Pre-encounter versus post-encounter inducible defenses in predator–prey model systems

It has been reported that, in order to reduce mortality, prey are able to change their phenotype in response to cues released from predators. These short-time responses constitute effective antipredator strategies in variable environments, and involve changes in morphology, behavior, physiology or life-history traits of prey individuals belonging to a wide spectrum of taxa. Defenses can be classified into pre-encounter and post-encounter, depending on the phase of the predation process in which they take place. Also, inducible defenses should be costly.Despite the current knowledge of inducible defenses at the individual level, our understanding of their dynamic consequences at the population and community level is limited. In this work we construct and analyze numerically a predator–prey system, parameterized from published experimental data, in which prey exhibit inducible defenses of the type pre-encounter (affecting attack rate) or post-encounter (affecting handling time) and entailing either metabolic or feeding costs. The above assumptions were analyzed over a gradient of resource availability.Our results indicated that both types of cost have a similar effect on the dynamics of the model system, but we expect that different costs will produce different outcomes in a more complex model community. Conversely, pre-encounter and post-encounter IDs define domains of attraction with different size and shape within the studied sections of the multidimensional parameter space. Roughly speaking, post-encounter IDs determine a more rich dynamics when plausible parameter values are chosen, and the effect of resource density is different if the ID is handling-time based or attack-rate based. In agreement with previous works, our analyses indicate that IDs can damp population oscillations and prevent the paradox of enrichment.     

Protected-Area Monitoring Dilemmas: a New Tool to Assess Success

Abstract:  Understanding factors that influence the success of protected areas in curbing unsustainable resource consumption is essential for determining best management strategies and allocating limited resources to those projects most likely to succeed. I used a law-enforcement and monitoring game-theory model from the political science literature to identify three key variables useful in predicting the success of a protected area: costs of monitoring for rule breakers, benefits of catching a rule breaker, and probability of catching a rule breaker if monitoring. Although assigning exact values for each of these variables was difficult, the variables had a strong predictive capacity even when coded as coarse ordinal values. A model in which such values were used correctly predicted the outcome of 88 of 116 protected areas sampled from the peer-reviewed literature. The model identified a critical zone of common mismatch between protected-area circumstances and management policies. In situations where the costs of monitoring were greater than the product of the probability of catching a rule breaker and the benefit of doing so, conservation was unlikely to succeed. Control of illegal use of protected resources was reported in only 8% of such cases, regardless of strategies to motivate potential users to cooperate with conservation. My model does not prescribe a best management policy for conserving natural resources; rather, it can be used as a tool to help predict whether a proposed management policy will likely succeed in a given situation.     

Emergy analysis applied to the estimation of the recovery of costs for water services under the European Water Framework Directive

In this paper, the European Union's Water Framework Directive 2000/60/EC (WFD) that is intended to foster protection of water resources is examined, focusing on the improvement of ecological and chemical quality of surface and groundwater. The WFD includes the concept of full cost recovery (FCR) in accordance with the Polluter-Pays Principle, as one of the tools of an adequate and sustainable water resource management system. The WFD defines three different costs associated with water: resource costs (RC), financial costs (FC), and environmental costs (ECs).The FCR of water is examined from a biophysical perspective using emergy evaluation to: (1) establish resource values of water from different sources, (2) establish the full economic costs associated with supplying water, and (3) the societal costs of water that is used incorrectly; from which the resource costs, financial costs, and environmental costs, respectively, can be computed. Financial costs are the costs associated with providing water including energy, materials, labor and infrastructure. The emergy based monetary values vary between 0.15 and 1.73 €/m³ depending on technology. The emergy based, global average resource value (from which resource costs can be computed) is derived from two aspects of water: its chemical potential and its geopotential. The chemical potential monetary value of different sources such as rain, groundwater, and surface water derived from global averages of emergy inputs varies from 0.03 to 0.18 €/m³, depending on source, and the geopotential values vary from 0.03 to 2.40 €/m³, depending on location in the watershed. The environmental costs of water were averaged for the county of Spain and were 1.42 €/m³.Time of year and spatial location within the watershed ultimately influence the resource costs (computed from emergy value of chemical potential and geopotential energy) of water. To demonstrate this spatial and temporal variability, a case study is presented using the Foix watershed in northeastern Spain. Throughout the year, the resource value of water varies from 0.21 to 3.17 €/m³, depending on location within the watershed. It is concluded that FCR would benefit from the evaluation of resource costs using spatially and temporally explicit emergy accounting.     

