Anthropogenic bottom-up and top-down impacts on boreal breeding waterbirds.

Wetland habitats are changing under multiple anthropogenic pressures. Nutrient leakage and pollution modify physico-chemical state of wetlands and affect the ecosystem through bottom-up processes, while alien predators affect the ecosystems in a top-down manner. Boreal wetlands are important breeding areas for several waterbird species, the abundances of which potentially reflect both bottom-up and top-down ecosystem processes. Here, we use long-term national monitoring data gathered from c. 130 waterbird breeding sites in Finland from the 1980s to the 2020s. We hypothesised that the physico-chemical state of the waters and increasing alien predator abundance both play a role in steering the waterbird population trends. We set out to test this hypothesis by relating population changes of 17 waterbird species to changes in water chemistry and to regional alien predator indices while allowing species-specific effects to vary with foraging niche (dabblers, invertivore divers, piscivorous divers, herbivores), nesting site, female mass and habitat (oligotrophic, eutrophic). We found niche and nesting site-specific, habitat-dependent changes in waterbird numbers. While the associations with higher phosphorus levels and browning water were in overall positive at the oligotrophic lakes, the numbers of invertivore and piscivore diving ducks were most strongly negatively associated with higher phosphorus levels and browning water at the eutrophic lakes. Furthermore, increased pH levels benefitted piscivores. Invertivore diving duck species nesting on the wetlands had declined most on sites with high alien predator indices. Large herbivorous species and species preferring oligotrophic lakes seem to be successful. We conclude that the large-scale breeding waterbird decline in Finland is closely connected to both bottom-up and top-down processes, where negative associations are emphasised especially at eutrophic lakes. Niche-, nest site- and habitat-specific management actions are required to conserve declining waterbird populations. Managing wetlands on catchments level together with alien predator control may provide important approaches to future wetland management.

Drivers of within- and among-individual variation in risk-taking behaviour during reproduction in a long-lived bird.

Plastic and selective mechanisms govern parental investment adjustments to predation threat. We investigated the relative importance of plasticity and selection in risk-taking propensity of incubating female common eiders Somateria mollissima facing unprecedented predation in SW Finland, Baltic Sea. Using a 12-year individual-based dataset, we examined within- and among-individual variation in flight initiation distance (FID), in relation to predation risk, nest detectability, individual traits and reproductive investment (NFID = 1009; Nindividual = 559). We expected females nesting in riskier environments (higher predation risk, lower nest concealment) to mitigate environmentally imposed risk by exhibiting longer FIDs, and females investing more in current reproduction (older, in better condition or laying larger clutches) to display shorter FIDs. The target of predation-adult or offspring-affected the mechanisms adapting risk-taking propensity; females plastically increased their FID under higher adult predation risk, while risk-avoiding breeders were predominant on islands with higher nest predation risk. Risk-taking females selected thicker nest cover, consistent with personality-matching habitat choice. Females plastically attenuated their anti-predator response (shorter FIDs) with advancing age, and females in better body condition were more risk-taking, a result explained by selection processes. Future research should consider predator type when investigating the fitness consequences of risk-taking strategies.

Temporal escape-adaptation to eutrophication by Skeletonema marinoi.

Diatoms commonly set off the spring-bloom in temperate coastal environments. However, their temporal offset may change in regions subject to nutrient enrichment, and by peaking earlier, such populations can maintain their position in the vernal plankton succession. We tested whether the marine keystone diatom Skeletonema marinoi can accomplish this through thermal evolutionary adaptation. Eight geographically separated subpopulations, representing hydromorphologically and climatologically similar inlets displaying a range of trophic states, were compared in a common-garden experiment. At early-spring temperatures, both doubling times and variation coefficients thereof, correlated negatively with the trophic state of the environment of origin, indicating selection for fast growth due to eutrophication. At mid-spring temperatures, the relationships were reversed, indicating selection in the opposite direction. At late-spring temperatures, no significant relationships were detected, suggesting relaxed selection. Subsequent field observations reflected these findings, where blooming temperatures decreased with trophic state. Natural selection thus moves along with eutrophication towards colder temperatures earlier in the spring, favouring genotypes with the capacity to grow fast. The thermal niche shift demonstrated herein may be an evolutionary mechanism essentially leading to trophic changes in the local ecosystem.

Mitigating impacts of invasive alien predators on an endangered sea duck amidst high native predation pressure.

