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Showing papers in "Behavioral Ecology and Sociobiology in 2006"


Journal ArticleDOI
Aldo Poiani1
TL;DR: The diversity of microorganismal, cellular and molecular components of seminal fluids can be interpreted in the light of emergence of co-adapted complexes, host–parasite coevolution, male–female arms races, sperm competition, pleiotropy and redundancy of function.
Abstract: The seminal fluid is a complex medium containing a great variety of molecules, mainly produced by sex accessory glands, and also cells other than spermatozoa (e.g. leucocytes). In this paper, I review current knowledge on composition of seminal fluid in both vertebrates (mainly mammals) and invertebrates (mainly insects) with internal fertilisation, in the light of possible benefits of seminal fluid components to males (e.g. sperm capacitation, sperm competition and fertilisation), possible costs to males (e.g. autoimmunity, antigenic effects), potential benefits to females being inseminated (e.g. food, immunostimulation and antibiotic effects) and potential costs to females (e.g. transmission of venereal diseases). The diversity of microorganismal, cellular and molecular components of seminal fluids can be interpreted in the light of emergence of co-adapted complexes, host–parasite coevolution, male–female arms races, sperm competition, pleiotropy and redundancy of function.

455 citations


Journal ArticleDOI
TL;DR: Testing a number of predictions derived from kin selection theory about the strength of social bonds among adult female baboons suggests that social bonds play a vital role in females’ lives, and the ability to establish and maintain strong social bonds may have important fitness consequences for females.
Abstract: Sociality has positive effects on female fitness in many mammalian species. Among female baboons, those who are most socially integrated reproduce most successfully. Here we test a number of predictions derived from kin selection theory about the strength of social bonds among adult female baboons. Our analyses are based on systematic observations of grooming and association patterns among 118 females living in seven different social groups in the Amboseli Basin of Kenya over a 16-year period. Females in these groups formed the strongest bonds with close kin, including their mothers, daughters, and maternal and paternal sisters. Females were also strongly attracted toward females who were close to their own age, perhaps because peers were often paternal sisters. Females’ bonds with their maternal sisters were strengthened after their mother’s deaths, whereas their relationships with their maternal aunts were weakened after their mother’s death. In addition, females formed stronger bonds with their paternal sisters when no close maternal kin were available, and they compensated for the absence of any close kin by forming strong bonds with nonrelatives. Taken together, these data suggest that social bonds play a vital role in females’ lives, and the ability to establish and maintain strong social bonds may have important fitness consequences for females.

414 citations


Journal ArticleDOI
TL;DR: The meta-analysis suggests that the effect of predation risk on foraging behavior in terrestrial systems is strongly dependent on the type ofpredation risk being utilized.
Abstract: Although there is ample evidence for the generality of foraging and predation trade-offs in aquatic systems, its application to terrestrial systems is less comprehensive. In this review, meta-analysis was used to analyze experiments on giving-up-densities in terrestrial systems to evaluate the overall magnitude of predation risk on foraging behavior and experimental conditions mediating its effect. Results indicate a large and significant decrease in foraging effort as a consequence of increased predation risk. Whether experiments were conducted under natural or artificial conditions produced no change in the overall effect predation had on foraging. Odor and live predators as a correlate of predation risk had weaker and nonsignificant effects compared to habitat characteristics. The meta-analysis suggests that the effect of predation risk on foraging behavior in terrestrial systems is strongly dependent on the type of predation risk being utilized.

381 citations


Journal ArticleDOI
TL;DR: This work draws on data derived from a 16-year study of baboons living in seven different social groups in the Amboseli basin of Kenya to evaluate the quality and stability of social bonds among females, and demonstrates that the quality of social Bonds directly affects their stability.
Abstract: A growing body of evidence suggests that social bonds have adaptive value for animals that live in social groups. Although these findings suggest that natural selection may favor the ability to cultivate and sustain social bonds, we know very little about the factors that influence the quality or stability of social bonds. Here, we draw on data derived from a 16-year study of baboons living in seven different social groups in the Amboseli basin of Kenya to evaluate the quality and stability of social bonds among females. Our results extend previous analyses, which demonstrate that females form the strongest bonds with close maternal and paternal kin, age mates (who may be paternal kin), and females who occupy similar ranks but are not maternal relatives. Here we show that the same factors influence the quality and strength of social bonds. Moreover, the results demonstrate that the quality of social bonds directly affects their stability.

