Female preferences for multiple attributes in the acoustic signals of the Italian treefrog, <Emphasis Type="Italic">Hyla intermedia</Emphasis>

The advertisement call of frogs and toads is an example of multiple message signal because different acoustic properties encode different kinds of biologically significant information. In the Italian treefrog, Hyla intermedia, pulse rate and frequency have been found to be under stabilizing female preferences and to encode information important for mate recognition, whereas the number of calls per call group have been found to be under directional preferences and, thus, to be important for mate quality assessment. In this study, we investigate preferences for calls that differ simultaneously in frequency, pulse rate, and number of calls per call group, and we ask how these properties interact with each other in influencing female mating decisions. Results of two-choice phonotaxis experiments provide no evidence to support the hypothesis that females process multi-attribute signals in a hierarchical way. In contrast, the pattern of preferences is consistent with the ‘preference function’ hypothesis, that is, with the hypothesis that females rank signals along an ordinal scale of values and choose accordingly. Pulse rate and frequency influence mating preferences more than does the number of calls per call group. The interaction between pulse rate and frequency is not additive but multiplicative: small differences in either pulse rate or frequency that, alone, have no effects on female choice, interact synergistically so that their combination has strong influence on female preferences. A preference repeatability test shows strong among-female differences in preference for multi-attribute signals. We suggest that this result reveals not only a variation in attribute values among females, but also a variation in the way females weight and combine attribute values into a single preference score.     

Shell preference in a hermit crab: comparison between a matrix of paired comparisons and a multiple-alternative experiment

The preference of the hermit crab, Calcinus californiensis, among six species of shells, was tested by two different experiments. The first experiment used pair-wise trials, analyzing the preference by Chi-square tests using two different constructions of the null hypothesis. One hypothesis was based on a no-preference among shell species, the second on comparing the number of crabs changing for a particular shell species when two options were given versus the changing when no options were offered. The second experiment was a multiple-alternative test based on a rank ordering of the shell preference. This method has both statistical and resource-saving advantages over the traditional pair-wise comparisons. The sequence of shell preference was similarly independent of the procedure used. The preferred shell species are heavy and might be associated with hydrodynamic advantages and with the protection against predation. The shell preference matches with the pattern of shell occupancy indicating that the shell use in nature is determined by the crab’s preference. The information generated may be used for further research on shell preference as a methodological alternative.     

Habitat preference increases territorial defence in brown trout (Salmo trutta)

The striking ability of territory owners to repel intruders has generated a number of theoretical explanations as well as experimental studies in many animal species. However, effects of individual habitat preferences on territorial defence have rarely been studied. From the territory value hypothesis, we predicted that owners of preferred habitats should invest more resources in defence than owners of non-preferred habitats. We tested this prediction with young territorial brown trout in a two-stage experiment. First, trout were allowed to choose individually between gravel and a uniform bright substrate. As expected, they showed a significant (79%) preference for gravel. However, there was considerable variation between individuals in substrate preference, with a few fish preferring the bright substrate. Half of the tested fish were then transferred to a gravel substrate and the rest to a bright substrate, manipulating habitat type in relation to preference. Territory owners were then staged against size-matched intruders whereupon contest aggression was observed and the winner of each contest determined. Overall, owners won most of the contests. Satisfied owners won 86% and owners of less preferred territories, 74% of the contests. Furthermore, more satisfied owners attacked sooner and were more aggressive relative to the intruders. We conclude that brown trout show individual variation in habitat preference, which appears to be linked with their investment in territorial defence. These results suggest that understanding and modelling of animal contests could benefit from considering how territorial defence is influenced by individual habitat preference and specialisation. Received: 25 February 2000 / Revised: 22 May 2000 / Accepted: 25 June 2000     

Trendanalysen mit Mann-Kendall-Test und Theil-Schätzer

Goal and Scope

The influence of the measurement uncertainty on the outcomes of statistical evaluations is shown.