Anthropogenically introduced invasive species represent a major threat to global biodiversity by causing population declines and extinctions of native species. The negative impacts of introduced predators are well documented, yet a fundamental knowledge gap exists regarding the efficiency of potential mitigation methods to restore the ecosystem. Other understudied aspects concern prey behavioural antipredator responses and the historical context of native predator-prey interactions, which may moderate invasion impacts on native prey. Invasion impacts of American mink (Neovison vison) and raccoon dog (Nyctereutes procyonoides) into the Baltic Sea archipelago are poorly understood, and the efficiency of removal efforts as a means to alleviate depredation pressure on native prey is debated. Here, we examine the effectiveness of invasive predator removal on ground-nesting female common eider (Somateria mollissima) mortality, breeding success and breeding propensity over a 9-year period, while controlling for predation risk imposed by the main native predator, the white-tailed eagle (Haliaeetus albicilla). Our results clearly show that intensified removal of American minks and raccoon dogs decreased the number of female eiders killed during nesting, while improving both nesting success and breeding propensity. Such obvious positive effects of invasive predator removal are particularly noteworthy against the backdrop of a soaring eagle population, indicating that the impacts of invasives may become accentuated when native predators differ taxonomically and by hunting mode. This study shows that invasive alien predator removal is an effective conservation measure clearly aiding native fauna even under severe native predation pressure. Such cost-effective conservation actions call for governmental deployment across large areas.

Detrimental impacts of climate change may be exacerbated by density-dependent population regulation in blue mussels.

The climate on our planet is changing and the range distributions of organisms are shifting in response. In aquatic environments, species might not be able to redistribute poleward or into deeper water when temperatures rise because of barriers, reduced light availability, altered water chemistry or any combination of these. How species respond to climate change may depend on physiological adaptability, but also on the population dynamics of the species. Density dependence is a ubiquitous force that governs population dynamics and regulates population growth, yet its connections to the impacts of climate change remain little known, especially in marine studies. Reductions in density below an environmental carrying capacity may cause compensatory increases in demographic parameters and population growth rate, hence masking the impacts of climate change on populations. On the other hand, climate-driven deterioration of conditions may reduce environmental carrying capacities, making compensation less likely and populations more susceptible to the effects of stochastic processes. Here we investigate the effects of climate change on Baltic blue mussels using a 17-year dataset on population density. Using a Bayesian modelling framework, we investigate the impacts of climate change, assess the magnitude and effects of density dependence, and project the likelihood of population decline by the year 2030. Our findings show negative impacts of warmer and less saline waters, both outcomes of climate change. We also show that density dependence increases the likelihood of population decline by subjecting the population to the detrimental effects of stochastic processes (i.e. low densities where random bad years can cause local extinction, negating the possibility for random good years to offset bad years). We highlight the importance of understanding, and accounting for both density dependence and climate variation when predicting the impact of climate change on keystone species, such as the Baltic blue mussel.

Sex-specific effects of the in ovo environment on early-life phenotypes in eiders.

Maternal effects affect offspring phenotype and fitness. However, the roles of offspring sex-specific sensitivity to maternal glucocorticoids and sex-biased maternal investment remain unclear. It is also uncertain whether telomere length (a marker associated with lifespan) depends on early growth in a sex-specific manner. We assessed whether maternal traits including corticosterone (CORT; the main avian glucocorticoid) and in ovo growth rate are sex-specifically related to offspring CORT exposure, relative telomere length (RTL) and body condition in eiders (Somateria mollissima). We measured feather CORT (fCORT), RTL and body condition of newly hatched ducklings, and growth rate in ovo was expressed as tarsus length at hatching per incubation duration. Maternal traits included baseline plasma CORT, RTL, body condition and breeding experience. We found that fCORT was negatively associated with growth rate in daughters, while it showed a positive association in sons. Lower offspring fCORT was associated with higher maternal baseline plasma CORT, and fCORT was higher in larger clutches and in those hatching later. The RTL of daughters was negatively associated with maternal RTL, whereas that of males was nearly independent of maternal RTL. Higher fCORT in ovo was associated with longer RTL at hatching in both sexes. Duckling body condition was mainly explained by egg weight, and sons had a slightly lower body condition. Our correlational results suggest that maternal effects may have heterogeneous and even diametrically opposed effects between the sexes during early development. Our findings also challenge the view that prenatal CORT exposure is invariably associated with shorter telomeres.

Annual variation in predation risk is related to the direction of selection for brain size in the wild.

