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Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 409-419
Trill performance components vary with age, season, and motivation in the banded wren
S. L. Vehrencamp, J. Yantachka, M. L. Hall, S. R. de Kort

Acoustic displays with difficult-to-execute sounds are often subject to strong sexual selection because performance levels are related to the sender’s condition or genetic quality. Performance may also vary with age, breeding stage, and motivation related to social context. We focused on within-male variation in four components of trill performance in banded wren (Thryophilus pleurostictus) songs: note consistency, frequency bandwidth, note rate, and vocal deviation. The latter is a composite measure reflecting deviation from the performance limit on simultaneously maximizing both frequency bandwidth and note rate. We compared the changes in these song parameters at three time scales: over the course of years, across the breeding season, and at different times of the day with contrasting agonistic contexts. Vocal deviation decreased and note consistency increased between years, suggesting that experience may improve individual proficiency at singing trills. Consistency also increased across the season, confirming that practice is important for this parameter. Although there was no significant seasonal change in vocal deviation, one of its components, note rate, increased during the season. Neither vocal deviation nor consistency varied with agonistic context. However, note rate increased during playback experiments simulating territorial intrusions compared to dawn chorus singing. The magnitude of a male’s increase in note rate was positively correlated with his aggressive behavior during the playback experiment. Thus consistency, bandwidth, and vocal deviation indicate age, whereas trill rate flexibly indicates the singer’s aggressive motivation. We also found evidence of a within-male trade-off between vocal deviation and consistency.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 349-359
Seasonal changes in the structure of rhesus macaque social networks
Lauren J. N. Brent, Ann MacLarnon, Michael L. Platt, Stuart Semple
Social structure emerges from the patterning of interactions between individuals and plays a critical role in shaping some of the main characteristics of animal populations. The topological features of social structure, such as the extent to which individuals interact in clusters, can influence many biologically important factors, including the persistence of cooperation, and the rate of spread of disease. Yet, the extent to which social structure topology fluctuates over relatively short periods of time in relation to social, demographic, or environmental events remains unclear. Here, we use social network analysis to examine seasonal changes in the topology of social structures that emerge from socio-positive associations in adult female rhesus macaques (Macaca mulatta). Behavioral data for two different association types (grooming and spatial proximity) were collected for females in two free-ranging groups during two seasons: the mating and birth seasons. Stronger dyadic bonds resulted in social structures that were more tightly connected (i.e., of greater density) in the mating season compared to the birth season. Social structures were also more centralized around a subset of individuals and more clustered in the mating season than those in the birth season, although the latter differences were mostly driven by differences in density alone. Our results suggest a degree of temporal variation in the topological features of social structure in this population. Such variation may feed back on interactions, hence affecting the behaviors of individuals, and may therefore be important to take into account in studies of animal behavior.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 471-479
Denning behaviour of the European badger (Meles meles) correlates with bovine tuberculosis infection status
Nicola Weber, Stuart Bearhop, Sasha R. X. Dall, Richard J. Delahay, Robbie A. McDonald, Stephen P. Carter
Heterogeneities in behaviours of individuals may underpin important processes in evolutionary biology and ecology, including the spread of disease. Modelling approaches can sometimes fail to predict disease spread, which may partly be due to the number of unknown sources of variation in host behaviour. The European badger is a wildlife reservoir for bovine tuberculosis (bTB) in Britain and Ireland, and individual behaviour has been demonstrated to be an important factor in the spread of bTB among badgers and to cattle. Radio-telemetry devices were deployed on 40 badgers from eight groups to investigate patterns of den (sett) use in a high-density population, where each group had one or two main and three to eight outlier setts in their territory. Badgers were located at their setts for 28 days per season for 1 year to investigate how patterns differed between individuals. Denning behaviour may have a strong influence on contact patterns and the transmission of disease. We found significant heterogeneity, influenced by season, sex and age. Also, when controlling for these, bTB infection status interacting with season was highly correlated with sett use. Test-positive badgers spent more time away from their main sett than those that tested negative. We speculate that wider-ranging behaviour of test-positive animals may result in them contacting sources of infection more frequently and/or that their behaviour may be influenced by their disease status. Measures to control infectious diseases might be improved by targeting functional groups, specific areas or times of year that may contribute disproportionately to disease spread.