Alula size signals male condition and predicts reproductive performance in an Arctic‐breeding passerine

While studies of achromatic plumage signaling are scarce relative to chromatic ornaments, achromatic ornaments have the potential to act as an efficient form of visual communication due to the highly conspicuous contrast between black and white body regions. Recently, achromatic plumage reflectance has been shown to indicate condition, yet the condition‐dependence of achromatic patch size remains unstudied. Here we show the first evidence that alula size, an achromatic plumage patch, has the potential to signal a male’s condition and predict reproductive performance. In Arctic‐breeding snow buntings Plectrophenax nivalis, the size of the alula simultaneously predicted pre‐breeding physiological health and the number of offspring produced, through an intermediate variable (lay date). Snow buntings appear to pair assortatively; males and females arriving earlier pair together, and changes in body condition over the breeding season are positively related within pairs. We suggest that simple achromatic plumage patches, like alula size, have the potential to act as condition‐dependent signals. Consequently, females may benefit from assessing these signals to reliably evaluate a male’s condition and reproductive potential as a means of maximizing their reproductive success.     

Genetics of plumage color in the Gyrfalcon (Falco rusticolus): analysis of the melanocortin-1 receptor gene

Genetic variation at the melanocortin-1 receptor (MC1R) gene is correlated with melanin color variation in a few reported vertebrates. In Gyrfalcon (Falco rusticolus), plumage color variation exists throughout their arctic and subarctic circumpolar distribution, from white to gray and almost black. Multiple color variants do exist within the majority of populations; however, a few areas (e.g., northern Greenland and Iceland) possess a single color variant. Here, we show that the white/melanic color pattern observed in Gyrfalcons is explained by allelic variation at MC1R. Six nucleotide substitutions in MC1R resulted in 9 alleles that differed in geographic frequency with at least 2 MC1R alleles observed in almost all sampled populations in Greenland, Iceland, Canada, and Alaska. In north Greenland, where white Gyrfalcons predominate, a single MC1R allele was observed at high frequency (>98%), whereas in Iceland, where only gray Gyrfalcons are known to breed, 7 alleles were observed. Of the 6 nucleotide substitutions, 3 resulted in amino acid substitutions, one of which (Val(128)Ile) was perfectly associated with the white/melanic polymorphism. Furthermore, the degree of melanism was correlated with number of MC1R variant alleles, with silver Gyrfalcons all heterozygous and the majority of dark gray individuals homozygous (Ile(128)). These results provide strong support that MC1R is associated with plumage color in this species.     

Seasonal movements of Gyrfalcons Falco rusticolus include extensive periods at sea

Little information exists on the movements of Gyrfalcons Falco rusticolus outside the breeding season, particularly amongst High Arctic populations, with almost all current knowledge based on Low Arctic populations. This study is the first to provide data on summer and winter ranges and migration distances. We highlight a behaviour previously unknown in Gyrfalcons, in which birds winter on sea ice far from land. During 2000–2004, data were collected from 48 Gyrfalcons tagged with satellite transmitters in three parts of Greenland: Thule (northwest), Kangerlussuaq (central‐west) and Scoresbysund (central‐east). Breeding home‐range size for seven adult females varied from 140 to 1197 km² and was 489 and 503 km² for two adult males. Complete outward migrations from breeding to wintering areas were recorded for three individuals: an adult male which travelled 3137 km over a 38‐day period (83 km/day) from northern Ellesmere Island to southern Greenland, an adult female which travelled 4234 km from Thule to southern Greenland (via eastern Canada) over an 83‐day period (51 km/day), and an adult female which travelled 391 km from Kangerlussuaq to southern Greenland over a 13‐day period (30 km/day). Significant differences were found in winter home‐range size between Falcons tagged on the west coast (383–6657 km²) and east coast (26 810–63 647 km²). Several Falcons had no obvious winter home‐ranges and travelled continually during the non‐breeding period, at times spending up to 40 consecutive days at sea, presumably resting on icebergs and feeding on seabirds. During the winter, one juvenile female travelled over 4548 km over an approximately 200‐day period, spending over half that time over the ocean between Greenland and Iceland. These are some of the largest winter home‐ranges ever documented in raptors and provide the first documentation of the long‐term use of pelagic habitats by any falcon. In general, return migrations were faster than outward ones. This study highlights the importance of sea ice and fjord regions in southwest Greenland as winter habitat for Gyrfalcons, and provides the first detailed insights into the complex and highly variable movement patterns of the species.     

