Thermoregulatory cost affects territorial behavior in hummingbirds: a model and its application

A common assumption in behavioral ecology is that the valuation of a resource by consumers depends on the energetic value of the resource itself. Nevertheless, the value of a resource may be relative to the condition of the organism, which is in turn related to the abiotic conditions such as ambient temperature. We developed a theoretical model—incorporating these untested assumptions—to predict a functional relationship between territorial aggression and ambient temperature for individuals sensitive to daily variations in energy availability. We evaluated our theoretical predictions against a field experiment carried out with the hummingbird Sephanoides sephaniodes. The model predicted a quadratic relation between aggression intensity and ambient temperature. Field data were better explained by a quadratic equation than a linear function, suggesting the existence of lower and upper thresholds of temperature which determine the intensity of territorial defense. Ambient temperature affects energy expenditure for thermoregulation, and therefore, it fixes the benefit level that must be produced by the territory to pay the costs of its defense. Our findings strongly suggest that abiotic conditions can change an animal evaluation of the yield of a resource and in turn influence the behavioral strategy which it adopts.     

Dissecting the tragedy: A spatial model of behavior in the commons

Much of the discussion of the tragedy of the commons focuses on aggregate impacts, often in data-poor developing country settings. Few non-experimental empirical studies shed light on contextual circumstances driving the extent of rent dissipation and overexploitation. We utilize a high-resolution data set to estimate a behavioral model of fishermen's spatial choices. A unique policy setting allows us to measure the degree to which individual fishermen's choices appear aimed at mitigating the tragedy of the commons in a small numbers setting. We find evidence of partial mitigation in excess of what we would expect under pure self-interest but short of what would occur under group-maximizing behavior. We also examine how contextual factors in the fishery shape the degree of cooperation within the fishing season and find evidence that competition for the common pool resource distorts fishermen's implicit cost of distance—creating a form of “common property inertia”.     

A robust-design formulation of the incidence function model of metapopulation dynamics applied to two species of rails

The incidence function model (IFM) uses area and connectivity to predict metapopulation dynamics. However, false absences and missing data can lead to underestimates of the number of sites contributing to connectivity, resulting in overestimates of dispersal ability and turnovers (extinctions plus colonizations). We extend estimation methods for the IFM by using a hierarchical Bayesian model to account both for false absences due to imperfect detection and for missing data due to sites not surveyed in some years. We compare parameter estimates, measures of metapopulation dynamics, and forecasts using stochastic patch occupancy models (SPOMs) among three IFM models: (1) a Bayesian formulation assuming no false absences and omitting site-year combinations with missing data; (2) a hierarchical Bayesian formulation assuming no false absences but incorporating missing data; and (3) a hierarchical Bayesian formulation allowing for imperfect detection and incorporating missing data. We fit the models to multiyear data sets of occupancy for two bird species that differ in body size and presumed dispersal ability but inhabit the same network of sites: the small Black Rail (Laterallus jamaicensis) and the medium-sized Virginia Rail (Rallus limicola). Incorporating missing data affected colonization parameters and led to lower estimates of dispersal ability for the Black Rail. Detection rates were high for the Black Rail in most years but moderate for the Virginia Rail. Incorporating imperfect detection resulted in higher occupancy and lower turnover rates for both species, with largest effects for the Virginia Rail. Forecasts using SPOMs were sensitive to both missing data and false absences; persistence in models assuming no false absences was more optimistic than from robust models. Our results suggest that incorporating false absences and missing data into the IFM can improve (1) estimates of dispersal ability and the effect of connectivity on colonization, (2) the scaling of extinction risk with patch area, and (3) forecasts of occupancy and turnover rates.     

