Testing optimal foraging theory models on benthic divers

Empirical testing of optimal foraging models on diving air-breathing animals is limited due to difficulties in quantifying the prey field through direct observations. Here we used accelerometers to detect rapid head movements during prey encounter events (PEE) of free-ranging benthic-divers, Australian fur seals, Arctocephalus pusillus doriferus. PEE signals from accelerometer data were validated by simultaneous video data. We then used PEEs as a measure of patch quality to test several optimal foraging model predictions. Seals had longer bottom durations in unfruitful dives (no PEE) than those with some foraging success (PEE. ≥. 1). However, when examined in greater detail, seals had longer bottom durations in dives with more PEEs, but shorter bottom durations in bouts (sequences of dives) with more PEEs. Our results suggest that seals were generally maximizing bottom durations in all foraging dives, characteristic of benthic divers. However, successful foraging dives might be more energetically costly (e.g. digestive costs), thus resulting in shorter bottom durations at the larger scale of bouts. Our study provides a case study of how the foraging behaviour of a central place forager foraging in a fairly homogeneous environment, with relatively high travel costs, may deviate from current foraging models under different situations. Future foraging models should aim to integrate other aspects (e.g. diet) of the foraging process for more accurate predictions.

From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Contrasting patterns of individual specialization and trophic coupling in two marine apex predators

1. Apex predators are often assumed to be dietary generalists and, by feeding on prey from multiple basal nutrient sources, serve to couple discrete food webs. But there is increasing evidence that individual level dietary specialization may be common in many species, and this has not been investigated for many marine apex predators. 2. Because of their position at or near the top of many marine food webs, and the possibility that they can affect populations of their prey and induce trophic cascades, it is important to understand patterns of dietary specialization in shark populations. 3. Stable isotope values from body tissues with different turnover rates were used to quantify patterns of individual specialization in two species of ‘generalist’ sharks (bull sharks, Carcharhinus leucas, and tiger sharks, Galeocerdo cuvier). 4. Despite wide population-level isotopic niche breadths in both species, isotopic values of individual tiger sharks varied across tissues with different turnover rates. The population niche breadth was explained mostly by variation within individuals suggesting tiger sharks are true generalists. In contrast, isotope values of individual bull sharks were stable through time and their wide population level niche breadth was explained by variation among specialist individuals. 5. Relative resource abundance and spatial variation in food-predation risk tradeoffs may explain the differences in patterns of specialization between shark species. 6. The differences in individual dietary specialization between tiger sharks and bull sharks results in different functional roles in coupling or compartmentalizing distinct food webs. 7. Individual specialization may be an important feature of trophic dynamics of highly mobile marine top predators and should be explicitly considered in studies of marine food webs and the ecological role of top predators.

Non-consumptive effects of predators in coral reef communities and the indirect consequences of marine protected areas

Predators exert strong direct and indirect effects on ecological communities by intimidating their prey. Non-consumptive effects (NCEs) of predators are important features of many ecosystems and have changed the way we understand predator-prey interactions, but are not well understood in some systems. For my dissertation research I combined a variety of approaches to examine the effect of predation risk on herbivore foraging and reproductive behaviors in a coral reef ecosystem. In the first part of my dissertation, I investigated how diet and territoriality of herbivorous fish varied across multiple reefs with different levels of predator biomass in the Florida Keys National Marine Sanctuary. I show that both predator and damselfish abundance impacted diet diversity within populations for two herbivores in different ways. Additionally, reef protection and the associated recovery of large predators appeared to shape the trade-off reef herbivores made between territory size and quality. In the second part of my dissertation, I investigated context-dependent causal linkages between predation risk, herbivore foraging behavior and resource consumption in multiple field experiments. I found that reef complexity, predator hunting mode, light availability and prey hunger influenced prey perception of threat and their willingness to feed. This research argues for more emphasis on the role of predation risk in affecting individual herbivore foraging behavior in order to understand the implications of human-mediated predator removal and recovery in coral reef ecosystems.^