331 citations


Journal ArticleDOI
TL;DR: Female guppies showed social preferences for stable partners, fulfilling a key assumption made by models of reciprocity, and the occurrence of highly interconnected social networks between stable partners suggests the existence of co-operation networks in free-ranging populations of the guppy.
Abstract: In contrast to the substantial number of theoretical papers that have examined the mechanisms by which cooperation may evolve, very few studies have investigated patterns of co-operation in natural animal populations. In the current study, we use a novel approach, social network analysis, to investigate the structure of co-operative interactions in the context of predator inspection in a wild population of guppies (Poecilia reticulata). Female guppies showed social preferences for stable partners, fulfilling a key assumption made by models of reciprocity. In the laboratory, wild female guppies disproportionately engaged in predator inspection with others with whom they had strong social associations. Furthermore, pairs of fish that frequently engaged in predator inspection did so in a particularly co-operative way, potentially reducing costs associated with predator inspection. Taken together, these results provide evidence for assortative interactions forming the basis of co-operation during predator inspection in a natural fish population. The occurrence of highly interconnected social networks between stable partners suggests the existence of co-operation networks in free-ranging populations of the guppy.

229 citations


Journal ArticleDOI
TL;DR: In noisy environments, serial redundancy in bird song may reflect a trade-off between successful signal transmission and preventing motor fatigue.
Abstract: Many animal signals are performed in a highly redundant manner as in some bird species where males sing several renditions of one song type before switching to another. However, differences in signal redundancy between contexts and between individuals are only poorly understood. We found that chaffinches (Fringilla coelebs) in noisier areas (i.e., close to waterfalls and torrents) sang longer bouts of the same song type before switching to a new type, suggesting that they use increased serial redundancy to get the message across in noisy conditions. This is the first evidence of a noise-dependent adjustment of signal redundancy in a songbird. In addition, we found that song types with faster trills were sung in shorter bouts suggesting that the performance of highly redundant song series is probably limited by motor constraints. Thus, in noisy environments, serial redundancy in bird song may reflect a trade-off between successful signal transmission and preventing motor fatigue.

189 citations


Journal ArticleDOI
TL;DR: Although most aggressive signals contained little information on attack likelihood, as some models suggest should be the case, the unreliability of these signals was not caused by convergence of individuals on a single signaling strategy, as those models argue should occur.
Abstract: A central question in animal communication research concerns the reliability of animal signals. The question is particularly relevant to aggressive communication, where there often may be advantages to signaling an exaggerated likelihood of attack. We tested whether aggressive signals are indeed reliable signals of attack in song sparrows (Melospiza melodia). We elicited aggressive signaling using a 1-min playback on a male’s territory, recorded the behavior of the male for 5 min, and then gave him the opportunity to attack a taxidermic mount of a song sparrow associated with further playback. Twenty subjects attacked the mount and 75 did not. Distance to the speaker was a significant predictor of attack for both the initial recording period and the 1 min before attack. For the initial recording period, none of the measures of singing behavior that we made was a significant predictor of attack, including song-type matching, type-switching frequency, and song rate. For the 1-min period immediately before attack, only the number of low amplitude “soft songs” was a significant predictor of attack. Although most aggressive signals contained little information on attack likelihood, as some models suggest should be the case, the unreliability of these signals was not caused by convergence of individuals on a single signaling strategy, as those models argue should occur.

183 citations


Journal ArticleDOI
TL;DR: It is interesting to note that the background level of risk over the past 3 days influenced the threshold level of response to varying concentrations of alarm cues, which illustrated a remarkable interplay between immediate (current) risk and background risk in shaping the intensity of antipredator responses.
Abstract: Prey animals often have to face a dynamic tradeoff between the costs of antipredator behavior and the benefits of other fitness-related activities such as foraging and reproduction. According to the threat-sensitive predator avoidance hypothesis, prey animals should match the intensity of their antipredator behavior to the degree of immediate threat posed by the predator. Moreover, longer-term temporal variability in predation risk (over days to weeks) can shape the intensity of antipredator behavior. According to the risk allocation hypothesis, changing the background level of risk for several days is often enough to change the response intensity of the prey to a given stimulus. As the background level of risk increases, the response intensity of the prey decreases. In this study, we tested for possible interactions between immediate threat-sensitive responses to varying levels of current perceived risk and temporal variability in background risk experienced over the past 3 days. Juvenile convict cichlids were preexposed to either low or high frequencies of predation risk (using conspecific chemical alarm cues) for 3 days and were then tested for a response to one of five concentrations (100, 50, 25, 12.5%, or a distilled water control). According to the threat-sensitive predator avoidance hypothesis, we found greater intensity responses to greater concentrations of alarm cues. Moreover, in accordance with the risk allocation hypothesis, we found that cichlids previously exposed to the high background level of risk exhibited a lower overall intensity response to each alarm cue concentration than those exposed to the low background level of risk. It is interesting to note that we found that the background level of risk over the past 3 days influenced the threshold level of response to varying concentrations of alarm cues. Indeed, the minimum stimulus concentration that evoked a behavioral response was lower for fish exposed to high background levels of predation than those exposed to low background levels of predation. These results illustrate a remarkable interplay between immediate (current) risk and background risk in shaping the intensity of antipredator responses.