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 519-528
Predator guild does not influence orangutan alarm call rates and combinations
Adriano R. Lameira, Han de Vries, Madeleine E. Hardus, Cedric P. A. Hall, Tatang Mitra-Setia, Berry M. Spruijt, Arik Kershenbaum, Elisabeth H. M. Sterck, Maria van Noordwijk, Carel van Schaik,Serge A. Wich
Monkey alarm calls have shown that in the primate clade, combinatorial rules in acoustic communication are not exclusive to humans. A recent hypothesis suggests that the number of different call combinations in monkeys increases with increased number of predator species. However, the existence of combinatorial rules in great ape alarm calls remains largely unstudied, despite its obvious relevance to ideas about the evolution of human speech. In this paper, we examine the potential use of combinatorial rules in the alarm calls of the only Asian great ape: the orangutan. Alarm calls in orangutans are composed of syllables (with either one or two distinct elements), which in turn are organized into sequences. Tigers and clouded leopards are predators for Sumatran orangutans, but in Borneo, tigers are extinct. Thus, orangutans make a suitable great ape model to assess alarm call composition in relation to the size of the predator guild. We exposed orangutans on both islands to a tiger and control model. Response compositionality was analyzed at two levels (i.e., syllable and syllable sequences) between models and populations. Results were corroborated using information theory algorithms. We made specific, directed predictions for the variation expected if orangutans used combinatorial rules. None of these predictions were met, indicating that monkey alarm call combinatorial rules do not have direct homologues in orangutans. If these results are replicated in other great apes, this indicates that predation did not drive selection towards ever more combinatorial rules in the human lineage.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 493-511
Environmental and social determinants of anuran lekking behavior: intraspecific variation in populations at thermal extremes
Diego Llusia, Rafael Márquez, Juan Francisco Beltrán, Catarina Moreira, José Pedro do Amaral
Environmental and social factors are critical to determine the timing and duration of lekking behavior since they provide species with signs to maximize benefits over costs in sexual displays. However, these factors have rarely been studied under different environmental conditions, and thus, it remains unclear whether exogenous factors affecting group displays show a general species-specific pattern or whether they are population-specific. Using audio-trapping techniques, we compared factors influencing the daily occurrence and duration of lekking behavior in two populations of Hyla molleri and two populations of Hyla meridionalis located at the thermal extremes (coldest vs. hottest) of their Iberian distribution range. From 12,240 hourly recordings over one season, multimodel inference revealed that the major determinants of chorus occurrence were similar between populations and species (i.e., chorus size the previous day, daytime air temperature, relative humidity, and barometric pressure), and accounted for 51–79 % of its deviance. In contrast, the major determinants of chorus duration differed between populations and species (i.e., chorus size, number of day, and air temperature and relative humidity at the onset of the chorus), and accounted for 38–69 % of its variance. Our findings suggest that the decision making related to lek attendance is environment-dependent, takes place at time between lekking events, and is associated with exogenous factors that may be both stable across species ranges and population-specific when populations are under different climatic conditions. This intraspecific variation might be underlain by plasticity mechanisms providing tree frogs with means to cope with changing environments. Moreover, social facilitation related to male-male acoustic competition seems to play a relevant role on the daily time invested by males in lek attendance.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 429-437
Socioecological correlates of inter-individual variation in orangutan diets at Ketambe, Sumatra
Madeleine E. Hardus, Han de Vries, David F. Dellatore, Adriano R. Lameira, Steph B. J. Menken, Serge A. Wich
