The impact of behavioral and physiological maternal effects on offspring sex ratio in the common snapping turtle,<Emphasis Type="Italic"> Chelydra serpentina</Emphasis>

Theory suggests that maternal effects are especially important in organisms with environmentally-sensitive sex-determining mechanisms. However, there is no substantive body of empirical evidence to confirm this conjecture. We integrated field and laboratory studies to jointly evaluate the significance of behavioral (nest-site choice) and physiological (yolk hormone allocation) maternal effects on offspring sex ratio in the common snapping turtle (Chelydra serpentina), a species with temperature-dependent sex determination (TSD). Of the 16 microhabitat variables measured, only three (south, east, and total overstory vegetation cover) were significantly correlated with nest temperature: cooler nests were located under more vegetation cover. In turn, these microhabitat predictors of nest temperature, and nest temperature itself, may influence nest sex ratio: shadier, cooler nests were more likely to produce a higher proportion of male offspring than less shady, warmer nests. Analysis of eggs from these same nests incubated in a common garden design in the laboratory revealed that clutch sex ratio was unaffected by levels of yolk estradiol, yolk testosterone, or their interaction. Examination of both behavioral and physiological maternal effects revealed no concordant impact on offspring sex ratio. However, eggs from nests that produced male-biased sex ratios in the field yielded higher proportions of males under constant-temperature conditions in the laboratory. Our study confirms the importance of behavioral maternal effects in nature on offspring sex ratios in species with TSD, while also revealing the potential presence of a predisposition for sex-ratio production underlying TSD in this system.Communicated by S. Krackow     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.     

Variation in nesting patterns affecting nest temperatures in two populations of painted turtles (<Emphasis Type="Italic">Chrysemys picta</Emphasis>) with temperature-dependent sex determination

Mechanisms maintaining sex ratios in populations with temperature-dependent sex determination (TSD) remain elusive. Although geographic variation in embryonic sex determination (i.e., pivotal temperature) has been widely investigated in reptiles exhibiting TSD, no previous studies have directly addressed geographic variation in maternal behavior affecting nest thermal conditions. I evaluated patterns of nest-site selection and its effects on thermal and hydric nest conditions for a population of painted turtles ( Chrysemys picta bellii) exhibiting TSD in New Mexico. These results are compared to data collected from a well-studied, conspecific population experiencing relatively cooler climatic conditions in Illinois. Since canopy vegetation cover reduces nest temperatures in Illinois, I expected females in New Mexico to nest under high amounts of canopy vegetation cover. However, females from New Mexico placed nests under significantly less canopy vegetation cover, but closer to standing water, than did females from Illinois. Experimental nests in New Mexico demonstrated that increased canopy vegetation cover and soil moisture reduced nest temperatures. By nesting close to standing water rather than under canopy vegetation cover, females in New Mexico nested in habitats more closely associated with maximizing moisture around nests rather than reducing nest temperatures through shading. Mean July nest temperatures were similar for both populations. Since nest hydric conditions affect hatching success and hatchling size in C. picta, nesting patterns in New Mexico may primarily reflect selection for microhabitats affecting offspring survivorship or size.     

Climate and predation dominate juvenile and adult recruitment in a turtle with temperature-dependent sex determination

Conditions experienced early in life can influence phenotypes in ecologically important ways, as exemplified by organisms with environmental sex determination. For organisms with temperature-dependent sex determination (TSD), variation in nest temperatures induces phenotypic variation that could impact population growth rates. In environments that vary over space and time, how does this variation influence key demographic parameters (cohort sex ratio and hatchling recruitment) in early life stages of populations exhibiting TSD? We leverage a 17-year data set on a population of painted turtles, Chrysemys picta, to investigate how spatial variation in nest vegetation cover and temporal variation in climate influence early life-history demography. We found that spatial variation in nest cover strongly influenced nest temperature and sex ratio, but was not correlated with clutch size, nest predation, total nest failure, or hatching success. Temporal variation in climate influenced percentage of total nest failure and cohort sex ratio, but not depredation rate, mean clutch size, or mean hatching success. Total hatchling recruitment in a year was influenced primarily by temporal variation in climate-independent factors, number of nests constructed, and depredation rate. Recruitment of female hatchlings was determined by stochastic variation in nest depredation and annual climate and also by the total nest production. Overall population demography depends more strongly on annual variation in climate and predation than it does on the intricacies of nest-specific biology. Finally, we demonstrate that recruitment of female hatchlings translates into recruitment of breeding females into the population, thus linking climate (and other) effects on early life stages to adult demographics.     