A kinematic model of the city of Miami ocean outfall plume behavior

A kinematic model is developed to predict the movement of effluent from the City of Miami ocean outfall. The outfall terminates at the 5 m isobath in the near vicinity of three tidal inlets opening into Biscayne Bay. Due to the shallow, uncomplicated topography and oscillatory coastal currents a linear bottom friction law is valid, which justifies the use of a potential flow model. The inlets are modeled as sinks and the outfall as a source discharging into a uniform coastal current. The model predicts that effluent will enter Biscayne Bay in approximately 3 h during flood tide for coastal current headings within an arc of 210° to 40°. Dye-tracking experiments were in good agreement with the model results. Local wind records suggest that sewage effluent enters Biscayne Bay on about 80% of the flood tides, or roughly 11 times per week.     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997     

A simple individual model for simulating weight gain dynamics in response to intake rate and daily behavior

It is often necessary to estimate the weight that an individual may be capable of gaining depending on its degree of activity. A simple individual-based model was developed for studying the dynamics of weight in terms of daily behavior and ingestion rate. It was based on the balance between the individual's energy intake and the cost of its daily activities. Costs depend on the weight of the individual and the photoperiod, as well as on the time spent on each activity. Different combinations of ingestion rate, individual's weight, photoperiod length, and time assigned to different activities were used for simulating the weight dynamics, taking the species Rhea americana as a study case. Estimations of energetic costs of the activities were obtained from specialized literature. Using different photoperiods and individual behaviors, the model yields field metabolic rate (FMR) values in agreement with those obtained from direct measurements for other omnivorous bird species.     

Effect of sperm depletion and starvation on female mating behavior in the water strider,Aquarius remigis

The resolution of intersexual conflict over mating should be dependent on the current state of each individual. In this study, I used a factorial design to examine the influence of two physiological factors, sperm depletion and food deprivation, on resistance to mating by females of the water strider, Aquarius remigis. Females employ several different mate-resisting tactics during an encounter with a male. Five measures of female resistance to mating were identified: jumping, rolling, dunking, time spent dunking, and struggle duration. Jump, roll, and dunk rates were highly correlated with each other and combined into one metric of resistance to mating (PC1) using principal components analysis. Time per dunk (T/D) and struggle duration were also analyzed. Discrete male behaviors during the struggle could not be identified. Two measures of female resistance, PC1 and T/D, were significantly lower in sperm-depleted females than in sperm-replenished females. Struggle duration did not differ between the two treatments. Starvation had no effect on any of the measures of resistance. Sperm depletion significantly enhanced the probability of mating (54% vs. 24% for replenished females), while starvation had no effect on the probability of mating. I pooled all the females and compared females that mated with those that did not mate. Nonmating females resisted significantly more than mating females in all three measures of resistance. Path analysis indicated that PC1 was the only measure of resistance that was significantly negatively related to the probability of mating. Almost half (46%) of sperm-depleted females showed no resistance to males, while only 3% of sperm-replenished females were nonresistant. When nonresisters were removed from the analysis, sperm depletion had no effect on any of the measures of female resistance to mating and no effect on the probability of mating. In A. remigis, female resistance appears to be a yes/no phenomenon with respect to sperm depletion and not affected directly by starvation. Received: 2 September 1994/Accepted after revision: 9 September 1995     

Application of the simple charge-discharge metabolic model to transient behavior of experimental marine microcosms

A simple charge-discharge circuit model conventionally used to simulate steady-state system metabolism was extended to simulate metabolic responses to perturbations in energy flow rates in balanced benthic marine laboratory microcosms. Empirical determination of the metabolic transfer coefficients using data collected with diel metabolic experiments indicated that these transfer coefficients are a property of the system and not of the environmental conditions to which the system had been acclimated.Used to simulate steady-state metabolic conditions based on various levels of light-energy input and temperature, the model gave values for integrated production and respiration that closely approximated experimental determinations. Moreover, the diel metabolic patterns produced were similar to those observed in the laboratory systems. The model's dynamic behavior upon loss of energy input was similar to that observed in the experimental systems for several days, but diverged after that period. Daytime respiration values were similar in magnitude but different in overall pattern from those observed in the microcosms.Simulations of a series of energy flow rate perturbation experiments produced response curves which were virtually indentical to final steady-state metabolic levels observed in the experiments, and which exhibited initial metabolic transients of the same pattern that had occurred in the laboratory. However, the transient response of the model was of greater amplitude and more highly damped.     