176 citations


Journal ArticleDOI
TL;DR: This review presents the results of basic theoretical models predicting female wasp search duration on a patch of hosts and analyzes all studies investigating the effect of different factors on parasitoid patch time allocation and patch-leaving decision rules.
Abstract: As most parasitoids are time limited, they usually die before they have laid all their eggs. In such cases, optimal foraging theory predicts that female parasitoids will adopt behavioral reproductive strategies enabling them to maximize progeny production per unit of time. One key situation in which parasitoid females must optimize their time budget is related to the fact that most of their hosts are distributed in discrete patches in the environment. In this review, I first present the results of basic theoretical models predicting female wasp search duration on a patch of hosts. I then compile and analyze all studies investigating the effect of different factors on parasitoid patch time allocation and patch-leaving decision rules. Different patch-leaving mechanisms that were proposed to explain the results obtained are discussed, along with statistical methods that should be used to estimate them from experimental data. Finally, ideas for future research are presented.

168 citations


Journal ArticleDOI
TL;DR: A model of nest-site selection in honeybees is developed and it is shown 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.
Abstract: 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.

149 citations


Journal ArticleDOI
TL;DR: The results provide support for the tug-of-war model in primates, and the approach developed here can be applied to study comparative patterns of skew in other biological systems.
Abstract: Reproductive skew models have been proposed as a unifying framework for understanding animal social systems, but few studies have investigated reproductive skew in a broad evolutionary context. We compiled data on the distribution of mating among males for 31 species of primates and calculated skew indices for each study. We analyzed the determinants of mating skew with phylogenetic comparative methods to investigate two models from reproductive skew theory, the concession model and the tug-of-war model. Mating skew decreased as the number of males increased in multimale groups, suggesting that monopolization of females becomes more difficult when there are more rivals, and therefore supporting the tug-of-war model. We predicted that single males are unable to monopolize receptive females as overlap in female receptivity increases (estrous synchrony) and, as a result, that mating skew decreases. However, we did not find any evidence for a link between female estrous synchrony and male mating skew. Finally, the concession model predicts high skew in male philopatric species relative to species in which males disperse, yet our measures of mating skew showed no significant associations with qualitative scores of male dispersal. More definitive tests of the concession model will require more quantitative measures of relatedness, which are presently unavailable for most primate species in our study. Overall, our results provide support for the tug-of-war model in primates, and the approach developed here can be applied to study comparative patterns of skew in other biological systems.

Journal ArticleDOI
TL;DR: A novel analysis of temporal polyethism in the ant Pheidole dentata is presented, demonstrating that task attendance by minor workers does not shift among spatially associated sets of behaviors that minimally overlap but rather expands with age, and that the number of tasks performed by older minors increases through the addition and retention of behaviors.
Abstract: The controversy concerning the extent to which the organization of division of labor in social insects is a developmental process or is based on task allocation dynamics that emerge from colony need independent of worker age and endocrine or neural state has yet to be resolved. We present a novel analysis of temporal polyethism in the ant Pheidole dentata, demonstrating that task attendance by minor workers does not shift among spatially associated sets of behaviors that minimally overlap but rather expands with age. Our results show that the number of tasks performed by older minors increases through the addition and retention of behaviors, with up to a sixfold increase in repertoire size from day 1 to day 20 of adult life. We also show that older minors respond to colony needs by performing significantly more brood care as its demand increases, indicating that they can quickly upregulate nursing according to labor requirements. This level of plasticity was absent in younger siblings. The breadth of responsiveness to task-related olfactory stimuli increased with age. In a binary choice test in which young and old minor workers could orient toward odorants from brood or food, older workers responded to both brood and food, whereas young workers responded only to brood. These dissimilar responses to stimuli associated with nursing and foraging indicate age-related differences in sensory ability and provide a physiological basis for the age-related repertoire expansion model. We discuss repertoire expansion in P. dentata in light of behavioral development and caste flexibility in ants.

