Integration of a fish bioenergetics model into a spatially explicit water quality model: Application to menhaden in Chesapeake Bay

Although fish are usually thought of as victims of water quality degradation, it has been proposed that some planktivorous species may improve water quality through consumption of algae and sequestering of nutrients via growth. Within most numerical water quality models, the highest trophic level modeled explicitly is zooplankton, prohibiting an investigation of the effect a fish species may be having on its environment. Conversely, numerical models of fish consumption do not typically include feedback mechanisms to capture the effects of fish on primary production and nutrient recycling. In the present study, a fish bioenergetics model is incorporated into CE-QUAL-ICM, a spatially explicit eutrophication model. In addition to fish consumption of algae, zooplankton, and detritus, fish biomass accumulation and nutrient recycling to the water column are explicitly accounted for. These developments advance prior modeling efforts of the impact of fish on water quality, many of which are based on integrated estimates over an entire system and which omit the feedback the fish have through nutrient recycling and excretion. To validate the developments, a pilot application was undertaken for Atlantic menhaden (Brevoortia tyrannus) in Chesapeake Bay. The model indicates menhaden may reduce the algal biomass while simultaneously increasing primary productivity.     

Pattern-oriented modeling of bird foraging and pest control in coffee farms

We develop a model of how land use and habitat diversity affect migratory bird populations and their ability to suppress an insect pest on Jamaican coffee farms. Bird foraging—choosing which habitat patch and prey to use as prey abundance changes over space and time—is the key process driving this system. Following the “pattern-oriented” modeling strategy, we identified nine observed patterns that characterize the real system's dynamics. The model was designed so that these patterns could potentially emerge from it. The resulting model is individual-based, has fine spatial and temporal resolutions, represents very simply the supply of the pest insect and other arthropod food in six habitat types, and includes foraging habitat selection as the only adaptive behavior of birds. Although there is an extensive heritage of bird foraging theory in ecology, most of it addresses only the individual level and is too simple for our context. We used pattern-oriented modeling to develop and test foraging theory for this across-scale problem: rules for individual bird foraging that cause the model to reproduce a variety of patterns observed at the system level. Four alternative foraging theories were contrasted by how well they caused the model to reproduce the nine characteristic patterns. Four of these patterns were clearly reproduced with the “null” theory that birds select habitat randomly. A version of classical theory in which birds stay in a patch until food is depleted to some threshold caused the model to reproduce five patterns; this theory caused lower, not higher, use of habitat experiencing an outbreak of prey insects. Assuming that birds select the nearby patch providing highest intake rate caused the model to reproduce all but one pattern, whereas assuming birds select the highest-intake patch over a large radius produced an unrealistic distribution of movement distances. The pattern reproduced under none of the theories, a negative relation between bird density and distance to trees, appears to result from a process not in the model: birds return to trees at night to roost. We conclude that a foraging model for small insectivorous birds in diverse habitat should assume birds can sense higher food supply but over short, not long, distances.     

Field evidence for pervasive indirect effects of fishing on prey foraging behavior

The indirect, ecosystem-level consequences of ocean fishing, and particularly the mechanisms driving them, are poorly understood. Most studies focus on density-mediated trophic cascades, where removal of predators alternately causes increases and decreases in abundances of lower trophic levels. However, cascades could also be driven by where and when prey forage rather than solely by prey abundance. Over a large gradient of fishing intensity in the central Pacific's remote northern Line Islands, including a nearly pristine, baseline coral reef system, we found that changes in predation risk elicit strong behavioral responses in foraging patterns across multiple prey fish species. These responses were observed as a function of both short-term ("acute") risk and longer-term ("chronic") risk, as well as when prey were exposed to model predators to isolate the effect of perceived predation risk from other potentially confounding factors. Compared to numerical prey responses, antipredator behavioral responses such as these can potentially have far greater net impacts (by occurring over entire assemblages) and operate over shorter temporal scales (with potentially instantaneous response times) in transmitting top-down effects. A rich body of literature exists on both the direct effects of human removal of predators from ecosystems and predators' effects on prey behavior. Our results draw together these lines of research and provide the first empirical evidence that large-scale human removal of predators from a natural ecosystem indirectly alters prey behavior. These behavioral changes may, in turn, drive previously unsuspected alterations in reef food webs.     