Morphological and plumage colour variation in the Réunion grey white‐eye (Aves: Zosterops borbonicus): assessing the role of selection

The Réunion grey white‐eye (Zosterops borbonicus), a small passerine endemic to the island of Réunion (Mascarene archipelago), constitutes an extraordinary case of phenotypic variation within a bird species, with conspicuous plumage colour differentiation at a microgeographical scale. To understand whether natural selection could explain such variability, we compared patterns of variation in morphological and plumage colour traits within and among populations. To quantify morphological variation, we used measurements obtained by Frank Gill in the 1960s from 239 individuals collected in 60 localities distributed over the entire island of Réunion. To quantify colour variation, we measured the reflectance spectra of plumage patches of 50 males from a subset of Gill's specimens belonging to the five recognized plumage colour variants and used a visual model to project these colours in an avian‐appropriate, tetrachromatic, colour space. We found that variants occupy different regions of the avian colour space and that between‐variant differences for most plumage patches could be discriminated by the birds. Differences in morphology were also detected, but these were, in general, smaller than colour differences. Overall, we found that variation in both plumage colour and morphology among variants is greater than would be expected if genetic drift alone was responsible for phenotypic divergence. As the plumage colour variants correspond to four geographical forms, our results suggest that phenotypic evolution in the Réunion grey white‐eye is at least partly explained by divergent selection in different habitats or regions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 459–473.     

Plumage colour and sexual selection in the pied flycatcher Ficedula hypoleuca

The plumage colour of male pied flycatchers varies on a continuum from brownish and female-like to a conspicuous black-and-white. Males have a conspicuous plumage only during the breeding season, when the mortality rate was low and not correlated with plumage colour. Brownish males breed successfully, and breeding males had no lower annual rate of return than non-breeders. Experiments on male intrusion showed that possession of a cryptic, female-like plumage did not facilitate nest site settlement by such males. Dark males were on average older, arrived earlier, and defended more nestboxes than brown males. However, experiments showed that plumage colour was not a reliable indicator of the outcome of competitive combats between males. These facts argue against several existing hypotheses about variation in male plumage colour. A new explanation is proposed: a bright colour serves to signal presence and thereby reduces territory holding costs. The brownish males arrive only shortly before the females and thus will not benefit as much by being conspicuous as do the black-and-white males that arrive earlier in spring.     

Investment in testes,sperm‐duct glands and lipid reserves differs between male morphs but not between early and late breeding season in Pomatoschistus minutus

This study of the sand goby Pomatoschistus minutus, a nest‐holding fish with paternal care, focused on gonadal investment among males of different sizes collected early and late in the breeding season. All males caught at the nest had breeding colour, whereas trawl‐caught fish consisted of males both with and without colour. The absence or presence of breeding colour was a good predictor of testes investment. Compared to males with breeding colour, males without colour were smaller in body size but had extraordinarily large testes. In absolute terms, testes mass of males without breeding colour was on average 3·4 times greater than those of males with breeding colour. Since small colourless males are known to reproduce as sneaker males, this heavy investment in testes probably reflects that they are forced to spawn under sperm competition. Contrary to testes size, sperm‐duct glands were largest among males with breeding colour. These glands produce mucins used for making sperm‐containing mucous trails that males place in the nest before and during spawning. Since both sneakers and nest‐holders potentially could benefit from having large glands, this result is intriguing. Yet, high mucus production may be more important for nest‐holders, because it also protects developing embryos from infections. There was no significant effect of season on body size, testes or sperm‐duct glands size, but colourless males tended to be less common late in the season. Possibly this may indicate that individual small colourless males develop into their more colourful counterparts within the breeding season.     