Non-Consumptive Effects of Predators in Coral Reef Communities and the Indirect Consequences of Marine Protected Areas

Predators exert strong direct and indirect effects on ecological communities by intimidating their prey. Non-consumptive effects (NCEs) of predators are important features of many ecosystems and have changed the way we understand predator-prey interactions, but are not well understood in some systems. For my dissertation research I combined a variety of approaches to examine the effect of predation risk on herbivore foraging and reproductive behaviors in a coral reef ecosystem. In the first part of my dissertation, I investigated how diet and territoriality of herbivorous fish varied across multiple reefs with different levels of predator biomass in the Florida Keys National Marine Sanctuary. I show that both predator and damselfish abundance impacted diet diversity within populations for two herbivores in different ways. Additionally, reef protection and the associated recovery of large predators appeared to shape the trade-off reef herbivores made between territory size and quality. In the second part of my dissertation, I investigated context-dependent causal linkages between predation risk, herbivore foraging behavior and resource consumption in multiple field experiments. I found that reef complexity, predator hunting mode, light availability and prey hunger influenced prey perception of threat and their willingness to feed. This research argues for more emphasis on the role of predation risk in affecting individual herbivore foraging behavior in order to understand the implications of human-mediated predator removal and recovery in coral reef ecosystems.

Acoustic telemetry and network analysis reveal the space use of multiple reef predators and enhance marine protected area design

Marine protected areas (MPAs) are commonly employed to protect ecosystems from threats like overfishing. Ideally, MPA design should incorporate movement data from multiple target species to ensure sufficient habitat is protected. We used long-term acoustic telemetry and network analysis to determine the fine-scale space use of five shark and one turtle species at a remote atoll in the Seychelles, Indian Ocean, and evaluate the efficacy of a proposed MPA. Results revealed strong, species-specific habitat use in both sharks and turtles, with corresponding variation in MPA use. Defining the MPA's boundary from the edge of the reef flat at low tide instead of the beach at high tide (the current best in Seychelles) significantly increased the MPA's coverage of predator movements by an average of 34%. Informed by these results, the larger MPA was adopted by the Seychelles government, demonstrating how telemetry data can improve shark spatial conservation by affecting policy directly.

Acoustic telemetry and network analysis reveal the space use of multiple reef predators and enhance marine protected area design

Marine protected areas (MPAs) are commonly employed to protect ecosystems from threats like overfishing. Ideally, MPA design should incorporate movement data from multiple target species to ensure sufficient habitat is protected. We used long-term acoustic telemetry and network analysis to determine the fine-scale space use of five shark and one turtle species at a remote atoll in the Seychelles, Indian Ocean, and evaluate the efficacy of a proposed MPA. Results revealed strong, species-specific habitat use in both sharks and turtles, with corresponding variation in MPA use. Defining the MPA's boundary from the edge of the reef flat at low tide instead of the beach at high tide (the current best in Seychelles) significantly increased the MPA's coverage of predator movements by an average of 34%. Informed by these results, the larger MPA was adopted by the Seychelles government, demonstrating how telemetry data can improve shark spatial conservation by affecting policy directly.

An overview on the role of Hexanchiformes in marine ecosystems: biology, ecology and conservation status of a primitive order of modern sharks

The large size, high trophic level and wide distribution of Hexanchiformes (cow and frilled sharks) should position this order as important apex predators in coastal and deep-water ecosystems. This review synthesizes available information on Hexanchiformes, including information not yet published, with the purpose of evaluating their conservation status and assessing their ecological roles in the dynamics of marine ecosystems. Comprising six species, this group has a wide global distribution, with members occurring from shallow coastal areas to depths of c. 2500 m. The limited information available on their reproductive biology suggests that they could be vulnerable to overexploitation (e.g. small litter sizes for most species and suspected long gestation periods). Most of the fishing pressure exerted on Hexanchiformes is in the form of commercial by-catch or recreational fishing. Comprehensive stock and impact assessments are unavailable for most species in most regions due to limited information on life history and catch and abundance time series. When hexanchiform species have been commercially harvested, however, they have been unable to sustain targeted fisheries for long periods. The potentially high vulnerability to intense fishing pressure warrants a conservative exploitation of this order until thorough quantitative assessments are conducted. At least some species have been shown to be significant apex predators in the systems they inhabit. Should Hexanchiformes be removed from coastal and deep-water systems, the lack of sympatric shark species that share the same resources suggests no other species would be capable of fulfilling their apex predator role in the short term. This has potential ecosystem consequences such as meso-predator release or trophic cascades. This review proposes some hypotheses on the ecology of Hexanchiformes and their role in ecosystem dynamics, highlighting the areas where critical information is required to stimulate research directions.

