Mate choice among sympatric fur seals: female preference for conphenotypic males

When closely related species breed in sympatry, and where hybrids have lower fitness, reinforcement theory predicts that selection should favour mechanisms that reduce the probability of interspecific matings. If this situation arises among species that exhibit resource defence polygyny where males and females of different species reside in the same territories, there may be some conflict between mate choice based on territory-holding ability (sexual selection) and mate choice for correct species. We investigated this in a population of fur seals where three species are sympatric and where some females breed in the territories of heterospecific males, and where interspecific matings and hybrid pups are observed. The territorial status of males and the birthing sites of females were determined during daily observations, as were the movements of males and females, the location of matings and mating partners. DNA extracted from skin samples was used to determine paternities using DNA fingerprinting and the mtDNA genotype of individuals. Individuals were also classed on the basis of species-typical phenotype. We found that extra-territory inseminations (ETIs) were significantly more prevalent (67%) when territorial males and resident females were of different phenotype than when of similar phenotype (27%), but mtDNA genotype had no effect on the rate of ETIs. ETIs were probably by males with the same phenotype, as pups born to these females in the following season had the same phenotype as their mothers, suggesting they were not hybrids. These results suggest that within the resource defence polygynous mating system of these sympatric fur seals, female mate choice is more influenced by male phenotype than genotype. Contrary to our predictions, our study indicates that potential conflict between mate choice based on sexual selection and species recognition is unlikely, because females have some capacity to discriminate between males both within and between species on phenotypic traits additional to those under sexual selection. Although at least 25% of the pups born in this study were hybrid, this study can only support reinforcement theory if hybrids have reduced fitness. The fitness of hybrids among the species studied is currently unknown. Received: 19 January 1998 / Accepted after revision: 12 September 1998     

Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus

We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.     

RIVPACS models for predicting the expected macroinvertebrate fauna and assessing the ecological quality of rivers

The European Union Water Framework Directive recognises the need for and value of biological monitoring. This paper reviews the modelling approach known as River Invertebrate Prediction and Classification System (RIVPACS for assessing the ecological quality of river sites using macroinvertebrate sampling. The RIVPACS philosophy is to develop statistical relationships between the fauna and the environmental characteristics of a large set of high quality reference sites which can be used to predict the macroinvertebrate fauna to be expected at any site in the absence of pollution or other environmental stress. The observed fauna at new test sites can then be compared with their site-specific expected fauna to derive indices of ecological quality. All methodological decisions in any such model development have implications for the reliability, precision and robustness of any resulting indices for assessing the ecological quality and ecological grade (‘status’) of individual river stretches. The choice of reference sites and environmental predictor variables, the site classification and discrimination methods, the estimation of the expected fauna, and indices for comparing the agreement, or lack of it, between the observed and expected fauna, are all discussed. The indices are assessed on the reference sites and on a separate test set of 340 sites, which are subject to a wide range of types and degrees of impairment.     

Copying and the repeatability of mate choice

Models of sexual selection by female choice require heritable variation in female mating preferences in order for sexual selection to operate. However, recent theoretical work shows that female preferences which are transmitted non-genetically can result in exaggeration of male ornamentation. Guppies exhibit both mate copying and considerable heritable variation in female preferences. I studied the importance of these phenomena by measuring repeatability of female mate choice, which acts as an estimate of the upper limit to which a feature may be heritable, and the incidence of mate choice reversal in paired-trial binary mate choice experiments. Mate choice was significantly repeatable except in the treatment where females were given the opportunity to copy a female that contradicted their original choice. Apart from this, I found no evidence that females copy the mate choice of others. The differences between males in ornamentation had no effect on the consistency of female mate choice or the probability that they would reverse their original choice decision (in both controls and the copying experiment). The interval between choice trials did not influence repeatability significantly, indicating that the independence of choice decisions is not related to the time interval between them. Received: 9 February 1996/Accepted after revision: 6 July 1996     

Improving multi-site benefit functions via Bayesian model averaging: A new approach to benefit transfer

