Measures Of Reproductive Success Research Articles

Perspective: Can early embryonic losses be reduced in lactating dairy cows?

Perspective: Can early embryonic losses be reduced in lactating dairy cows?

Community associations of birds with amphibians and fish in wetlands created for biodiversity

Conservation initiatives to support declining water-related biodiversity through wetland creation have increased during the last decades. Multiple studies have evaluated the suitability of created wetlands for birds and amphibians, but only a few have considered the species associations that might also affect the outcome. Using joint species distribution models, we explored species associations of birds, amphibians and fish in 52 created biodiversity wetlands in Sweden. As most of these wetlands were primarily created for increasing bird diversity, we asked whether the occurrence of fish and amphibians relates to bird species richness, pair abundance and chick abundance (as a measure of reproductive success) and whether conservation conflicts or synergies between birds, amphibians and fish can be observed. In general, we found positive bird-amphibian association patterns and negative bird-fish association patterns, although the uncertainties were high for these estimates. In line with previous research, the generally negative bird-fish co-variance indicates potential conservation conflicts between wetland creation for birds and fish, where fish might be introduced for recreational fishing or other ecosystem services. Therefore, our results suggest that it can be hard to benefit bird and fish communities with the same wetland, and separate wetland creation with different goals may be needed. The generally positive bird-amphibian species-species associations and the lack of previous studies revealing conflicts indicate synergies between wetland creation for birds and amphibians. However, research needs to further consolidate such synergies, including amphibian reproductive output from bird-rich wetlands.

Group social structure has limited impact on reproductive success in a wild mammal

Abstract The frequency and type of dyadic social interactions individuals partake in has important fitness consequences. Social network analysis is an effective tool to quantify the complexity and consequences of these behaviors on the individual level. Less work has used social networks to quantify the social structure—specific attributes of the pattern of all social interactions in a network—of animal social groups, and its fitness consequences for those individuals who comprise the group. We studied the association between social structure, quantified via five network measures, and annual reproductive success in wild, free-living female yellow-bellied marmots (Marmota flaviventer). We quantified reproductive success in two ways: (1) if an individual successfully weaned a litter and (2) how many pups were weaned. Networks were constructed from 38 968 interactions between 726 unique individuals in 137 social groups across 19 years. Using generalized linear mixed models, we found largely no relationship between either measure of reproductive success and social structure. We found a modest relationship that females residing in more fragmentable social groups (i.e., groups breakable into two or more separate groups of two or more individuals) weaned larger litters. Prior work showed that yellow-bellied marmots residing in more fragmentable groups gained body mass faster—another important fitness correlate. Interestingly, we found no strong relationships between other attributes of social group structure, suggesting that in this facultatively social mammal, the position of individuals within their group, the individual social phenotype, may be more important for fitness than the emergent group social phenotype.

Genetic Association between Farrowing Rates and Swine Leukocyte Antigen Alleles or Haplotypes in Microminipigs.

We have previously reported specific swine leukocyte antigen (SLA) haplotype associations with significant effects on several reproduction performance traits in a highly inbred miniature pig population of Microminipigs (MMPs). In this study, to clarify the effects on farrowing rates of SLA similarity between mating partners in the MMP population, we compared the farrowing rates as a measure of reproductive success after 1063-cumulative matings among the following three groups of mating partners: (1) completely sharing SLA class I or class II haplotypes or alleles between partners (CS), (2) only one sharing the haplotypes or alleles (OS), and (3) non-sharing the haplotypes or alleles (NS). Average farrowing rates in CS groups consisting of completely sharing SLA class II haplotypes or DRBI and DQB1 alleles were lowest in the three groups. Moreover, lower farrowing rates were indicated in mating pairs with smaller amino acid pairwise genetic distances of SLA-1, SLA-3, DRB1 and DQB1 alleles between the pairs. These results suggested that the dissimilarity of SLA class I and class II alleles between mating partners markedly improved reproductive performance; therefore, SLA alleles or haplotypes are potentially useful genetic markers for the selection of mating pairs in breeding programs and epistatic studies of reproductive traits of MMPs.

Factors limiting reproductive success in urban Greylag Geese (Anser anser).

