Brood Parasitism and Nest Takeover in Common Eiders

Conspecific brood parasitism (CBP) is an alternative breeding tactic that occurs in many brood‐tending animals and can have important fitness effects for both host and parasite. We use protein fingerprinting of egg albumen to distinguish the eggs from different females and to estimate the frequency, pattern and tactics of CBP and other forms of mixed maternity in a Hudson Bay population of common eiders (Somateria mollissima sedentaria). Mixed clutches, containing eggs from more than one female, occurred in 31% of the 86 nests studied that progressed to clutch completion. Other females than the host laid 8% of the eggs. In 11 (41%) of the mixed clutches another female laid before the host started laying, corroborating the hypothesis that takeover of nests started by other females accounts for many of the mixed clutches in this population. Our results also indicate that traditional non‐molecular methods of identifying foreign eggs may considerably underestimate the frequency of mixed clutches.     

Blue–Green Eggs in Pied Flycatchers: An Experimental Demonstration that a Supernormal Stimulus Elicits Improved Nestling Condition

It has been proposed that blue–green egg colouration in certain avian species has evolved as a signal of female and egg quality to males, affecting their investment in the brood according to the differential allocation scenario. A recent experiment has successfully manipulated male investment by placing dummy eggs simulating the extremes in the blue–green natural reflectance. Here we have substituted one egg in certain pied flycatcher clutches with a single deep‐blue (blue) or light‐blue (pale) dummy egg and used other clutches as controls. Blue dummy eggs reflected more in the blue–green part of the spectrum than natural and pale dummy eggs, which were similar in reflectance. Nestlings in nests with one blue dummy egg during laying and incubation attained a significantly higher mass and condition than those in nests with one pale dummy egg or in control broods, when controlling for phenology, brood size and blue–green chroma of the clutch. Only a shift in parental investment can have induced this highly significant effect. Preference for deeply coloured clutches associated with high‐quality females and their maternal effects may explain stimulation by a single supernormal stimulus emanating from multiple objects like clutches.     

Within-nest spawning-site preferences of female bluefin damselfish: the effect of early-stage eggs

In some fish species with exclusive paternal care females prefer to deposit their eggs in nests containing early-stage eggs and avoid those containing only late-stage eggs. Such female preference may also be shown when it comes to deciding where to lay within a given nest; that is, females may prefer to lay contiguously to younger rather than to older eggs. To understand if females benefit from such preferences by obtaining higher offspring survival, we conducted field studies with the bluefin damselfish, Abudefduf luridus. We investigated (1) female between-nest and within-nest spawning-site preferences relative to brood characters and (2) egg-hatching success related to male filial cannibalism. We found evidence that males entirely ate some of their clutches, especially the eggs received near the end of the breeding season. These were also more susceptible to being eaten when laid in nests containing only eggs in a late stage of development. Theoretically, these patterns of clutch loss should result in adaptive female preference for nests containing early-stage eggs, but we found no evidence for such a preference in female A. luridus. However, once females had chosen a nest they strongly preferred to lay contiguously to early-stage eggs than to late-stage eggs. Moreover, females benefited from this decision by obtaining higher survival rates for their clutches. We conclude that within-nest spawning-site preference of the female for early-stage eggs may have evolved as an adaptive response to a dilution effect on the risk of egg loss.     

Multiyear multiple paternity and mate fidelity in the American alligator, Alligator mississippiensis

We examined multiple paternity during eight breeding events within a 10-year period (1995–2005) for a total of 114 wild American alligator nests in Rockefeller Wildlife Refuge in south-west Louisiana. Our goals included examining (i) within population variation in multiple paternity among years, (ii) variation in multiple paternity in individual females and (iii) the potential for mate fidelity. To accomplish this, in the current study, eggs were sampled from 92 nests over 6 years and analysed along with 22 nests from a previous 2-year study. Genotypes at five microsatellite loci were generated for 1802 alligator hatchlings. Multiple paternity was found in 51% of clutches and paternal contributions to these clutches were highly skewed. Rates of multiple paternity varied widely among years and were consistently higher in the current study than previously reported for the same population. Larger females have larger clutches, but are not more likely to have multiply sired nests. However, small females are unlikely to have clutches with more than two sires. For 10 females, nests from multiple years were examined. Seven (70%) of these females exhibited long-term mate fidelity, with one female mating with the same male in 1997, 2002 and 2005. Five females exhibiting partial mate fidelity (71%) had at least one multiple paternity nest and thus mated with the same male, but not exclusively. These patterns of mate fidelity suggest a potential role for mate choice in alligators.     