Methods

Examplarily using data from the Environmental Specimen Bank the influence of measurement uncertainties on results of trned analyses (Mann Kendall test and Theil estimator) was tested. Both, the simple and the double measurement uncertainty were tested, as well as the standard deviation of repeated measurements.

Results

Considering measurement uncertainty means that not points but intervals are used. A value y_j is then considered larger [smaller] than y_i if applies: y_j-u(y_j>y_i+u(y_i) [y_j+u(y_j)<y_i-u(y_j)], where u(y_j) and u(y_i) are measurement uncertainties of y_j and y_i. Mann Kendall test of trend will react more conservatively (in sense of the maintenance of null hypothesis: no trend), when measurement uncertainties are considered. Thus, the resulting trend statements will be safer.

Outlook

For future application a decision is neccessary about the kind of the measurement uncertainty to be used (none, simple, double, standard deviation). Part 2 will deal with weighting of the data.     

The evolution of food sharing in primates

The aim of this study is to explain the occurrence of food sharing across primates. Defined as the unresisted transfer of food, evolutionary hypotheses have to explain why possessors should relinquish food rather than keep it. While sharing with offspring can be explained by kin selection, explanations for sharing among unrelated adults are more controversial. Here we test the hypothesis that sharing occurs with social partners that have leverage over food possessors due to the opportunity for partner choice in other contexts. Thus, we predict that possessors should relinquish food to potential mates or allies, who could provide or withhold matings or coalitionary support in the future. We used phylogenetic analyses based on both maximum likelihood and Bayesian approaches in a sample of 68 primate species to test these predictions. The analyses strongly indicate that (1) sharing with offspring is predicted by the relative processing difficulty of the diet, as measured by the degree of extractive foraging, but not overall diet quality, (2) food sharing among adults only evolved in species already sharing with offspring, regardless of diet, and (3) male–female sharing co-evolved with the opportunity for female mate choice and sharing within the sexes with coalition formation. These results provide comparative support for the hypothesis that sharing is “traded” for matings and coalitionary support in the sense that these services are statistically associated and can thus be selected for. Based on this, we predict that sharing should occur in any species with opportunities for partner choice.     

An economic analysis of air pollution from coal-fired power plants

The benefits of air pollution control techniques on a coal-fired power plant are Simulated with a scientifically based environmental model. Air pollution abatement techniques are assessed in terms of their resource cost (measured in dollars) and their effectiveness in reducing environmental damage (measured in dollars and healthy days lost). Which air pollution techniques are most efficient depend upon how much a day of health should be valued. Other factors of potential interest such as uncertainty and equity are also simulated through the model. The paper demonstrates that scientific evidence can be organized around economic principles in order to develop more rational and effective environmental policies.     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

No relationship between long-distance acoustic mate attraction signals and male fertility or female preference in spring field crickets

To father offspring, a male must succeed at two processes of sexual selection: (1) mate with a female and (2) fertilize her eggs. We investigated the relationships between pre- and post-copulatory male traits and female mating responses in wild-captured and laboratory-reared spring field crickets, Gryllus veletis. The phenotype-linked fertility hypothesis suggests that females may receive a direct benefit, enhanced fertilization efficiency, by mating with males that signal attractively. We measured fine-scale components of male acoustic mate attraction signals as well as how much time males spent signalling, measured female preference for males in mating trials and then quantified sperm number and viability. We found no relationship between male signalling traits and male fertility or female preference, providing no evidence for the phenotype-linked fertility hypothesis. We also found no difference in sperm metrics between wild-captured and laboratory-reared males. While female crickets may receive benefits by choosing males based on acoustic signal characteristics, whether the benefits are a result of genetic quality, seminal fluid contents or some other male trait remains unknown.     