In the late eighties, Greylag Geese (Anser anser) started to colonise an urban area previously void of geese in southwestern Germany. Between 2004 and 2020, in a period of steady population increase with subsequent population stagnation, we analysed two measures of reproductive success: (1) the relation between freshly hatched to fledged young for each brood and (2) the probability of a hatchling to survive to fledging. We were able to show that the dispersal of pairs from the nesting site to a different brood rearing area resulted in higher reproductive success. However, the increasing population size of Greylag Geese and the number of breeding pairs of recently immigrated Egyptian Geese (Alopochen aegyptiaca) had a negative impact on reproductive success, indicating density dependence. Our results show that newly established populations in urban settings do not grow indefinitely, which is an important fact that should be taken into account by wildlife managers.

Nestling Savannah Sparrows and Tree Swallows differ in their sensitivity to weather

AbstractSavannah Sparrows (Passerculus sandwichensis) and Tree Swallows (Tachycineta bicolor) breed and forage in the same habitat on Kent Island, a boreal island in the Bay of Fundy, New Brunswick, but respond differently to the same weather conditions. The 2 passerines are similar in body size but because Tree Swallows depend upon small flying insects captured on the wing, they may be more sensitive to weather than Savannah Sparrows, which forage on insects and seeds on the ground and in shrubs and trees. To compare how reproductive success in the 2 species was affected by weather conditions, we took advantage of an 18-year dataset and used a model-building approach that controlled for year, adult sex and age, and field where they nested. We focused on 3 measures of reproductive success (hatching success, fledging success, and nestling condition) and different time periods (3- to 18-day time windows) before hatching or fledging. The responses of the 2 species differed in magnitude and direction. In Tree Swallows, adding weather variables to the basic model increased the explanatory power of fixed effects by 19.1%, illustrating the swallows’ sensitivity to weather. In contrast, in Savannah Sparrows, the addition of weather variables only increased the model’s explanatory power by 0.4% and the proportion of variation attributed to fixed factors by only 1.5%, which reflected the species’ hardiness in the face of inclement weather. Our results suggest that how a bird species forages and the nature of its prey may influence its sensitivity to weather and indicate that increased rainfall, strong winds and other events associated with climate change may affect Tree Swallows and other aerial insectivores more than ground-foraging birds such as Savannah Sparrows.

Antagonistic selection on body size and sword length in a wild population of the swordtail fish, Xiphophorus multilineatus: Potential for intralocus tactical conflict.

Alternative reproductive tactics (ARTs) have provided valuable insights into how sexual selection and life history trade‐offs can lead to variation within a sex. However, the possibility that tactics may constrain evolution through intralocus tactical conflict (IATC) is rarely considered. In addition, when IATC has been considered, the focus has often been on the genetic correlations between the ARTs, while evidence that the ARTs have different optima for associated traits and that at least one of the tactics is not at its optimum is often missing. Here, we investigate selection on three traits associated with the ARTs in the swordtail fish Xiphophorus multilineatus; body size, body shape, and the sexually selected trait for which these fishes were named, sword length (elongation of the caudal fin). All three traits are tactically dimorphic, with courter males being larger, deeper bodied and having longer swords, and the sneaker males being smaller, more fusiform and having shorter swords. Using measures of reproductive success in a wild population we calculated selection differentials, as well as linear and quadratic gradients. We demonstrated that the tactics have different optima and at least one of the tactics is not at its optimum for body size and sword length. Our results provide the first evidence of selection in the wild on the sword, an iconic trait for sexual selection. In addition, given the high probability that these traits are genetically correlated to some extent between the two tactics, our study suggests that IATC is constraining both body size and the sword from reaching their phenotypic optima. We discuss the importance of considering the role of IATC in the evolution of tactical dimorphism, how this conflict can be present despite tactical dimorphism, and how it is important to consider this conflict when explaining not only variation within a species but differences across species as well.

Using behavior indices and vital rates to determine the conservation impact of wildlife tourism: Guided sea turtle watch programs in Florida

Recent rapid growth in wildlife tourism necessitates understanding its biologically-significant impacts on targeted species. To evaluate these effects, we studied how loggerhead sea turtles (Caretta caretta) responded to public sea turtle watch programs on a nesting beach in southeast Florida. We introduced two new measures of behavior in addition to other well-known behavior parameters, by which to measure behavioral responses of turtles during the nesting process. Five of ten behavior parameters we measured were significantly influenced by the presence of a turtle watch group. However, we found no relationship between observed changes in nesting behavior and measures of reproductive success (estimated clutch size, hatching success, and hatchling emergence success). We conclude that either the measured behavioral changes did not affect the outcome of reproduction, or their effect was undetectable. This study highlights the importance of pairing behavioral and demographic variables when evaluating the conservation impact of wildlife tourism programs. When wildlife tourism programs prioritize animal welfare and are periodically evaluated for their biological impact on target species, they are an effective tool for educating the general public and promoting conservation and environmental stewardship.