Foraging impact on zooplankton by age-0 walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) around a front in the southeast Bering Sea

The waters around the Pribilof Islands in the southeast Bering Sea are a center of abundance for age-0 walleye pollock (Theragra chalcogramma). Each spring and summer a tidal front is formed around the islands separating a well-mixed inshore habitat from a stratified offshore habitat. The objective of this study was to assess the foraging impact on zooplankton by age-0 pollock in the vicinity of this frontal structure. A bioenergetic model was used to estimate age-0 pollock food consumption from field estimates of water temperature, age-0 pollock density, diet and growth. Sampling of field variables took place over three hydrographic habitats along an inshore–offshore transect located north of the islands. The bioenergetics analysis was applied for a 2-week period during the late summer of four consecutive years, 1994–1997. Model results of age-0 pollock food consumption indicated variable levels of food depletion, changing with prey type, year and habitat. The foraging impact of age-0 pollock on copepods and euphausiids (most common prey) ranged from about 3% to 77% of the biomass available at the start of the simulation. Copepod depletion was typically greater than euphausiid depletion. Consequently, juvenile pollock <60 mm in standard length were more likely to experience food limitation due to the greater proportion of copepods in their diet. We present evidence of severe foraging impact during 1996, when one of the primary prey items of juvenile pollock (i.e. large copepods) was scarcely represented both in their diet and in the water column. In all years, most instances of prey depletion were found at the inshore and front habitats; age-0 pollock densities were too low relative to their prey to severely impact the offshore zooplankton populations. We discuss these results with respect to modeling assumptions and in the context of previously acquired knowledge of fish behavior around frontal regions.Communicated by T. Ikeda, Hakodate     

Models of fish school size in relation to environmental productivity

A simple energy-balance model, relating energetic requirements of fish schools to food production, was used to predict shoal sizes. Lower limits to school size are unlikely to be set by food but rather by predation. Upper limits depend on both food and school behavior, being greater for schools that break up to feed than for schools that remain continuously cohesive. Faced with a decreasing food supply, a school could either break into smaller schools or change behavior, increasing the area available for foraging. The models suggest that environmental productivity needs to be considered when examining fishery statistics such as (catch per unit effort), where maximum catch may be limited by maximum school size.     

Modeling individual and population dynamics in a consumer–resource system: Behavior under food limitation and crowding and the effect on population cycling in Daphnia

In population modeling, a considerable level of complexity is often required to provide trustworthy results, comparable with field observations. By assuring sufficient detail at the individual level while preserving the potential to explore the consequences at higher levels, individual-based modeling may thus provide a useful tool to investigate dynamics at different levels of organization. Still, population dynamics resulting from such models are often at odds with observations from the field. This may be partly caused by a lack of focus on the individual dynamics under conditions of food stress and starvation. I developed a physiologically structured, individual-based simulation model to investigate life history of Daphnia and its effect on population dynamics in response to the productivity of the system. In verifying model behavior with available literature data on life history and physiology, I paid special attention to the dynamics of food intake and the verification of individual level results under conditions of food limitation and starvation. I show that the maximum filtering rates under low food levels used in the current model are much closer to measured filtering rates than the ones used in other models. Being consistent with results on physiology and life history from experiments at a wide range of food availability (including starvation), the model generates low amplitude or high amplitude population density cycles depending on the productivity of the system, as observed in field and experimental populations of Daphnia and with the minimum population densities being one to two orders of magnitude lower in the high amplitude than in the low amplitude cycles. To generate results which are not only qualitatively but also quantitatively comparable to experimental and field observations, however, a crowding effect on the filtering response has to be incorporated in the model.     

