Road traffic has a major impact on the environment and numerous studies have shown how various species respond to road traffic. However, a clear picture based on large-scale studies of how complete species communities in different environments are affected by road traffic is lacking. We assessed the distance within which road traffic negatively impacts breeding bird densities and the extent to which effect distances differ between species and habitats. We also tested how traffic intensity is associated with effect distances. We used a large dataset from protocolled breeding bird surveys of > 140,000 territories of complete breeding bird communities within 1500 m from motorways across the Netherlands. We examined effect distances of road traffic on breeding density of individual species, for various ecological species groups and for all species together. Finally, we tested how road traffic intensity influences effect distances. Road traffic had a consistent negative effect on territory densities up to approximately 650 m distance from the road for different species and ecological species groups. Within road-effect zones, densities of breeding birds were significantly lower (on average 25

Context Road infrastructure construction is integral to economic development, but negatively affects biodiversity. To mitigate the negative impacts of infrastructure, various types of wildlife crossings are realized worldwide, but little is known about their effectiveness, and cost-effectiveness. Objective The paper contributes to the methodological and empirical discussion on the effectiveness of wildlife crossings for enhancing the quality of surrounding nature and its cost-effectiveness by analyzing a large-scale wildlife-crossings program in the Netherlands. Method A multi-criteria cost–benefit analysis is applied, comprised of monetary and non-monetary measures, and a mixed-method approach is used to determine ecological effects. Ecological effects are expressed in the standardized weighted hectare measurement of threat-weighted ecological quality area (1 T-EQA = 1 ha of 100% ecological quality, averagely threatened). Cost-effectiveness is calculated comparing the monetary costs of intervention with ecological benefits (Euro costs/T-EQA), for different types of wildlife crossings and for two other nature policies. Results The Dutch habitat defragmentation program has induced an increase in nature value of 1734 T-EQA at a cost of Euro 283 million. Ecological gains per hierarchically ordered groups of measures differ strongly: The most effective are ecoducts (wildlife crossing bridges) followed by shared-use viaducts and large fauna tunnels. Ecoducts generated the largest gain in nature value, but were also the most costly measures. In terms of cost-effectiveness, both large fauna tunnels and shared-use viaducts for traffic and animals outperformed ecoducts. Conclusions Ecoducts deliver ecologically, but their cost-effectiveness appears modest. Purchasing agricultural land for restoration of nature appears more cost-effective than building wildlife crossings. Yet, reducing environmental pressures or their effects on existing nature areas is likely to be most cost-effective.

In most passerine birds, individuals attempt to maximize their fitness by providing parental care while also mating outside their pair bond. A sex-specific trade-off between these two behaviours is predicted to occur, as the fitness benefits of extra-pair mating differ between the sexes. We use nest observations and parentage analysis to reveal a negative association between male care and the incidence of extra-pair paternity across three species of penduline tit (Remizidae). This provides evidence of a trade-off between these two behaviours, possibly due to the devaluing of paternal care by extra-pair offspring.

Penduline tits (Remiz spp.) are renowned for their diverse mating and parenting strategies, and are a well‐studied system by behavioural ecologists. However, the phylogenetic relationships and species delimitations within this genus are poorly understood. Here, we investigate phylogenetic relationships within the genus Remiz by examining the genetic variation in the mitochondrial cytochrome‐b gene of 64 individuals and in ten autosomal microsatellite markers from 44 individuals. The taxon sampling includes individuals from all currently recognized species (R. pendulinus, R. macronyx, R. coronatus, and R. consobrinus) and most subspecies in the Palearctic region. We showed that R. coronatus and R. consobrinus are genetically well differentiated and constitute independent evolutionary lineages, separated from each other and from R. pendulinus/macronyx. However, we found no evidence for significant differentiation among R. pendulinus/macronyx individuals in mtDNA haplotypes and only marginal differences between R. pendulinus and R. macronyx in microsatellite markers. Hence, based on present data our recommendation is to treat R. pendulinus and R. macronyx as conspecific and R. coronatus and R. consobrinus as separate species.

