Linking movement, diving, and habitat to foraging success in a large marine predator

Establishing where and when predators forage is essential to understanding trophic interactions, yet foraging behavior remains poorly understood in large marine carnivores. We investigated the factors leading to foraging success in gray seals (Halichoerus grypus) in the Northwest Atlantic in the first study to use simultaneous deployments of satellite transmitters, time depth recorders, and stomach-temperature loggers on a free-ranging marine mammal. Thirty-two seals were each fitted with the three types of instrumentation; however, complete records from all three instruments were obtained from only 13 individuals, underscoring the difficulty of such a multi-instrument approach. Our goal was to determine the characteristics of diving, habitat, and movement that predict feeding. We linked diving behavior to foraging success at two temporal scales: trips (days) and bouts (hours) to test models of optimal diving, which indicate that feeding can be predicted by time spent at the bottom of a dive. Using an information-theoretic approach, a Generalized Linear Mixed Model with trip duration and accumulated bottom time per day best explained the number of feeding events per trip, whereas the best predictor of the number of feeding events per bout was accumulated bottom time. We then tested whether characteristics of movement were predictive of feeding. Significant predictors of the number of feeding events per trip were angular variance (i.e., path tortuosity) and distance traveled per day. Finally, we integrated measures of diving, movement, and habitat at four temporal scales to determine overall predictors of feeding. At the 3-h scale, mean bottom time and distance traveled were the most important predictors of feeding frequency, whereas at the 6-h and 24-h time scales, distance traveled alone was most important. Bathymetry was the most significant predictor of feeding at the 12-h interval, with feeding more likely to occur at deeper depths. Our findings indicate that several factors predict feeding in gray seals, but predictor variables differ across temporal scales such that environmental variation becomes important at some scales and not others. Overall, our results illustrate the value of simultaneously recording and integrating multiple types of information to better understand the circumstances leading to foraging success.     

Individual- and population-level responses of a keystone predator to geographic variation in prey

Investigating how food supply regulates the behavior and population structure of predators remains a central focus of population and community ecology. These responses will determine the strength of bottom-up processes through the food web, which can potentially lead to coupled top-down regulation of local communities. However, characterizing the bottom-up effects of prey is difficult in the case of generalist predators and particularly with predators that have large dispersal scales, attributes that characterize most marine top predators. Here we use long-term data on mussel, barnacle, limpet, and other adult prey abundance and recruitment at sites spread over 970 km to investigate individual- and population-level responses of the keystone intertidal sunstar Heliaster helianthus on the coast of Chile. Our results show that this generalist predator responds to changes in the supply of an apparently preferred prey, the competitively dominant mussel Perumytilus purpuratus. Individual-level parameters (diet composition, per capita prey consumption, predator size) positively responded to increased mussel abundance and recruitment, whereas population-level parameters (density, biomass, size structure) did not respond to bottom-up prey variation among sites separated by a few kilometers. No other intertidal prey elicited positive individual predator responses in this species, even though a large number of other prey species was always included in the diet. Moreover, examining predator-prey correlations at approximately 80, 160, and 200 km did not change this pattern, suggesting that positive prey feedback could occur over even larger spatial scales or as a geographically unstructured process. Thus individual-level responses were not transferred to population changes over the range of spatial scales examined here, highlighting the need to examine community regulation processes over multiple spatial scales.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Calanoid copepod escape behavior in response to a visual predator

Calanoid copepods typically exhibit escape reactions to hydrodynamic stimuli such as those generated by the approach of a predator. During the summers of 2000, 2001 and 2004, two small calanoid species, Temora turbinata Dana, 1849 and Paracalanus parvus Claus, 1863 were exposed to a visual predatory fish, the blenny Acanthemblemaria spinosa Metzelaar, 1919, and their predator–prey interactions were recorded using both high-speed and standard videographic techniques. Copepod escape reaction components, including swimming pattern, reactive distance, turning rate, and jump kinetics, were quantified from individual predation events using motion analysis techniques. Among the observed escape reaction components, differences were noted between the species’ swimming patterns prior to attack and their response latencies. Temora turbinata was a continuous cruiser and P. parvus exhibited a hop-and-sink swimming pattern. During periods of sinking, P. parvus stopped beating its appendages, which presumably reduced any self-generated hydrodynamic signals and increased perceptual abilities to detect an approaching predator. Response latency was determined for each copepod species using a hydrodynamic stimulus produced by a 1 ms acoustic signal. Response latencies of T. turbinata were significantly longer than those of P. parvus. Despite some apparent perceptual advantages of P. parvus, the blenny successfully captured both species by modifying its attack behavior for the targeted prey.     