Simulating spatiotemporal dynamics of urbanization with multi-agent systems—A case study of the Phoenix metropolitan region, USA

Urbanization is a human-dominated process and has greatly impacted biodiversity, ecosystem processes, and regional climate. To understand the socioeconomic drivers of urbanization and project future urban landscape changes, multi-agent systems provide a powerful tool. We develop an agent-based model of urban growth for the Phoenix metropolitan region of the United States, which simulates the behavior of regional authorities, real estate developers, residents, and environmentalists. The BDI (Beliefs-Desires-Intentions) structure is employed to simulate the agents behavior and decision models. The heterogeneity of agents is reflected by adjusting parameters according to the agents’ beliefs, desires and preferences. Three scenarios, baseline, economic development priority and environmental protection, are developed and analyzed. The combination of multi-agent system and spatial regression model is employed to predict the future urban development of the Phoenix metropolitan region. Landscape metrics are used to compare the spatial patterns of the urban landscape resulting from different scenarios in different times. In general, with the rapid urban expansion, the shape of urban patches will become more regular as many of them become coalesced. The spatial analysis of urban development through modeling individual and group decisions and human-environment interactions with a multi-agent systems approach can enhance our understanding of the socioeconomic driving forces and mechanisms of urban development.     

Modeling Human Factors That Affect the Loss of Biodiversity

Within conservation biology human factors are treated as driving forces of biodiversity loss, yet there are few empirical studies on how human actions affect biodiversity. We developed and tested an interdisciplinary model of biodiversity loss using socioeconomic and ecological data from 107 countries and structural equation modeling techniques. Some portions of the model fit the data well, other parts were less predictive. Counterintuitive results may be a result of the quality and availability of cross-national data and statistical limitations in testing a model of such complex processes. This model test provides insight into future research needs for examining human impacts on biodiversity. Issues including data quality, temporal and spatial scale, and model refinement are outlined. The results highlight the importance of relations between human social systems and biodiversity and the potential of interdisciplinary research.     

Relationship between the growth and survival of larval Pacific bluefin tuna, Thunnus orientalis

We tested the hypothesis that a large body size and rapid growth rate affect the survival of larval Pacific bluefin tuna, Thunnus orientalis (PBT), and analyzed larval growth in relation to environmental conditions. Seven high density larval patches of PBT were tracked with reference buoys in the northwestern Pacific Ocean for 28–171 h in May–June from 2004 to 2008. The otolith radii and daily growth rates of the survivor larvae (collected on later tracking days of each tracking session) tended to be larger and more rapid, respectively, than those of original larvae (collected on earlier tracking days). A large body size was found to positively affect the survival of larval PBT, as did a rapid growth rate, even at an early larval stage (7 days after hatching). Generalized linear modeling showed that the otolith radius was influenced positively by the sea temperature, stratification parameter and food density, while the growth rate was influenced positively by the sea temperature and food density.     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.     

Habitat use and group size of pied cormorants (<Emphasis Type="Italic">Phalacrocorax varius</Emphasis>) in a seagrass ecosystem: possible effects of food abundance and predation risk