Traits used in communication, such as colour signals, are expected to have positive consequences for reproductive success, but their associations with survival are little understood. Previous studies have mainly investigated linear relationships between signals and survival, but both hump-shaped and U-shaped relationships can also be predicted, depending on the main costs involved in trait expression. Furthermore, few studies have taken the plasticity of signals into account in viability selection analyses. The relationship between signal expression and survival is of particular interest in melanin-based traits, because their main costs are still debated. Here, we first determined the main factors explaining variability in a melanin-based trait linked to dominance: the bib size of a colonial bird, the sociable weaver Philetairus socius. We then used these analyses to obtain a measure representative of the individual mean expression of bib size. Finally, we used capture-recapture models to study how survival varied in relation to bib size. Variation in bib size was strongly affected by year and moderately affected by age, body condition and colony size. In addition, individuals bearing small and large bibs had higher survival than those with intermediate bibs, and this U-shaped relationship between survival and bib size appeared to be more pronounced in some years than others. These results constitute a rare example of disruptive viability selection, and point towards the potential importance of social costs incurred by the dominance signalling function of badges of status.

Sexual selection determines the elaboration of morphological and behavioural traits and thus drives the evolution of phenotypes. Sexual selection on males and females can differ between populations, especially when populations exhibit different breeding systems. A substantial body of literature describes how breeding systems shape ornamentation across species, with a strong emphasis on male ornamentation and female preference. However, whether breeding system predicts ornamentation within species and whether similar mechanisms as in males also shape the phenotype of females remains unclear. Here, we investigate how different breeding systems are associated with male and female ornamentation in five geographically distinct populations of Kentish plovers Charadrius alexandrinus. We predicted that polygamous populations would exhibit more elaborate ornaments and stronger sexual dimorphism than monogamous populations. By estimating the size and intensity of male (n = 162) and female (n = 174) melanin-based plumage ornaments, i.e. breast bands and ear coverts, we show that plumage ornamentation is predicted by breeding system in both sexes. A difference in especially male ornamentation between polygamous (darker and smaller ornaments) and monogamous (lighter and larger) populations causes the greatest sexual dimorphism to be associated with polygamy. The non-social environment, however, may also influence the degree of ornamentation, for instance through availability of food. We found that, in addition to breeding system, a key environmental parameter, rainfall, predicted a seasonal change of ornamentation in a sex-specific manner. Our results emphasise that to understand the phenotype of animals, it is important to consider both natural and sexual selection acting on both males and females.

Melanin‐based plumage ornaments have been shown to play an important role in male–male competition, but also influence inter‐sexual communication. Consequently, ornaments may be associated with reproductive effort of both males and females. Females mated to males with larger melanin ornaments may acquire access to better territories or benefit from increased paternal care. Here we investigated whether the melanin‐based breast‐band of male and female Bar‐throated ApalisApalis thoracicais a signal of information about its bearer and is associated with male and female reproductive effort. Breast‐band size was a highly variable morphometric trait in both sexes, but only in males was it associated with body mass. We then assessed whether male and female breast‐band size predicted maternal and paternal investment. Egg mass increased with male breast‐band size, but decreased with female breast‐band size. Whether females adjust maternal hormone allocation in response to their partner's ornamentation remains a contentious issue. We found that yolk testosterone and androstenedione concentrations were not predicted by male ornamentation or body mass. Finally, males with larger breast‐bands provided their mates with more food, allowing those females to spend more time incubating. Reproductive effort of both parents is therefore predicted by their own and their mate's ornamentation in Bar‐throated Apalis, and thus breast‐band size potentially acts as a signal of reproductive performance in both sexes. These results highlight the need for more comprehensive analyses of a relationship between melanin‐based ornaments and fitness, incorporating multiple behavioural variables associated with reproductive effort.

Dispersal is a critical driver of gene flow, with important consequences for population genetic structure, social interactions and other biological processes. Limited dispersal may result in kin-structured populations in which kin selection may operate, but it may also increase the risk of kin competition and inbreeding. Here, we use a combination of long-term field data and molecular genetics to examine dispersal patterns and their consequences for the population genetics of a highly social bird, the sociable weaver (Philetairus socius), which exhibits cooperation at various levels of sociality from nuclear family groups to its unique communal nests. Using 20 years of data, involving capture of 6508 birds and 3151 recaptures at 48 colonies, we found that both sexes exhibit philopatry and that any dispersal occurs over relatively short distances. Dispersal is female-biased, with females dispersing earlier, further, and to less closely related destination colonies than males. Genotyping data from 30 colonies showed that this pattern of dispersal is reflected by fine-scale genetic structure for both sexes, revealed by isolation by distance in terms of genetic relatedness and significant genetic variance among colonies. Both relationships were stronger among males than females. Crucially, significant relatedness extended beyond the level of the colony for both sexes. Such fine-scale population genetic structure may have played an important role in the evolution of cooperative behaviour in this species, but it may also result in a significant inbreeding risk, against which female-biased dispersal alone is unlikely to be an effective strategy.