The multidimensionality of behavioural defences against brood parasites: evidence for a behavioural syndrome in magpies?

Studies of antiparasite defences against cuckoo parasites have largely neglected the possibility that behavioural components of host defence may correlate giving rise to a behavioural syndrome. Furthermore, the different contribution of the host’s sex in nest defence has traditionally been disregarded. Here, we studied magpie (Pica pica) mobbing behaviour towards dummies of great spotted cuckoo (Clamator glandarius) and non-harmful hoopoes (Upupa epops) and egg rejection of parasite eggs in a population of colour-banded magpies. We predicted a positive correlation between the intensity of nest defence and egg rejection within each sex and that females respond more intensely than males to the threat of brood parasitism as they undertake incubation. Magpie males, but not females, defended their nests more intensely in those nests in which cuckoo model eggs were rejected. Individual magpies did significantly differ in their baseline level of nest attentiveness; however, there were no individual differences once pair identity was considered. Males and females defended their nests more intensely when it was exposed to the presence of a great spotted cuckoo dummy. Males, but not females, were more prone to appear at their nests, and females, but not males, were more prone to defend more intensely when their nests were challenged by a parasite threat. Our results thus agree with the view that mobbing behaviour and egg rejection in magpies may actually constitute a pseudosyndrome and highlight the necessity to integrate interindividual variation and the sex of the host in studies of the evolution of host defences.     

Confronting two-sex demographic models with data

Most population dynamics models explicitly track the density of a single sex. When the operational sex ratio can vary, two-sex models may be needed to understand and predict population trajectories. Various functions have been proposed to describe the relative contributions of females and males to recruitment, and these functions can differ qualitatively in the patterns that they generate. Which mating function best describes the dynamics of real populations is not known, since alternative two-sex models have not been confronted with experimental data. We conducted the first such comparison, using laboratory populations of the bean beetle Callosobruchus maculatus. Manipulations of the operational sex ratio and total density provided strong support for a demographic model in which the birth rate was proportional to the harmonic mean of female and male densities, and females, males, and their offspring made unique contributions to density dependence. We offer guidelines for transferring this approach to other, less tractable systems in which possibilities for sex ratio manipulations are more limited. We show that informative experimental designs require strong perturbations of the operational sex ratio. The functional form of density dependence (saturating vs. over-compensatory) and the relative contributions of each sex to density dependence can both determine in which direction and at which population densities such perturbations would be most informative. Our experimental results and guidelines for design strategies promote synthesis of two-sex population dynamics theory with empirical data.     

Deforestation: risk of sex ratio distortion in hawksbill sea turtles.

Phenotypic sex in sea turtles is determined by nest incubation temperatures, with warmer temperatures producing females and cooler temperatures producing males. The common finding of highly skewed female-biased hatchling sex ratios in sea turtle populations could have serious repercussions for the long-term survival of these species and prompted us to examine the thermal profile of a relatively pristine hawksbill nesting beach in Guadeloupe, French West Indies. Data loggers placed at nest depth revealed that temperatures in the forested areas were significantly cooler than temperatures in the more open, deforested areas. Using these temperatures as a predictor of sex ratio, we were able to assess the relative contributions of the different beach zones to the primary sex ratio: significantly more males were likely to be produced in the forested areas. Coastal forests are therefore important male-producing areas for the hawksbill sea turtle, and this has urgent conservation implications. On Guadeloupe, as on many Caribbean islands, deforestation rates are high and show few signs of slowing, as there is continual pressure to develop beachfront areas. The destruction of coastal forest could have serious consequences both in terms of local nesting behavior and of regional demography through the effects on population sex ratios. Human alterations to nesting habitat in other reptile taxa have been shown to modify the thermal properties of nest sites in ways that can disrupt their ecology by allowing parasite transmission, increasing vulnerability to climate change, or rendering existing habitat unsuitable.     