Area and mammalian elevational diversity

Elevational gradients hold enormous potential for understanding general properties of biodiversity. Like latitudinal gradients, the hypotheses for diversity patterns can be grouped into historical explanations, climatic drivers, and spatial hypotheses. The spatial hypotheses include the species-area effect and spatial constraint (mid-domain effect null models). I test these two spatial hypotheses using regional diversity patterns for mammals (non-volant small mammals and bats) along 34 elevational gradients spanning 24.4 degrees S-40.4 degrees N latitude. There was high variability in the fit to the species-area hypothesis and the mid-domain effect. Both hypotheses can be eliminated as primary drivers of elevational diversity. Area and spatial constraint both represent sources of error rather than mechanisms underlying these mammalian diversity patterns. Similar results are expected for other vertebrate taxa, plants, and invertebrates since they show comparable distributions of elevational diversity patterns to mammalian patterns.     

Herbivory-mediated pollinator limitation: negative impacts of induced volatiles on plant-pollinator interactions

Although induced plant responses to herbivory are well studied as mechanisms of resistance, how induction shapes community interactions and ultimately plant fitness is still relatively unknown. Using a wild tomato, Solanum peruvianum, native to the Peruvian Andes, we evaluated the disruption of pollination as a potential ecological cost of induced responses. More specifically, we tested the hypothesis that metabolic changes in herbivore-attacked plants, such as the herbivore-induced emission of volatile organic compounds (VOCs), alter pollinator behavior and consequentially affect plant fitness. We conducted a series of manipulative field experiments to evaluate the role of herbivore-induced vegetative and floral VOC emissions as mechanisms by which herbivory affects pollinator behavior. In field surveys and bioassays in the plants' native habitat, we found that real and simulated herbivory (methyl jasmonate application) reduced attractiveness of S. peruvianum flowers to their native pollinators. We show that reduced pollinator preference, not resource limitation due to leaf tissue removal, resulted in reduced seed set. Solitary bee pollinators use floral plant volatiles, emitted in response to herbivory or methyl jasmonate treatment, as cues to avoid inflorescences on damaged plants. This herbivory-induced pollinator limitation can be viewed as a general cost of induced plant responses as well as a specific cost of herbivory-induced volatile emission.     

Reconciling competing ecological explanations for sexual segregation in ungulates

Sexual segregation in ungulates has important conservation and theoretical implications, but despite numerous studies, the impetus for this behavioral pattern remains a topic of debate. Sexual segregation hypotheses can be broadly grouped into social and ecological explanations, but only ecological explanations can adequately describe why sexes use different areas and habitats. The reproductive strategy hypothesis (RSH) and forage selection hypothesis (FSH) are leading ecological explanations, and although both have received support in the literature, neither the collective basis for that support nor overlap between these hypotheses has been adequately evaluated. This review analyzed seasonal sex comparisons of habitat forage quantity (n=66), quality (n=67), and diet (n=63) from peer-reviewed studies of north temperate ruminants to test predictions of each hypothesis. Empirical data supported predictions of the RSH, but did not support two of three key predictions of the FSH. Males used habitats with greater forage quantity significantly more than females. But, contrary to predictions of the FSH, females did not use habitats with superior forage quality nor consume higher-quality diets more than males. Sexes typically used habitats and consumed diets of similar quality, and when differences were reported, males used higher-quality habitats significantly more than females. Results refute FSH arguments that differences in dietary requirements associated with sexual dimorphism are a universal explanation for sexual segregation in ungulates, but the physiological mechanism on which the FSH is predicated may explain why males consume poorer-quality diets when high-quality forage is scarce. The FSH, therefore, operates at a proximate level because it explains diet and habitat selection by males under certain environmental conditions, but multiple environmental factors may influence sexual segregation, and no single proximate explanation adequately describes this behavioral pattern. The RSH explains sexual segregation as the evolutionary response to differences in reproductive strategies: males choose habitats to maximize energy gains in preparation for rut, and females select habitats with combinations of resources that contribute to offspring survival. Consequently, the RSH provides an ultimate explanation that can be used to explain and interpret studies of sexual segregation in ungulates.     