In animal societies, individuals face the dilemma of whether to cooperate or to compete over a shared resource. Two intertwined mechanisms may help to resolve this enduring evolutionary dilemma by preventing conflicts and thereby mediating the costs of living in groups: the establishment of dominance hierarchies and the use of 'badge-of-status' for signalling dominance. We investigated these two mechanisms in the sociable weaver (Philetairus socius), a colonial and social passerine which cooperates over multiple tasks. We examined the sociable weavers' dominance structure in 2 years by recording 2563 agonistic interactions between 152 individuals observed at a feeder at eight colonies. We tested which individual traits, including sex, age, relatedness and two melanin-based plumage traits, predicted variation in social status. First, using social network analysis, we found that colonies were structured by strongly ordered hierarchies which were stable between years. Second, medium-ranked birds engaged more in aggressive interactions than highly ranking individuals, suggesting that competition over food is most pronounced among birds of intermediate social status. Third, we found that colony size and kinship influenced agonistic interactions, so aggression was less pronounced in smaller colonies and among relatives. Finally, within- and between-individual variation in social status and the presence of an individual at the feeder were associated with variation in bib size, as predicted by the badge-of-status hypothesis. These results suggest that dominance hierarchies and bib size mediate conflicts in sociable weaver societies.

The tragedy of the commons predicts social collapse when public goods are jointly exploited by individuals attempting to maximize their fitness at the expense of other social group members. However, animal societies have evolved many times despite this vulnerability to exploitation by selfish individuals. Kin selection offers a solution to this social dilemma, but in large social groups mean relatedness is often low. Sociable weavers (Philetairus socius) live in large colonies that share the benefits of a massive communal nest, which requires individual investment for construction and maintenance. Here, we show that despite low mean kinship within colonies, relatives are spatially and socially clustered and that nest-building males have higher local relatedness to other colony members than do non-building males. Alternative hypotheses received little support, so we conclude that the benefits of the public good are shared with kin and that cooperative investment is, despite the large size and low relatedness of these communities, kin directed.

Maternal effects can influence offspring phenotype with short- and long-term consequences. Yet, how the social environment may influence egg composition is not well understood. Here, we investigate how laying order and social environment predict maternal effects in the sociable weaver, Philetairus socius, a species that lives in massive communal nests which may be occupied by only a few to 100+ individuals in a single nest. This range of social environments is associated with variation in a number of phenotypic and life-history traits. We investigate whether maternal effects are adjusted accordingly. We found no evidence for the prediction that females might benefit from modifying brood hierarchies through an increased deposition of androgens with laying order. Instead, females appear to exacerbate brood reduction by decreasing the costly production of yolk mass and antioxidants with laying order. Additionally, we found that this effect did not depend on colony size. Finally, in accordance with an expected increased intensity of environmental stress with increasing colony size, we found that yolk androgen concentration increased with colony size. This result suggests that females may enhance the competitive ability of offspring raised in larger colonies, possibly preparing the offspring for a competitive social environment.

Structures built by animals, such as nests, mounds and burrows, are often the product of cooperative investment by more than one individual. Such structures may be viewed as a public good, since all individuals that occupy them share the benefits they provide. However, access to the benefits generated by the structure may vary among individuals and is likely to be an important determinant of social organisation. Here we use the massive, communal nests of sociable weavers Philetairus socius, to investigate whether their thermoregulatory function varies in relation to the size of communal nests, and the position of individual nest chambers within the communal structure. We then examine whether this spatial variation in thermoregulatory function predicts the social organisation of colonies. First, we show that the sociable weavers' communal nests buffer variation in ambient temperature, and reduce temperature variability within nest chambers. The extent of this buffering effect depends significantly on the position of nest chambers within the communal structure, and on the depth to which chambers are embedded within the nest mass. We detected no effect of nest volume on thermoregulatory benefits, suggesting that there are likely to be additional, non-thermoregulatory benefits leading to communal nests. Finally, our results indicate that there may be competition for access to the benefits of the public good, since older birds occupied the chambers with the highest thermoregulatory benefits, where breeding activity was also more common. We discuss how the spatial structure of the benefits of the public good might influence social organisation in the unique communal lifestyle of sociable weavers.