Both food abundance and predation risk may influence habitat use decisions. However, studies of habitat use by birds in marine environments have focused only on food abundance. I investigated the possible influences of food abundance and predation risk from tiger sharks (Galeocerdo cuvier) on habitat use by pied cormorants (Phalacrocorax varius) over two spatial scales and on cormorant group size. Cormorants were usually solitary, but group size was highest in shallow habitats during months when shark density was low. Regardless of season, cormorant density within shallow habitats was higher over seagrass than sand, and cormorants were distributed between these two microhabitats proportional to prey density. Therefore, cormorants appear to respond to prey abundance at a relatively narrow spatial scale (i.e., tens of meters). At the habitat-patch scale (~1 km), the density of cormorants and their prey (teleosts) was higher in shallow habitats than in deep ones, but the density of cormorants was influenced by an interaction between water temperature (i.e., season) and habitat. There was decreased use of shallow habitats as water temperature, and the density of tiger sharks, increased. When shark density was low, cormorants were distributed across habitats roughly in proportion to the abundance of fish, suggesting that cormorants respond to food abundance at the scale of habitat patches. However, as shark abundance increased, the relative density of cormorants dropped in the dangerous shallow habitats such that there was a greater density of cormorants relative to their food in deep habitats when sharks were abundant. This suggests that pied cormorants trade-off food and risk by accepting lower energetic returns to forage in safer habitats. This study provides the first evidence that marine habitat selection by birds may be influenced by such a trade-off, and provides further evidence that tiger sharks are important in determining habitat use of their prey and mediating indirect interactions within Shark Bay.Communicated by P. W. Sammarco, Chauvin     

Linking patch-use behavior, resource density, and growth expectations in fish

Optimality theory rests on the assumptions that short-term foraging decisions are driven by variation in environmental quality, and that these decisions have important implications for long-term fitness. These assumptions, however, are rarely tested in a field setting. We linked behavioral foraging decisions in food patches with measures of environmental quality covering larger spatial (resource density) or temporal (growth parameters) scales. In 10 lakes, we measured the food density at which benthic fish give up foraging in experimental food patches (giving-up density, GUD), quantified the biomass of benthic invertebrates, and calculated the maximum individual size (L(infinity)) of bream (Abramis brama L.), a typical benthivore in these lakes. We found positive relationships between resource density and both GUD and L(infinity), and a positive relationship between L(infinity) and GUD. Prey characterized as vulnerable to predation contributed most to the relationships between resource density and either GUD or L(infinity). A path analysis showed that resource density and L(infinity) directly explained 54% and 28%, respectively, of the variation in GUD, whereas 86% of the variation in L(infinity) was explained by resource density, with mostly indirect contribution from GUD. We conclude that the short-term foraging behavior of benthivores matched our expectations based on optimality theory by being positively linked to variables on environmental quality operating at both a larger spatial scale and a longer temporal scale.     

Wind- and tide-induced currents in the Stagnone lagoon (Sicily)

The hydrodynamic circulation is analyzed in the coastal lagoon of Stagnone di Marsala, a natural reserve located in the north-western part of Sicily, using both experimental measurements and numerical simulations. Field measurements of velocities and water levels, carried out using an ultrasound sensor (3D), are used to validate the numerical model. A 3D finite-volume model is used to solve the Reynolds-averaged momentum and mass balance differential equations on a curvilinear structured grid, employing the k–e{\varepsilon} turbulence model for the Reynolds stresses. The numerical analysis allows to identify the relative contribution of the forces affecting the hydrodynamic circulation inside the lagoon. In the simulations only wind and tide forces are considered, neglecting the effects of water density changes. Two different conditions are considered. In the first both the wind stress over the free-surface and the tidal motion are imposed. In the second the wind action is neglected, to separately analyze the tide-induced circulation. The comparison between the two test cases highlights the fundamental role of the wind on the hydrodynamics of the Stagnone lagoon, producing a strong vertical recirculation pattern that is not observed when the flow is driven by tides only.     

A framework for evaluating regional-scale numerical photochemical modeling systems