The diet of great apes consists of several hundred plant species. The factors determining diet differences have been examined between populations but not within a population, probably due to the confounding effect of seasonal fluctuations on fruit availability. In Sumatran orangutans (Pongo abelii), fruit availability appears to be sufficiently high year round to have little influence on diet composition, which in turn allows for addressing this question. We examined the diet of eight adult female orangutans at Ketambe, Sumatra, and investigated whether fig and non-fig fruit availability, association time, and/or home range measures influenced dietary overlap between female dyads. Between most pairs, females’ diets were different: 16 out of 23 pairs had a significantly low diet species overlap. Only fig diet overlap was influenced (negatively) by the availability of non-fig fruit. Association time only influenced (positively) fig diet overlap. Hence, orangutans gathered in fig trees when non-fig fruit availability was low. Home range measures did not influence overall diet overlap. To our knowledge, this is the first study showing that, while controlling for confounding factors, individuals with similar energetic requirements, from the same population and sharing the same area, make different dietary choices relatively to their preferred (non-fig) fruit constituting the majority of their diet. Social transmission, with putative matrilineal diet traditions, suitably explains these results. We discuss the implications of the findings for orangutan conservation, namely on reintroduction and the felling of fig trees.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 361-372
Long-lasting effects of yolk androgens on phenotype in the pied flycatcher (Ficedula hypoleuca)
Suvi Ruuskanen, Esa Lehikoinen, Mikko Nikinmaa, Heli Siitari, Wolfgang Waser, Toni Laaksonen
The hormonal environment during early development, such as maternally derived androgens in bird eggs, shapes the development of the offspring in ways that may have important long-term consequences for phenotype and behavior and, ultimately, fitness. We studied the long-term effects of yolk androgens on several phenotypic and physiological traits in male and female pied flycatchers (Ficedula hypoleuca) by experimentally elevating yolk androgen levels and rearing birds in common-garden environment in captivity. We found that high yolk androgen levels increased the basal metabolic rates in both females and males in adulthood. High yolk androgen levels did not affect male melanin coloration or plumage ornaments, or timing or speed of moult in either sex. No effect of androgen treatment on cell-mediated or humoral immune response was found in either sex. Covariation among the measured phenotypic traits was further not altered by androgen treatment. Our results suggest that exposure to high androgen levels can have long-lasting effects on some offspring traits, but do not seem to lead to different phenotypes. Furthermore, the role of yolk androgens affecting sexually selected male traits in our study species seems to be minor. The fitness consequences of yolk androgen-induced higher metabolic rates remain to be studied.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 481-492
Are white-crowned sparrow badges reliable signals?
Zachary M. Laubach, Daniel T. Blumstein, L. Michael Romero, Greg Sampson, Johannes Foufopoulos
Status badges, such as crown plumage, mediate intraspecific interactions. The reliability of crown morphology as a status badge in male mountain white-crowned sparrows (Zonotrichia leucophrys oriantha) is uncertain. We examined morphological and physiological correlates of the proportion of crown that was white (“crown-white”) in 178 male mountain white-crowned sparrows during the 2008–2009 breeding seasons. Using a paired experimental design, we presented territory-holding males with white-enhanced and white-reduced decoys and recorded aggressive behaviors. To assess physiological constraints on signal bluffing, a subsample of birds was captured and released after manipulating natural crowns to simulate bluffed white-enhanced or white-reduced crowns; corticosterone concentrations were assayed from blood drawn upon recapture and after a restraint-induced stressor. We found a significant positive association between crown-white and a measure of body size—tarsus length—which is an established indicator of resource-holding potential. In the decoy challenge, males responded more aggressively toward white-enhanced than white-reduced decoys. In the hormone experiment, white-enhanced birds had higher baseline corticosterone levels, whereas white-reduced birds had similar concentrations to controls. Furthermore, white-enhanced birds had an attenuated restraint-induced corticosterone response, while white-reduced birds mounted a significantly larger increase in corticosterone than controls. Taken together, these findings indicate that crown-white is a reliable status badge of resource-holding potential in male mountain white-crowned sparrows during the breeding season.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 383-393