The direction of predator-mediated selection on brain size is debated. However, the speed and the accuracy of performing a task cannot be simultaneously maximized. Large-brained individuals may be predisposed to accurate but slow decision-making, beneficial under high predation risk, but costly under low risk. This creates the possibility of temporally fluctuating selection on brain size depending on overall predation risk. We test this idea in nesting wild eider females (Somateria mollissima), in which head volume is tightly linked to brain mass (r2 = 0.73). We determined how female relative head volume relates to survival, and characterized the seasonal timing of predation. Previous work suggests that relatively large-brained and small-brained females make slow versus fast nest-site decisions, respectively, and that predation events occur seasonally earlier when predation is severe. Large-brained, late-breeding females may therefore have higher survival during high-predation years, but lower survival during safe years, assuming that predation disproportionately affects late breeders in such years. Relatively large-headed females outsurvived smaller-headed females during dangerous years, whereas the opposite was true in safer years. Predation events occurred relatively later during safe years. Fluctuations in the direction of survival selection on relative brain size may therefore arise due to brain-size dependent breeding phenology.

Protected areas act as a buffer against detrimental effects of climate change-Evidence from large-scale, long-term abundance data.

Climate change is driving species to shift their distributions toward high altitudes and latitudes, while habitat loss and fragmentation may hamper species ability to follow their climatic envelope. These two drivers of change may act in synergy, with particularly disastrous impacts on biodiversity. Protected areas, PAs, may thus represent crucial buffers against the compounded effects of climate change and habitat loss. However, large-scale studies assessing the performance of PAs as such buffers remain scarce and are largely based on species occurrence data. Conversely, abundance data have proven to be more reliable for addressing changes in wildlife populations under climate change. We evaluated changes in bird abundance from the 1970s-80s to the 2000s inside and outside PAs at the trailing range edge of 30 northern bird species and at the leading range edge of 70 southern species. Abundances of retracting northern species were higher and declined less inside PAs at their trailing range edge. The positive effect of PAs on bird abundances was particularly marked in northern species that rely strongly on PAs, that is, their density distribution is largely confined within PAs. These species were nearly absent outside PAs in the 2000s. The abundances of southern species were in general lower inside PAs and increased less from the 70s-80s to 2000s. Nonetheless, species with high reliance on PAs had much higher abundances inside than outside PAs in the 2000s. These results show that PAs are essential in mitigating the retraction of northern species, but also facilitate northward expansions of southern species highly reliant on PAs. Our study provides empirical evidence documenting the role of PAs in facilitating species to adjust to rapidly changing climatic conditions, thereby contributing to the mitigation of impending biodiversity loss. PAs may thus allow time for initiating wider conservation programs on currently unprotected land.

Counteracting wetland overgrowth increases breeding and staging bird abundances.

Human actions have led to loss and degradation of wetlands, impairing their suitability as habitat especially for waterbirds. Such negative effects may be mitigated through habitat management. To date scientific evidence regarding the impacts of these actions remains scarce. We studied guild specific abundances of breeding and staging birds in response to habitat management on 15 Finnish wetlands. In this study management actions comprised several means of vegetation removal to thwart overgrowth. Management cost efficiency was assessed by examining the association between site-specific costs and bird abundances. Several bird guilds exhibited positive connections with both habitat management as well as with invested funds. Most importantly, however, red-listed species and species with special conservation concern as outlined by the EU showed positive correlations with management actions, underlining the conservation value of wetland management. The results suggest that grazing was especially efficient in restoring overgrown wetlands. As a whole this study makes it clear that wetland habitat management constitutes a feasible conservation tool. The marked association between invested funds and bird abundance may prove to be a valuable tool for decision makers when balancing costs and impact of conservation measures against one another.

Blood and feather concentrations of toxic elements in a Baltic and an Arctic seabird population.

We report blood and feather concentrations of elements in the Baltic Sea and Arctic population of common eiders (Somateria mollissima). The endangered Baltic Sea population of eiders was demonstrably affected by element pollution in the 1990s. While blood concentrations of Hg were higher in Baltic breeding eiders, blood Se, As and Cd concentrations were higher in Arctic eiders. Blood concentrations of Pb, Cr, Zn and Cu did not differ between the two populations. While blood Pb concentrations had declined in Baltic eiders since the 1990s, Hg concentrations had not declined, and were above concentrations associated with adverse oxidative effects in other bird species. Inconsistent with blood concentrations, feather concentrations suggested that Pb, Zn, and Cd exposure was higher in Baltic eiders, and that Hg exposure was higher in Arctic eiders. Our study thus emphasizes the need for comprehensive evaluation of toxic element status, covering the annual cycle of a species.

