Using the sonar equation, the biosonar threshold detection range of spinner dolphins was estimated to be approximately 100 plus m, more than sufficient range for the animals to formulate their prey herding behavior.

Central-place foraging and ecological effects of an invasive predator across multiple habitats.

Cross-habitat foraging movements of predators can have widespread implications for predator and prey populations, community structure, nutrient transfer, and ecosystem function. Although central-place foraging models and other aspects of optimal foraging theory focus on individual predator behavior, they also provide useful frameworks for understanding the effects of predators on prey populations across multiple habitats. However, few studies have examined both the foraging behavior and ecological effects of nonnative predators across multiple habitats, and none has tested whether nonnative predators deplete prey in a manner predicted by these foraging models. I conducted behavioral observations of invasive lionfish (Pterois volitans) to determine whether they exhibit foraging movements similar to other central-place consumers. Then, I used a manipulative field experiment to test whether their effects on prey populations are consistent with three qualitative predictions from optimal foraging models. Specifically, I predicted that the effects of invasive lionfish on native prey will (1) occur at central sites first and then in surrounding habitats, (2) decrease with increasing distance away from their shelter site, and (3) extend to greater distances when prey patches are spaced closer together. Approximately 40% of lionfish exhibited short-term crepuscular foraging movements into surrounding habitats from the coral patch reefs where they shelter during daylight hours. Over the course of 7weeks, lionfish depleted native fish populations on the coral patch reefs where they reside, and subsequently on small structures in the surrounding habitat. However, their effects did not decrease with increasing distance from the central shelter site and the influence of patch spacing was opposite the prediction. Instead, lionfish always had the greatest effects in areas with the highest prey densities. The differences between the predicted and observed effects of lionfish foraging are likely due to different constraints faced by invasive predators compared to native predators, namely that lionfish do not face increased predation risk with increased movement away from shelter sites. By foraging at greater distances from patch reefs than native predators, lionfish eliminated a spatial refuge from predation used by juveniles of many commercially and ecologically important reef fishes.

Interaction of ecological and angler processes: experimental stocking in an open access, spatially structured fishery.

Effective management of socioecological systems requires an understanding of the complex interactions between people and the environment. In recreational fisheries, which are prime examples of socioecological systems, anglers are analogous to mobile predators in natural predator-prey systems, and individual fisheries in lakes across a region are analogous to a spatially structured landscape of prey patches. Hence, effective management of recreational fisheries across large spatial scales requires an understanding of the dynamic interactions among ecological density dependent processes, landscape-level characteristics, and angler behaviors. We focused on the stocked component of the open access rainbow trout (Oncorhynchus mykiss) fishery in British Columbia (BC), and we used an experimental approach wherein we manipulated stocking densities in a subset of 34 lakes in which we monitored angler effort, fish abundance, and fish size for up to seven consecutive years. We used an empirically derived relationship between fish abundance and fish size across rainbow trout populations in BC to provide a measure of catch-based fishing quality that accounts for the size-abundance trade off in this system. We replicated our experimental manipulation in two regions known to have different angler populations and broad-scale access costs. We hypothesized that angler effort would respond to variation in stocking density, resulting in spatial heterogeneity in angler effort but homogeneity in catch-based fishing quality within regions. We found that there is an intermediate stocking density for a given lake or region at which angler effort is maximized (i.e., an optimal stocking density), and that this stocking density depends on latent effort and lake accessibility. Furthermore, we found no clear effect of stocking density on our measure of catch-based fishing quality, suggesting that angler effort homogenizes catch-related attributes leading to an eroded relationship between stocking density and catch-based fishing quality at the timescale of annual surveys. We conclude that declines in fishing quality resulting from understocking (due to declines in catch rate with low fish abundance) and overstocking (due to suppressed growth and limited recruitment at high density) give an optimal stocking rate that depends on accessibility and latent effort.