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1.  Lethal combat over limited resources: testing the importance of competitors and kin 
Although most animals employ strategies to avoid costly escalation of conflict, the limitation of critical resources may lead to extreme contests and fatal fighting. Evolutionary theories predict that the occurrence and intensity of fights can be explained by resource value and the density and relatedness of competitors. However, the interaction between these factors and their relative importance often remains unclear; moreover, few systems allow all variables to be experimentally investigated, making tests of these theoretical predictions rare. Here, we use the parasitoid wasp Melittobia to test the importance of all these factors. In contrast to predictions, variation in contested resource value (female mates) and the relatedness of competitors do not influence levels of aggression. However, as predicted, fight intensity increased with competitor density and was not influenced by the greater cost of fighting at high density. Our results suggest that in the absence of kin recognition, indirectly altruistic behavior (spite) is unlikely to evolve, and in such circumstances, the scale of competition will strongly influence the amount of kin discrimination in the form of level of aggression as observed in Melittobia species.
doi:10.1093/beheco/arq209
PMCID: PMC3947730  PMID: 24619384
fatal fighting; kin discrimination; Melittobia; relatedness; resource competition; spite
2.  Spatial movements and social networks in juvenile male song sparrows 
Behavioral Ecology  2011;23(1):141-152.
The time between fledging and breeding is a critical period in songbird ontogeny, but the behavior of young songbirds in the wild is relatively unstudied. The types of social relationships juveniles form with other individuals can provide insight into the process through which they learn complex behaviors crucial for survival, territory establishment, and mate attraction. We used radio telemetry to observe social associations of young male song sparrows (Melospiza melodia) from May to November. Juvenile song sparrows were frequently observed in social flocks and generally associated with more birds in the summer than in the autumn months. Most juvenile subjects formed stable social relationships with other birds and were seen with the same individual on up to 60% of the days observed. The strongest associations occurred with other juvenile males, and these individuals were often seen <1 m from the subject, even when the subject moved large distances between tracking observations. Associations also had long-term behavioral consequences as subjects were more likely to establish territories near their associates and learn shared song types. Our results indicate that male song sparrows spend a large percentage of the juvenile life stage forming social relationships and suggest that these associations may be important for the ecology of young birds and the ontogeny of their behaviors.
doi:10.1093/beheco/arr167
PMCID: PMC3242974  PMID: 22479140
3.  Intrasexual competition in females: evidence for sexual selection? 
Behavioral Ecology  2011;22(6):1131-1140.
In spite of recent interest in sexual selection in females, debate exists over whether traits that influence female–female competition are sexually selected. This review uses female–female aggressive behavior as a model behavioral trait for understanding the evolutionary mechanisms promoting intrasexual competition, focusing especially on sexual selection. I employ a broad definition of sexual selection, whereby traits that influence competition for mates are sexually selected, whereas those that directly influence fecundity or offspring survival are naturally selected. Drawing examples from across animal taxa, including humans, I examine 4 predictions about female intrasexual competition based on the abundance of resources, the availability of males, and the direct or indirect benefits those males provide. These patterns reveal a key sex difference in sexual selection: Although females may compete for the number of mates, they appear to compete more so for access to high-quality mates that provide direct and indirect (genetic) benefits. As is the case in males, intrasexual selection in females also includes competition for essential resources required for access to mates. If mate quality affects the magnitude of mating success, then restricting sexual selection to competition for quantity of mates may ignore important components of fitness in females and underestimate the role of sexual selection in shaping female phenotype. In the future, understanding sex differences in sexual selection will require further exploration of the extent of mutual intrasexual competition and the incorporation of quality of mating success into the study of sexual selection in both sexes.
doi:10.1093/beheco/arr106
PMCID: PMC3199163  PMID: 22479137
aggression; female competition; intrasexual selection; mating success; sexual selection
4.  By any name, female–female competition yields differential mating success 
Behavioral Ecology  2011;22(6):1144-1146.
doi:10.1093/beheco/arr111
PMCID: PMC3199164  PMID: 22479138
female aggression; female competition; sexual selection
5.  What is sexual selection and the short herstory of female trait variation 
Behavioral Ecology  2011;22(6):1146-1147.
doi:10.1093/beheco/arr113
PMCID: PMC3199165  PMID: 22479139
sexual selection
6.  Resource quality or competition: why increase resource acceptance in the presence of conspecifics? 
Behavioral Ecology  2011;22(4):730-737.