Determinants of reproductive success and offspring sex in a turtle with environmental sex determination

Despite the importance of maternal effects in evolution, and knowledge of links among nest site choice, timing of nesting, offspring sex, and reproductive success in animals with environmental sex determination, these attributes have not been rigorously studied in a combined and natural context. To address this need we studied the relationships between three maternal traits (nest site choice, lay date, and nest depth) and two fitness‐related attributes of offspring (hatchling sex and embryonic survival) in the riverine turtle Carettochelys insculpta, a species with temperature‐dependent sex determination, for four years. Predation and flooding were the major sources of embryonic mortality in 191 nests. Embryonic survival was influenced by both lay date and nest site choice: in one year when nesting began later than average, nests laid later and at lower elevations were destroyed by early wet season river rises. In other years early nesting precluded flood mortality. However, turtles did not nest at the highest available elevations, and a field experiment confirmed that turtles were constrained to nest at lower elevations where they could construct a nest chamber. The principal determinant of hatchling sex in 140 nests was lay date, which in turn was apparently related to the magnitude of the previous wet season(s). Clutches laid earlier in the season (a female's first clutch) produced mainly males, while later clutches (her second clutch) yielded mostly females, due to seasonal increases in air temperatures. Accordingly, later nesting produced female‐biased hatchling sex ratios in 1996, while earlier nesting resulted in sex ratios near unity in the other years. However, all‐female nests were more likely to be flooded than mixed‐sex or all‐male nests in years when nesting was late. In conclusion, we found evidence that the position of two maternal trait distributions (elevation of the nest site and lay date), associated with the reproductive strategy of C. insculpta, reflect a combination of natural selection, physical constraints, and phenotypic plasticity. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 1–16.     

Reproductive ecology of the jacky dragon (Amphibolurus muricatus): an agamid lizard with temperature‐dependent sex determination

Abstract The jacky dragon, Amphibolurus muricatus (White, ex Shaw 1790) is a medium sized agamid lizard from the southeast of Australia. Laboratory incubation trials show that this species possesses temperature‐dependent sex determination. Both high and low incubation temperatures produced all female offspring, while varying proportions of males hatched at intermediate temperatures. Females may lay several clutches containing from three to nine eggs during the spring and summer. We report the first field nest temperature recordings for a squamate reptile with temperature‐dependent sex determination. Hatchling sex is determined by nest temperatures that are due to the combination of daily and seasonal weather conditions, together with maternal nest site selection. Over the prolonged egg‐laying season, mean nest temperatures steadily increase. This suggests that hatchling sex is best predicted by the date of egg laying, and that sex ratios from field nests will vary over the course of the breeding season. Lizards hatching from eggs laid in the spring (October) experience a longer growing season and should reach a larger body size by the beginning of their first reproductive season, compared to lizards from eggs laid in late summer (February). Adult male A. muricatus attain a greater maximum body size and have relatively larger heads than females, possibly as a consequence of sexual selection due to male‐male competition for territories and mates. If reproductive success in males increases with larger body size, then early hatching males may obtain a greater fitness benefit as adults, compared to males that hatch in late summer. We hypothesize that early season nests should produce male‐biased sex ratios, and that this provides an adaptive explanation for temperature‐dependent sex determination in A. muricatus.     

Temporal Variation in Decisions about Parental Care in Bluegill, Lepomis macrochirus

Parental investment theory states that an individual will trade‐off present and future reproductive potential to maximize lifetime reproductive success. Only when parental care is costly in terms of reduced future reproductive potential should individuals be sensitive to changes in the value of current offspring and adjust their care. Here, we examine temporal variation in parental care decision‐making in bluegill (Lepomis macrochirus), in which care is provided by males called ‘parentals’. Previous research has shown that parentals that nest early in the breeding season are in higher energetic condition than those that nest later, and early nesting males appear not to pay an opportunity cost to their care in terms of reduced future reproductive potential. Early nesting males also may have higher paternity in their broods than later nesting males. To examine the parental care decisions made by early and mid‐season nesting parentals, we experimentally reduced males’ perceived paternity by swapping eggs between nests. We found that experimental males that nested early in the breeding season adjusted their brood defence behaviour similarly to control males, which had sham egg swaps performed. Conversely, experimental males that nested mid‐season significantly decreased their brood defence behaviour after the manipulation as compared with control males. Thus, unlike mid‐season nesting males, early nesting males appear relatively insensitive to changes in brood value (paternity), possibly because early nesting males pay little cost in terms of reduced future reproductive potential to providing full care or because these males have a predisposition to high paternity.     