Journal ArticleDOI
TL;DR: The hypothesis that differences in sensory access to prey contribute to resource partitioning between potentially competing species using two sympatric, similar-sized, congeneric bat species as a model system is tested.
Abstract: Coexisting animal species frequently differ in resource use in at least one niche dimension and thus avoid competition. While a range of morphological differences that lead to differentiation in animals' mechanical access to food have been identified, the role of sensory differences in within-guild niche differentiation has received less attention. We tested the hypothesis that differences in sensory access to prey contribute to resource partitioning between potentially competing species using two sympatric, similar-sized, congeneric bat species as a model system. Nursery colonies of Natterer's bat (Myotis nattereri) and Bechstein's bat (Myotis bechsteinii) roost in bat boxes in the same orchard and forage in forests and orchards nearby. In observations and behavioural experiments with freshly captured M. bechsteinii, we showed that individuals are able to hunt using prey-generated sound alone. In contrast, M. nattereri rarely uses prey-generated sound, but instead is able to find prey by echolocation very close to vegetation. In accordance with these behavioural data, we showed that M. bechsteinii has significantly larger ears than M. nattereri, providing it with superior detection and localization abilities for relatively low-frequency prey rustling sounds. We hypothesized that these differences in sensory ecology of the two syntopic, congeneric species would contribute to resource partitioning, so that M. bechsteinii would find more noisy prey taxa, possibly hidden in vegetation, by listening for prey sounds, while M. nattereri would have better access to still prey using echolocation or associative learning. Analysis of faecal samples collected on the same nights from bat boxes occupied by each species corroborated this prediction. The diets of the two species differed significantly, reflecting their different prey perception techniques and thereby supporting the hypothesis that differences in sensory ecology contribute to niche differentiation.

Journal ArticleDOI
TL;DR: The results support the challenge hypothesis with males exhibiting elevated testosterone in association with the acquisition of high rank (ensuring access to mates), rather than with mating itself, and testosterone levels predicted future, rather than current, mating activity.
Abstract: Despite the many benefits that testosterone has on male reproduction, sustaining high levels of testosterone for long periods can be costly. The challenge hypothesis predicts that males will show temporarily sustained elevations of testosterone at critical periods, counterbalanced by decreased levels during noncritical periods. We investigated male testosterone measures extracted from fecal samples in a group of chacma baboons (Papio hamadryas ursinus) living in the Okavango Delta, Botswana. Because rank serves as a proxy for competition for mates, we examined how male testosterone was related to dominance rank, age, aggression, and mating activity. Males showed an elevation in testosterone at maturity; young adult males had the highest testosterone levels followed by a steady decline with age. Among dispersing males, testosterone was temporarily elevated in the month following dispersal. After controlling for age, testosterone and rank were unrelated, but testosterone and changes in rank were positively correlated, such that males rising in rank had higher testosterone than males falling in rank. Thus, for males in this group, testosterone was predictive of a male's rank trajectory, or future rank. Similarly, male testosterone levels predicted future, rather than current, mating activity. Finally, male testosterone and aggression rates were unrelated during stable periods in the dominance hierarchy but positively related during unstable periods when high ranks were being contested. In general, our results support the challenge hypothesis with males exhibiting elevated testosterone in association with the acquisition of high rank (ensuring access to mates), rather than with mating itself.

Journal ArticleDOI
TL;DR: Evidence is provided for cooperative nursing and adoption by plural-breeding females in a nocturnal primate, the gray mouse lemur, in which females forage solitarily during the night, but form day-time sleeping groups with one to two other females.
Abstract: Lactation imposes substantial physiological costs on mothers and should therefore not be directed towards foreign offspring. Such allonursing, however, is common in mammal species that share roosts. Hypotheses to explain allonursing among such plural breeders include misdirected parental care, milk evacuation, brood parasitism, reciprocity, and kin selection. The necessary behavioral data, in combination with data on kinship and kin recognition, have rarely been available to distinguish among these explanations, however. In this study, we provide evidence for cooperative nursing and adoption by plural-breeding females in a nocturnal primate, the gray mouse lemur (Microcebus murinus), in which females forage solitarily during the night, but form day-time sleeping groups with one to two other females. We observed 34 resident females in an 8 ha study area in Kirindy Forest, Madagascar, over three consecutive annual breeding seasons and determined genetic relationships among all members of this population. Five sleeping groups of adult females were filmed inside their roosts during one breeding season after females gave birth. The composition of groups changed substantially across years, but they always consisted of close maternal relatives. All females within a group gave birth to one to three infants. They regularly transferred only their own offspring among roosting sites, demonstrating an ability to discriminate between their own and other’s offspring, but they regularly groomed and nursed related offspring other than their own and adopted related dependent young after their mother’s death. Kin selection may therefore be the main selective force behind cooperative breeding among these closely related females with a high mortality risk, providing each of them with family insurance.