A benefit function transfer obtains estimates of willingness-to-pay (WTP) for the evaluation of a given policy at a site by combining existing information from different study sites. This has the advantage that more efficient estimates are obtained, but it relies on the assumption that the heterogeneity between sites is appropriately captured in the benefit transfer model. A more expensive alternative to estimate WTP is to analyze only data from the policy site in question while ignoring information from other sites. We make use of the fact that these two choices can be viewed as a model selection problem and extend the set of models to allow for the hypothesis that the benefit function is only applicable to a subset of sites. We show how Bayesian model averaging (BMA) techniques can be used to optimally combine information from all models.The Bayesian algorithm searches for the set of sites that can form the basis for estimating a benefit function and reveals whether such information can be transferred to new sites for which only a small data set is available. We illustrate the method with a sample of 42 forests from U.K. and Ireland. We find that BMA benefit function transfer produces reliable estimates and can increase about 8 times the information content of a small sample when the forest is ‘poolable’.     

Modulation of individual behavior and collective decision-making during aggregation site selection by the ant<Emphasis Type="Italic"> Messor barbarus</Emphasis>

Insect societies are often confronted with choices among several options such as food sources of different richness or potential nest sites with different qualities. The mechanisms by which a colony as a whole evaluates these situations and takes the appropriate decision are of crucial importance for its survival. Here we studied how collective decisions arise from individual behaviors when a group of workers of the ant Messor barbarus is given a choice between two aggregation sites. Two hundred ants were introduced into an arena and given a choice between two tubes connected to the arena. The tubes had different physical properties: dry and transparent (termed as dry), humid and transparent (termed as humid), or dry and dark (termed as dark). After 30 min, most ants were found to be aggregated in a humid tube when paired with a dry tube, or in a dark tube when paired with a humid one. When two humid tubes were in competition, ants aggregate more in one of the sites. The choice of ants was consistent throughout experiments. An analysis of individual behaviors shows that the probability of an ant recruiting and the intensity of its trail-laying behavior strongly depend on the quality of the tubes. Our study suggests that the selection of an aggregation site does not require that individual ants directly compare sites, but rather relies on the synergy between amplification processes involving recruitment by chemical trails, and a modulation of the individual resting time in a site as a function of its population.Communicated by L. Sundström     

An evaluation of three statistical methods used to model resource selection

The performance of statistical methods for modeling resource selection by animals is difficult to evaluate with field data because true selection patterns are unknown. Simulated data based on a known probability distribution, though, can be used to evaluate statistical methods. Models should estimate true selection patterns if they are to be useful in analyzing and interpreting field data. We used simulation techniques to evaluate the effectiveness of three statistical methods used in modeling resource selection. We generated 25 use locations per animal and included 10, 20, 40, or 80 animals in samples of use locations. To simulate species of different mobility, we generated use locations at four levels according to a known probability distribution across DeSoto National Wildlife Refuge (DNWR) in eastern Nebraska and western Iowa, USA. We either generated 5 random locations per use location or 10,000 random locations (total) within 4 predetermined areas around use locations to determine how the definition of availability and the number of random locations affected results. We analyzed simulated data using discrete choice, logistic-regression, and a maximum entropy method (Maxent). We used a simple linear regression of estimated and known probability distributions and area under receiver operating characteristic curves (AUC) to evaluate the performance of each method. Each statistical method was affected differently by number of animals and random locations used in analyses, level at which selection of resources occurred, and area considered available. Discrete-choice modeling resulted in precise and accurate estimates of the true probability distribution when the area in which use locations were generated was ≥ the area defined to be available. Logistic-regression models were unbiased and precise when the area in which use locations were generated and the area defined to be available were the same size; the fit of these models improved with increased numbers of random locations. Maxent resulted in unbiased and precise estimates of the known probability distribution when the area in which use locations were generated was small (home-range level) and the area defined to be available was large (study area). Based on AUC analyses, all models estimated the selection distribution better than random chance. Results from AUC analyses, however, often contradicted results of the linear regression method used to evaluate model performance. Discrete-choice modeling was best able to estimate the known selection distribution in our study area regardless of sample size or number of random locations used in the analyses, but we recommend further studies using simulated data over different landscapes and different resource metrics to confirm our results. Our study offers an approach and guidance for others interested in assessing the utility of techniques for modeling resource selection in their study area.     