Accounting for Uncertainty in Making Species Protection Decisions

Abstract:  Uncertainty gives rise to two decision errors in implementing the U.S. Endangered Species Act: listing species that are not in danger of extinction and delisting species that are in danger of extinction. I evaluated four methods (minimum standard, precautionary principle, minimax regret criterion, adaptive management) for deciding whether to list or delist a species when there is uncertainty about how those decisions are likely to influence survival of the species. A safe minimum standard criterion preserves some minimum amount or safe standard (population) of a species unless maintaining that amount generates unacceptable social cost. The precautionary principle favors not delisting a species when there is insufficient evidence on the efficacy of state management plans for protecting them. A minimax regret criterion selects the delisting decision that minimizes the maximum loss likely to occur under alternative ecosystem states. When the cost of making a correct decision is less than the cost of making an incorrect decision, the minimax regret criteria indicates that delisting is the optimal decision. Active adaptive management employs statistically valid experiments to test hypotheses about the likely impacts of delisting decisions. Safe minimum standard and minimax regret criterion are not compatible with the U.S. Endangered Species Act. The precautionary principle comes closest to describing how federal agencies make delisting decisions. Active adaptive management is scientifically superior to the other methods but is costly and time consuming and may not be compatible with the U.S. National Environmental Policy Act.     

Incorporating Uncertainty of Management Costs in Sensitivity Analyses of Matrix Population Models

The importance of accounting for economic costs when making environmental‐management decisions subject to resource constraints has been increasingly recognized in recent years. In contrast, uncertainty associated with such costs has often been ignored. We developed a method, on the basis of economic theory, that accounts for the uncertainty in population‐management decisions. We considered the case where, rather than taking fixed values, model parameters are random variables that represent the situation when parameters are not precisely known. Hence, the outcome is not precisely known either. Instead of maximizing the expected outcome, we maximized the probability of obtaining an outcome above a threshold of acceptability. We derived explicit analytical expressions for the optimal allocation and its associated probability, as a function of the threshold of acceptability, where the model parameters were distributed according to normal and uniform distributions. To illustrate our approach we revisited a previous study that incorporated cost‐efficiency analyses in management decisions that were based on perturbation analyses of matrix population models. Incorporating derivations from this study into our framework, we extended the model to address potential uncertainties. We then applied these results to 2 case studies: management of a Koala (Phascolarctos cinereus) population and conservation of an olive ridley sea turtle (Lepidochelys olivacea) population. For low aspirations, that is, when the threshold of acceptability is relatively low, the optimal strategy was obtained by diversifying the allocation of funds. Conversely, for high aspirations, the budget was directed toward management actions with the highest potential effect on the population. The exact optimal allocation was sensitive to the choice of uncertainty model. Our results highlight the importance of accounting for uncertainty when making decisions and suggest that more effort should be placed on understanding the distributional characteristics of such uncertainty. Our approach provides a tool to improve decision making.     

Cost‐effective conservation of an endangered frog under uncertainty

How should managers choose among conservation options when resources are scarce and there is uncertainty regarding the effectiveness of actions? Well‐developed tools exist for prioritizing areas for one‐time and binary actions (e.g., protect vs. not protect), but methods for prioritizing incremental or ongoing actions (such as habitat creation and maintenance) remain uncommon. We devised an approach that combines metapopulation viability and cost‐effectiveness analyses to select among alternative conservation actions while accounting for uncertainty. In our study, cost‐effectiveness is the ratio between the benefit of an action and its economic cost, where benefit is the change in metapopulation viability. We applied the approach to the case of the endangered growling grass frog (Litoria raniformis), which is threatened by urban development. We extended a Bayesian model to predict metapopulation viability under 9 urbanization and management scenarios and incorporated the full probability distribution of possible outcomes for each scenario into the cost‐effectiveness analysis. This allowed us to discern between cost‐effective alternatives that were robust to uncertainty and those with a relatively high risk of failure. We found a relatively high risk of extinction following urbanization if the only action was reservation of core habitat; habitat creation actions performed better than enhancement actions; and cost‐effectiveness ranking changed depending on the consideration of uncertainty. Our results suggest that creation and maintenance of wetlands dedicated to L. raniformis is the only cost‐effective action likely to result in a sufficiently low risk of extinction. To our knowledge we are the first study to use Bayesian metapopulation viability analysis to explicitly incorporate parametric and demographic uncertainty into a cost‐effective evaluation of conservation actions. The approach offers guidance to decision makers aiming to achieve cost‐effective conservation under uncertainty.     