Windows of embryonic sexual lability in two lizard species with environmental sex determination

Temperature-dependent sex determination (TSD) occurs in all major reptile lineages, but the selective forces and physiological mechanisms that link sex to incubation temperature may differ among and within those groups. Different models for TSD evolution make different predictions about when offspring sex will respond to environmental cues. Although TSD has evolved in several lizard lineages, there is less detailed information on these taxa than in turtles and crocodilians with TSD. We incubated eggs of an agamid lizard (Amphibolurus muricatus) and a scincid lizard (Bassiana duperreyi), two species with TSD. Rather than manipulate incubation temperature to identify periods of sexual lability (as in most previous studies of this topic), we topically applied the aromatase inhibitor fadrozole to eggs at a variety of times through the incubation period. Fadrozole application sex-reversed the resultant hatchlings if applied from the time of oviposition until at least 60% of the way through incubation. In all of the TSD lizard species studied so far, offspring sex is determined either while the eggs are held inside the mother's body or soon after oviposition, providing substantial maternal control over incubation temperatures at this critical period. Hence, the hypothesis that TSD evolves because it enables offspring sex to be matched to conditions that are unpredictable at the time of laying is less likely to apply to squamates than to turtles, sphenodontians, and (especially) crocodiles, in which the period of sexual lability is delayed until long after oviposition.     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Female nest defense in a coral-reef fish, Dascyllus albisella, with uniparental male care

Intraspecific variation in the patterns of parental care has been observed in a variety of animals; however, the possibility of parental care by a non-caregiving parent of uniparental species has not been thoroughly explored. In the coral-reef damselfish, Dascyllus albisella, only males normally exhibit parental care. In this study, we examined the response of females of this species to egg predators after experimental male removal and an elevated level of egg predation, at two small patch reefs (reefs 1 and 2) in Hawaii. We tested theoretical expectations that a nest was defended only by females which had spawned in the nest, and that larger females had a higher likelihood of defense than smaller females. A nest was defended against egg predators more frequently by females that had spawned in that nest than would be expected by chance. Not all females that had spawned in a given nest participated in defense. There was a positive association between female body length and the likelihood of defense at reef 2, but not at reef 1. Within a set of females that had spawned in the same nest during the same nesting cycle, defending females had larger body lengths than non-defending females at reef 2 but not at reef 1. Lack of association between female size and likelihood of defense at reef 1 was unexpected, but may correlate with the smaller average female size and smaller size differences among females on that reef.     

Hidden demographic impacts of fishing and environmental drivers of fecundity in a sea turtle population

Fisheries bycatch is a critical threat to sea turtle populations worldwide, particularly because turtles are vulnerable to multiple gear types. The Canary Current is an intensely fished region, yet there has been no demographic assessment integrating bycatch and population management information of the globally significant Cabo Verde loggerhead turtle (Caretta caretta) population. Using Boa Vista island (Eastern Cabo Verde) subpopulation data from capture–recapture and nest monitoring (2013–2019), we evaluated population viability and estimated regional bycatch rates (2016–2020) in longline, trawl, purse-seine, and artisanal fisheries. We further evaluated current nesting trends in the context of bycatch estimates, existing hatchery conservation measures, and environmental (net primary productivity) variability in turtle foraging grounds. We projected that current bycatch mortality rates would lead to the near extinction of the Boa Vista subpopulation. Bycatch reduction in longline fisheries and all fisheries combined would increase finite population growth rate by 1.76% and 1.95%, respectively. Hatchery conservation increased hatchling production and reduced extinction risk, but alone it could not achieve population growth. Short-term increases in nest counts (2013–2021), putatively driven by temporary increases in net primary productivity, may be masking ongoing long-term population declines. When fecundity was linked to net primary productivity, our hindcast models simultaneously predicted these opposing long-term and short-term trends. Consequently, our results showed conservation management must diversify from land-based management. The masking effect we found has broad-reaching implications for monitoring sea turtle populations worldwide, demonstrating the importance of directly estimating adult survival and that nest counts might inadequately reflect underlying population trends.     