Fish Habitat Utilization Patterns and Evaluation of the Efficacy of Marine Protected Areas in Hawaii: Integration of NOAA Digital Benthic Habitat Mapping and Coral Reef Ecological Studies

Over the past four decades, the state of Hawaii has developed a system of eleven Marine Life Conservation Districts (MLCDs) to conserve and replenish marine resources around the state. Initially established to provide opportunities for public interaction with the marine environment, these MLCDs vary in size, habitat quality, and management regimes, providing an excellent opportunity to test hypotheses concerning marine protected area (MPA) design and function using multiple discreet sampling units. NOAA/NOS/NCCOS/Center for Coastal Monitoring and Assessment’s Biogeography Team developed digital benthic habitat maps for all MLCD and adjacent habitats. These maps were used to evaluate the efficacy of existing MLCDs for biodiversity conservation and fisheries replenishment, using a spatially explicit stratified random sampling design. Coupling the distribution of habitats and species habitat affinities using GIS technology elucidates species habitat utilization patterns at scales that are commensurate with ecosystem processes and is useful in defining essential fish habitat and biologically relevant boundaries for MPAs. Analysis of benthic cover validated the a priori classification of habitat types and provided justification for using these habitat strata to conduct stratified random sampling and analyses of fish habitat utilization patterns. Results showed that the abundance and distribution of species and assemblages exhibited strong correlations with habitat types. Fish assemblages in the colonized and uncolonized hardbottom habitats were found to be most similar among all of the habitat types. Much of the macroalgae habitat sampled was macroalgae growing on hard substrate, and as a result showed similarities with the other hardbottom assemblages. The fish assemblages in the sand habitats were highly variable but distinct from the other habitat types. Management regime also played an important role in the abundance and distribution of fish assemblages. MLCDs had higher values for most fish assemblage characteristics (e.g. biomass, size, diversity) compared with adjacent fished areas and Fisheries Management Areas (FMAs) across all habitat types. In addition, apex predators and other targeted resources species were more abundant and larger in the MLCDs, illustrating the effectiveness of these closures in conserving fish populations. Habitat complexity, quality, size and level of protection from fishing were important determinates of MLCD effectiveness with respect to their associated fish assemblages. (PDF contains 217 pages)

Anadromous fish as marine nutrient vectors

The tidal freshwater of Virginia supports anadromous herring (Alosa spp.) spawning runs in the spring; however, their importance as nutrient delivery vectors to the freshwater fish food web remains unknown. The stable isotope signatures of fishes from 21 species and four different guilds (predators, carnivores, generalists, and planktivores) were examined in this study to test the hypothesis that marine derived nutrients (MDNs) brought by anadromous fish would be traced into the guilds that incorporated them. Spawning anadromous fish were 13C and 34S-enriched (δ13C and δ34S of approximately 18‰ and 17.7‰, respectively) relative to resident freshwater fish. Of the guilds examined, only predators showed 13C and 34S-enrichment similar to the anadromous fish; however, some generalist catfish also showed enriched signatures. Specific fatty acid δ13C signatures for gizzard shad (Dorosoma cepedianum), blue catfish (Ictalurus furcatus), and alewife (Alosa pseudoharengus), show a 10‰ range among fishes, clearly reflecting isotopically distinct dietary sources. The δ13C and δ34S distribution and range among the freshwater fishes suggest that both autochthonous and allochthonous (terrestrial C3 photosynthetic production and MDN) nutrient sources are important to the tidal freshwater fish community.