Colony choice in birds: models based on temporally invariant site quality

We propose two stochastic models to explain how birds choose colonies. In the resource choice model, birds settle at each site at a rate proportional to the total resources the site contains. In the reduced resource choice model, a smaller cohort of birds enters sites at a rate determined by the total resources at each, and the remaining individuals enter sites at a rate that is linearly proportional to the total number of birds already nesting at each site. Thus, a fraction of birds chooses sites based on the resources present, and the remainder are attracted to a site by the presence of other birds. Colony site quality is assumed not to vary between years. Both models result, on average, in an ideal free distribution of colony sizes if the birds' settlement rate is linearly related to the resources in a site, if resources are distributed equally among individuals within sites, and if individuals with equal resources have equal fitness. We applied these models to long-term data on colony sizes and site usage of cliff swallows in south-western Nebraska. A test of the resource choice model suggested that the swallow population as a whole did not choose sites based strictly on site quality or the total resources contained at each site. However, a test of the reduced resource choice model suggested that a smaller fraction of the individuals in each colony may have based their choice of site on local resource availability, with the remaining birds aggregating at those sites based on the number of birds already settled there. Tests of these models may provide insight into how individuals choose colony sites and why colonies vary in size.     

Consequences of experimentally elevated yolk testosterone levels for intra- and inter-sexual selection in canaries

Hormones of maternal origin transferred to the eggs of oviparous species have been shown to significantly affect offspring development. Furthermore, there is now increasing evidence that these effects may last into adulthood. This underlines the persistence of yolk hormone-mediated maternal effects as well as their trans-generational potential as these changes may involve fitness-related traits such as mate choice behaviour, reproductive traits and longevity. Here, we tested the potential of yolk testosterone to affect sexual selection by experimentally increasing the yolk testosterone levels via egg injections. We focused on two central axes of sexual selection, male–male competition for access to a female (intra-sexual selection) and female mate choice behaviour (inter-sexual selection), using canaries (Serinus canaria) as a model species. Neither male agonistic behaviour nor access to the opposite sex, as measured in staged male–male encounters in the presence of a female, were affected by experimentally elevating yolk testosterone levels. We did not find any evidence for effects on female mate choice behaviour either, given the lack of significant effects on mate choice activity, consistency in female mate choice or choosiness. In conclusion, our results indicate that the consequences of yolk testosterone for sexual selection through changes in behavioural traits, which are expressed during pair formation or male–male competition, are probably limited.     

Nest-Site Selection in Individual Loggerhead Turtles and Consequences for Doomed-Egg Relocation

Abstract:  Relocation of eggs is a common strategy for conservation of declining reptilian populations around the world. If individuals exhibit consistency in their nest-site selection and if nest-site selection is a heritable trait, relocating eggs deposited in vulnerable locations may impose artificial selection that would maintain traits favoring unsuccessful nest-site selection. Conversely, if most individuals scatter their nesting effort and individuals that consistently select unsuccessful nest sites are uncommon, then artificial selection would be less of a concern. During the 2005 nesting season of loggerhead turtles ( Caretta caretta ) at Mon Repos beach, Queensland, Australia, we measured the perpendicular distance from the original nest site to a stationary dune baseline for in situ (unrelocated) and relocated clutches of eggs. We observed the fate of in situ clutches and predicted what would have been the fate of relocated clutches if they had not been moved by mapping tidal inundation and storm erosion lines. In 2005 turtles deposited an average of 3.84 nests and did not consistently select nest sites at particular distances from the stationary dune baseline. Selection of unsuccessful nest sites was distributed across the nesting population; 80.3% of the turtles selected at least one unsuccessful nest site and when previous breeding seasons were included, 97% selected at least one unsuccessful nest site. Females with nesting experience selected more successful nest sites than females with little or no experience. Relocating eggs vulnerable to tidal inundation and erosion saves the progeny from a large percentage of the population and the progeny from individuals who may in subsequent years nest successfully. Our results suggest that doomed-egg relocation does not substantially distort the gene pool in the eastern Australian loggerhead stock and should not be abandoned as a strategy for the conservation of marine turtle populations.     