This paper discusses the need for critically evaluating regional-scale (~200–2,000 km) three-dimensional numerical photochemical air quality modeling systems to establish a model’s credibility in simulating the spatio-temporal features embedded in the observations. Because of limitations of currently used approaches for evaluating regional air quality models, a framework for model evaluation is introduced here for determining the suitability of a modeling system for a given application, distinguishing the performance between different models through confidence-testing of model results, guiding model development and analyzing the impacts of regulatory policy options. The framework identifies operational, diagnostic, dynamic, and probabilistic types of model evaluation. Operational evaluation techniques include statistical and graphical analyses aimed at determining whether model estimates are in agreement with the observations in an overall sense. Diagnostic evaluation focuses on process-oriented analyses to determine whether the individual processes and components of the model system are working correctly, both independently and in combination. Dynamic evaluation assesses the ability of the air quality model to simulate changes in air quality stemming from changes in source emissions and/or meteorology, the principal forces that drive the air quality model. Probabilistic evaluation attempts to assess the confidence that can be placed in model predictions using techniques such as ensemble modeling and Bayesian model averaging. The advantages of these types of model evaluation approaches are discussed in this paper.     

Costs and benefits of pocket gopher foraging: linking behavior and physiology

Animals can attain fitness benefits by maintaining a positive net energy balance, including costs of movement during resource acquisition and the profits from foraging. Subterranean rodent burrowing provides an excellent system in which to examine the effects of movement costs on foraging behavior because it is energetically expensive to excavate burrows. We used an individual-based modeling approach to study pocket gopher foraging and its relationship to digging cost, food abundance, and food distribution. We used a unique combination of an individual-based foraging-behavior model and an energetic model to assess survival, body mass dynamics, and burrow configurations. Our model revealed that even the extreme cost of digging is not as costly as it appears when compared to metabolic costs. Concentrating digging in the area where food was found, or area-restricted search (ARS), was the most energetically efficient digging strategy compared to a random strategy. Field data show that natural burrow configurations were more closely approximated by the animals we modeled using ARS compared to random diggers. By using behavior and simple physiological principles in our model, we were able to observe realistic body mass dynamics and recreate natural movement patterns.     

Fish dispersal in fragmented landscapes: a modeling framework for quantifying the permeability of structural barriers

Dispersal is a key determinant of the spatial distribution and abundance of populations, but human-made fragmentation can create barriers that hinder dispersal and reduce population viability. This study presents a modeling framework based on dispersal kernels (modified Laplace distributions) that describe stream fish dispersal in the presence of obstacles to passage. We used mark-recapture trials to quantify summer dispersal of brook trout (Salvelinus fontinalis) in four streams crossed by a highway. The analysis identified population heterogeneity in dispersal behavior, as revealed by the presence of a dominant sedentary component (48-72% of all individuals) characterized by short mean dispersal distance (<10 m), and a secondary mobile component characterized by longer mean dispersal distance (56-1086 m). We did not detect evidence of barrier effects on dispersal through highway crossings. Simulation of various plausible scenarios indicated that detectability of barrier effects was strongly dependent on features of sampling design, such as spatial configuration of the sampling area, barrier extent, and sample size. The proposed modeling framework extends conventional dispersal kernels by incorporating structural barriers. A major strength of the approach is that ecological process (dispersal model) and sampling design (observation model) are incorporated simultaneously into the analysis. This feature can facilitate the use of prior knowledge to improve sampling efficiency of mark-recapture trials in movement studies. Model-based estimation of barrier permeability and its associated uncertainty provides a rigorous approach for quantifying the effect of barriers on stream fish dispersal and assessing population dynamics of stream fish in fragmented landscapes.     

Relative growth and production of the estuary grouper Epinephelus salmoides under different stocking densities in floating net-cages

Studies on the relative growth and production of the estuary grouper Epinephelus salmoides Maxwell were conducted in floating net-cages at 5 different stocking densities to determine the optimal level for stocking for commercial culture. The fish were stocked at densities of 15, 30, 60, 90 and 120 fish per m³, and reared for a period of 8 months. Results of the present study indicated that fish stocked at a density of 60 fish m^-3 grew equally fast and showed a food conversion ratio, mortality rate, and condition factor comparable to those at the lower stocking densities of 15 and 30 fish m^-3. At the end of the experiments, net-yield and production at this level of stocking density (60 fish m^-3) were not significantly different from those at the higher stocking densities of 90 and 120 fish m^-3, but showed increases of 86.7% over those at a stocking density of 30 fish m^-3 and 294.2% over those at a stocking density of 15 fish m^-3. The stocking density of 60 fish m^-3 is therefore taken as the optimal stocking rate for economical production of estuary groupers in floating net-cages. The estuary groupers take 7 to 8 months to grow from 15–16 g to marketable size (>500 g) at a stocking density of 15 fish m^-3, 8 to 9 months at a stocking density of 30 to 60 fish m^-3 and 11 to 12 months at 90 to 120 fish m^-3.     