Elaborate male song may restrain competitors in various songbirds, although the exact mechanism, information content and information flow of acoustic signals are not completely understood. Here we focus on the interactions between resident and intruder males using the Eurasian penduline tit Remiz pendulinus. The breeding system of this small passerine bird is unusually variable including sequential polygamy by both sexes, and appears to be driven by intense sexual selection and sexual conflict over parental care. We had two objectives in this study: (i) to investigate whether male repertoire size is an intra-sexual signal in penduline tits, and (ii) to determine emitter and receiver roles during acoustic communication between residents and intruders. We tested these objectives in a natural population by recording the resident males' song and then challenging them by song playbacks. The residents' approach to intruder stimuli and behavioural responses were monitored. We found that intruder repertoire size did not evoke different responses by the residents. Rather, the resident's approach distance and behavioural response was predicted by his own repertoire size, suggesting that song in penduline tits is involved in male-male communication, and repertoire size may function as a reliable signal of the resident male's ability and willingness to defend his nest and avert possible intruders.

Diverse patterns of parental care, including uniparental care by either the male or the female, provide excellent opportunities to investigate how variation in social traits is maintained in wild populations. Coexistence of different parental strategies within the same population is expected when they exhibit similar cost-benefit ratios. We investigated one of the most diverse avian breeding systems and compared parental behavior and reproductive output between nests that are uniparentally cared for by male or female Eurasian Penduline Tits (Remiz pendulinus). In this small passerine bird, full care (incubation and brood care) is provided by the male only (7-18% of nests) or the female only (48-65% of nests). Additionally, a third of all nests are deserted by both parents as a result of intense sexual conflict over care. We show that parental effort is similar between male and female parents, and they achieve similar offspring survival after controlling for clutch-size differences. Females care for larger clutches than males (5.7 +/- 0.1 [SE] and 3.5 +/- 0.2 eggs, respectively), and they produce more nestlings (3.6 +/- 0.2 and 2.4 +/- 0.3 nestlings, respectively), although this difference is not significant at fledging of the chicks (3.1 +/- 0.2 and 2.4 +/- 0.3 young, respectively). Therefore, sex difference in care does not explain the bias toward female-only care in Eurasian Penduline Tits. We propose that the differential benefits of desertion and/or the differential costs of care for males and females facilitate the existence of both uniparental care types in the population. Received 18 April 2012, accepted 2 July 2012.

Parental care is costly, and in many organisms, the male or the female parent benefits from reducing its own care which may be compensated for by its mate. One of the parents may even face all costs of parental care if its mate deserts and leaves him/her to care for the offspring alone. Theoretical models have generated contrasting predictions as to how parents negotiate a resolution of this sexual conflict over care, although empirical tests are largely lacking. We investigated pre-desertion behaviour (nest attendance) of a highly polygamous passerine, the Eurasian penduline tit ( Remiz pendulinus) that exhibits intense sexual conflict over care culminating in clutch desertion by the male, by the female or by both parents. We conjectured that nest attendance of parents should predict desertion, so that the lack of care provided by the deserting parent may be compensated for. By analysing over 200 000 video frames (2.50 +/- 1.36 d per pair, 302 +/- 170 min/d) of 20 pairs, we show that nest desertion is not a gradual process and cannot be predicted by nest attendance. These results are consistent with the argument that the predictable desertion may not be evolutionarily stable, and suggest that male and female penduline tits do not negotiate clutch desertion.

Supplementary data tml http://rspb.royalsocietypublishing.org/content/suppl/2011/12/15/rspb.2011.2297.DC1.h "Data Supplement" References ml#ref-list-1 http://rspb.royalsocietypublishing.org/content/early/2011/12/15/rspb.2011.2297.full.ht This article cites 44 articles, 10 of which can be accessed free P<P Published online 21 December 2011 in advance of the print journal. Email alerting service here right-hand corner of the article or click Receive free email alerts when new articles cite this article sign up in the box at the top

Eurasian penduline tits (Remiz pendulinus) have an unusually diverse breeding system consisting of frequent male and female polygamy, and uniparental care by the male or the female. Intriguingly, 30 to 40 per cent of all nests are deserted by both parents. To understand the evolution of this diverse breeding system and frequent clutch desertion, we use 6 years of field data to derive fitness expectations for males and females depending on whether or not they care for their offspring. The resulting payoff matrix corresponds to an asymmetric Snowdrift Game with two alternative evolutionarily stable strategies (ESSs): female-only and male-only care. This, however, does not explain the polymorphism in care strategies and frequent biparental desertion, because theory predicts that one of the two ESSs should have spread to fixation. Using a bootstrapping approach, we demonstrate that taking account of individual variation in payoffs explains the patterns of care better than a model based on the average population payoff matrix. In particular, a model incorporating differences in male attractiveness closely predicts the observed frequencies of male and female desertion. Our work highlights the need for a new generation of individual-based evolutionary game-theoretic models.