Mate choice tactics and swarm size: a model and a test in a dance fly

In the dance flyEmpis borealis (L.) (Diptera: Empididae) females gather to swarm and males visit swarms for mating. A model was constructed, based on previously published data, simulating how males may choose among females of different sizes in swarms of different sizes. The focal question was, what influences the number of individuals in the swarm in this and possibly other swarming insects? The relationships between original swarm size and both the number of males arriving per minute and the proportion of males mating are both logarithmic. The model predicted that if these relationships were linear, or if males were able to judge absolute female size, the mean swarm size should increase and be at least four times as large as those found in the field. The only type of male mate choice strategy that gave rise to very large swarms (>25) was size-related choice (if males are able to assess the size of a female in relation to the entire population and not merely to the swarm). Furthermore, no swarming behaviour would occur if males mate independently of swarm size. Thus, the numbers of females attending a given swarm site are influenced by male arrival pattern, male preference for larger swarms, the inability of males to judge the absolute body size of females, and female polyandry. Males searching for mates seem to prefer larger swarms than females searching for a swarm to join, but the mean swarm size is primarily set by the swarm size preference of females. Optimal swarm size predicted from the model was 4.68±0.53 females. In order to test model predictions, 69 natural swarm sites were studied during one season. The mean swarm size was 4.85±4.54 females (median 4.03), and about 90% of swarms consisted of 11 females or fewer. Predicted and observed swarm size did not differ significantly.     

The effect of perceived female parasite load on post-copulatory male choice in a sex-role-reversed pipefish

The last several decades of research in behavioral ecology have resulted in a deeper appreciation of post-mating processes and sexual conflict in sexual selection. One of the most controversial aspects of sexual selection is cryptic mate choice. Here, we take advantage of male pregnancy in a sex-role-reversed pipefish (Syngnathus typhle) to quantify cryptic choice based on perceived parasite load and other sources of variance in female fitness. Studies have shown that S. typhle males preferentially mate with females with lower parasite loads and that a male’s perception of female parasite load can be altered by tattooing females. We manipulated the apparent parasite load of females in controlled mating experiments to test the hypothesis that post-copulatory sexual selection is dependent on a male’s perception of female parasite load in pipefish. Our results provided no evidence for cryptic male choice based on perceived female parasite load. However, we found evidence that eggs from larger females were more likely to result in viable offspring than eggs from smaller females and that the first female to mate with a male transferred more eggs per copulation on average. Overall, our results show that potential for post-copulatory sexual selection does exist in pipefish, but the male’s perception of female parasite load does not play a major role in this process.     

Suitability of short or long conservation contracts under ecological and socio-economic uncertainty

Acquisition of land rights has become a primary tool used to protect terrestrial biodiversity. Fixed length contracts are often used when trying to secure conservation benefits on private land in agri-environment schemes and payment for environmental services schemes, but the duration of the conservation contracts used in different programmes varies. To date, very little research has been undertaken to determine the situations in which contracts of differing lengths are optimal or when conservation agencies or groups should use a portfolio of different contract lengths rather than relying on a single type. Using stochastic dynamic programming and related heuristic methods, we investigate how the choice between short or long conservation contracts is affected by uncertainty regarding the future availability of sites and their ecological condition. We also examine the benefits offered by using a portfolio of different contract lengths. Conservation agencies must pay private landowners a premium to secure longer agreements and because of this, shorter contracts are advantageous if sites are likely to remain available for conservation in the future. Long contracts are preferred when future site availability becomes more unlikely. In contrast to uncertainty over site availability, uncertainty over future ecological conditions has little effect on contract selection and only markedly influences the choice between short and long contracts when there is heterogeneity across sites in expected conservation outcomes and future availability of sites is also uncertain. Finally, when future site availability is unlikely, the use of a portfolio of short and long contracts would offer greater conservation gains than using either type in isolation, even though this option is not yet one that is commonly found in conservation practice.     