Risk‐taking behaviour relates to timing of breeding in a sub‐Antarctic rainforest bird

Within‐population variation in timing of breeding may be linked to a trade‐off between the risks and benefits of breeding earlier. This trade‐off may be mediated by individual risk‐taking behaviour, but this needs to be assessed in detail in wild populations. Here, we recorded timing of breeding and risk‐taking behaviour during three consecutive breeding seasons in a resident population of Thorn‐tailed Rayadito Aphrastura spinicauda located in Navarino Island (55°S), southern Chile. Navarino is a high‐latitude, highly seasonal continental island in southern Chile where early breeding may be risky for rayaditos, given the presence of low temperatures, storms and relatively low food abundance during early spring. We used novel environment tests to assess exploratory behaviour, which in turn was used as a proxy of risk‐taking behaviour. In addition, we evaluated the potential consequences of timing of breeding and risk‐taking behaviour on three measures of reproductive success: clutch size, number of fledglings produced and body condition of the offspring. We found that risk‐prone individuals started breeding earlier than risk‐averse individuals but we did not find evidence for an effect of timing of breeding on any of the variables of seasonal breeding success. However, we observed that fast‐exploring females tended to lay smaller clutches. Measurements reflecting lifetime reproductive success may better reflect an effect of timing of breeding and risk‐taking behaviour, but this needs to be studied. Our results support the idea that risk‐taking behaviour is linked to early breeding in high‐latitude environments. We suggest that, despite the risks that low temperatures and snowstorms pose to breeding rayaditos during early spring, it is possible that early breeders increase the probability of occupying better nesting cavities and they may adjust the timing of breeding to match brood provisioning with peak resource abundance.

Evolutionary graph theory derived from eco-evolutionary dynamics

A biologically motivated individual-based framework for evolution in network-structured populations is developed that can accommodate eco-evolutionary dynamics. This framework is used to construct a network birth and death model. The evolutionary graph theory model, which considers evolutionary dynamics only, is derived as a special case, highlighting additional assumptions that diverge from real biological processes. This is achieved by introducing a negative ecological feedback loop that suppresses ecological dynamics by forcing births and deaths to be coupled. We also investigate how fitness, a measure of reproductive success used in evolutionary graph theory, is related to the life-history of individuals in terms of their birth and death rates. In simple networks, these ecologically motivated dynamics are used to provide new insight into the spread of adaptive mutations, both with and without clonal interference. For example, the star network, which is known to be an amplifier of selection in evolutionary graph theory, can inhibit the spread of adaptive mutations when individuals can die naturally.

Heterozygosity-Fitness Correlations in a Continental Island Population of Thorn-Tailed Rayadito.

Heterozygosity-fitness correlations (HFCs) have been used to monitor the effects of inbreeding in threatened populations. HFCs can also be useful to investigate the potential effects of inbreeding in isolated relict populations of long-term persistence and to better understand the role of inbreeding and outbreeding as drivers of changes in genetic diversity. We studied a continental island population of thorn-tailed rayadito (Aphrastura spinicauda) inhabiting the relict forest of Fray Jorge National Park, north-central Chile. This population has experienced a long-term, gradual process of isolation since the end of the Tertiary. Using 10 years of field data in combination with molecular techniques, we tested for HFCs to assess the importance of inbreeding depression. If inbreeding depression is important, we predict a positive relationship between individual heterozygosity and fitness-related traits. We genotyped 183 individuals at 12 polymorphic microsatellite loci and used 7 measures of reproductive success and estimates of apparent survival to calculate HFCs. We found weak to moderate statistical support (P-values between 0.05 and 0.01) for a linear effect of female multi-locus heterozygosity (MLH) on clutch size and nonlinear effects on laying date and fledging success. While more heterozygous females laid smaller clutches, nonlinear effects indicated that females with intermediate values of MLH started laying earlier and had higher fledging success. We found no evidence for effects of MLH on annual fecundity or on apparent survival. Our results along with the long-term demographic stability of the study population contradict the hypothesis that inbreeding depression occurs in this population.