Population size and density may influence various aspects of breeding systems. For example, the number of potential mates may influence the mating system and parental care strategies. The breeding system of Eurasian penduline fits, Remiz pendulinus, involves sequential polygamy by both sexes and uniparental care by either male or female. Additionally, about one third of the clutches is deserted by both parents. Mating opportunities and breeding behaviour were studied at two populations: a low-density population (The Netherlands) and a high-density population (Hungary). We expected that higher breeding density is associated with higher incidence of polygamy. However, despite the substantial differences in breeding density and mating opportunities between Hungary and The Netherlands, there was no difference in the frequency of parental care types. We further investigated the plasticity of the breeding system of penduline tits using data from five breeding populations in Europe, and found that care patterns did not differ between these populations. We conclude that the type of parental care is not related to local breeding density. Proximate mechanisms of breeding system variation (e.g., gene expression) may be inflexible and, thus, not adjusted to local conditions. Alternatively, long-distance dispersal of penduline fits may mix the gene pool, so that local adaptation cannot possibly occur.

Engagement in extra-pair copulations is an example of the abundant conflicting interests between males and females over reproduction. Potential benefits for females and the risk of cuckoldry for males are expected to have important implications on the evolution of parental care. However, whether parents adjust parental care in response to parentage remains unclear. In Eurasian penduline tits Remiz pendulinus, which are small polygamous songbirds, parental care is carried out either by the male or by the female. In addition, one third of clutches is deserted by both male and female. Desertion takes place during the egg-laying phase. Using genotypes of nine microsatellite loci of 443 offspring and 211 adults, we test whether extra-pair paternity predicts parental care. We expect males to be more likely to desert cuckolded broods, whereas we expect females, if they obtain benefits from having multiple sires, to be more likely to care for broods with multiple paternity. Our results suggest that parental care is not adjusted to parentage on an ecological timescale. Furthermore, we found that male attractiveness does not predict cuckoldry, and we found no evidence for indirect benefits for females (i.e., increased growth rates or heterozygosity of extra-pair offspring). We argue that male Eurasian penduline tits may not be able to assess the risk of cuckoldry; thus, a direct association with parental care is unlikely to evolve. However, timing of desertion (i.e., when to desert during the egg-laying phase) may be influenced by the risk of cuckoldry. Future work applying extensive gene sequencing and quantitative genetics is likely to further our understanding of how selection may influence the association between parentage and parental care.

BACKGROUND:The evolutionary interests of males and females rarely coincide (sexual conflict), and these conflicting interests influence morphology, behavior and speciation in various organisms. We examined consequences of variation in sexual conflict in two closely-related passerine birds with contrasting breeding systems: the Eurasian penduline tit Remiz pendulinus (EPT) exhibiting a highly polygamous breeding system with sexually antagonistic interests over parental care, and the socially monogamous Cape penduline tit Anthoscopus minutus (CPT). We derived four a priori predictions from sexual conflict theory and tested these using data collected in Central Europe (EPT) and South Africa (CPT). Firstly, we predicted that EPTs exhibit more sexually dimorphic plumage than CPTs due to more intense sexual selection. Secondly, we expected brighter EPT males to provide less care than duller males. Thirdly, since song is a sexually selected trait in many birds, male EPTs were expected to exhibit more complex songs than CPT males. Finally, intense sexual conflict in EPT was expected to lead to low nest attendance as an indication of sexually antagonistic interests, whereas we expected more cooperation between parents in CPT consistent with their socially monogamous breeding system.RESULTS:Consistent with our predictions EPTs exhibited greater sexual dimorphism in plumage and more complex song than CPTs, and brighter EPT males provided less care than duller ones. EPT parents attended the nest less frequently and less simultaneously than CPT parents.CONCLUSIONS:These results are consistent with sexual conflict theory: species in which sexual conflict is more manifested (EPT) exhibited a stronger sexual dimorphism and more elaborated sexually selected traits than species with less intense sexual conflict (CPT). Our results are also consistent with the notion that EPTs attempt to force their partner to work harder as expected under sexual conflict: each member of the breeding pair attempts to shift the costs of care to the other parent. More brightly colored males benefit more from desertion than dull ones, because they are more likely to remate with a new female. Taken together, the comparison between two closely related species with contrasting breeding systems suggest that sexual conflict over care has influenced the evolution of behavior and morphology in penduline tits.