Female choice in sage grouse: the roles of attraction and active comparison

Previous studies of female choice in sage grouse Centrocercus urophasianus have implicated both the acoustic quality and repetition rate of the stereotyped strut display as putative cues for female choice. Stages in the choice process at which specific components of male courtship display influence female decisions were investigated using field observations of female pre-mating behavior. Females visited a subset of territorial males and then actively chose one of these as a mate. The order in which males were visited suggested that females searched until an acceptable mate was found, rather than employing a “best-of-n” tactic. Numbers of females visiting a male were related to differences in an acoustical component of display (inter-pop interval) whereas the probability that a visiting female mated was related to display rate (Table 3), indicating that initial attraction and active choice are influenced by different components of display. In addition, inter-pop interval and display rate tended to covary inversely (Fig. 1), suggesting that attraction and active choice may impose conflicting selection pressures on display performance. Received: 11 November 1995/Accepted after revision: 16 March 1996     

Modeling and analysis of nest-site selection by honeybee swarms: the speed and accuracy trade-off

Nest-site selection in honeybees is a process of social decision making in which the scout bees in a swarm locate several potential nest sites, evaluate them, and select the best one by means of competitive signaling. We develop a model of this process and validate that the model possesses the key features of the bees' decision-making process, as revealed by prior empirical studies. Next, we use the model to study the “design” of the nest-site selection process, with a focus on how certain behavioral parameters have been tuned by natural selection to achieve a balance between speed and accuracy. First, we study the effects of the quorum threshold and the dance decay rate. We show that evolution seems to have settled on values for these two parameters that seek a balance between speed and accuracy of decision making by minimizing the time needed to achieve a consensus and maximizing the probability that the best site is chosen. Second, we study the adaptive tuning of the tendency of bees to explore for vs be recruited to a site. We show that this tendency appears to be tuned to regulate the positive feedback process of recruitment to ensure both a reasonably rapid choice and a low probability of a poor choice. Finally we show that the probability of choosing the best site is proportional to its quality, but that this proportionality depends on its quality relative to other discovered sites.

Genetic evidence for the “good genes” process of sexual selection

The adaptive nature of female mate choice remains one of the most contentious issues in the study of sexual selection. Here, I provide evidence that mate choice by females of the ovoviviparous cockroach Nauphoeta cinerea influences the rate at which offspring develop and provides both direct and indirect benefits to the female. Males that are more attractive to females produce offspring with shorter development times than less preferred males. Development time is heritable and apparently unconstrained by antagonistic pleiotropy. Male attractiveness and rate of offspring development are genetically correlated. Offspring gain an indirect benefit from their mother's mate choice because, on average, individuals that hatch faster reach sexual maturity more quickly. Females that discriminate among males gain a direct benefit because N. cinerea is ovoviviparous and the time between clutches is decreased by producing offspring with shorter development. In addition to providing evidence for beneficial consequences of mate choice, this study highlights how genetic data provide insights into the process of sexual selection not gained in a purely phenotypic study.Communicated by P. Pamilo     

Habitat selection increases the isolating effect of habitat fragmentation on the gopher tortoise