Optimum swimming pathways of fish spawning migrations in rivers

Fishes that swim upstream in rivers to spawn must navigate complex fluvial velocity fields to arrive at their ultimate locations. One hypothesis with substantial implications is that fish traverse pathways that minimize their energy expenditure during migration. Here we present the methodological and theoretical developments necessary to test this and similar hypotheses. First, a cost function is derived for upstream migration that relates work done by a fish to swimming drag. The energetic cost scales with the cube of a fish's relative velocity integrated along its path. By normalizing to the energy requirements of holding a position in the slowest waters at the path's origin, a cost function is derived that depends only on the physical environment and not on specifics of individual fish. Then, as an example, we demonstrate the analysis of a migration pathway of a telemetrically tracked pallid sturgeon (Scaphirhynchus albus) in the Missouri River (USA). The actual pathway cost is lower than 10(5) random paths through the surveyed reach and is consistent with the optimization hypothesis. The implication--subject to more extensive validation--is that reproductive success in managed rivers could be increased through manipulation of reservoir releases or channel morphology to increase abundance of lower-cost migration pathways.     

Male choice and sperm competition as constraints on polyandry in the red-necked phalarope Phalaropus lobatus

Summary The mating system of the red-necked phalarope Phalaropus lobatus is usually classed as polyandrous, but studies show that the majority of females are monogamous. Polyandry appears to be constrained because later arriving males do not usually nest with females that have already laid a clutch that season (C1-females), despite the sexual receptiveness of these females. Rather they nest with females that have not previously laid a clutch that season (C0-females). Five potential explanations of this constraint on polyandry were tested. Two hypotheses, invoking female-female competition, were not supported as female agonistic success appeared to be unrelated to priority of access to males, and there was no apparent difference between C0- and C1-females in their ability to find unpaired males. Two hypotheses suggest that C0-females are preferred by males either because they can initiate a clutch more rapidly or because they lay larger eggs. Neither hypothesis was supported by the available data. The data upheld a third male choice hypothesis that C1-females are discriminated against because they have copulated previously with another male and this may compromise paternity. Secondary males copulated and attempted to copulate more often when paired with C1-females, suggesting that sperm competition was greater and confidence of paternity was lower. Corroborative evidence of active male choice showed that males were more aggressive towards associating C1-females than towards C0-females, although how males discriminated between the two classes of females was not determined.     

Costing reproduction: effects of mating opportunity on mating success in male Bicyclus anynana butterflies

It is frequently assumed that males have an almost unlimited reproductive capacity, while access to receptive females is typically limiting. Consequently, sexual selection is expected to favor vigorous courtship behavior in males. If such behavior is associated with non-trivial costs, ample current mating opportunities should be accompanied by a reduction in future mating vigor. To test this hypothesis, three treatments differing in sex ratio were established using the tropical butterfly Bicyclus anynana: 50 males each were housed either with 0, 25, or 50 females. Mating trials involving the competition among males from each treatment for a single virgin female were carried out on days 3, 9, and 15 after allocation to treatments. While there was no difference on day 3, prior mating opportunity clearly reduced mating success on days 9 and 15, being lowest if identical numbers of males and females were housed together. This finding suggests accumulating costs associated with high courtship and/or mating activity. Further, older males were more active and initiated copulation earlier than the younger ones, consistent with the residual reproductive value hypothesis. We found no evidence for a survival or fertility cost of mating.     