Temperature-dependent ranges of coexistence in a model of a two-prey-one-predator microbial food web

The objective of our study was to analyze the effects of temperature on the population dynamics of a three-species food web consisting of two prey bacteria (Pedobacter sp. and Acinetobacter johnsonii) and a protozoan predator (Tetrahymena pyriformis) as model organisms. We assessed the effects of temperature on the growth rates of all three species with the objective of developing a model with four differential equations based on the experimental data. The following hypotheses were tested at a theoretical level: Firstly, temperature changes can affect the dynamic behavior of a system by temperature-dependent parameters and interactions and secondly, food web response to temperature cannot be derived from the single species temperature response. The main outcome of the study is that temperature changes affect the parameter range where coexistence is possible within all three species. This has significant consequences on our ideas regarding the evaluation of effects of global warming.     

Adaptive behaviour of chironomid larvae (Chironomus riparius) in response to chemical stimuli from predators and resource density

Chemical-mediated effects of predatory fish on chironomid larvae behaviour have been ignored so far. Sediment-dwelling chironomid larvae inhabit protective burrows from which they extend their bodies only to feed on deposited detritus and microalgae from the surrounding sediment. Here, we performed factorial laboratory experiments to study whether fish-borne chemical cues (kairomones) are responsible for behavioural trait changes of chironomid larvae, and whether chironomid larvae are able to assess the densities of fish predators and food resources and the trade-off between them. We exposed naïve Chironomus riparius larvae to the chemical presence of zero, one, and ten predator fish (Rutilus rutilus) and offered two resource levels (low food, high food) for each treatment. Kairomones induced significant inherent behavioural trait changes in chironomid larvae. During the first 120 min after exposing chironomids to fish-conditioned water, we found a significant increase in digging activity with increasing predator density. After 3 days of exposure, the deepest chironomid burrows were found in treatments with the highest predator density. Chironomid larvae were significantly able to adjust their foraging behaviour to different predator densities and food concentrations and trade off between them; that is, when fish predators were more abundant or when more food resources were available, the foraging activities of larvae were significantly reduced. Our data suggest that chemically mediated trait changes (burrowing and foraging behaviour) may cascade through the littoral food web.     

Adaptive peaks and alternative foraging tactics in brook charr: evidence of short-term divergent selection for sitting-and-waiting and actively searching

Some recently emerged brook charr (Salvelinus fontinalis) inhabiting still-water pools along the sides of streams are sedentary and eat crustaceans from the lower portion of the water column. Others are more active and eat insects from the upper portion of the water column. We provide evidence that this divergent foraging behavior reflects short-term divergent selection brought about by intraspecific competition in the presence of alternative food sources. Rates of encounters and interactions between individuals were density dependent, and encounter and interaction events were closely timed with prey capture attempts. In addition, aggressive fish made more foraging attempts per minute than nonaggressive fish. Aggressive fish were also either inactive or very active, while nonaggressive fish exhibited intermediate levels of activity. Growth rate potential, an important component of fitness during the early life stages of brook charr, was assessed using tissue concentrations of RNA and found to be highest for sedentary fish and for active fish making frequent foraging attempts, and lower for fish exhibiting intermediate levels of activity. Our findings support contentions that individual behavior plays an important role during initial steps in the evolution of resource polymorphisms. Received: 27 July 1998 / Accepted after revision: 16 November 1998