Negative effects of individual heterozygosity on reproductive success in a wild bird population.

The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity-fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding and outbreeding depression are lacking. We studied a genetically diverse population of blue tits in southern Germany using a genome-wide set of 87 microsatellites to investigate the relationship between proxies of reproductive success and measures of multilocus and single-locus individual heterozygosity (MLH and SLH). We used complimentary measures of MLH and partitioned markers into functional categories according to their position in the blue tit genome. HFCs based on MLH were consistently negative for functional loci, whereas correlations were rather inconsistent for loci found in nonfunctional areas of the genome. Clutch size was the only reproductive variable showing a general effect. We found evidence for local effects for three measures of reproductive success: arrival date at the breeding site, the probability of breeding at the study site and male reproductive success. For these, we observed consistent, and relatively strong, negative effects at one functional locus. Remarkably, this marker had a similar effect in another blue tit population from Austria (~400km to the east). We suggest that a genetic local effect on timing of arrival might be responsible for most negative HFCs detected, with carry-over effects on other reproductive traits. This effect could reflect individual differences in the distance between overwintering areas and breeding sites.

Oceanographic drivers of winter habitat use in Cassin's Auklets.

Reduced prey abundance and severe weather can lead to a greater risk of mortality for seabirds during the non-breeding winter months. Resource patterns in some regions are shifting and becoming more variable in relation to past conditions, potentially further impacting survival and carryover to the breeding season. As animal tracking technologies and methods to analyze movement data have advanced, it has become increasingly feasible to draw fine-scale inference about how environmental variation affects foraging behavior and habitat use of seabirds during this critical period. Here, we used archival light-sensing tags to evaluate how interannual variation in oceanography affected the winter distribution of Cassin's Auklets from Southeast Farallon Island, California. Thirty-five out of 93 geolocators deployed from 2015 to 2017 were recovered and successfully recorded light-level data, from which geographic positions were estimated. Step-selection functions were applied to identify environmental covariates that best explained winter movement decisions and habitat use, revealing Cassin's Auklets dispersed farther from the colony during a winter with warm SST anomalies, but remained more centralized near the breeding colony during two average winters. Movement patterns were driven by avoidance of areas with higher sea surface temperatures and possible limits of dispersal from the breeding colony, and selection for areas with well-defined mesoscale fronts and cooler surface waters. Through multiple years of tagging and the application of step-selection functions, a robust and widely applied approach for analyzing animal movement in terrestrial species, we show how interannual differences in the movement patterns of a small seabird are driven by oceanographic variability across years. Understanding the winter habitat use of seabirds can help inform changes in population structure and measures of reproductive success, aiding managers in determining potential causes of breeding failures.

Correlates of maternal glucocorticoid levels in a socially flexible rodent

While it is generally accepted that social isolation has detrimental effects on social species, little is known about the importance of social interactions in less social species–particularly for wild reproductive females. We studied socially-flexible yellow-bellied marmots (Marmota flaviventer) and asked whether features of the social environment are associated with maternal fecal glucocorticoid metabolite (FGM) concentrations. Since changes in maternal baseline glucocorticoids may have positive or negative consequences for offspring fitness, we were also interested in estimating their relationship with measures of reproductive success. We fitted generalized linear mixed effects models to a dataset including maternal FGM measurements, social network metrics, maternal/alloparental care, and pup FGM and survival. Agonistic interactions were positively associated with maternal FGM levels, while mothers that engaged in relatively more affiliative interactions had reduced FGM levels when living in environments with low predator pressure. Pups associated with mothers exhibiting high FGM levels had low annual survival rates, received less maternal/alloparental care and had higher FGM levels. Interestingly, offspring from mothers with high FGM levels were more likely to survive the summer when born in small litters. In sum, social interactions likely influence and are influenced by glucocorticoid levels of facultatively social females. Potential benefits of social bonds may be context-specific, and agonistic interactions may be tightly correlated with fitness. Female marmots exhibiting high FGM levels had overall low reproductive success, which is predicted by the cort-fitness hypothesis. However, under adverse conditions, offspring summer survival can be maximized if pups are born in small litters.

