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L-M.C. and P.d.V. acknowledge support from the European Research Council (ERC) (Grant 678140-FluctEvol). The Montpellier tit group acknowledges the long-term support of the Observatoire des Sciences de l’Univers – Obser-vatoire de REcherche Montpelliérain de l’Environnement (OSU-OREME). The bighorn, mountain goat, and eastern gray kangaroo studies were supported by Natural Sciences and Engineering Research Council (NSERC) of Canada. Recent data collection for Wytham has been provided by grants from Biotechnology and Biological Sciences Research Council (BB/L006081/1), ERC (AdG250164), and the UK Natural Environment Research Council (NERC) (NE/K006274/1, NE/S010335/1). The Columbian ground squirrel study was supported by the National Science Foundation (Grant DEB-0089473). Trait and fitness data for hihi were collected/managed by John Ewen under New Zealand Department of Conservation hihi management contracts and research permits AK/15073/RES, AK-24128-FAU, 36186-FAU, and 44300-FAU and with additional financial support via NERC UK, The Leverhulme Trust UK, Marsden Fund New Zealand, and the Hihi Conservation Charitable Trust. The data on reindeer were made available through the Reindeer husbandry in a Globalizing North Nordic Center of Excellence, and the crew at Kutuharju Experimental Reindeer Research Station in the Reindeer Herder’s Association are thanked for their valuable assistance and logistic support in data collection. The red deer, Silwood blue tit, and Soay sheep datasets were supported by UK NERC. Lundy sparrow data were supported by NERC, a Marie Skłodowska-Curie Action, and Volkswagenstiftung. The red squirrel project was funded by NSERC of Canada and the National Science Foundation. J.C.S. was supported by a grant from the Ministry of Economy and Competitivity, Spanish Research Council (CGL-2016-79568-C3-3-P). J.T., T.K., and M.G. were supported by the Research Council of Norway through its Centers for Excellence funding scheme, Project 223257. Research on fairy wrens has been supported by the Australian Research Council. The Northern wheatear and the flycatcher studies were supported by grants from the Swedish Research Council VR. Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection. Postprint

Inbreeding depression is usually quantified by regressing individual phenotypic values on inbreeding coefficients, implicitly assuming there is no correlation between an individual's phenotype and the kinship coefficient to its mate. If such an association between parental phenotype and parental kinship exists, and if the trait of interest is heritable, estimates of inbreeding depression can be biased. Here we first derive the expected bias as a function of the covariance between mean parental breeding value and parental kinship. Subsequently, we use simulated data to confirm the existence of this bias, and show that it can be accounted for in a quantitative genetic animal model. Finally, we use long-term individual-based data for white-throated dippers (Cinclus cinclus), a bird species in which inbreeding is relatively common, to obtain an empirical estimate of this bias. We show that during part of the study period, parents of inbred birds had shorter wings than those of outbred birds, and as wing length is heritable, inbred individuals were smaller, independent of any inbreeding effects. This resulted in the overestimation of inbreeding effects. Similarly, during a period when parents of inbred birds had longer wings, we found that inbreeding effects were underestimated. We discuss how such associations may have arisen in this system, and why they are likely to occur in others, too. Overall, we demonstrate how less biased estimates of inbreeding depression can be obtained within a quantitative genetic framework, and suggest that inbreeding and additive genetic effects should be accounted for simultaneously whenever possible.

Telomeres are protective DNA–protein complexes located at the ends of eukaryotic chromosomes, whose length has been shown to predict life-history parameters in various species. Although this suggests that telomere length is subject to natural selection, its evolutionary dynamics crucially depends on its heritability. Using pedigree data for a population of white-throated dippers ( Cinclus cinclus ), we test whether and how variation in early-life relative telomere length (RTL, measured as the number of telomeric repeats relative to a control gene using qPCR) is transmitted across generations. We disentangle the relative effects of genes and environment and test for sex-specific patterns of inheritance. There was strong and significant resemblance among offspring sharing the same nest and offspring of the same cohort. Furthermore, although offspring resemble their mother, and there is some indication for an effect of inbreeding, additive genetic variance and heritability are close to zero. We find no evidence for a role of either maternal imprinting or Z-linked inheritance in generating these patterns, suggesting they are due to non-genetic maternal and common environment effects instead. We conclude that in this wild bird population, environmental factors are the main drivers of variation in early-life RTL, which will severely bias estimates of heritability when not modelled explicitly.