Reputation is valuable within and outside one’s own social group

To find conditions under which humans cooperate within groups of unrelated individuals has been of major interest in the behavioral sciences. The experimental paradigm for studying potential cooperation in social dilemmas is the public goods game. Here humans regularly fail to sustain a public resource cooperatively. However, the need to maintain good reputation for other social interactions, such as indirect reciprocity, has been identified as an effective mechanism to sustain cooperation in public goods situations. As a side effect of building a good reputation through cooperative actions, an individual provides direct benefits to members of his/her own social group. These benefits could be an incentive to reward a good reputation of group members. Here we show experimentally that building a good reputation through cooperative behavior in a public goods situation is rewarded in future social interactions, not only within ones own social group but also, at a similar level, in other social groups: humans regard cooperative behavior of others as an honest signal irrespective of past direct personal benefits. Reputation gained within as well as outside ones own social group can be a driving force for selfish individuals to cooperate in public goods situations, and thereby sustain any public resource.Communicated by T. Czeschlik     

Seasonal decline in reproductive performance varies with colony size in the fairy martin, <Emphasis Type="Italic">Petrochelidon ariel</Emphasis>

Reproductive success within populations often varies with the timing of breeding, typically declining over the season. This variation is usually attributed to seasonal changes in resource availability and/or differences in the quality or experience of breeders. In colonial species, the timing of breeding may be of particular importance because the costs and benefits of colonial breeding are likely to vary over the season and also with colony size. In this study, we examine the relationship between timing of breeding and reproductive performance (clutch size and nest success) both within and between variable sized colonies (n = 18) of fairy martins, Petrochelidon ariel. In four of these colonies, we also experimentally delayed laying in selected nests to disentangle the effects of laying date and individual quality/experience on reproductive success. Within colonies, later laying birds produced smaller clutches, but only in larger colonies. The general seasonal decline in nest success was also more pronounced in larger colonies. Late laying birds were generally smaller than earlier laying birds, but morphological differences were also related to colony size, suggesting optimal colony size also varies with phenotype. Experimentally delayed clutches were larger than concurrently produced non-delayed clutches, but only in larger colonies. Similarly, delayed clutches were more likely to produce fledglings, particularly later in the season and in larger colonies. We suggest that the reduced performance of late breeding pairs in larger colonies resulted primarily from inexperienced/low quality birds preferring to settle in larger colonies, possibly exacerbated by an increase in the costs of coloniality (e.g., resource depletion and ectoparasite infestations) with date and colony size. These findings highlight the importance of phenotype-related differences in settlement decisions and reproductive performance to an improved understanding of colonial breeding and variation in colony size.     

Risk-averse foraging by bananaquits on negative energy budgets

Summary Adult bananaquits on negative energy budgets were presented with a patch containing two flower types with identical mean rewards, but different variances. The flower patch contained a random array of 85 yellow and 85 red artificial flowers. Flowers of one color were filled with the same quantity of nectar (constant flowers); flowers of the other color were filled with variable quantities of nectar (variable flowers). In the first series of experiments the birds were given three presentations, followed by three more presentations with the flower colors reversed, to control for color preferences. Some individuals were occasionally indifferent during a presentation, but overall the birds significantly preferred the constant flowers. In the second series of experiments two birds were give five presentations of the floral patch during a day at a rate less than minimally required to meet all 24-h energy costs. In all experiments, bananaquits on negative energy budgets were either indifferent or risk-averse, but never risk-prone. The absence of risk-prone foraging might be attributed to resource dispersion pattern, reward skew, or a species characteristic.     