Some animal species increase resource acceptance rates in the presence of conspecifics. Such responses may be adaptive if the presence of conspecifics is a reliable indicator of resource quality. Similarly, these responses could represent an adaptive reduction in choosiness under high levels of scramble competition. Although high resource quality and high levels of scramble competition should both favor increased resource acceptance, the contexts in which the increase occurs should differ. In this paper, we tested the effect of social environment on egg-laying and aggressive behavior in the walnut fly, Rhagoletis juglandis, in multiple contexts to determine whether increased resource acceptance in the presence of conspecifics was better viewed as a response to increased host quality or increased competition. We found that grouped females oviposit more readily than isolated females when provided small (low-quality) artificial hosts but not when provided large (high-quality) artificial hosts, indicating that conspecific presence reduces choosiness. Increased resource acceptance was observed even when exposure to conspecifics was temporally or spatially separate from exposure to the resource. Finally, we found that individuals showed reduced aggression after being housed in groups, as expected under high levels of scramble competition. These results indicate that the pattern of resource acceptance in the presence of conspecifics may be better viewed as a response to increased scramble competition rather than as a response to public information about resource quality.
doi:10.1093/beheco/arr042
PMCID: PMC3117901  PMID: 22479135
conspecific attraction; experience; host choice; Rhagoletis; social facilitation; social information
7.  Progressive parenting behavior in wild golden lion tamarins 
Behavioral Ecology  2011;22(4):745-754.
Young primates in the family Callitrichidae (the marmosets and tamarins) receive extensive and relatively prolonged care from adults. Of particular note, callitrichid young are routinely provisioned until well after weaning by parents and helpers, which is in stark contrast to typical juvenile primates, who must acquire most of their food independently once they are weaned. Adults of some callitrichid species produce a specialized vocalization that encourages immature group members to take proffered food from the caller. Here, I report that wild adult golden lion tamarins (Leontopithecus rosalia) not only used this food-offering call to encourage young, mobile offspring to approach and take captured prey from them, but as the young began to spend significant time foraging for themselves and to acquire prey by independent means, the frequency of these vocalizations in the context of food transfer declined. Adults then began to use food-offering calls in a novel context: to direct juveniles to foraging sites that contained hidden prey that the adults had found but not captured. During the period of these most frequent adult-directed prey captures, the independent prey-capture success rates of juveniles improved. Thus, adults modified their provisioning behavior in a progressive developmentally sensitive manner that may have facilitated learning how to find food. I hypothesize that as a result of these demonstrations by adults, juveniles either may be encouraged to continue foraging despite low return rates or to learn the properties of productive prey-foraging substrates in a complex environment.
doi:10.1093/beheco/arr055
PMCID: PMC3117902  PMID: 22479136
golden lion tamarin; infant development; parenting behavior; prey foraging; provisioning; teaching
8.  Linking amphibian call structure to the environment: the interplay between phenotypic flexibility and individual attributes 
Behavioral Ecology  2011;22(3):520-526.
The structure of the environment surrounding signal emission produces different patterns of degradation and attenuation. The expected adjustment of calls to ensure signal transmission in an environment was formalized in the acoustic adaptation hypothesis. Within this framework, most studies considered anuran calls as fixed attributes determined by local adaptations. However, variability in vocalizations as a product of phenotypic expression has also been reported. Empirical evidence supporting the association between environment and call structure has been inconsistent, particularly in anurans. Here, we identify a plausible causal structure connecting environment, individual attributes, and temporal and spectral adjustments as direct or indirect determinants of the observed variation in call attributes of the frog Hypsiboas pulchellus. For that purpose, we recorded the calls of 40 males in the field, together with vegetation density and other environmental descriptors of the calling site. Path analysis revealed a strong effect of habitat structure on the temporal parameters of the call, and an effect of site temperature conditioning the size of organisms calling at each site and thus indirectly affecting the dominant frequency of the call. Experimental habitat modification with a styrofoam enclosure yielded results consistent with field observations, highlighting the potential role of call flexibility on detected call patterns. Both, experimental and correlative results indicate the need to incorporate the so far poorly considered role of phenotypic plasticity in the complex connection between environmental structure and individual call attributes.
doi:10.1093/beheco/arr011
PMCID: PMC3078827  PMID: 22479134
acoustic adaptation hypothesis; call adjustment; Hypsiboas pulchellus; local adaptation; phenotypic plasticity; scale
9.  Colony variation in the collective regulation of foraging by harvester ants 
Behavioral Ecology  2011;22(2):429-435.