Experimental evidence that brighter males sire more extra‐pair young in tree swallows

Across taxa, extra‐pair mating is widespread among socially monogamous species, but few studies have identified male ornamental traits associated with extra‐pair mating success, and even fewer studies have experimentally manipulated male traits to determine whether they are related directly to paternity. As a consequence, there is little experimental evidence to support the widespread hypothesis that females choose more ornamented males as extra‐pair mates. Here, we conducted an experimental study of the relationship between male plumage colour and fertilization success in tree swallows (Tachycineta bicolor), which have one of the highest levels of extra‐pair mating in birds. In this study, we experimentally dulled the bright blue plumage on the back of males (with nontoxic ink markers) early in the breeding season prior to most mating. Compared with control males, dulled males sired fewer extra‐pair young, and, as a result, fewer young overall. Among untreated males, brighter blue males also sired more extra‐pair young, and in paired comparisons, extra‐pair sires had brighter blue plumage than the within‐pair male they cuckolded. These results, together with previous work on tree swallows, suggest that extra‐pair mating behaviour is driven by benefits to both males and females.     

Mate choice in Darwin's Finches

Female Geospiza conirostris on Isla Genovesa, Galapagos, pair preferentially with males who have had previous breeding experience. They choose mates on the basis of courtship behaviour and black adult plumage. By mating with experienced black males, they gain a fitness advantage in terms of fledgling production and recruitment of young into the breeding population. Behavioural signs of past breeding experience and black plumage are reliable age- and condition-dependent traits. We suggest that females use conspicuous black plumage to identify old males at a distance, then interactions through courtship to modify initial assessments. Females paired with inferior males may increase the genetic quality of their offspring by extra-pair copulations; results of heritability analysis of morphology are consistent with this suggestion. Females change mates at a frequency of 12–27% per breeding season. They re-pair with males who are generally old, experienced, and hold territories adjacent to the deserted male. Females that re-pair gain a benefit, whereas males who are deserted within a breeding season incur a cost of more than 50% of their future potential production for that season. We conclude that females in choosing males seek reliable indicators of potential parental care, and in addition they may seek indicators of genetic quality.     

Supplemental food increases ornamentation of male nestling Eastern Bluebirds

Although the condition‐dependence and signaling function of ornamental plumage coloration among adult males is well studied, less research has focused on the information content of ornamental coloration among juvenile birds. Eastern Bluebird (Sialia sialis) nestlings grow their nuptial plumage while in the nest and dependent on parents for food, making them an ideal species for studying the development and function of elaborate plumage. Previous research suggests that plumage brightness of Eastern Bluebirds functions, in the juvenile stage, in parent–offspring interactions as a sexually selected trait in adults. Using an experimental approach, we tested the effects of supplemental food on the structural plumage coloration (i.e., tips of primary feathers) of Eastern Bluebird nestlings in Watauga County, North Carolina, during the 2011 breeding season. We provided supplemental mealworms daily to breeding pairs from the onset of incubation through the nestling period, and measured plumage brightness, UV chroma, and mass of nestlings (N = 89 males and 71 females). Male nestlings of supplementally fed parents exhibited brighter plumage. The mass and UV chroma of young bluebirds were not significantly affected by food supplementation. However, the relationship between mass and brightness differed between male nestlings in the control and supplementally fed treatments. Males reared in food‐supplemented territories exhibited a positive relationship between color and mass. Nestlings in control territories, however, exhibited a negative relationship between size and brightness, suggesting that reduced food availability results in a tradeoff between allocating resources toward somatic growth and development of bright plumage. Our results suggest that UV‐blue structural plumage in male juvenile Eastern Bluebirds is at least partially condition‐dependent and may help to explain why plumage color can influence social interactions in Eastern Bluebirds.     