Assessing the benefits of cooperation in honeybee foraging: search costs,forage quality,and competitive ability

Summary The foragers in honeybee colonies cooperate by sharing information about rich sources of food. This study examines three hypotheses about the benefits of this cooperation: (H1) it decreases foragers' costs in finding new food sources, (H2) it increases the quality of the food sources located by foragers, and (H3) it increases the ability of a colony's foragers to compete for high-quality food sources. To test each hypothesis, we identified a critical pattern in the foraging process which, if observed, would cast doubt on that hypothesis, and then gathered data to check for these patterns. Our observations do not support the first hypothesis, but do support the second and third. These results, in addition to helping us understand the functional significance of the honeybee's dance language, provide insights into the colonial organization of foraging by honeybees.     

Different habitats, different habits? Response to foraging information in the parasitic wasp Venturia canescens

The parasitic wasp, Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), has two reproductive modes, namely, thelytoky or arrhenotoky, and occurs in habitats with highly variable or relatively stable host abundances, respectively. Since information processing is costly, we expected that information indicating resource availability would be mainly used by the thelytokous wasps and less so by the arrhenotokous type. This idea was explored by two different approaches. In a study on patch-time allocation, we used females from ten populations and measured patch-residence times of individuals that visited multiple patches at different encounter rates. In a more detailed approach, thelytokous and arrhenotokous females from a single location were observed continuously while foraging, and all behaviors were recorded. Wasps of both reproductive modes (i.e., both habitat types) used information for the assessment of habitat quality. However, the way that the information was used differed between them. Whereas thelytokous females used foraging information to maximize their efficiency at high patch-encounter rates, arrhenotokous females merely reduced the number of offspring produced without changing patch times. The behavior of the arrhenotokous females should result in a spreading of offspring across the habitat and, thus, reduced sib-mating. The foraging strategy of these wasps might therefore be an adaptation to reduce costs associated with inbreeding.     

Ecological constraints on group size: an analysis of spider monkey and chimpanzee subgroups

The social organization of spider monkeys (Ateles geoffroyi) and chimpanzees (Pan troglodytes) appear remarkably similar. In this paper, field studies of these two species were used to (1) test a model of ecological constraints on animal group size which suggests that group size is a function of travel costs and (2) assess ecological and social factors underlying the social organization of these two species. Spider monkeys were studied over a 6-year period in Santa Rosa National Park, Costa Rica, and chimpanzees were studied for 6 years in Kibale National Park, Uganda. Adults of both species spent their time in small subgroups that frequently changed size and composition. Thus, unlike most primate species, spider monkeys and chimpanzees were not always in a spatially cohesive social group; each individual had the option of associating in subgroups of a different size or composition. Both species relied on ripe fruit from trees that could be depleted through their feeding activity. However, spider monkey food resources tended to occur at higher densities, were more common, less temporally variable, and did not reach the low levels experienced by chimpanzees. Analyses of the relationship between subgroup size and the density and distribution of their food resources suggested that travel costs limit subgroup size. However, these ecological factors did not influence all age/sex classes equally. For example, the number of adult males in a subgroup was a function of food density and travel costs. However, this was not the case for female chimpanzees, suggesting that the benefits of being in a subgroup for females did not exceed the costs, even when ecological conditions appeared to minimize subgroup foraging costs. Therefore, it seems likely that social strategies influenced the relationship between food resource variables and subgroup size.     

Territorial defence and intra-pair cooperation in the carrion crow (Corvus corone)

Summary Mates of breeding crow pairs are both involved in territorial defence. They cooperate by operating simultaneously, by choosing certain spatial positions relative to one another and by encouraging one another. Evidence is presented for the view that these types of intra-pair cooperation can be looked upon as adaptations increasing the effectiveness of territorial defence. They may have been evolved under such pressures as nest robbery by, and resource competition with, conspecifics.     