Non-consumptive effects of predator presence on copepod reproduction: insights from a mesocosm experiment

Reproduction in planktonic animals depends on numerous biotic and abiotic factors. One of them is predation pressure, which can have both direct consumptive effects on population density and sex ratio, and non-consumptive effects, for example on mating and migration behaviour. In copepods, predator vulnerability depends on their sex, motility pattern and mating behaviour. Therefore, copepods can be affected at multiple stages during the mating process. We investigated the reproductive dynamics of the estuarine copepod Eurytemora affinis in the presence and absence of its predator the mysid Neomysis integer in a mesocosm experiment. We found that the proportion of ovigerous females decreased in the presence of predators. This shift was not caused by differential predation as the absolute number of females was unaffected by mysid presence. Presence of predators reduced the ratio of males to non-ovigerous females, but not by predation of males. Our combined results suggest that the shift from ovigerous to non-ovigerous females under the presence of predators was caused by either actively delayed egg production or by shedding of egg sacs. Nauplii production was initially suppressed in the predation treatment, but increased towards the end of the experiment. The proportion of fertilized females was similar in both treatments, but constantly fell behind model predictions using a random mating model. Our results highlight the importance of non-consumptive effects of predators on copepod reproduction and hence on population dynamics.     

Causes of couspecific nest parasitism in the northern masked weaver

Summary Conspecific nest parasitism is a common reproductive strategy in the northern masked weaver (Ploceus taeniopterus). Parasites appear to be females with nests of their own who lay an additional egg in another female's nest as a way to enhance their reproductive success. Brood size in the northern masked weaver is practically constrained to three: starvation in four-chick broods is very common. The constraint on brood size is probably imposed by the extreme hatching asynchrony characteristic of this species: the last egg in a four-egg clutch hatches 72 h after the first egg, and when a chick starves, it is almost always the youngest chick. Late-hatching chicks also grow more slowly than do early-hatching chicks. If a female were to lay a fourth egg in her own nest, there is little chance that it would succeed. However; if she places it in another female's nest before that host lays her third egg, then it may have a greater chance of success.     

Philopatry and nesting phenology of pine snakes Pituophis melanoleucus in the New Jersey Pine Barrens

Summary We studied nesting behavior of pine snakes Pituophis melanoleucus from 1977 to 1989. Female pine snakes in the New Jersey Pine Barrens excavate tunnels and a nest chamber for egg deposition. Over 70% of all nests had old, hatched shells remaining in the nests from previous years. The presence of hatched shells and the incidence of recaptured females in the same nest indicates nest site tenacity for some females. From 1977 to 1988 we branded 98 females and 45 were recaptured in the same general nesting area. All 45 recaptured females were recaptured one to four times at the same nest site; two of these snakes were captured in three years at the same site and in one year at a different site. We conclude that females show a high degree of nest site philopatry. We suggest that high nest site philopatry is selected for because it promotes location of sites that are safe from predators (buried, hatched shells) and contain suitable temperature conditions. Offprint requests to: J. Burger     

Demographic Evidence of Illegal Harvesting of an Endangered Asian Turtle

Harvesting pressure on Asian freshwater turtles is severe, and dramatic population declines of these turtles are being driven by unsustainable collection for food markets, pet trade, and traditional Chinese medicine. Populations of big‐headed turtle (Platysternon megacephalum) have declined substantially across its distribution, particularly in China, because of overcollection. To understand the effects of chronic harvesting pressure on big‐headed turtle populations, we examined the effects of illegal harvesting on the demography of populations in Hong Kong, where some populations still exist. We used mark‐recapture methods to compare demographic characteristics between sites with harvesting histories and one site in a fully protected area. Sites with a history of illegal turtle harvesting were characterized by the absence of large adults and skewed ratios of juveniles to adults, which may have negative implications for the long‐term viability of populations. These sites also had lower densities of adults and smaller adult body sizes than the protected site. Given that populations throughout most of the species’ range are heavily harvested and individuals are increasingly difficult to find in mainland China, the illegal collection of turtles from populations in Hong Kong may increase over time. Long‐term monitoring of populations is essential to track effects of illegal collection, and increased patrolling is needed to help control illegal harvesting of populations, particularly in national parks. Because few, if any, other completely protected populations remain in the region, our data on an unharvested population of big‐headed turtles serve as an important reference for assessing the negative consequences of harvesting on populations of stream turtles. Evidencia Demográfica de la Captura Ilegal de una Tortuga Asiática en Peligro     