Harvest policies and nonmarket valuation in a predator — prey system

Although prey may not have commercial value, their economic value can be ascertained in a predator-prey model if the predator has a harvest value. The economic optimal (recovery) path of the predator and prey are carefully described when growth is quadratic in the predator (prey) and linear in prey (predator). Parameter values, in part, resembling Pacific halibut are used to provide numerical illustrations.     

Behavioral response of a generalist predator to chemotactile cues of two taxonomically distinct prey species

Chemotactile cues unintentionally left by animals can play a major role in predator–prey interactions. Specialized predators can use them to find their prey, while prey individuals can assess predation risk. However, little is known to date about the importance of chemotactile cues for generalist predators such as ants. Here, we investigated the response of a generalized predatory ant, Formica polyctena, to cues of two taxonomically distinct prey: a spider (Pisaura mirabilis) and a cricket (Nemobius sylvestris). In analogy, we studied whether crickets and spiders showed antipredator behavior in response to ant cues. When confronted with cues of the two prey species, Formica polyctena workers showed increased residence time and reduced movement speed, which suggests success-motivated searching behavior and thus increased foraging effort. The ants’ response did not differ between cues of the two prey species, coinciding with similar aggression and consumption rates of dead prey. However, the cuticular hydrocarbons, which likely resemble part of the potential cues, differed strongly between the species, with only few methyl-branched alkanes in common. This suggests that ants respond to multiple compounds left by other organisms with prey-search behavior. The two prey species, in turn, showed no detectable antipredator behavior in response to ant cues. Our study shows that ants can detect and respond to chemotactile cues of taxonomically and ecologically distinct prey species, probably to raise their foraging success. Using such chemotactile cues for prey detection may drastically increase their foraging efficiency and thus contribute to the high ecological success of ants.     

Using time-depth-light recorders to measure light levels experienced by a diving marine mammal

Acoustic telemetry was used to track vertical and horizontal movement patterns and to monitor the stomach temperatures of seven juvenile shortfin mako sharks (Isurus oxyrinchus Rafinesque) in the Southern California Bight from July to November 2002. Makos (80–145 cm fork length, FL) were attracted to the tracking vessel, where they were fed a mackerel containing an acoustic transmitter that reported temperature and pressure. Tracks ranged from 6.8–45.4 h. Collectively, the mako sharks spent 80% of the track record at 0–12 m, 15% at 12–24 m, and 5% at depths >24 m. The average horizontal swimming speed was 2.3 km h^–1 or 0.55 FLs s^–1, and the greatest distance traveled was 145 km in 45.4 h. For the six tracks >21 h, there was a positive correlation between body size and maximum depth. Makos used more of the water column during daylight hours. Mean stomach temperature was 3.8±1.5°C above ambient, and body size was positively correlated with both maximum and average stomach temperature. Stomach content analyses of four makos captured at the end of tracking verified the occurrence of feeding events as indicated by changes in stomach temperature.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick     

Marine habitat mapping in support of Marine Protected Area management in a subarctic fjord: Gilbert Bay, Labrador, Canada

This paper presents an approach that allows production of benthic substrate and habitat maps in fjord environments. This approach is used to support the management of the Gilbert Bay Marine Protected Area (MPA) in southeastern Labrador, Atlantic Canada. Multibeam sonar-derived bathymetry, seabed slope, and acoustic reflectance (backscatter) were combined using supervised classification methods and GIS with ground-truthed benthic sampling in order to derive maps of the substrates and main benthic habitats. Six acoustically distinct substrate types were identified in the fjord, and three additional substrate types without a unique acoustic signature were recognized. Ordination by multidimensional scaling and analysis of similarity generalized these to four acoustically distinct habitat types. Greatest within-habitat (alpha) diversity was found in the coralline-algae encrusted gravel habitat. Greatest between-habitat (beta) diversity was found in the management Zones 1 and 2, which have the highest level of protection. The study confirmed that the zoning plan for the MPA, which was designed to protect spawning and juvenile fish habitat for a local genetically distinct population of Atlantic cod, afforded highest levels of protection to areas with highest habitat diversity.     