Journal ArticleDOI
TL;DR: Foraging time decreased after the winter solstice and was correlated with subsequent breeding phenology, such that individuals that spent less time foraging in February bred earlier, demonstrating that pre-breeding intrinsic foraging ability is critical in determiningbreeding phenology.
Abstract: In temperate regions, winter presents animals with a number of challenges including depressed food abundance, increased daily energy requirements, higher frequency of extreme weather events and shortened day length. Overcoming these constraints is critical for overwintering survival and scheduling of future breeding of long-lived species and is likely to be state dependent, associated with intrinsic abilities such as food acquisition rates. We examined the relationship between environmental and intrinsic factors on overwintering foraging and subsequent breeding phenology of the European shag Phalacrocorax aristotelis, a diurnal marine predator. We tested a range of hypotheses relating to overwintering foraging time and location. We found that individuals greatly increased their foraging time in winter to a peak of more than 90% of available daylight at the winter solstice. The seasonal patterns of foraging time appear to be driven by a combination of light levels and weather conditions and may be linked to the availability of the shag's principal prey, the lesser sandeel Ammodytes marinus. There was no evidence that shags dispersed south in winter to increase potential foraging time. Foraging time decreased after the winter solstice and, crucially, was correlated with subsequent breeding phenology, such that individuals that spent less time foraging in February bred earlier. The relationship was much stronger in females than males, in line with their more direct control of timing of breeding. Our results demonstrate that pre-breeding intrinsic foraging ability is critical in determining breeding phenology.

Journal ArticleDOI
TL;DR: It is shown that T acts to upregulate lipoprotein production and allows male zebra finches to display flashy, sexually attractive coloration, one of the more detailed physiological mechanisms underlying the development of a sex-steroid-dependent trait in animals.
Abstract: A salient feature of many secondary sexual characteristics in animals is that their expression is controlled by sex-steroid hormones. However, for only a few types of ornaments do we know the precise molecular mechanism by which androgens like testosterone (T) enhance trait production. We studied the red carotenoid-based beak of male zebra finches (Taeniopygia guttata), which serves as a sexually selected trait and is thought to be T dependent. In previous research, we demonstrated that the extent to which male finches produce lipoproteins in the bloodstream, which bind carotenoid pigments acquired from the diet and transport them to peripheral tissues, regulates the accumulation of carotenoids in the body and beak pigmentation. Here we show that T acts to upregulate lipoprotein production and allows male zebra finches to display flashy, sexually attractive coloration. Levels of circulating T in blood positively and significantly predicted lipoprotein profile (as measured by cholesterol levels), blood carotenoid concentration, and beak color. Exogenous T administration elevated cholesterol and carotenoid status as well as beak redness. Last, experimental inhibition of T (using the anti-androgen flutamide) downregulated lipoprotein production and carotenoid circulation and faded the beak. This androgen- and lipoprotein-mediated system represents one of the more detailed physiological mechanisms underlying the development of a sex-steroid-dependent trait in animals.

Journal ArticleDOI
TL;DR: Testing the hypothesis that expression of blue-green coloration of eastern bluebird eggs positively correlates to female condition indicates that egg coloration covaries with female condition, suggesting that eggcoloration could function as a reliable signal of female quality or that egg Coloration may allow females to recognize eggs laid by conspecific brood parasites.
Abstract: Egg coloration has been hypothesized to reflect female condition. Because of the proposed physiological costs associated with deposition of biliverdin pigments and because of their conspicuousness, eggs with blue-green coloration mayreliably convey information aboutfemale or brood quality. We tested the hypothesis that expression of blue-greencolorationofeasternbluebird(Sialiasialis)eggs positively correlates to female condition. First, we doc- umented the incidence of egg color polymorphism within the population. We observed that 98% of females laid blue- green eggs while less than 2% laid white eggs and less than 1% laid pink eggs. In a subset of clutches, we used full spectrum reflectance spectrometry (300-700 nm) to com- pare eggshell coloration to measures of female condition. We found that the color of eggs within clutches was more similar than the color of eggs from different clutches, and that the blue-green eggs have spectral peaks that are con- sistent with the characteristic absorbance spectra of bili- verdin pigmentation. Females in better body condition and older females laid more colorful eggs. Moreover, individual females laid morecolorful eggslater inthe layingsequence. Overall, these data indicate that egg coloration covaries with female condition, suggesting that egg coloration could function as a reliable signal of female quality or that egg coloration may allow females to recognize eggs laid by conspecific brood parasites.