Cycles in natural-resource commodity prices: An analysis of the frequency domain

A model is derived that relates price cycles to investment behavior in the mineral industries and predicts that these cycles will be roughly twice as long as the investment period (the period between the investment decision and the project completion). Spectral techniques are used to estimate the principal cycles in metal and fuel prices. For the metals, the estimated cycle lengths reinforce the model predictions. However, no significant cycles are found in the prices of the fuels. The lead-lag relationships between prices of different commodities are also investigated and are found to be significant for those that are jointly produced.     

Trends in natural-resource commodity prices: An analysis of the time domain

This paper attempts to reconcile the theoretical predictions of increasing real prices for nonrenewable natural-resource commodities obtained from Hotelling-style models with the empirical findings of falling prices for these commodities. A theoretical model for relative-price movements is derived for the case of exogenous technical change and endogenous change in the grade of ores mined. The model suggests a U-shaped time path for relative prices. The implied price movements are tested for all the major metals and fuels and the model parameters are found to be statistically significant for 11 out of the 12 commodities tested.     

Correlation analysis as a tool to investigate the bioaccessibility of nickel, vanadium and zinc in Northern Ireland soils

Correlation analyses were conducted on nickel (Ni), vanadium (V) and zinc (Zn) oral bioaccessible fractions (BAFs) and selected geochemistry parameters to identify specific controls exerted over trace element bioaccessibility. BAFs were determined by previous research using the unified BARGE method. Total trace element concentrations and soil geochemical parameters were analysed as part of the Geological Survey of Northern Ireland Tellus Project. Correlation analysis included Ni, V and Zn BAFs against their total concentrations, pH, estimated soil organic carbon (SOC) and a further eight element oxides. BAF data were divided into three separate generic bedrock classifications of basalt, lithic arenite and mudstone prior to analysis, resulting in an increase in average correlation coefficients between BAFs and geochemical parameters. Sulphur trioxide and SOC, spatially correlated with upland peat soils, exhibited significant positive correlations with all BAFs in gastric and gastro-intestinal digestion phases, with such effects being strongest in the lithic arenite bedrock group. Significant negative relationships with bioaccessible Ni, V and Zn and their associated total concentrations were observed for the basalt group. Major element oxides were associated with reduced oral trace element bioaccessibility, with Al₂O₃ resulting in the highest number of significant negative correlations followed by Fe₂O₃. spatial mapping showed that metal oxides were present at reduced levels in peat soils. The findings illustrate how specific geology and soil geochemistry exert controls over trace element bioaccessibility, with soil chemical factors having a stronger influence on BAF results than relative geogenic abundance. In general, higher Ni, V and Zn bioaccessibility is expected in peat soil types.     

Influence of support on catalytic behavior of nickel catalysts in the steam reforming of ethanol for hydrogen production

Al₂O₃, MgO, SiO₂ and ZnO-supported nickel catalysts were prepared and evaluated in the ethanol steam reforming for hydrogen production. It is shown that the catalytic behavior can be influenced depending on the experimental conditions employed and chemical composition of the catalyst.     

A linked model of animal ecology and human behavior for the management of wildlife tourism

Wildlife tourism attractions are characterized as having intricately coupled human-wildlife interactions. Accordingly, the ability to mitigate negative impacts of tourism on wildlife necessitates research into the ecology of the system and of the human dimensions, since plans aimed at optimizing wildlife fitness must also be acceptable to tourists. We developed an integrated systems dynamics model for the management of tourist-stingray interactions at ‘Stingray City Sandbar’ (SCS), Cayman Islands. The model predicts the state of the tourism attraction over time in relation to stingray population size, stingray life expectancy, and tourist visitation under various management scenarios. Stingray population data in the model comprised growth rates and survival estimates (from mark-and-recapture data) and mortality estimates. Inputted changes in their respective rates under different management scenarios were informed by previous research. Original research on the demand of heterogeneous tourist segments for management regulations via a stated choice model was used to calculate changes in the tourist population growth rate from data supplied by the Caymanian government. The management attributes to which tourists were responsive also have anticipated effects on stingray ecology (migration and mortality), and vice versa, thus linking the two components. We found that the model's predictions over a 25-year time span were sensitive to the stingray population growth rate and alternate management options. Under certain management scenarios, it was possible to maximize both the tourist segment in favor of no management and stingray numbers while reducing stingray health. However, the most effective relative strategy included a reduction in visitor density, restricted stingray interactions, and an imposition of a small fee. Over time, although fewer stingrays were predicted to remain at SCS, they would live longer and experience fewer stochastic disease events, and the desirable tourist segment was predicted to predominate. By understanding how management will affect tourist activities and their subsequent impacts on both wildlife health and visitor satisfaction, one can explore the management alternatives that would optimize both.     