This study investigates variation in collective behavior in a natural population of colonies of the harvester ant, Pogonomyrmex barbatus. Harvester ant colonies regulate foraging activity to adjust to current food availability; the rate at which inactive foragers leave the nest on the next trip depends on the rate at which successful foragers return with food. This study investigates differences among colonies in foraging activity and how these differences are associated with variation among colonies in the regulation of foraging. Colonies differ in the baseline rate at which patrollers leave the nest, without stimulation from returning ants. This baseline rate predicts a colony's foraging activity, suggesting there is a colony-specific activity level that influences how quickly any ant leaves the nest. When a colony's foraging activity is high, the colony is more likely to regulate foraging. Moreover, colonies differ in the propensity to adjust the rate of outgoing foragers to the rate of forager return. Naturally occurring variation in the regulation of foraging may lead to variation in colony survival and reproductive success.
doi:10.1093/beheco/arq218
PMCID: PMC3071749  PMID: 22479133
behavioral reaction norm; behavioral syndrome; individual variation
10.  Birds flee en mass from New Year’s Eve fireworks 
Behavioral Ecology  2011;22(6):1173-1177.
Anthropogenic disturbances of wildlife, such as noise, human presence, hunting activity, and motor vehicles, are becoming an increasing concern in conservation biology. Fireworks are an important part of celebrations worldwide, and although humans often find fireworks spectacular, fireworks are probably perceived quite differently by wild animals. Behavioral responses to fireworks are difficult to study at night, and little is known about the negative effects fireworks may have on wildlife. Every year, thousands of tons of fireworks are lit by civilians on New Year’s Eve in the Netherlands. Using an operational weather radar, we quantified the reaction of birds to fireworks in 3 consecutive years. Thousands of birds took flight shortly after midnight, with high aerial movements lasting at least 45 min and peak densities measured at 500 m altitude. The highest densities were observed over grasslands and wetlands, including nature conservation sites, where thousands of waterfowl rest and feed. The Netherlands is the most important winter staging area for several species of waterfowl in Europe. We estimate that hundreds of thousands of birds in the Netherlands take flight due to fireworks. The spatial and temporal extent of disturbance is substantial, and potential consequences are discussed. Weather radar provides a unique opportunity to study the reaction of birds to fireworks, which has otherwise remained elusive.
doi:10.1093/beheco/arr102
PMCID: PMC3199162  PMID: 22476363
birds; disturbance; fireworks; flight; Natura 2000; radar; waterfowl
11.  Water-seeking behavior in worm-infected crickets and reversibility of parasitic manipulation 
Behavioral Ecology  2011;22(2):392-400.
One of the most fascinating examples of parasite-induced host manipulation is that of hairworms, first, because they induce a spectacular “suicide” water-seeking behavior in their terrestrial insect hosts and, second, because the emergence of the parasite is not lethal per se for the host that can live several months following parasite release. The mechanisms hairworms use to increase the encounter rate between their host and water remain, however, poorly understood. Considering the selective landscape in which nematomorph manipulation has evolved as well as previously obtained proteomics data, we predicted that crickets harboring mature hairworms would display a modified behavioral response to light. Since following parasite emergence in water, the cricket host and parasitic worm do not interact physiologically anymore, we also predicted that the host would recover from the modified behaviors. We examined the effect of hairworm infection on different behavioral responses of the host when stimulated by light to record responses from uninfected, infected, and ex-infected crickets. We showed that hairworm infection fundamentally modifies cricket behavior by inducing directed responses to light, a condition from which they mostly recover once the parasite is released. This study supports the idea that host manipulation by parasites is subtle, complex, and multidimensional.
doi:10.1093/beheco/arq215
PMCID: PMC3071748  PMID: 22476265
behavior; insects; nematomorph; parasite manipulation; parasitism; phototaxis
12.  Increased host aggression as an induced defense against slave-making ants 
Behavioral Ecology  2011;22(2):255-260.
Slave-making ants reduce the fitness of surrounding host colonies through regular raids, causing the loss of brood and frequently queen and worker death. Consequently, hosts developed defenses against slave raids such as specific recognition and aggression toward social parasites, and indeed, we show that host ants react more aggressively toward slavemakers than toward nonparasitic competitors. Permanent behavioral defenses can be costly, and if social parasite impact varies in time and space, inducible defenses, which are only expressed after slavemaker detection, can be adaptive. We demonstrate for the first time an induced defense against slave-making ants: Cues from the slavemaker Protomognathus americanus caused an unspecific but long-lasting behavioral response in Temnothorax host ants. A 5-min within-nest encounter with a dead slavemaker raised the aggression level in T. longispinosus host colonies. Contrarily, encounters with nonparasitic competitors did not elicit aggressive responses toward non-nestmates. Increased aggression can be adaptive if a slavemaker encounter reliably indicates a forthcoming attack and if aggression increases postraid survival. Host aggression was elevated over 3 days, showing the ability of host ants to remember parasite encounters. The response disappeared after 2 weeks, possibly because by then the benefits of increased aggression counterbalance potential costs associated with it.
doi:10.1093/beheco/arq191
PMCID: PMC3071747  PMID: 22476194
aggression; behavior; parasites; phenotypic plasticity; social insects

Results 1-12 (12)