Journal ArticleDOI
TL;DR: This study provides good evidence that BGC competition occurs in primates but it also reinforces the idea that folivore food resources may be worth defending, and provides evidence that adult male black and white colobus monkeys defend the food their mates need against other groups.
Abstract: In theory, between-group contest (BGC) competition for food can greatly influence female social relationships and reproductive success in primates, but few studies have investigated whether such effects occur and, if so, under what ecological conditions. There is evidence that adult male black and white colobus monkeys (Colobus guereza, “guerezas”) defend the food their mates need against other groups, suggesting that BGC competition is important in this species. Using data on feeding, ranging, vegetation patterns, and intergroup encounters between six neighboring guereza groups, I provide evidence that the highly folivorous guerezas at Kanyawara, Kibale National Park, Uganda, engaged in BGC competition over unevenly dispersed, relatively high-quality feeding areas or “core areas”. Intergroup aggression was common, and groups’ home ranges overlapped. Groups were more likely to initiate high-level aggression if they encountered another group within or near their core area, and groups that initiated and won encounters often fed in the same areas in which losing groups had fed. Guerezas fed selectively on species with contagious (clumped) distributions and concentrated their feeding efforts in areas of the forest that contained the most food (core areas). Groups could be ranked in a linear dominance hierarchy, and group rank number was inversely related to the quantity and quality of food in groups’ core areas. This study not only provides good evidence that BGC competition occurs in primates but it also reinforces the idea that folivore food resources may be worth defending.

Journal ArticleDOI
TL;DR: Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events, and a review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees.
Abstract: Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.

Journal ArticleDOI
TL;DR: It is demonstrated that kinships correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters.
Abstract: Kinship has been shown to be an important correlate of group membership and associations among many female mammals. In this study, we investigate association patterns in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting an embayment in southeastern Australia. We combine the behavioral data with microsatellite DNA and mitochondrial DNA data to test the hypotheses that genetic relatedness and maternal kinship correlate with associations and social clusters. Mean association between females was not significantly different from a random mean, but the standard deviation was significantly higher than a random standard deviation, indicating the presence of nonrandom associates in the dataset. A neighbor-joining tree, based on the distance of associations between females, identified four main social clusters in the area. Mean genetic relatedness between pairs of frequent female associates was significantly higher than that between pairs of infrequent associates. There was also a significant correlation between mtDNA haplotype sharing and the degree of female association. However, the mean genetic relatedness of female pairs within and between social clusters and the proportion of female pairs with the same and different mtDNA haplotypes within and between clusters were not significantly different. This study demonstrates that kinship correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters. We hypothesize that different forces acting on female bottlenose dolphin sociality appear to promote the formation of flexible groups which include both kin and nonkin.

Journal ArticleDOI
TL;DR: The results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent.
Abstract: Aposematic species exploit the ability of predators to associate, for example, conspicuous colouration with the unprofitability of prey. We tested the importance of colour for avoidance learning, memory and generalisation in wild-caught great tits (Parus major). First, we determined the birds’ initial colour preferences for red, yellow, orange and grey artificial prey items. The birds showed some preferences, as they were more willing to eat grey prey as their first choice, but these were not strong preferences. We then trained birds to discriminate red, yellow or variable (red and yellow) signals from grey where colours signalled palatable and unpalatable food. In general, the birds learned the discrimination task equally well, irrespective of which colours signalled unpalatability, and subsequently remembered the distinction between previously palatable and previously unpalatable colours in the memorability test. We did not find strong evidence that variability in the signal affected learning or memory. Our results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent. To study whether training also affects responses to unconditioned stimuli, we included orange prey items in the memorability test. Although orange had been palatable in the initial preference test, the birds ate fewer orange prey items after they had been trained to avoid red, yellow or both colours (variable signal) as unpalatable prey, but did not change their preference when trained that these colourful signals were palatable. This indicates that generalisation occurred more readily after a negative experience than a positive experience, a situation that would potentially allow imperfect mimicry to occur.