Mercury Concentration in Nestling Feathers Better Predicts Individual Reproductive Success than Egg or Nestling Blood in a Piscivorous Bird.

Piscivorous birds are at high risk of mercury exposure in aquatic food webs, and their reproductive success is sensitive to methylmercury exposure. Although effects are convincingly shown in a handful of lab studies, sublethal effects at environmentally relevant concentrations in the field, where there is a range of other natural stressors, are not well delineated. Part of that uncertainty arises because mercury concentration (hereafter, [Hg]) in adult tissues used to assess Hg risk can be influenced by Hg values in wintering grounds or other nonrelated areas. Several studies have used nestling tissues under the assumption that they better represent local risk to breeding since nestlings consume locally derived food. However, the correlation between [Hg] in nestling tissues and local breeding success remains unassessed. We analyzed great egret ( Ardea alba) breeding parameters collected over 3 years (2015-2017) in the Everglades (Florida, USA). The Everglades is a large contiguous wetland with geographically dispersed wading bird breeding sites exposed to variable and biologically relevant ranges of mercury concentrations. We examined mercury concentrations in albumen and nestling blood and feathers as predictors of 6 measures of reproductive success at individual nests. Albumen [Hg] did not correlate with reproductive end points, and correlations with blood [Hg] were weak. Feather [Hg] correlated negatively with all of the posthatching endpoints and explained 8.3% of the variance in the probability of a laid egg resulting in a fledged chick. However, most of the observed failures were hatching failures, which were not explained by albumen [Hg], and other nestling tissues could therefore not be evaluated. While our results support the use of nestling feathers as indicators of site-specific mercury exposure, we discuss both advantages and possible limitations of using nestling feathers as indicators of local mercury exposure.

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

ABSTRACTCapsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.

On the adaptive value of monomorphic versus dimorphic enantiostyly in Solanum rostratum.

Enantiostyly is a reproductive system with heteromorphic flowers characterized by asymmetrical deflection of the style, either to the left or to the right of the floral axis. There are two types of enantiostyly. In monomorphic enantiostyly, plants produce the two types of flowers in the same individual. Dimorphic enantiostyly is restricted to only seven species and their populations consist of individuals producing either the right or the left flower type. It is hypothesized that the dimorphic form is derived from monomorphic ancestors because it functions as an outcrossing mechanism. We tested this latter hypothesis and investigated if monomorphic enantiostyly is resistant to invasion by individuals with dimorphic enantiostyly, because it functions as a reproductive assurance mechanism. To determine the conditions favouring the invasion of dimorphic enantiostyly, measurements of reproductive success and outcrossing rates in 15 natural flowering patches of Solanum rostratum were made. To test if monomorphic enantiostyly provides a reproductive assurance mechanism, experimental plants with either manually created dimorphic or natural monomorphic reproductive systems were exposed to two different pollination scenarios (flower density treatments), and reproductive success and outcrossing rates were measured. Naturally flowering patches experienced severe pollination limitation, showed marked differences in reproductive success and had relatively high outcrossing rates. Plants in the experimental patches also showed pollination limitation and high outcrossing rates. Individuals with dimorphic enantiostyly expressed higher reproductive and outcrossing advantages under high-density conditions. These advantages disappeared in the low-density treatment, where the monomorphic form attained a higher reproductive success and no differences in outcrossing rates were detected. Monomorphic enantiostyly should be resistant to invasion of the dimorphic form because the prevalent ecological conditions favour the maintenance of geitonogamous individuals that are able to take advantage of ecological heterogeneity and generalized pollination limitation.