Towards an ecological solution to the folivore paradox: patch depletion as an indicator of within-group scramble competition in red colobus monkeys (Piliocolobus tephrosceles)

A number of socioecological models assume that within-group food competition is either weak or absent among folivorous primates. This assumption is made because their food resources are presumed to be superabundant and evenly dispersed. However, recent evidence increasingly suggests that folivore group size is food-limited, that the primates prefer patchily distributed high-quality foods, and display some of the expected responses to within-group scramble competition. To investigate this apparent contradiction between theoretical models and recent empirical data, we examined the foraging behaviour of red colobus monkeys (Piliocolobus tephrosceles) in Kibale National Park, Uganda. We found that red colobus monkeys foraged in a manner that suggests they deplete patches of preferred foods: intake rate slowed significantly during patch occupancy while movement rate, an index of foraging effort, increased. Furthermore, patch occupancy was related to the size of the feeding group and the size of the patch. These results suggest that within-group scramble competition occurs, may limit folivore group size, and should be considered in models of folivore behavioural ecology.     

An econometric model of the U.S. secondary copper industry: Recycling versus disposal

In this paper, a theoretical model of secondary recovery is developed that integrates microeconomic theories of production and cost with a dynamic model of scrap generation and accumulation. The model equations are estimated for the U.S. secondary copper industry and used to assess the impacts that various policies and future events have on copper recycling rates. The alternatives considered are: subsidies for secondary production, differing energy costs, and varying ore quality in primary production.     

Diversity of swimming behavior in hydromedusae as related to feeding and utilization of space

Feeding behaviors of the following 4 species of hydromedusae are described from field and laboratory observations: Probosidactyla flavicirrata, Stomotoca atra, Phialidium gregarium and Polyorchis penicillatus. Feeding efficiency of medusae has previously been considered equivalent to fishing with a given amount (combined tentacle length) of adhesive fishing line; however, detailed observation shows that behavior of medusae greatly modifies the fishing capacity of each species. It is hypothesized that in addition to (1) tentacle number and length, the following factors strongly influence feeding efficiency: (2) tentacle posture, (3) velocity of tentacles moving through water (4) swimming pattern of medusa, (5) streamlining effects of medusa bell on water flow, (6) diameter of prey, (7) swimming pattern and velocity of prey. Each species of hydromedusa utilizes the above factors in different combinations.Mailing address     

A model for energetics and bioaccumulation in marine mammals with applications to the right whale.

We present a dynamic energy budget (DEB) model for marine mammals, coupled with a pharmacokinetic model of a lipophilic persistent toxicant. Inputs to the model are energy availability and lipid-normalized toxicant concentration in the environment. The model predicts individual growth, reproduction, bioaccumulation, and transfer of energy and toxicant from mothers to their young. We estimated all model parameters for the right whale; with these parameters, reduction in energy availability increases the age at first parturition, increases intervals between reproductive events, reduces the organisms' ability to buffer seasonal fluctuations, and increases its susceptibility to temporal shifts in the seasonal peak of energy availability. Reduction in energy intake increases bioaccumulation and the amount of toxicant transferred from mother to each offspring. With high energy availability, the toxicant load of offspring decreases with birth order. Contrary to expectations, this ordering may be reversed with lower energy availability. Although demonstrated with parameters for the right whale, these relationships between energy intake and energetics and pharmacokinetics of organisms are likely to be much more general. Results specific to right whales include energy assimilation estimates for the North Atlantic and southern right whale, influences of history of energy availability on reproduction, and a relationship between ages at first parturition and calving intervals. Our model provides a platform for further analyses of both individual and population responses of marine mammals to pollution, and to changes in energy availability, including those likely to arise through climate change.