Achromatic plumage reflectance predicts reproductive success in male black-capped chickadees

The size of achromatic (black, white, gray) plumage patchesserves as a male status signal in many species of birds, butvariation in the colors of these patches has received littleattention. We assessed the relation between achromatic plumagereflectance, dominance rank, body condition, and reproductivesuccess in male black-capped chickadees, Poecile atricapillus.We measured plumage reflectance for five body regions of 40male chickadees in late winter and monitored these males throughoutthe following breeding season to determine whether they survivedto breed, whether they successfully paired, whether their partnerlaid eggs, and both their apparent and realized reproductivesuccess. As expected from past studies, a male's dominance ranksignificantly predicted whether his partner laid eggs. However,only achromatic plumage reflectance significantly predictedother measures of male reproductive performance. Among maleswho fledged at least one offspring, both the brightness of whiteplumage regions and the UV-chroma of melanin-based plumage regionswere significant predictors of the proportion of within-pairyoung in their nests. When we consider all males we measured,assigning zero values to males who failed to sire any offspring,the UV-chroma of melanin-based plumage regions was a significantpredictor of realized reproductive success. Bib size was alsorelated to male realized reproductive success. Our findingssuggest that individual variation in achromatic plumage mayplay an important role in sexual signaling in chickadees.     

Parental care does not vary with age‐dependent plumage in male Saffron Finches Sicalis flaveola

The status signalling hypothesis states that conspicuous male plumage varies among males and serves as an honest signal of male quality and competitive ability. We expected immature‐plumaged males of the Saffron Finch Sicalis flaveola to feed their incubating mates and nestlings and remove faecal sacs at lower rates than those of mature‐plumaged males. We also predicted that females paired with immature‐plumaged males would compensate for their mates’ lower contribution. We found no differences in either feeding rates or sanitation rates between immature‐ and mature‐plumaged males. Similarly, females mated to immature‐ and mature‐plumaged males fed nestlings at equivalent rates. Apparently, male plumage colour and age are not reliable signals of the ability of a male Saffron Finch to provide for his mate and offspring.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Double brooding and offspring desertion in the barn owl Tyto alba

Many bird species produce two annual broods during a single breeding season. However, not all individuals reproduce twice in the same year suggesting that double brooding is condition‐dependent. In contrast to most raptors and owls, the barn owl Tyto alba produces two annual clutches in most worldwide distributed populations. Nevertheless, the determinants of double brooding are still poorly studied. We performed such a study in a Swiss barn owl population monitored between 1990 and 2014. The annual frequency of double brooding varied from 0 to 14% for males and 0 to 59% for females. The likelihood of double brooding was higher when individuals initiated their first clutch early rather than late in the season and when males had few rather than many offspring at the first nest. Despite the reproductive benefits of double brooding (single‐ and double‐brooded individuals produced 3.97 ± 0.11 and 7.07 ± 0.24 fledglings, respectively), double brooding appears to be traded off against offspring quality because at the first nest double‐brooded males produced poorer quality offspring than single‐brooded males. This might explain why females desert their first mate to produce a second brood with another male without jeopardizing reproductive success at the first nest. Furthermore, the reproductive cycle being very long in the barn owl (120 d from start of laying to offspring independence), selection may have favoured behaviours that accelerate the initiation of a second annual brood. Accordingly, half of the double‐brooded females abandoned their young offspring to look for a new partner in order to initiate the second breeding attempt, 9.48 d earlier than when producing the second brood with the same partner. We conclude that male and female barn owls adopt different reproductive strategies. Females have more opportunities to reproduce twice in a single season than males because mothers are not strictly required during the entire rearing period in contrast to fathers. A high proportion of male floaters may also encourage females to desert their first brood to re‐nest with a new male who is free of parental care duties.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

Colour Polymorphism and Alternative Breeding Strategies: Effects of Parent’s Colour Morph on Fitness Traits in the Common Wall Lizard