Relative costs of conserving threatened species across taxonomic groups

Bias toward legally protecting and prioritizing charismatic taxonomic groups, such as mammals and birds, and against others, such as insects and plants, is well documented. However, the relative costs of conserving various taxonomic groups and the potential of these costs to interact with existing biases have been much less explored. We analyzed conservation programs across more than 2,000 species in 3 countries to investigate the costs of conserving species within taxonomic groups and how these costs might affect conservation planning. For each data set, we tested for differences in mean annual cost among taxonomic groups. For the data set from the United States, recovery plans differed in duration, so we also tested for differences in total costs among taxonomic groups. Although the costs for individual species varied widely, there were strong international consistencies. For example, mammals cost 8–26 times more on average to conserve than plants and 13–19 times more to conserve than aquatic invertebrates. On average, bird species cost 5–30 times more to conserve than plants and 6–14 times more to conserve than aquatic invertebrates. These cost differences could exacerbate unequal resource allocation among taxonomic groups such that more charismatic groups both receive more attention and require more resources, leading to neglect of other taxonomic groups.     

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.     

Modeling Recreation Demand in the Presence of Unobservable Travel Costs: Toward a Travel Price Model

An important issue in the application of travel cost models is the construction of a travel cost variable. This paper develops an econometric approach that views travel costs as an unobserved latent variable. The latent variable approach utilizes indicators to capture the role of individual travel costs in recreational demand models. The latent variables approach has at least two advantages over conventional approaches. One, the indicators can include both traditional components such as time and distance and non-traditional components such as the scenic beauty. Second, the estimation procedure results in each indicator being valued in dollar terms.     

Conservation Goals and the Relative Importance of Costs and Benefits in Reserve Selection

Abstract: Including both economic costs and biological benefits of sites in systematic reserve selection greatly increases cost‐efficiency. Nevertheless, limited funding generally forces conservation planners to choose which data to focus the most resources on; therefore, the relative importance of different types of data must be carefully assessed. We investigated the relative importance of including information about costs and benefits for 3 different commonly used conservation goals: 2 in which biological benefits were measured per site (species number and conservation value scores) and 1 in which benefits were measured on the basis of site complementarity (total species number in the reserve network). For each goal, we used site‐selection models with data on benefits only, costs only, and benefits and costs together, and we compared the efficiency of each model. Costs were more important to include than benefits for the goals in which benefits were measured per site. By contrast, for the complementarity‐based goal, benefits were more important to include. To understand this pattern, we compared the variability in benefits and in costs for each goal. By comparing the best and the worst possible selection of sites with regard to costs alone and benefits alone for each conservation goal, we introduced a simple and consistent variability measure that is applicable to all kinds of reserve‐selection situations. In our study, benefit variability depended strongly on how the conservation goal was formulated and was largest for the complementarity‐based conservation goal. We argue that from a cost‐efficiency point of view, most resources should be spent on collecting the most variable type of data for the conservation goal at hand.     

A free lunch in the commons

We derive conditions under which raising costs through a regulatory constraint or a fully expropriated tax can increase the profits arising from a common-pool resource. The basic model assumes a fixed number of identical agents with linear costs selling in a single period at an exogenous price. A necessary and sufficient condition for a cost increase to be profitable is that aggregate output from the resource be locally convex in aggregate effort. We also show that cost increases can be profitable even if price is endogenous, agents are heterogeneous, entry is costless, or agents are playing a Markov-perfect equilibrium of a dynamic game. We also discuss more general welfare implications of the result along with its relation to existing results for a Cournot oligopoly.     

Energy budgets,risk and foraging preferences in dark-eyed juncos (Junco hyemalis)

Summary Juncos' preferences for constant versus variable food rewards were tested in three series of aviary experiments. Deprivation and feeding rates were varied across the three treatments, but the mean of the variable reward equalled the constant reward in every experiment. When the birds gained energy faster than required to meet all 24-h costs, they preferred the constant reward. When the birds' energy intake was less than the minimally required rate, they preferred the variable reward. When energy intake just balanced average daily costs, the birds preferred the constant reward or were indifferent, and their response depended on the particular mean-variance combination presented.     