Seasonal fluctuations in sand temperature: effects on the incubation period and mortality of loggerhead sea turtle (<Emphasis Type="Italic">Caretta</Emphasis><Emphasis Type="Italic">caretta</Emphasis>) pre-emergent hatchlings in Minabe,Japan

Incubation period, hatching success, and emergence percentage in loggerhead (Caretta caretta) nests were quantified during the 1993 and 1995 nesting seasons and following incubation seasons in Minabe, Wakayama, Japan. Sand and nest temperatures were also monitored. Over the seasons, daily mean sand temperature at nest depth fluctuated between 18.0°C and 33.3°C, with a steep increase in the second week of July and a peak in late August. Temperatures inside the nest chambers were a few degrees above those of the surrounding sand at the end of incubation. The incubation period ranged from 46 to 82 days. A significant negative correlation was found between mean sand temperature and incubation period. The relationship conformed to the day-degree concept. There was no significant seasonal trend in hatching success, but many pre-emergent hatchlings were found dead in most of the clutches during the warmest part of the season. Emergence percentage was correlated with mean sand temperature calculated for 4 days before emergence, suggesting that mortality may be due to heat. This heat-related mortality is considered to be a common phenomenon at our study site, because the peak in emergences coincides with the peak in high temperatures. These temperature effects on hatchling mortality must be taken into account in estimates of hatchling sex ratios. Because sand temperatures already exceed the optimal thermal range for incubation, this population is vulnerable to even small temperature increases resulting from global warming.     

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

A Contemporary,Sex‐Limited Change in Body Size of an Estuarine Turtle in Response to Commercial Fishing

Abstract: Juvenile growth rate and adult body size are important components of life‐history strategies because of their direct impact on fitness. The diamondback terrapin (Malaclemys terrapin) is a sexually dimorphic, long‐lived turtle inhabiting brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of its range, terrapins face anthropogenically imposed mortality: juveniles of both sexes inadvertently enter commercial crab traps and drown. For adult females, the carapace eventually grows large enough that they cannot enter traps, whereas males almost never reach that critical size. We compared age structure, carapace dimensions, growth curves, and indices of sexual dimorphism for a Chesapeake Bay population of terrapins (where mortality of turtles is high due to crab traps) with contemporary terrapins from Long Island Sound and museum specimens from Chesapeake Bay (neither group subject to commercial crab traps). We also calculated the allochronic and synchronic rates of evolutionary change (haldanes) for males and females to measure the rate of trait change in a population or between populations, respectively. We found a dramatic shift to a younger male age structure, a decrease in the length of time to terminal female carapace size, a 15% increase in female carapace width, and an increase in sexual dimorphism in Chesapeake Bay. In a new twist, our results implicate a fishery in the selective increase in size of a reptilian bycatch species. These sex‐specific changes in life history and demography have implications for population viability that need to be considered when addressing conservation of this threatened turtle.     

Are blue eggs a sexually selected signal of female collared flycatchers? A cross-fostering experiment

Impressive variation in egg colouration among birds has puzzled evolutionary biologists for a long time. The most frequently studied selective forces moulding egg colouration—predation and brood parasitism—have either received little empirical support or may play a role in only a minority of species. A novel hypothesis has suggested that egg colour may be significantly influenced by sexual selection. Females may deposit a blue-green pigment biliverdin into eggshells instead of using it for themselves as a powerful antioxidant. By this handicap, females may signal their quality to males, which are then hypothesized to increase their paternal effort. We experimentally tested the hypothesis in the collared flycatcher (Ficedula albicollis), a species laying blue-green eggs. We cross-fostered clutches between nests to disentangle effects of female/territory quality and egg colour on paternal effort and nestling quality. The results supported two assumptions of sexual signalling through egg colour hypothesis: Blue pigment seems to be a limited resource for females, and female quality is positively correlated with the intensity of the blue-green colour. However, we did not find support for the main prediction of the hypothesis, as male parental effort parameters (feeding frequencies to nestlings and intensity of nest defence) were unrelated to egg colour. We discuss possible reasons for the discrepancy between our results and previous correlative analyses that supported the hypothesis that blue egg colour may be a postmating, sexually selected signal in females.