DNA double-strand breaks in incubating female common eiders (Somateria mollissima): Comparison between a low and a high polluted area.

Alterations in the genetic material may have severe consequences for individuals and populations. Hence, genotoxic effects of environmental exposure to pollutants are of great concern. We assessed the impact of blood concentrations of persistent organic pollutants (POPs) and mercury (Hg) on DNA double-strand break (DSB) frequency, in blood cells of a high-exposed Baltic, and lower exposed Arctic population of common eiders (Somateria mollissima). Furthermore, we examined whether the genotoxic response was influenced by antioxidant concentration (plasma total glutathione (tGSH) and total antioxidant capacity) and female body mass. The DNA DSB frequency did not differ between the two populations. We found significant positive relationships between Hg and DNA DSB frequency in Baltic, but not in Arctic eiders. Although both p,p'-DDE and PCB 118 had a lesser effect than Hg, they exhibited a positive association with DNA DSB frequency in Baltic eiders. Antioxidant levels were not important for the genotoxic effect, suggesting alternative mechanisms other than GSH depletion for the relationship between Hg and DNA DSBs. Hence, the Baltic population, which is considered to be endangered and is under the influence of several environmental stressors, may be more susceptible to genotoxic effects of environmental exposure to Hg than the Arctic population.

State-dependent capital and income breeding: a novel approach to evaluating individual strategies with stable isotopes.

BACKGROUND: Species-specific strategies for financing the costs of reproduction are well understood, forming a continuum ranging from high to low reliance on stored nutrients. Animals relying mostly on stored reserves are termed 'capital breeders', whereas 'income breeders' rely mostly on concurrent intake when financing the costs of reproduction. The role and adaptive value of individual variation in these strategies remain elusive. Life-history theory posits that capital breeding should be favoured when offspring reproductive value peaks, typically occurring early in the season, and that current income should increasingly be used with progressing season. Because resource limitation may hamper flexible resource allocation, a corollary prediction is that only good-condition individuals may show the expected seasonal shift in resource use. To test this prediction, we examined stable isotopes (δ(13)C and δ(15)N) in blood and lipid-free egg yolk of breeding eider females (Somateria mollissima) from the Baltic Sea to assess the role of individual variation in the use of proteins from local diet vs. stored reserves. RESULTS: We show for the first time that individuals from a single population differ in their utilization of stored reserves and concurrent intake to finance the costs of reproduction. Consistent with our prediction, heavy females predominantly used stored reserves for producing egg yolks early in the season, increasingly relying on local feeding with later onset of breeding, whereas light females showed no seasonal change in allocation strategy. CONCLUSIONS: Stable isotope profiling at the individual level is a powerful tool for monitoring relative changes in investment strategies through time, showing promise as an early warning indicator of ecological change in food webs.

Reproductive investment is connected to innate immunity in a long-lived animal.

Life-history theory predicts that organisms optimize their resource allocation strategy to maximize lifetime reproductive success. Individuals can flexibly reallocate resources depending on their life-history stage, and environmental and physiological factors, which lead to variable life-history strategies even within species. Physiological trade-offs between immunity and reproduction are particularly relevant for long-lived species that need to balance current reproduction against future survival and reproduction, but their underlying mechanisms are poorly understood. A major unresolved issue is whether the first-line innate immune function is suppressed by reproductive investment. In this paper, we tested if reproductive investment is associated with the suppression of innate immunity, and how this potential trade-off is resolved depending on physiological state and residual reproductive value. We used long-lived capital-breeding female eiders (Somateria mollissima) as a model. We showed that the innate immune response, measured by plasma bacteria-killing capacity (BKC), was negatively associated with increasing reproductive investment, i.e., with increasing clutch size and advancing incubation stage. Females in a better physiological state, as indexed by low heterophil-to-lymphocyte (H/L) ratios, showed higher BKC during early incubation, but this capacity decreased as incubation progressed, whereas females in poorer state showed low BKC capacity throughout incubation. Although plasma BKC generally declined with increasing H/L ratios, this decrease was most pronounced in young females. Our results demonstrate that reproductive investment can suppress constitutive first-line immune defence in a long-lived bird, but the degree of immunosuppression depends on physiological state and age.