Habitat-dependent effects of personality on survival and reproduction in red squirrels

Variation in intrinsic (body mass, reproductive condition) and extrinsic factors (habitat quality, spatio-temporal variation in food availability) can affect the costs and benefits of personality traits. Relationships between personality and fitness components can vary with changes in population density and/or habitat quality. Here, using capture-mark-recapture data of individually marked Eurasian red squirrels (Sciurus vulgaris), we explored whether indices of boldness and exploration are correlated with local survival in either of the sexes and with a measure of reproductive success in females, under different conditions of food availability. PCA scores derived from the number of captures (trappability) and number of different traps where an individual was captured (trap diversity), in the first 1 or 2 years of presence, were used as indices of personality (boldness and pure exploration). The relationships between boldness and local survival differed between habitats in both sexes. Bold squirrels survived better than shy ones in Norway spruce forest, while in the other two study areas (Scots pine or mixed spruce-fir forest), survival was lower for bolder squirrels. Pure exploration behavior was negatively correlated with local survival in all habitats and in both sexes. Female reproductive success increased with body mass and decreased with the tendency to explore, a relationship consistent across habitat types. Bolder females did not have a reproductive advantage in years of poor food availability. We suggest that costs and benefits of boldness vary with spatio-temporal differences in availability of high-quality food resources which may help to maintain variation in personality in red squirrels in heterogeneous landscapes. Differences in personality among individuals can affect their survival or reproductive success. We investigated the relationship between personality traits and fitness components in Eurasian red squirrels under different food availability. We scored boldness and exploration using indices derived from capture-mark-recapture data. Bold red squirrels survived longer than shy ones in spruce forest where food availability varied strongly between years. Instead, in mixed forests with more stable food supplies, shy individuals survived longer. Heavy females produced more litters in their lifetime than those with lower body mass. Explorative squirrels were more likely to die young and to produce no or few litters, independent of the habitat they lived in. Our results suggest that variation in personality traits within populations of Eurasian red squirrels was maintained by changing fitness benefits in relation to habitat type in pulsed resource systems.

Evolutionary significance of antiparasite, antipredator and learning phenotypes of avian nest defence

Avian nest defence, which is expected to serve both antiparasite and antipredator functions, may benefit or be detrimental to birds, although selective forces that potentially operate on nest defence have not been quantified as a whole. Together with fitness values, we analysed two traits of nest defence, intensity and plasticity, in two distantly related passerine species, yellow warbler (Setophaga petechia) in North America and reed warbler (Acrocephalus scirpaceus) in Europe, both favourite host species for brood parasites. Breeders that escaped parasitism were the most vocal among reed warblers, whereas there was no specific defence phenotype that predicted prevention of parasitism in yellow warblers. Breeders that escaped nest predation were, in both species, those with the most distractive response at the first exposure to a nest-threatening event, such as the experimental predation or parasitism simulated at the nest. However, increasing defence intensity benefited yellow warblers but was detrimental to reed warblers, because intense defence responses attracted predators. Adaptiveness of nest defence was revealed by nest defence phenotypes when examined in concert with the seasonal fitness (i.e. measures of reproductive success). Results revealed selective forces favoured yellow warblers with strong defence phenotypes. Opposite forces were instead revealed among reed warblers whose favoured phenotypes were strong, yet less flexible, defenders.

Patterns of female nest attendance and male feeding throughout the incubation period in Blue Tits Cyanistes caeruleus

Most bird species exhibit biparental care, but the type of care provided by each sex may differ substantially. In particular, during the incubation phase in passerines, females perform most or all of the incubation, while the male cares for the brood indirectly by feeding the female. However, detailed descriptions of this male investment during the incubation period are missing. Here, we quantitatively describe female nest attendance and male incubation feeding throughout the ~ 14‐day incubation period in a population of Eurasian Blue Tits Cyanistes caeruleus breeding in nestboxes. Males and females progressively increased their daily activity at the nest over the incubation period. The amount of day‐time incubation, measured as the proportion of the active day (time interval between first nestbox exit in the morning and last entry in the evening) a female spent inside the nestbox, varied between 52 and 60% with an average of 55% per day. The frequency of male incubation feeding varied between 0 and 74 times per day with an average of 12 feeds per day. Both male feeding rate and female nest attendance were highest in the morning and declined rapidly throughout the day. Females were more likely to be off the nest during the warmest periods (15–21 °C), as expected based on thermal needs of the developing embryos, but also during the coldest periods (2–5 °C), presumably due to the energetic needs of the female. This was despite the fact that males fed their females more often at the nest when ambient temperatures were low. Females that received more feeds incubated more and their off‐nest bouts were shorter after a feed. The observed variation in female incubation and in male feeding rate was not linked to individual age or to variation in measures of reproductive success. However, direct observations showed that in some pairs a substantial amount of feeding by males occurred outside the nestbox. This suggests that the true male investment might have been underestimated, here and in previous studies.