Gergely Hegyi

Using information theory as a substitute for stepwise regression in ecology and behavior

In ecological and behavioral research, drawing reliable conclusions from statistical models with multiple predictors is usually difficult if all predictors are simultaneously in the model. The traditional way of handling multiple predictors has been the use of threshold-based removal-introduction algorithms, that is, stepwise regression, which currently receives considerable criticism. A more recent and increasingly propagated modelling method for multiple predictors is the information theoretic (IT) approach that quantifies the relative suitability of multiple, potentially non-nested models based on a balance of model fit and the accuracy of estimates. Here, we examine three shortcomings of stepwise regression, subjective critical values, model uncertainty, and parameter estimation bias, which have been suggested to be avoided by applying information theory. We argue that, in certain circumstances, the IT approach may be sensitive to these issues as well. We point to areas where further testing and development could enhance the performance of IT methods and ultimately lead to robust inferences in behavioral ecology.

Determinants of male territorial behavior in a Hungarian collared flycatcher population: plumage traits of residents and challengers

In intrasexual conflicts, contestants can rely on relative or absolute size of status badges to mediate aggressive behavior. Most studies focus on the response of focal animals to variation in status badges of their competitors; few have simultaneously considered the traits of both participants under experimental conditions. By simulating territorial intrusions, we tested the importance of two sexual traits [forehead patch size (FPS) and wing patch size (WPS)] in territorial behavior of males in a Hungarian population of the collared flycatcher, Ficedula albicollis. We presented a stimulus male to an unpaired resident male to examine whether the characteristics of the territory owner or those of the challenger were associated with the latency of the first attack of the owner, which is a good predictor of the territorial behavior in general. WPS of the stimulus male was a significant determinant of the latency of the first attack, as males with a larger WPS elicited quicker attacks from the residents than males with a small WPS. From the residents’ perspective, age appeared to influence their territorial behavior, as yearlings had shorter attack latencies than older males. Additionally, latency could be considered an individual-specific attribute because it varied consistently among males, even when the WPS of the stimulus male was controlled, and it was associated with pairing success. Contrary to findings in a Swedish population, FPS seemed to be unimportant in male–male competition in our population, which suggests population differences in the role of the two plumage traits. Our results indicate that in a territorial conflict, the characteristics of both participants are important.

Qualitative population divergence in proximate determination of a sexually selected trait in the collared flycatcher

We examined proximate determination of sexually selected forehead patch size in a Central‐European population of Ficedula albicollis, the collared flycatcher, using a 9‐year database, and compared our results with those obtained in other populations of the same and the sister species. Between‐individual variation of forehead patch size was large, its repeatability larger than, and heritability similar to the Swedish population. Unlike in the other populations, the trait proved unaffected by body condition, and only very slightly influenced by age. There was no relationship between forehead patch size and breeding lifespan, and a marginal negative association with survivorship in adult males. Our results suggest that additive genetic variance of the trait in this population is large, but genes act independently of body condition, and there is no viability indicator value of the trait. This is the first report of a qualitative intraspecific difference in proximate determination of a sexually selected trait.

Carotenoids in the egg yolks of collared flycatchers (Ficedula albicollis) in relation to parental quality, environmental factors and laying order

Birds may influence the fitness of their offspring by transmission of different amounts of carotenoids to their eggs. Carotenoids play crucial roles in antioxidant protection and immune defence mechanisms, but they may be available to females in limiting amounts. Therefore, their allocation to the eggs may be influenced by the female’s condition, age and environmental circumstances. Furthermore, the quality of the male parent, which affects the reproductive value of the offspring, may also influence this investment. In this correlational study, we investigated proximate and ultimate factors that may lead to variation in yolk lutein, zeaxanthin and β-carotene concentrations among and within clutches of a wild passerine, the collared flycatcher (Ficedula albicollis). We found that carotenoid concentration was positively associated with caterpillar supply at the time of egg formation, which suggests a proximate constraint of carotenoid availability on yolk composition. Neither female condition, body size, age, nor male plumage ornamentation, age and body size correlated with carotenoid deposition. Yolk β-carotene concentration was found to be positively linked to yolk testosterone concentration. We suggest that females allocated more β-carotene to their eggs to mitigate the potentially detrimental effects of elevated steroid concentration. We found that concentration of β-carotene increased with laying order. The possible function of this pattern may be to enhance the resistance to oxidative stress and pathogens of the disadvantaged last-hatching nestling, suggesting that collared flycatchers pursue a compensatory, “brood survival” strategy.

EFFECTS OF ENVIRONMENTAL CONDITIONS AND PARENTAL QUALITY ON INTER- AND INTRACLUTCH EGG-SIZE VARIATION IN THE COLLARED FLYCATCHER (FICEDULA ALBICOLLIS)

Abstract Egg size is a particularly important life-history trait mediating maternal influences on offspring phenotype. Females can vary their egg-size investment in relation to environmental circumstances, their own breeding condition, and the quality of their mate. Here we analyzed inter- and intraclutch variation in egg size in the Collared Flycatcher (Ficedula albicollis) on the basis of eight years of data. According to our results, mean egg size increased with female condition, but did not differ among young, middle-aged, and old females. The male’s age, body size, and forehead patch size did not influence egg size; thus, we found no evidence for differential investment in egg size in relation to male quality. We found no effect of laying date on egg size when controlling for ambient temperature during the egg formation period, yet temperature had a significant effect on egg size. That result indicates proximate constraints on egg formation. Furthermore, we report on annual differences in intraclutch egg-size variation. Egg size increased within clutches in years with a warm prelaying period; whereas in years when the weather during that period was cold, there was no significant intraclutch trend. Proximate considerations seem to explain the observed patterns of intraclutch egg-size variation; however, we cannot reject the adaptive explanation. Mean egg size and intraclutch egg-size variation were unrelated to clutch size. Therefore, we found no evidence for a trade-off between size and number of eggs within a clutch.