Difference in the net value of ecological services between natural and artificial forests in China

Land degradation is a global problem that seriously threatens human society. However, in China and elsewhere, ecological restoration still largely relies on a traditional approach that focuses only on ecological factors and ignores socioeconomic factors. To improve the effectiveness of ecological restoration and maximize its economic and ecological benefits, a more efficient approach is needed that provides support for policy development and land management and thereby promotes environmental conservation. We devised a framework for assessing the value of ecosystem services that remain after subtracting costs, such as the opportunity costs, costs of forest protection, and costs for the people who are affected by the program; that is, the net value of ecosystem services (NVES). To understand the difference between the value of a resource and the net value of the ecosystem service it provides, we used data on VES, timber sales, and afforestation costs from China's massive national afforestation programs to calculate the net value of forest ecosystem services in China. Accounting for the abovementioned costs revealed an NVES of ¥6.1 × 10¹² for forests in 2014, which was 35.9% less than the value calculated without accounting for costs. As a result, the NVES associated with afforestation was 55.9% less than the NVES of natural forests. In some regions, NVES was negative because of the huge costs of human-made plantations, high evapotranspiration rates (thus, high water opportunity costs), and low forest survival rates. To maximize the ecological benefits of conservation, it is necessary to account for as many costs as possible so that management decisions can be based on NVES, thereby helping managers choose projects that maximize both economic and ecological benefits.     

Microbial productivity in variable resource environments

The rate, timing, and quality of resource supply exert strong controls on a wide range of ecological processes. In particular, resource-mediated changes in microbial activity have the potential to alter ecosystem processes, including the production and respiration of organic matter. In this study, we used field experiments and simulation modeling to explore how aquatic heterotrophic bacteria respond to variation in resource quality (low vs. high) and resource schedule (pulse vs. press). Field experiments revealed that one-time pulse additions of resources in the form of dissolved organic carbon (DOC) caused short-lived (< or =48 h) peaks in bacterial productivity (BP), which translated into large differences across treatments: cumulative BP was twice as high in the pulse vs. press treatment under low resource quality, and five times as high under high resource quality. To gain a more mechanistic understanding of microbial productivity in variable resource environments, we constructed a mathematical model to explore the attributes of bacterial physiology and DOC supply that might explain the patterns observed in our field experiments. Model results suggest that the mobilization rate of refractory to labile carbon, an index of resource quality, was critical in determining cumulative differences in BP between pulse and press resource environments (BPPu:Pr ratios). Moreover, BPPu:Pr ratios were substantially larger when our model allowed for realistic changes in bacterial growth efficiency as a function of bacterial carbon consumption. Together, our field and modeling results imply that resource schedule is important in determining the flow of material and energy from microbes to higher trophic levels in aquatic food webs, and that the effects of resource quality are conditional upon resource schedule. An improved understanding of the effects of resource variability on microorganisms is therefore critical for predicting potential changes in ecosystem functioning in response to environmental change, such as altered DOC fluxes from terrestrial to aquatic ecosystems.     

The cost of endangered species protection: Evidence from auctions for natural resources

This paper examines the effect that endangered species regulation has on natural resource development. Specifically, we use data from competitive auctions to estimate the effect that land-use regulation protecting endangered caribou in the Canadian province of Alberta has on the price producers pay for the right to extract oil. We exploit a regression discontinuity design to evaluate how prices differ along regulation boundaries that constrain resource development. The auction format and the regulation discontinuity allow us to measure the total cost of the regulation. We find that producers pay 24% less on average for oil leases that are regulated and that the total net present value cost of the regulation exceeds $1.15 billion for leases sold between 2003 and 2012, all of which is borne by the government. In spite of these costs, the populations of endangered caribou remain in widespread decline.     

Extraction capacity and the optimal order of extraction

The least-cost-first extraction rule for deposits with different extraction costs previously has been shown to be invalid in general equilibrium. This paper demonstrates that this rule also does not hold in partial equilibrium when extraction capacity is limited. Necessary and sufficient conditions for several surprising extraction orders are presented. If extraction from a high-cost resource is constrained, it may be optimal to begin extraction from a high-cost deposit (or backstop) strictly before extracting from a lower-cost deposit. If extraction from a low-cost resource is limited, it may be optimal to exhaust a high-cost deposit strictly before the low-cost deposit is exhausted or to abandon extraction temporarily from a high-cost deposit and then to exhaust it later. The analysis demonstrates how extraction constraints affect the order of extraction and shows that certain cost reversals are caused by limited extraction capacity rather than by the general equilibrium definition of extraction costs.