Antioxidant Responses in Relation to Persistent Organic Pollutants and Metals in a Low- and a High-Exposure Population of Seabirds.

Oxidative stress occurs when there is an imbalance between the production of reactive oxygen species (ROS) and antioxidant defense. Exposure to pollutants may increase ROS and affect antioxidant levels, and the resulting oxidative stress may negatively affect both reproduction and survival. We measured concentrations of 18 persistent organic pollutants (POPs) and 9 toxic elements in blood, as well as total antioxidant capacity (TAC), total glutathione (tGSH), and carotenoids in plasma of Baltic and Arctic female common eiders (Somateria mollissima) (N = 54) at the end of their incubation-related fasting. The more polluted Baltic population had higher TAC and tGSH concentrations compared to the Arctic population. Carotenoid levels did not differ between populations. The effect of mixtures of pollutants on the antioxidants was assessed, and the summed molar blood concentrations of 14 POPs were positively related to TAC. There was no significant relationship between the analyzed pollutants and tGSH concentrations. The adaptive improvement of the antioxidant defense system in the Baltic population may be a consequence of increased oxidative stress. However, both increased oxidative stress and energy allocation toward antioxidant defense may have adverse consequences for Baltic eiders at the incubation stage, when energy resources reach an annual minimum due to incubation-related fasting.

Persistent organic pollutant levels and the importance of source proximity in Baltic and Svalbard breeding common eiders.

The distance to sources and the long-range transport potential of persistent organic pollutants (POPs) are important in understanding the impact of anthropogenic pollution on natural seabird populations. The present study documented blood concentrations of POPs in the Baltic Sea (Tvärminne, Finland) population of common eiders (Somateria mollissima) in 2009 and in 2011 and compared the concentrations with the presumably less exposed Arctic population in Svalbard (Kongsfjorden, Norway). The Baltic population had 26, 10, and 5 times greater concentrations of hexachlorocyclohexane, polychlorinated biphenyls, and p,p'-dichlorodiphenyldichloroethylene than the Svalbard population. Unexpectedly, concentrations of chlordanes were higher in Svalbard eiders, whereas concentrations of hexachlorobenzenes (HCBs) did not differ between the 2 populations. Although the similar HCB levels may partly be explained by the high transport potential of HCBs, unknown factors may have been more important than distance to sources and long-range transport potential for the chlordanes. One plausible explanation may be that the fasting-related redistribution of POPs from fat to blood was greater throughout the incubation in Arctic eiders, causing them to have higher blood levels of these POPs at the end of incubation. The blood concentrations of POPs in Baltic eiders were higher than documented in any other eider population and were comparable to levels in seabirds feeding at higher trophic positions in the food chain. Environ Toxicol Chem 2016;35:1526-1533. © 2015 SETAC.

Brain size-related breeding strategies in a seabird.

The optimal compromise between decision speed and accuracy may depend on cognitive ability, associated with the degree of encephalization: larger brain size may select for accurate but slow decision-making, beneficial under challenging conditions but costly under benign ones. How this brain size-dependent selection pressure shapes avian breeding phenology and reproductive performance remains largely unexplored. We predicted that (1) large-brained individuals have a delayed breeding schedule due to thorough nest-site selection and/or prolonged resource acquisition, (2) good condition facilitates early breeding independent of relative brain size, and (3) large brain size accrues benefits mainly to individuals challenged by environmental or intrinsic constraints. To test these predictions, we examined how the relative head volume of female eiders (Somateria mollissima) of variable body condition correlated with their breeding schedule, hatching success and offspring quality. The results were consistent with our predictions. First, large head size was associated with a progressively later onset of breeding with increasing breeding dispersal distance. Second, increasing body condition advanced the timing of breeding, but this effect was significantly weaker in large-brained females. Third, larger head volume was associated with increased hatching success mainly among late breeders and those in poor body condition, and duckling body condition was positively related to maternal head volume, but only in poor-condition mothers. Our study is, to our knowledge, the first to demonstrate the presence of brain size-related differences in reproductive strategies within a single natural population.

Context dependency of baseline glucocorticoids as indicators of individual quality in a capital breeder.