Colour polymorphism (CP) is widespread in animals, but mechanisms underlying morph evolution and maintenance are not completely resolved. In reptiles, CP is often genetically based and associated with alternative behavioural strategies, mainly in males for most cases. However, female colour morphs also display alternative reproductive strategies associated with behavioural and physiological traits, which may contribute to maintain CP in the population. Both sexes of the common wall lizard (Podarcis muralis) show three pure colour morphs, white, yellow and red. Here, we looked for the effects of male and female colour morphs on fitness traits of captive-breeding pairs. All yellow-throated females laid clutches of many small eggs and produced many light offspring, behaving as r-strategists, whereas white-throated females laid clutches of few large eggs and produced few heavy offspring, behaving as K-strategists. Red-throated females adopted a conditional Kr-strategy depending on their size/age. These basic female strategies were modulated in relation to mate morph: white females had the best fitness gain in terms of viable offspring when mated to red males; mating between yellow morphs yielded a greater breeding success than all other morph crosses, but also lighter offspring; finally, red females produced heavy progeny when paired with red or white males, and light offspring in pair with yellow males. Thus, correlation between CP and traits relevant to fitness combined with non-random mating, either assortative or disassortative, could increase the potential for CP to contribute to divergent evolution in the common wall lizard.     

Offspring sex ratio allocation in the parasitic jaeger: selection for pale females and melanic males?

The maintenance of plumage color polymorphism in the parasiticjaeger (Stercorarius parasiticus) is still not well understood.Earlier studies indicated that selection may favor pale femalesand melanic males. If so, females would maximize their fitness,producing pale female and melanic male offspring. We thereforepredicted that females might bias their offspring sex ratiotoward daughters in pale pairs and toward sons in melanic pairs.Females might also choose to mate assortatively in relationto plumage color, thereby maximizing the probability of producingeither pale or melanic offspring. Because females are largerthan males, differential rearing costs may affect the offspringsex ratio independent of parental plumage color. We examinedoffspring sex ratio allocation, breeding variables indicativeof parental quality, and mating pattern in relation to plumagecolor in a colony of parasitic jaegers in northern Norway. Jaegerstended to mate assortatively in relation to plumage color. Thereproductive performance declined with season, and matched pairsappeared to be of lower quality than mixed pairs. The proportionof male offspring increased with hatching date in matched paleand mixed pairs, whereas the situation was reversed in matchedmelanic pairs. Matched pale pairs produced an overall surplusof favorable pale but costly daughters despite their lower quality,while melanic pairs produced a surplus of favorable melanicsons. However, differential offspring rearing costs and parentalrearing capacity may have additionally affected the realizedoffspring sex ratio. Mixed pairs producing an overall surplusof pale and melanic daughters allocated their resources accordingto differential rearing costs and parental quality only. Wesuggest that both strategies of sex ratio allocation togetherwith differences in reproductive success in matched versus mixedpairs may have a balancing effect on the mating pattern betweenplumage morphs and may contribute to the maintenance of thecolor polymorphism in this species.     

Telomere length in relation to colour polymorphism across life stages in the tawny owl

Telomere erosion has been proposed to be tightly associated with senescence, environmental stressors and life history trade‐offs. How telomere dynamics vary across life stages and especially in relation to (heritable) phenotypic traits is still unclear. The tawny owl Strix aluco display a highly heritable melanin‐based colour polymorphism, a grey and a brown morph, linked to several fitness traits including morph‐specific telomere dynamics. As adults, brown tawny owls have shorter relative telomere length (RTL) and exhibit faster telomere shortening rate than grey owls. Here we test if these morph‐specific telomere dynamics emerge already during growth, or if they are induced only in adult life through differential physiological costs associated with the life history of the morphs. We analysed RTL from 287 tawny owl offspring and 81 first breeding adults to evaluate at what life stage morph‐specific patterns emerge. We found no differences in RTL between the two morphs during the nestling period nor at the first breeding attempt. Sex, brood size or size rank in the nest did not affect offspring RTL. Among first‐breeders, females had shorter telomeres than males suggesting a sampling‐time dependent difference in reproductive costs between sexes, due to the prominent sex roles in tawny owls in the early nestling period. The probability to return to breed after the first breeding attempt was not affected by RTL, sex or colour morph. The lack of morph‐specific difference in RTL among nestlings and first breeders suggests that previously observed morph‐specific differences in RTL dynamics in adults emerge at the onset of the breeding career and is likely due to different physiological profiles and life‐history strategies adopted by adults. We conclude that different telomere dynamics and senescence patterns among highly heritable phenotypes (colour morphs) are likely to be a result of differential costs of reproduction and self‐maintenance.