Foraging behavior of a generalist marine top predator,Japanese cormorants (<Emphasis Type="Italic">Phalacrocorax filamentosus</Emphasis>), in years of demersal versus epipelagic prey

To examine the behavioral adjustment of a generalist marine top predator to variability of their prey, we studied the foraging behavior of Japanese cormorants (Phalacrocorax filamentosus) breeding at Teuri Island, Hokkaido, in years of contrasting demersal and epipelagic prey composition. We used radio telemetry and ship-based surveys to determine behavior and at-sea distribution during three summers (1996–1998). The cormorants fed on epipelagic anchovy (Engraulis japonicus) and sandlance (Ammodytes personatus) in 1998 (year of epipelagic diet), while they fed on benthic rock fish (Sebastes spp.) and flatfish (Pleuronectidae) and nearshore-living naked sandlance (Hypophychus dybowskii), as well as epibenthic greenling (Hexagrammidae) in 1996 and 1997 (year of demersal diet). Cormorants engaged in larger feeding groups, visited more feeding sites, and stayed at each feeding site for a shorter period in the year of epipelagic diet than in the years of demersal diet. The cormorants made long foraging trips and fed in the mainland coastal habitat, distant from the colony, in the years of demersal diet. Individual radio-tracked birds fed over the wide area between the islands and mainland, in the year of epipelagic diet, while most individuals specialized in mainland or island coastal habitats in the years of demersal diet. Behavioral adjustment of Japanese cormorants might allow them to exploit both unpredictable epipelagic and predictable benthic prey efficiently.Communicated by T. Ikeda, Hakodate     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Seasonal changes in microbial heterotrophic activity in subarctic marine waters as related to phytoplankton primary productivity

Water samples were collected in and near Kasitsna Bay (Cook Inlet), Alaska over 18 mo (February 1979–August 1980). Seasonal changes in glucose and glutamate uptake rates were measured in these samples. During the second year of the study, the uptake rates of glycolate and acetate and primary productivity were also measured. Of the substrates tested, significant positive linear correlations were observed between glucose and glutamate uptake and primary productivity. A higher correlation was observed between glucose to glutamate uptake ratios and primary productivity. The seasonal patterns of glucose uptake, glucose-to-glutamate uptake ratios and primary productivity rates show that the ratios changes simultaneously with fluctuations in primary productivity rates. The glucose uptake patterns reveal a delay in the response of the glucose-utilizing microorganisms to the onset of the bloom. It is suggested that by measuring the uptake rates of the simple compounds by microheterotrophs, and comparing these rates with each other, one can determine the relative flux of these compounds through the system. If the major source of these compounds is material released into the seawater by phytoplankton, it is possible that this approach could characterize the flux of extracellular products. As more is known about how the composition of extracellular material relates to the physiological state of phytoplankton, such an approach may eventually be a useful indirect assessment of the physiological state of natural phytoplankton populations.Published as Technical Paper No. 6397, Oregon Agricultural Experiment Station     

Linking local to global properties in branching modular networks: gorgonian coral colonies

Branching growth is present both in plants and animals, either marine or terrestrial. Although cellular and other modular levels of organization in plants and animals have evolved through different molecular and physiological mechanisms, several aspects of their branching modular system and morphology are similar. We studied vessel organization and colony integration, in order to comprehend underlying relationships between different structural components in a gorgonian coral network. The theoretical formalism was validated in the gorgonian coral Eunicea mammosa (Plexauridae, Octocorallia) in Belize. As in vascular plants, these colonial animals create a complex network of connections among modular branches integrated in stem canals downstream toward the base. A new formalism is proposed for describing gorgonian branching. A global property of a colony is for instance the size of its base or its weight whereas a local property is the size of branch in a particular place of the colony. However, a global property is not the simple addition of local modular properties, as the case of stem canals in the colony base. Theoretically, the process of branching is tightly intertwined with the internal network organization. The colony network centralization is driven by a linear relationship between the total number of branches and the stem canals at the base of the colony. If stem canals play important roles in the transport of nutrients throughout the colony and the biomechanical support from the base up to the tips, we can assume that there is an underlying association between the number of stem canals at the base and the number of for example, terminal branches. These associations may provide new findings that extend our understanding of the functional organization of tree-like networks in octocorals and their vascular systems. The idea that the external components of a tree-like plant network are directly correlated and connected down to the main trunk seems to be analogous in an animal system.     

Predatory behavior and prey selection by army ants in a desert-grassland habitat

Summary Colonies of Neivamyrmex nigrescens conduct extensive nocturnal raids on other ants and termites in the desert-grassland of Arizona-New Mexico. We collected quantitative data on several aspects of raiding to pinpoint differences due to colony size and behavioral phase. In the nomadic phase, colonies began raiding at sunset and continued until dawn. Larger colonies covered more area, discovered more prey sites, and collected more booty than smaller colonies, but there were no systematic changes in raid intensity over the course of the nomadic phase. In the statary phase, raiding occurred less frequently and was less intense when it occurred; however, at the end of this phase, raiding was similar to nomadic phase raids in extent, duration, and booty captured. N. nigrescens preyed exclusively on termites and ants, and appeared to select certain species of Pheidole in preference to other ants. Pheidole was the most abundant genus, but was preyed upon twice as often as expected based on relative colony density. Pheidole attempted to avoid predation by fleeing or defending their nest, but rarely succeeded. Because they are about the same size as army ants and lack defensive chemicals, Pheidole made comparatively easy prey. N. nigrescens ignored or was repelled by other ants (Pogonomyrmex, Novomessor, Iridomyrmex, Myrmecocystus) during the early summer, when Pheidole was abundant; however, in late summer when Pheidole was less available, the army ants preyed upon Novomessor cockerelli. N. harrisi raided in close proximity to N. nigrescens, but preyed exclusively on Solenopsis xyloni. Selection of prey and partitioning of resources are now indicated in several army ant species; these processes have probably been important factors in the evolution of the ants' predatory behavior.     

