Tradeoffs and synergies in wetland multifunctionality: A scaling issue.

Wetland area in agricultural landscapes has been heavily reduced to gain land for crop production, but in recent years there is increased societal recognition of the negative consequences from wetland loss on nutrient retention, biodiversity and a range of other benefits to humans. The current trend is therefore to re-establish wetlands, often with an aim to achieve the simultaneous delivery of multiple ecosystem services, i.e., multifunctionality. Here we review the literature on key objectives used to motivate wetland re-establishment in temperate agricultural landscapes (provision of flow regulation, nutrient retention, climate mitigation, biodiversity conservation and cultural ecosystem services), and their relationships to environmental properties, in order to identify potential for tradeoffs and synergies concerning the development of multifunctional wetlands. Through this process, we find that there is a need for a change in scale from a focus on single wetlands to wetlandscapes (multiple neighboring wetlands including their catchments and surrounding landscape features) if multiple societal and environmental goals are to be achieved. Finally, we discuss the key factors to be considered when planning for re-establishment of wetlands that can support achievement of a wide range of objectives at the landscape scale.

Song similarity predicts hybridization in flycatchers.

Given that population divergence in sexual signals is an important prerequisite for reproductive isolation, a key prediction is that cases of signal convergence should lead to hybridization. However, empirical studies that quantitatively demonstrate links between phenotypic characters of individuals and their likelihood to hybridize are rare. Here we show that song convergence between sympatric pied (Ficedula hypoleuca) and collared flycatchers (F. albicollis) influence social and sexual interactions between the two species. In sympatry, the majority of male pied flycatchers (65%) include various parts of collared flycatcher song in their song repertoire (but not vice versa). Playback experiments on male interactions demonstrate that male collared flycatchers respond similarly to this 'mixed' song as to conspecific song. Long-term data on pairing patterns show that males singing a converged song attract females of the other species: female collared flycatchers only pair with male pied flycatchers if the males sing the mixed song type. From the perspective of a male pied flycatcher, singing a mixed song type is associated with 30% likelihood of hybridization. This result, combined with our estimates of the frequency of mixed singers, accurately predicts the observed occurrence of hybridization among male pied flycatchers in our study populations (20.45% of 484 pairs; predicted 19.5%). Our results support the suggestion that song functions as the most important prezygotic isolation mechanism in many birds.

Experimental evidence of environmental effects on age-specific reproductive success: the importance of resource quality.

Age-specific access to high-quality resources (e.g. territory or nest site) might be an important determinant for improved reproductive performance with increasing age. I experimentally investigated the effects of territory quality versus other age-related improvements in breeding competence (e.g. foraging skills, breeding experience and local knowledge) on age-specific reproductive success. Territory quality (i.e. territory field layer height) was manipulated in year 2 of northern wheatears (Oenanthe oenanthe) that were breeding in the same territory in two consecutive years. Changing territory quality by changing field layer height had a strong effect on within-individual change in the reproductive success of wheatears. This effect was mainly due to a corresponding change in nest predation risk. When territory quality was kept constant (i.e. no between-year change in territory field layer height), within-individual reproductive success did not change between subsequent years. Thus, age-related improvements in foraging skills, breeding experience and local familiarity had no significant effect on within-individual changes in reproductive success. Increased reproductive success with increased age in northern wheatears is therefore mainly explained by an improved access to high-quality territories with increasing age. I conclude that age-dependent access to high-quality breeding resources might be a widespread phenomenon in nature.

Adaptive plasticity in mate preference linked to differences in reproductive effort.

There is abundant evidence for the existence of marked mate preferences in natural populations, but the occurrence of within-population variation in mate preferences has received little attention and is often regarded as nonadaptive deviation from the optimal norm. Here we show experimentally that the preference of female collared flycatchers Ficedula albicollis for male forehead patch size, a sexually selected trait, varies with the time of breeding, an environmental factor with strong effects on reproductive success. Contrary to expectations based on time-constrained choice models, only late-breeding females prefer males with a large patch size. The variation in mate preference matches a seasonal change in female reproductive success: long-term data reveal a positive relationship between female reproductive success and male patch size exclusively in late breeders. In addition, female reproductive effort, as assessed by clutch size, appears to be adjusted relative to both timing of breeding and male phenotype. We conclude that not only can mate preferences display adaptive plasticity within populations, but this plasticity can also be linked to differences in reproductive investment.

Age and reproduction in birds - hypotheses and tests.

It is well known that reproductive performance improves with age in birds. Many hypotheses, involving factors such as differential survival, delayed breeding, breeding experience, foraging ability and reproductive effort, have been proposed to explain this pattern. Although these hypotheses are not mutually exclusive, they can be classified in three major groups relating to progressive appearance or disappearance of phenotypes, age-related improvements of competence, and optimization of reproductive effort. However, a closer examination of the literature reveals that only few studies have rigorously tested the hypotheses. Future work should focus on carefully designed tests that critically investigate the hypotheses.

"Terminal investment" and a sexual conflict in the Collared Flycatcher (Ficedula albicollis).

When the expectation of future reproduction is reduced by senescence, life-history theory predicts that reproductive effort will increase with increasing age. This idea was examined in the collared flycatcher by estimating whether reproductive costs increase with female age, comparing feeding rates and weight losses of old, "senescent" females (i.e., >=5 yr old) and middle-aged females (2-3 yr old) with the same breeding phenology and the same brood size, and testing whether feeding rate was correlated with daily energy expenditure and with weight loss of females during the nestling period. There was a negative relationship between fledgling production and subsequent survival among old females (>=5 yr old), but not among younger age classes, which suggests that reproductive effort increases with age. Also, old females fed their nestlings more often and lost more weight during the nestling period than did middle-aged females Observed feeding rates were positively correlated with daily energy expenditure and weight loss. Since there was no evidence that individuals that survived to old ages were better at all ages, the results strongly suggest that old collared flycatcher females increase their reproductive effort at the cost of a decreased probability of surviving to the next year. However, the payoff of the increased reproductive effort of old females seemed to be small. We suggest that this is a consequence of a conflict between the sexes over the division of work, because old females generally are mated to younger males that probably have better future prospects. Data on male feeding rates in relation to female feeding rates support this idea.