Alternative strategies in avian scavengers: how subordinate species foil the despotic distribution
Corinne J. Kendall
Trade-offs in species’ traits can mediate competition and enable coexistence. A key challenge in ecology is understanding the role of species’ trade-offs in maintaining diversity, and evolutionary trade-offs between the abilities of competing species are best understood by considering how competitive advantages change along an environmental gradient. Previous studies of such trade-offs are generally limited to two-species systems and a single trade-off. In this study, I consider the effect of trade-offs in search efficiency and competitive abilities on habitat use patterns among a diverse avian scavenger guild. I hypothesize that species’ dominance status and search efficiency will both be correlated with patch quality. Using counts of searching birds in areas that vary in habitat quality in terms of both wildlife and human settlement density and observations at experimental carcasses, I assess the competitive ability, search efficiency, and habitat use of seven avian scavenger species in Masai Mara National Reserve, Kenya. Findings support the hypothesis with Bateleurs, a species with high search efficiency, and Ruppell’s, Lappet-faced, and White-backed vultures, species with high individual or social dominance, preferentially exploiting habitats of high quality, while Tawny eagles and Hooded vultures, species with low search efficiency and competitive ability, prefer habitats of low quality. This paper demonstrates the importance of considering multiple strategies for assessing the effect of competition on habitat use within complex communities.

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 421-428
Mate availability accelerates male filial cannibalism in a nest brooding fish: effects of number and fecundity of females
Tomohiro Takeyama, Naoko Namizaki, Masanori Kohda
Theoretical models predict that increased mate availability accelerates filial cannibalism by the parental male, but we do not yet fully understand how the various aspects of mate availability contribute to this effect. We examined the effects of two elements of mate availability—female fecundity and sex ratio—on filial cannibalism by the lizard goby, Rhinogobius flumineus, which is a paternal nest brooding fish. We used three types of females (stimulus-females): a single female with slim belly (not ready to spawn), a single gravid female (ready to spawn), and two gravid females. Stimulus-females were housed in a transparent cage and shown to subject males just before and after spawning with a separate female, after which males cared for the brood. A single gravid stimulus-female accelerated filial cannibalism, compared with a control-stimulus consisting of an empty cage, but only during the early care period. In contrast, a single slim-bellied stimulus-female did not accelerate cannibalism. A stimulus of two gravid females accelerated cannibalism to the same degree as a single gravid female stimulus. Our results suggest that in lizard gobies, filial cannibalism by parental males is accelerated by female quality (fecundity) in the early care period, but not by a higher number of available females (sex ratio).

Behavioral Ecology and Sociobiology
March 2013, Volume 67, Issue 3, pp 513-518
Consequences of paternal care on pectoral fin allometry in a desert-dwelling fish
Emile van Lieshout, P. Andreas Svensson, Bob B. M. Wong
Positive static allometry is a scaling relationship where the relative size of traits covaries with adult body size. Traditionally, positive allometry is thought to result from either altered physiological requirements at larger body size or from strongly condition-dependent allocation under sexual selection. Yet, there are no theoretical reasons why positive allometry cannot evolve in fitness-related traits that are solely under the influence of natural selection. We investigated scaling and sexual dimorphism of a naturally selected trait, pectoral fin size, in comparison to a trait important in male–male combat, head width in natural populations of a fish, the desert goby Chlamydogobius eremius. Male desert gobies provide uniparental care and use their pectoral fins to fan the brood (often under hypoxic conditions); hence, larger fins are expected to be more efficient. Male pectoral fins do not appear to fulfil a signalling function in this species. We found that, for both pectoral fin size and head width, males exhibited positive allometric slopes and greater relative trait size (allometric elevation) than females. However, for head width, females also showed positive allometry, albeit to a lesser degree than males. Because fin locomotory function typically does not result in positive allometry, our findings indicate that other naturally selected uses, such as paternal care, can exaggerate trait scaling relationships.

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