Nephtys hombergii, a free-living predator in marine sediments: energy production under environmental stress

Nephtys hombergii is a free-living, burrowing predator in marine sediments. The worm is, therefore, exposed to various environmental conditions which tube-dwelling polychaetes of the same habitat most likely do not encounter. The worms have to survive periods of severe hypoxia and sulphide exposure, while at the same time, they have to maintain agility in order to feed on other invertebrates. N. hombergii is adapted to these conditions by utilising several strategies. The species has a remarkably high content of phosphagen (phosphoglycocyamine), which is the primary energy source during periods of environmental stress. With increasing hypoxia, energy is also provided via anaerobic glycolysis (pO₂<7 kPa), with strombine as the main end-product. Energy production via the succinate pathway becomes important only under severe hypoxia (<2 kPa), suggesting a biphasic response to low oxygen conditions which probably is related to the worm's mode of life. The presence of sulphide resulted in a higher anaerobic energy flux and a more pronounced energy production via glycolysis than in anoxia alone. Nevertheless, after sulphide exposure under anaerobic conditions of <24 h, N. hombergii is able to recover completely. Although N. hombergii appears to be well adapted to a habitat with short-term fluctuations in oxygen and appearance of hydrogen sulphide, its high energy demand as a predator renders it likely to limit its survival in an environment with longer lasting anoxia and concomitant sulphide exposure. Received: 28 May 1997 / Accepted: 21 June 1997     

Constancy and change in marine predator diets across a shift in oceanographic conditions in the Northern California Current

Variable ocean conditions can greatly impact prey assemblages and predator foraging in marine ecosystems. Our goal was to better understand how a change in ocean conditions influenced dietary niche overlap among a suite of midtrophic-level predators. We examined the diets of three fishes and one seabird off central Oregon during two boreal summer upwelling periods with contrasting El Niño (2010) and La Niña (2011) conditions. We found greater niche specialization during El Niño and increased niche overlap during La Niña in both the nekton and micronekton diet components, especially in the larger, more offshore predators. However, only the two smaller, more nearshore predators exhibited interannual variation in diet composition. Concurrent trawl surveys confirmed that changes in components of predator diets reflected changes in the prey community. Using multiple predators across diverse taxa and life histories provided a comprehensive understanding of food-web dynamics during changing ocean conditions.     

Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system

In variable environments, organisms are bound to track environmental changes if they are to survive. Most marine mammals and seabirds are colonial, central-place foragers with long-term breeding-site fidelity. When confronted with environmental change, such species are potentially constrained in their ability to respond to these changes. For example, if environmental conditions deteriorate within their limited foraging range, long-lived species favour adult survival and abandon their current breeding effort, which ultimately influences population dynamics. Should poor conditions persist over several seasons, breeding-site fidelity may force animals to continue breeding in low-quality habitats instead of emigrating towards more profitable grounds. We assessed the behavioural response of a site-faithful central-place forager, the Cape gannet Morus capensis, endemic to the Benguela upwelling system, to a rapid shift in the distribution and abundance of its preferred prey, small pelagic shoaling fish. We studied the distribution and the abundance of prey species, and the diet, foraging distribution, foraging effort, energy requirements, and breeding success of gannets at Malgas Island (South Africa) over four consecutive breeding seasons. Facing a rapid depletion of preferred food within their foraging range, Cape gannets initially increased their foraging effort in search of their natural prey. However, as pelagic fish became progressively scarcer, breeding birds resorted to scavenging readily available discards from a nearby demersal fishery. Their chicks cannot survive on such a diet, and during our 4-year study, numbers of breeding birds at the colony decreased by 40% and breeding success of the remaining birds was very low. Such behavioural inflexibility caused numbers of Cape gannets breeding in Namibia to crash by 95% following over-fishing of pelagic fish in the 1970s. In the context of rapid environmental changes, breeding-site fidelity of long-lived species may increase the risk of local or even global extinction, rendering these species particularly vulnerable to global change.