Journal ArticleDOI
TL;DR: The results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.
Abstract: Androgen hormones of maternal origin contained in the eggs of avian species are considered to have positive effects on offspring characteristics and performance. However, negative consequences have also been reported, suggesting that mothers may experience a trade-off between beneficial and detrimental effects of egg androgens to offspring fitness. We studied the effects of elevated yolk testosterone (T) concentration on survival, development and phenotype of male and female yellow-legged gull (Larus michahellis) chicks by injecting egg yolks with physiological doses of the hormone. Elevated yolk T resulted in a male-biased post-hatching sex ratio, T-treated clutches producing a greater proportion of males compared to control ones at day 4 post-hatching, likely resulting from a reduction of female embryonic survival, whereas no effect of hormone treatment on hatching success or short-term chick survival was observed. In addition, T depressed post-hatching body mass in both sexes but had no effects on the intensity of the cell-mediated immune response or skeletal growth. No sex differences in egg characteristics or chick phenotype were detected. Time to hatching was not affected by T, but females originating from first laid eggs hatched earlier than males of the same laying order, independently of hormone treatment. However, the implications of sex differences in hatching times are unclear in the study species. Taken together, our results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.

Journal ArticleDOI
TL;DR: It is concluded that hypotheses about the ecological causes of fission–fusion and the origin of complex social structures should consider the heterogeneity and complexity of the environment in which social animals live.
Abstract: Precisely how ecological factors influence animal social structure is far from clear. We explore this question using an agent-based model inspired by the fission–fusion society of spider monkeys (Ateles spp). Our model introduces a realistic, complex foraging environment composed of many resource patches with size varying as an inverse power law frequency distribution with exponent β. Foragers do not interact among them and start from random initial locations. They have either a complete or a partial knowledge of the environment and maximize the ratio between the size of the next visited patch and the distance traveled to it, ignoring previously visited patches. At intermediate values of β, when large patches are neither too scarce nor too abundant, foragers form groups (coincide at the same patch) with a similar size frequency distribution as the spider monkey’s subgroups. Fission–fusion events create a network of associations that contains weak bonds among foragers that meet only rarely and strong bonds among those that repeat associations more frequently than would be expected by chance. The latter form subnetworks with the highest number of bonds and a high clustering coefficient at intermediate values of β. The weak bonds enable the whole social network to percolate. Some of our results are similar to those found in long-term field studies of spider monkeys and other fission–fusion species. We conclude that hypotheses about the ecological causes of fission–fusion and the origin of complex social structures should consider the heterogeneity and complexity of the environment in which social animals live.

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TL;DR: Manipulating oxygen level and predator presence for small and large males found support for a parental trade-off between fanning and nest defense, and surprisingly, filial cannibalism did not differ between these groups.
Abstract: Brood guarding animals face many critical trade-offs. Sand goby males (Pomatoschistus minutus) build nests with larger openings during low oxygen conditions, presumably to enhance ventilation. However, this may make the nest easier for egg predators to detect and harder for guarding males to defend. Manipulating oxygen level and predator presence (a small crab) for small and large males, we found support for a parental trade-off between fanning and nest defense. An increased fanning activity resulted in less time for guarding. Small males and males in low oxygen showed a higher fanning expenditure than large males and males in high oxygen, but surprisingly, filial cannibalism did not differ between these groups. Males built larger nest openings in low than high oxygen. However, males in both high and low oxygen treatments reduced their nest opening size in the presence of a predator, again indicating an important trade-off between ventilation and nest defense.

Journal ArticleDOI
TL;DR: It is shown experimentally that in the sand goby, Pomatoschistus minutus, females prefer to mate with males that provide higher levels of parental care, demonstrating thatmale 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.
Abstract: 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.

Journal ArticleDOI
TL;DR: It is shown that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments, as well as to show a clear bimodal activity pattern for breeding birds only.
Abstract: Feral pigeons are descendants of wild rock pigeons that have adapted to the urban habitat. They have partially conserved the foraging behaviour of their wild ancestors (flights to agricultural areas) but have also developed new habits. Previous studies on the foraging strategies of feral pigeons have given various results, e.g. maximum distances reached by the pigeons (measured in a straight line from the resting places) differed between 0.3–0.5 km and 18–25 km. This study focuses on the spatio-temporal activity of feral pigeons in the urban habitat. We equipped 80 free-living feral pigeons from Basel, Switzerland with GPS receivers. We found three different foraging strategies for pigeons in Basel: (1) in the streets, squares and parks near the home loft, (2) in agricultural areas surrounding the city, (3) on docks and railway lines in harbours. The maximum distance reached by a pigeon was 5.29 km. More than 32% of the pigeons remained within 0.3 km of the home lofts and only 7.5% flew distances of more than 2 km. Females covered significantly longer distances than males, preferring to fly to more abundant and predictable food sources. Temporal activity patterns showed to be influenced by sex, breeding state and season. In contrast to wild rock pigeons and to feral pigeons in other cities, pigeons in Basel showed a clear bimodal activity pattern for breeding birds only. The differences between our results and those of other studies seem to be partly method-dependent, as the GPS-technique allows to record the pigeons’ localisations continuously in contrast to other methods. Other differences might be due to different kinds of food supply in the various cities. Our study shows that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments.