Von 1992 bis 2000 wurde an den Fliessgewassern rund um das Becken des Zurichsees im schweizerischen Mittelland der Brutbeginn der Wasseramsel untersucht. Diese Wasserlaufe haben eine Gesamtlange von 40 km und werden von durchschnittlich 46 Paaren besiedelt. Die Nester liegen auf einer Hohe von 410 – 680 m u.M. In den neun Untersuchungsjahren registrierten wir 552 Bruten (Nestlingsalter 8 – 12 Tage). Die Wasseramsel brutet in dieser Region fruh, aber mit jahrlich sehr variablem Brutbeginn: Absolut Fruhester Legebeginn war der 9. Februar in den Jahren 1995 und 1997, spatester der 9. Marz im Jahr 1992. Als Vergleichsbasis zwischen den Brutjahren wurde der Mediantag des Legebeginns der ersten 25 % aller Bruten des entsprechenden Jahres gewahlt. Dieses Datum korrelierte hochsignifikant (Rangkorrelation nach Spearman; p

Der Zurichbergwald ist eine 350 ha grose, bewaldete Kuppe auf 480–680 m u.M. am Rande der Stadt Zurich. Hier untersuchten wir von 1989 bis 1995 die jahreszeitliche Gesangsaktivitat der Singdrossel mit zwei verschiedenen Ansatzen. 1989 und 1991–1995 zahlte JH an 123 Tagen auf zwei festgelegten Strecken von 6,1 und 7,1 km die Sanger jeweils in der Stunde der abendlichen Dammerung. Im gleichen Gebiet zahlte RS 1990 an 46 Tagen auf einer 6,7 km langen Strecke am Morgen in der Stunde nach Sonnenaufgang. Ebenfalls 1990 sammelte RS Daten zur Brutbiologie.

Eighteen polymorphic microsatellite DNA loci were isolated in the Central European subspecies of the European Dipper (Cinclus cinclus aquaticus). The loci were tested for polymorphism using a test panel of 24 breeding birds. Numbers of alleles ranged from 2 to 21 per locus and expected heterozygosities varied between 0.47 and 0.83. Two loci (Cici10 and Cici12) proved to be Z-linked. Some pairs of loci exhibited significant linkage disequilibrium but not the two loci that are located on the Z-chromosome. This pattern suggests that demographic effects rather than physical linkage are likely responsible for the observed levels of linkage disequilibrium. These loci will be useful for applied conservation projects and for investigations of the dispersal and mating patterns of European and other dippers.

Knowledge of the ranging behavior and spatial requirements of a species is fundamental for establishing meaningful conservation strategies. Such information is lacking for the middle spotted woodpecker (Dendrocopos medius), a species endangered throughout its westpalearctic range. By radiotracking, we studied spacing behavior of this habitat specialist in a lowland oak forest of northeastern Switzerland from 1992-1996. Home range and core area size decreased from winter to late spring, with males and females having home ranges of similar size. Overlap of male home ranges was highest in winter (up to 40%) and lowest in late spring, whereas core area overlap remained low. For both home ranges and core areas, overlapping parts were used randomly in winter but more often than expected in early spring. Overlap of female ranges and of core areas did not change from early to late spring, and the shared parts of these home ranges were used as expected in both seasons. Aggressive interactions were most common in March and April and occurred mainly between individuals of the same sex. Our results suggest that the middle spotted woodpecker is not territorial in winter but defends nearly exclusive territories during spring, with both sexes participating to similar degrees in territorial defense. Based on this seasonal territoriality, we propose consideration of core areas in early spring (Mar and Apr) as a reliable estimate of the area requirements of the species to be used in management plans.