Habitat selection requires choice, which differentiates it from habitat use, and choice, in turn, is dependent upon the responses of organisms to the environmental, social, and other cues that they perceive. Habitat selection by the gopher tortoise (Gopherus polyphemus) was investigated by translocating tortoises and monitoring their movements within two sites in central Florida. The first site supported a stable preponderance of high-quality habitat, and tortoises avoided areas with a dense tree canopy cover caused by fire exclusion. The second site was badly invaded by an introduced weed, and tortoises avoided areas where the weed had formed a dense monoculture. At both sites, individuals appeared to be responding to visual cues to avoid areas that were relatively dark. In landscapes with relatively large amounts of high-quality habitat, this avoidance behavior serves the gopher tortoise well by keeping individuals within the dominant habitat type. In degraded areas, high-quality habitat often becomes increasingly uncommon, and the avoidance behavior exhibited by the tortoises will result in individuals becoming confined to small patches, causing a significant reduction in fitness and hence questioning their long-term survival in such areas. The results from our study show that in order to maintain viable tortoise populations in areas increasingly subjected to human fragmentation and degradation, it is crucial not only to suppress tree canopy cover continually and prevent invasion by exotic weeds, but also to be mindful that the avoidance behavior of the gopher tortoise could prevent individuals from fully occupying a high-quality habitat in response to restoration and management efforts.     

Sexual selection for male parental care in the sand goby, Pomatoschistus minutus

Male parental care is typically thought to come at a cost to mate attraction and future mating success. However, it has also been hypothesized that paternal care may be under sexual, as well as natural, selection, such that good fathers actually attract more mates. Here we show experimentally that in the sand goby, Pomatoschistus minutus, females prefer to mate with males that provide higher levels of parental care. We manipulated male behavior using (1) different nest sizes and (2) an application of low-O₂ water in the nests, and found that females consistently preferred males with elevated levels of care in dichotomous mate choice tests. This complements our earlier study in which we showed that males increase the amount and quality of care they provide in the presence of females. Our results demonstrate that male care may have evolved as a result of sexual selection rather than natural selection alone, and furthermore, that male care may not necessarily be in conflict with mate attraction.     

Effects of male dominance and courtship display on female choice in the ring-necked pheasant

Male traits and behaviours acting in mate choice and intrasexual competition are expected to be congruent. When studying their evolution, this often makes it difficult to differentiate between these two components of sexual selection. Studies are therefore needed on mate choice in conjunction with the role of displays and dominance. We present the results from two experiments conducted to investigate the effects of male dominance and courtship displays on female choice in the ring-necked pheasant, Phasianus colchicus, controlling for differences in morphological male traits. We found: (1) different courtship behaviours had different effects on female choice: females were mainly attracted by the feeding courtship behaviour, while another courtship display (the lateral display) was effective in producing the copulation-acceptance response by the females; (2) subordinate males performed the courtship behaviour before females less frequently than dominant males, and females reinforced intrasexual selection by choosing dominant males, and (3) subordinate males in visual contact with a dominant became less attractive to females. The results support the idea (armament-ornament model) that female pheasants may benefit from using traits selected in male-male competition as clues for mate choice. Received: 23 October 1997 / Accepted after revision: 7 October 1998     

Consensus building during nest-site selection in honey bee swarms: the expiration of dissent

This study addresses a question that lies at the heart of understanding how the scouts in a honey bee swarm achieve unanimity in their dances, and so reach agreement in their choice of a future nest site: what causes the scouts that perform dances for the non-chosen sites to stop dancing for these sites? One possibility is that a scout stops dancing for a non-chosen site only after she follows a lively dance for another site, such as the site that is ultimately chosen. This hypothesis is contradicted by the finding that 23 out of 27 scouts (in 6 swarms) that danced initially for a non-chosen site stopped their dancing before they followed a dance for another site. Evidently, a scout that supports initially one of the non-chosen sites is likely to withdraw her support for this site even before she learns about another site. What causes her to do so? Close examination of the behavior of scouts revealed that they reduce the strength of their dancing (waggle runs/return to the swarm) for a given site over consecutive returns to the swarm. On average, the pattern of this reduction in dancing is strikingly linear, which suggests that it arises from an internal, neurophysiological process that automatically drives down a scout's motivation to dance for a site. Other results suggest that scouts from inferior sites start their dancing less strongly, and so cease their dancing more rapidly, than do scouts from superior sites. If so, then during the consensus-building process of the scouts, it is the support (the dancing) for inferior sites that is most likely to die out while it is the support for a superior site that is most likely to prevail.