Identifying markers of individual quality is a central goal of life-history theory and conservation biology. The 'corticosterone (CORT)-fitness hypothesis' postulates that low fitness signals impaired ability to cope with the environment, resulting in elevated baseline CORT levels. CORT can, however, be negatively, positively or neutrally related to fitness, depending on the context. In order to clarify this controversial issue, we elucidate the utility of using baseline CORT as a correlate of individual fitness in incubating female eiders across variable environments. An increase in serum CORT with decreasing body condition was evident in older, more experienced breeders, while increased clutch mass was associated with elevated serum CORT in females breeding late in the season. For faecal CORT, the expected negative association with body condition was observed only in early breeders. We found a strong increase in faecal CORT with increasing baseline body temperature, indicating the utility of body temperature as a complementary stress indicator. Females in good body condition had a lower baseline body temperature, but this effect was only observed on open islands, a harsher breeding habitat less buffered against weather variability. Females with higher reproductive investment also maintained a lower baseline body temperature. Nest success strongly decreased with increasing serum and faecal CORT concentrations, and individual stress hormone and body temperature profiles were repeatable over years. Although our data support the tenet that baseline CORT is negatively related to fitness, the complex context-dependent effects call for cautious interpretation of relationships between stress physiology and phenotypic quality.

Relative importance of social status and physiological need in determining leadership in a social forager.

Group decisions on the timing of mutually exclusive activities pose a dilemma: monopolized decision-making by a single leader compromises the optimal timing of activities by the others, while independent decision-making by all group members undermines group coherence. Theory suggests that initiation of foraging should be determined by physiological demand in social foragers, thereby resolving the dilemma of group coordination. However, empirical support is scant, perhaps because intrinsic qualities predisposing individuals to leadership (social status, experience or personality), or their interactions with satiation level, have seldom been simultaneously considered. Here, we examine which females initiated foraging in eider (Somateria mollissima) brood-rearing coalitions, characterized by female dominance hierarchies and potentially large individual differences in energy requirements due to strenuous breeding effort. Several physiological and social factors, except for female breeding experience and boldness towards predators, explained foraging initiation. Initiators spent a larger proportion of time submerged during foraging bouts, had poorer body condition and smaller structural size, but they were also aggressive and occupied central positions. Initiation probability also declined with female group size as expected given random assignment of initiators. However, the relative importance of physiological predictors of leadership propensity (active foraging time, body condition, structural size) exceeded those of social predictors (aggressiveness, spatial position) by an order of magnitude. These results confirm recent theoretical work suggesting that 'leading according to need' is an evolutionary viable strategy regardless of group heterogeneity or underlying dominance structure.

Rapid climate driven shifts in wintering distributions of three common waterbird species.

Climate change is predicted to cause changes in species distributions and several studies report margin range shifts in some species. However, the reported changes rarely concern a species' entire distribution and are not always linked to climate change. Here, we demonstrate strong north-eastwards shifts in the centres of gravity of the entire wintering range of three common waterbird species along the North-West Europe flyway during the past three decades. These shifts correlate with an increase of 3.8 °C in early winter temperature in the north-eastern part of the wintering areas, where bird abundance increased exponentially, corresponding with decreases in abundance at the south-western margin of the wintering ranges. This confirms the need to re-evaluate conservation site safeguard networks and associated biodiversity monitoring along the flyway, as new important wintering areas are established further north and east, and highlights the general urgency of conservation planning in a changing world. Range shifts in wintering waterbirds may also affect hunting pressure, which may alter bag sizes and lead to population-level consequences.

Brood size matching: a novel perspective on predator dilution.

A primary benefit of grouping is diluting the individual risk of attack by predators. However, the fact that groups are formed not always by solitary adults but also by subgroups (e.g., families) has been overlooked. The subgroup-specific benefit of predator dilution depends on its relative contribution to total group size. Therefore, the willingness of a subgroup to merge with others should increase the less it contributes to total group size, but the conflicting preferences of partners may result in the preferential merger of similar-sized subgroups. Here, we evaluate how the proportional contribution of subgroups to diluting risk affects group formation. We generate predictions using a bidding game over parental care and test them using data on common eiders (Somateria mollissima), in which females with variable-sized broods may form brood-rearing coalitions. The predictions (1) that size-matched subgroups should have a higher propensity to merge, (2) that predation should increase group formation propensity, and (3) that increased bargaining power, as proxied by female body condition, should increase the time needed to establish partnerships were all supported. Partners do negotiate over their relative contributions to predator dilution, accepting or rejecting partnerships on the basis of this criterion. Our results show that consideration of the size of subgroups before merger is critical in understanding the process of group formation under the threat of predation.