Diurnal and nocturnal temperatures for Atlantic salmon postsmolts (Salmo salar L.) during their early marine life

Data storage tags (DSTs) were applied to Atlantic salmon (Salmo salar L.) smolts during their seaward migration in the spring of 2002 at a fish counting fence on Campbellton River, Newfoundland. Our objectives were to discover whether or not salmon smolts could carry DSTs and survive, whether or not useful data on thermal habitat could be obtained and interpreted, and whether or not salmon smolts moved vertically in the water column. Data were downloaded from 15 of the recovered tags and revealed the hourly water temperatures experienced by the fish for periods of 3 to 71 days. The data on the DSTs were analyzed for temperature patterns in relation to migration behavior and diurnal movement of the fish. While in the sea, the DSTs recorded night temperatures of 12.5°C, which were higher than day temperatures of 11.6°C; the record from moored recorders, however, indicated that sea temperatures actually declined at night. It is hypothesized that posts-molts avoid avian predators during daylight hours by positioning themselves deeper in the water column and that they were pursuing prey during the deeper vertical descents or ascents noted during the periods of more rapid changes in temperature.

Diet composition and food habits of demersal and pelagic marine fishes from Terengganu waters, east coast of Peninsular Malaysia

Fish stomachs from 18 demersal and pelagic fishes from the coast of Terengganu in Malaysia were examined. The components of the fishes’ diets varied in number, weight, and their frequency of occurrence. The major food items in the stomachs of each species were determined using an Index of Relative Importance. A conceptual food web structure indicates that fish species in the study area can be classified into three predatory groups: (1) predators on largely planktivorous or pelagic species; (2) predators on largely benthophagous or demersal species; and (3) mixed feeders that consume both pelagic and demersal species.

The Southeast Bering Sea Ecosystem: implications for marine resource management (Final Report: Southeast Bering Sea Carrying Capacity).

Southeast Bering Sea Carrying Capacity (SEBSCC, 1996–2002) was a NOAA Coastal Ocean Program project that investigated the marine ecosystem of the southeastern Bering Sea. SEBSCC was co-managed by the University of Alaska Fairbanks, NOAA Alaska Fisheries Science Center, and NOAA Pacific Marine Environmental Laboratory. Project goals were to understand the changing physical environment and its relationship to the biota of the region, to relate that understanding to natural variations in year-class strength of walleye pollock (Theragra chalcogramma), and to improve the flow of ecosystem information to fishery managers. In addition to SEBSCC, the Inner Front study (1997–2000), supported by the National Science Foundation (Prolonged Production and Trophic Transfer to Predators: Processes at the Inner Front of the S.E. Bering Sea), was active in the southeastern Bering Sea from 1997 to 1999. The SEBSCC and Inner Front studies were complementary. SEBSCC focused on the middle and outer shelf. Inner Front worked the middle and inner shelf. Collaboration between investigators in the two programs was strong, and the joint results yielded a substantially increased understanding of the regional ecosystem. SEBSCC focused on four central scientific issues: (1) How does climate variability influence the marine ecosystem of the Bering Sea? (2) What determines the timing, amount, and fate of primary and secondary production? (3) How do oceanographic conditions on the shelf influence distributions of fish and other species? (4) What limits the growth of fish populations on the eastern Bering Sea shelf? Underlying these broad questions was a narrower focus on walleye pollock, particularly a desire to understand ecological factors that affect year-class strength and the ability to predict the potential of a year class at the earliest possible time. The Inner Front program focused on the role of the structural front between the well-mixed waters of the coastal domain and the two-layer system of the middle domain. Of special interest was the potential for prolonged post-spring-bloom production at the front and its role in supporting upper trophic level organisms such as juvenile pollock and seabirds. Of concern to both programs was the role of interannual and longer-term variability in marine climates and their effects on the function of sub-arctic marine ecosystems and their ability to support upper trophic level organisms.