Non-renewable resource prices: Deterministic or stochastic trends?

In this paper, we examine temporal properties of 11 natural resource real price series from 1870 to 1990. Recent studies by Ahrens and Sharma [Trends in natural resource commodity prices: deterministic or stochastic? J. Environ. Econom. Manage. 33(1997)59–74], Berck and Roberts [Natural resource prices: will they ever turn up? J. Environ. Econom. Manage. 31(1996)65–78], and Slade [Grade selection under uncertainty: least cost last and other anomalies, J. Environ. Econom. Manage. 15(1988)189–205], among others, find that many non-renewable resource prices have a stochastic trend. We revisit this issue by employing a Lagrangian multiplier unit root test that allows for two endogenously determined structural breaks with and without a quadratic trend. Contrary to previous research, we find evidence against the unit root hypothesis for all price series. Our findings support characterizing natural resource prices as stationary around deterministic trends with structural breaks. We additionally show that both pre-testing for unit roots with breaks and allowing for breaks in the forecast model can improve forecast accuracy. Overall, the results in this paper are important in both a positive and normative sense; without an appropriate understanding of the dynamics of a time series, empirical verification of theories, forecasting, and proper inference are potentially fruitless.     

Nest defence by song sparrows: methodological and life history considerations

Summary Several aspects of nest defence behavior were investigated in song sparrows (Melospiza melodia) in eastern Ontario. Two independent tests were made of the hypothesis that the increase in nest defence observed through a nesting attempt is due to the birds becoming familiar with the nest threat, rather than because the nest contents increase in value to the parents. Neither test supported the hypothesis. As predicted by life history theory for species with age-independent mortality, males did not defend their nest more vigorously as they become older. Parents defended their nests less vigorously through the breeding season, contrary to the expected pattern of increased nest defence in response to declining renesting potential. This result may be attributable to a decline in offspring value through the breeding season. Nest defence behavior of mated individuals was positively correlated, independent of factors such as offspring age, renesting potential and brood size. From this result it is proposed that a source of variation in nest defence behavior may be individuals basing their own response on their mate's response in a positive feedback fashion. Males defended nests less vigorously than females, consistent with the expectation that males have lower certainty of parentage in the offspring. It is proposed that variation in paternal uncertainty could contribute to the unexplained variation reported in nest defence studies.     

Kin preference behavior in Bufo boreas tadpoles

Summary The ontogeny of sibling recognition behavior was studied in the laboratory in tadpoles of the western toad (Bufo boreas boreas) to test the hypothesis that Bufo tadpoles associate with siblings and to compare this behavior with two species of anurans previously studied. Tadpoles reared exclusively with sibs demonstrated a preference to associate with sibs over non-sibs both early and late in development but tadpoles reared with sibs and non-sibs (mixed rearing groups) exhibited no preference. Larvae that developed a preference for sibs after being reared with them for 75 days lost this preference following exposure to a mixed group for 2 to 6 days. Additionally, larvae reared in a mixed group did not develop a preference for the familiar mixed group nor was a preference exhibited by individual larvae that were reared exclusively with 5 non-sibs. These results suggest that larvae learn by prior association to discriminate sibs from non-sibs and that preferences are rapidly modifiable following exposure to non-sibs. However, the latter two experiments suggest that social preferences are not totally labile and are not based entirely on familiarity. Field observations of larval activity, aggregation behavior, and dispersal patterns in B. boreas indicate that tadpoles mix with non-sibs from early larval stages throughout development. Because even short term exposure to nonsibs resulted in a loss of sib preference in our experiments, we suggest that a kin selection interpretation of B. boreas aggregation behavior is not parsimonious. Results differ with those of other studies of sib recognition in larval anurans.