Journal ArticleDOI
TL;DR: It is predicted that the spatial segregation revealed by the present study is a result of selection favouring the avoidance of competition in these species through differential habitat use.
Abstract: The extent of spatial partitioning in insectivorous bats, whose prey is patchily distributed and transient in nature, remains a contentious issue. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and Pipistrellus pygmaeus, which are morphologically similar and sympatric, provides an opportunity to examine this question. The present study used radio telemetry to address the spatial distribution and foraging characteristics of P. pipistrellus and P. pygmaeus in northeast Scotland, to test the hypothesis that coexistence between these species is facilitated through spatial segregation. We reveal large and significant differences in the spatial distribution and foraging characteristics of these two cryptic species. Individual P. pipistrellus home ranges were on average three times as large as that of P. pygmaeus, and they foraged for approximately an hour longer each night. Inter-specific spatial overlap was minimal (<5%) and core foraging areas of either species were essentially mutually exclusive despite the proximity of the two roosts. Inter-specific differences in range size were associated with the spatial dispersion of productive foraging sites within individual foraging ranges. P. pipistrellus foraging sites were highly dispersed, necessitating larger ranges. It is predicted that the spatial segregation revealed by the present study is a result of selection favouring the avoidance of competition in these species through differential habitat use.

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TL;DR: It is concluded that surface hydrocarbons of eggs and the cuticular profiles of queens both signal queen fertility, suggesting a major role of fertility signals in the regulation of reproduction in social insects.
Abstract: In ant societies, workers do not usually reproduce but gain indirect fitness benefits from raising related offspring produced by the queen. One of the preconditions of this worker self-restraint is sufficient fertility of the queen. The queen is, therefore, expected to signal her fertility. In Camponotus floridanus, workers can recognize the presence of a highly fertile queen via her eggs, which are marked with the queen's specific hydrocarbon profile. If information on fertility is encoded in the hydrocarbon profile of eggs, we expect workers to be able to differentiate between eggs from highly and weakly fertile queens. We found that workers discriminate between these eggs solely on the basis of their hydrocarbon profiles which differ both qualitatively and quantitatively. This pattern is further supported by the similarity of the egg profiles of workers and weakly fertile queens and the similar treatment of both kinds of eggs. Profiles of queen eggs correspond to the cuticular hydrocarbon profiles of the respective queens. Changes in the cuticular profiles are associated with the size of the colony the queen originates from and her current egg-laying rate. However, partial correlation analysis indicates that only colony size predicts the cuticular profile. Colony size is a buffered indicator of queen fertility as it is a consequence of queen productivity within a certain period of time, whereas daily egg-laying rate varies due to cyclical oviposition. We conclude that surface hydrocarbons of eggs and the cuticular profiles of queens both signal queen fertility, suggesting a major role of fertility signals in the regulation of reproduction in social insects.

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TL;DR: The data suggests that wave structure has evolved through female choice in the fiddler crab Uca perplexa, and that male display rate and body size were not female choice cues.
Abstract: The fiddler crab Uca perplexa has a conspicuous male courtship signal that is directed at females to attract them to the male’s burrow for mating. The signal involves waving the unflexed large claw up and down. To determine whether the spatiotemporal structure of the wave is under selection by female choice, we examined whether females had a preference for any particular features of the wave. Females respond to a waving display by either visiting the male’s burrow entrance or by electing to pass without visiting the burrow. We filmed mate-searching females and the waving males that they visited or passed. We documented the wave structure of these males using frame-by-frame analysis. Males produce a two-part wave with component A preceding component B. Both components have an upstroke, a pause at the apex and a downstroke. The tip of the claw was raised much higher in B than in A. Visited males had a shorter delay between the two wave components than did males that the females passed without visiting. Visited males also produced component B waves that had a slower upstroke than those of passed males. There was a significant correlation between the relative height of the raised claw and the duration of the upstroke of component B. Females were selecting males that raised their major claw to the highest position (two to three times as high as the carapace width). Passed males brought down their major claw earlier and from a lower position than did visited males. The data suggests that wave structure has evolved through female choice. Male display rate and body size were not female choice cues. An earlier study showed that display duration was also not used by females in selecting mates.