Rapid temporal change in the expression and age-related information content of a sexually selected trait.

The expression of sexual signals is often phenotypically plastic and also evolves rapidly. Few studies have considered the possibility that proximate determination – the pathway between genes and trait expression – may also be subject to both phenotypic plasticity and evolutionary change. We examined long‐term patterns in size, condition‐ and age‐dependence, repeatability and heritability of forehead patch size, a sexually selected plumage trait in male collared flycatchers. We also estimated survival and sexual selection on the phenotypic value of the trait. Forehead patch size linearly declined during the 15 years, probably due to the significantly negative survival selection. In addition, the expression of genetic variation for the ornament apparently underwent an age‐limited change, which implies a change in the information content of the signal to receivers. The persistent lack of condition‐dependence makes phenotypic plasticity an unlikely explanation to our results. This raises the possibility of a microevolutionary change of both expression and proximate determination during the study period.

Female ornamentation and territorial conflicts in collared flycatchers (Ficedula albicollis)

Female ornaments in species with conventional sex roles often indicate individual quality, but the evolutionary forces maintaining them are less clear. Sexual competition for breeding opportunities may represent an important role for female signals, especially in polygynous species, but there is little experimental evidence for this. The wing patch size (WPS) of female collared flycatchers indicates age and body condition and predicts social mating patterns. We challenged nest-building females with decoy females of varying WPS and found that the aggressive response of residents increased with decoy WPS, suggesting a role for this female ornament in territorial competition. Our results explain why female WPS predicts territorial distances when mated to a polygynous male and indicate that the role of WPS in female competitive interactions is similar to that in males of the same population.

Paternal age and offspring growth: separating the intrinsic quality of young from rearing effects

Younger individuals are often less successful in reproduction than older ones. This might be because of improving breeding skills with age or because the genetic quality of young or early maternal effects on them vary with parental age. However, no attempt has been made to experimentally separate these processes in vertebrates. We conducted a cross-fostering experiment in collared flycatchers (Ficedula albicollis) in three breeding seasons to disentangle origin- and rearing-related effects of paternal age on chick growth, while controlling for date-specific environmental conditions as well as differences in clutch and brood size. The age of the male at the nest of origin, but not that of the rearing male, had a year-dependent effect on nestling body mass and tarsus length. In two seasons, young of subadult males grew slower in the early linear phase of growth than young of adult males. There was no compensatory growth in the final asymptotic phase, so both body mass and tarsus length before fledging reflected the differential early development. In the remaining year, the age of the male at the nest of origin had no significant effect on chick growth. The environment-dependent origin effect we detected was unexplained by incubation times, hatching asynchrony, chick masses at swapping or previously described age-dependent egg quality patterns. Our results therefore suggest a genotype × environment interaction on the relative development of offspring sired by subadult and adult males. Our results also raise the possibility that female birds may gain genetic benefits by mating with older males. Further studies should identify general patterns of male age-dependent female mate choice and offspring quality in different environmental conditions.

Phenotypic plasticity in a conspicuous female plumage trait: information content and mating patterns

Explaining sexual ornamentation in the limiting sex, usually females, requires information on the proximate background of ornaments and their consequences for sexual selection. Phenotypic variation within individuals has received little attention in either of these research directions. We used 6 years of data to examine the information content and potential role of white wing patch size in female collared flycatchers, Ficedula albicollis. Female wing patch size differed among years. Yearling females had smaller wing patches than older females. The negative effect of original patch size on intraindividual patch size change was stronger in yearling than in older birds, which may reflect an age-dependent trade-off. Change in wing patch size was strongly positively related to the summer North Atlantic Oscillation (NAO) index. Clutch size laid in the previous breeding season had a negative effect on patch size change, but only in high-NAO years. Thus, the immediate effect of poor climate during moult apparently overrode the more indirect influence of reproductive effort. Two sexually selected ornaments of mates were unrelated to female wing patch size at the population level, but intraindividual changes of female wing patch size significantly predicted differences in wing patch size between mates obtained in the 2 years. Our data suggest that significant mating advantages to more ornamented females may not be detected from population-level mating patterns. Research on potential female ornaments should also pay more attention to age-dependent phenotypic plasticity, the trade-off between current and future ornament size and the costs of reproduction to apparent future attractiveness.

Nutritional correlates and mate acquisition role of multiple sexual traits in male collared flycatchers

The information content of a sexual signal may predict its importance in a multiple signal system. Many studies have correlated sexual signal expression with the absolute levels of nutrient reserves. In contrast, the changes of nutrient reserves associated with signal expression are largely unknown in the wild due to technical limitations although they are important determinants of signal information content. We compared two visual and eight acoustic sexual traits in male collared flycatchers to see whether the nutritional correlates of expression predict the role of the signal in sexual selection. We used single point assays of plasma lipid metabolites to estimate short-term changes in nutritional state in relation to sexual trait expression during courtship. As a measure of sexual selection, we estimated the relationship with pairing latency after arrival in a 4-year dataset. Males which found a mate rapidly were characterized by large wing and forehead patches, but small song strophe complexity and small figure repertoire size. Traits more strongly related to pairing latency were also more closely related to changes in nutrient reserves. This indicates a link between signal role and information content. Small wing patches and, surprisingly, complex songs seemed to indicate poor phenotypic quality and were apparently disfavoured at mate acquisition in our population. Future studies of the information content of sexual traits, especially dynamic traits such as song, may benefit from the use of plasma metabolite profiles as non-invasive indicators of short-term changes in body condition.