Adaptations of Polygynous Breeding in the Bobolink, Dolichonyx oryzivorus

In some species of polygynous songbirds, males assist in feedingnestlings of their first (primary) female but ignore offspringof subsequent (secondary) mates. A number of adaptations importantin minimizing the disadvantages of a secondary status in femaleBobolinks (Dolichonyx oryzivorus) have been discovered. In Bobolinks, clutches of secondary females are significantlysmaller than those of primary females, thereby adjusting broodsize to correspond better with food delivery capabilities ofsecondary hens. Commencement of incubation with deposition ofthe penultimate egg allows additional brood size adjustment,for the resulting hatching asynchronism promotes efficient cullingof the late-hatching chick in times of food shortage, withoutendangering the entire brood. Secondary females differ fromprimary pairs in food exploitation techniques by capitalizingon insects found close to the nest and by showing less discriminationin food selection. Such behavior permits compensation in therate of food delivery trips to secondary nestlings so as toequal the tempo maintained by both members of the primary pair,yet results in quantitative disparities in food brought to primaryand secondary nestlings. Flexibility in male feeding responses,resulting in partial male assistance at secondary nests in situationswhere secondary brood size is exceptionally large, is also importantin helping to maximize the reproductive performance of birdsengaging in this mating strategy.     

Parental care of Tilapia mariae in the field and in aquaria

Synopsis Parental care of Tilapia mariae was observed in nature (Ethiop River, Nigeria) and in aquaria with or without intruders present. In the field, 25–30% of nests are guarded by one parent, normally the female. It is assumed that most missing males have deserted. Males who participate in brood care exhibit both close brood guarding and brood defence at a lower level than females, and hence seem to invest less than females. Broods were guarded under three distinct types: (1) female at the brood, male in surroundings, (2) parents take turns, or (3) parents stay together at the brood. Each pair used predominantly one type until the young swam freely, thereafter type 3. Females defended most in type 3, but male attack rate did not differ among the types. Type 3 seems related to increased risk of brood predation and type 2 to the female's foraging needs, being more common when she is small and the mates do not differ much in size. The unequal guarding times of type 1 indicate rather a low parental investment by the male (and thus risk of desertion) than specialization in roles on equal investment basis. Parental behaviour exhibited in aquaria differed in many ways from that in nature. The role types were indistinct and there were more signs of motivational conflict between the mates. Isolated pairs avoided joint guarding in the embryo period and while switching, female turns were much longer than male turns, unlike in nature. When intruders were added, males attacked them more than did females.     

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

Low relatedness and frequent queen turnover in the stenogastrine wasp <Emphasis Type="Italic">Eustenogaster fraterna</Emphasis> favor the life insurance over the haplodiploid hypothesis for the origin of eusociality

Summary The origin of sociality is best studied in taxa with rudimentary social development, like the stenogastrine wasp, Eustenogaster fraterna. Our study of colony structure and relatedness in this species found very small colonies aver-aging only 2.7 adult females, including a single mated, re-productive female. Microsatellite genotyping showed that adult females were related to each other but not generally as full sisters (r = 0.427 ± 0.131, 95% confidence interval). Microsatellite genotyping showed that sociality is not favored by the haplodiploid hypothesis, because relatedness of unmated females to female brood is low (r = 0.210 ± 0.171, 95% confidence interval), far lower than that among sisters in either haplodiploid or diploid species. Relatedness of unmated females to female brood is significantly lower than that of mated females to female brood (r = 0.374 ± 0.266). Mated females are also significantly more related to the male brood (r = 0.871 ± 0.168) than are unmated females (r = 0.588 ± 0.339), suggesting that unmated helpers do not generally produce sons. These results argue against an important role for exceptionally high relatedness in the origin of eusociality. One quarter of the brood could not have been the progeny of any collected female, suggesting high rates of queen turnover. In all, 7/17 nests had some brood that could not be assigned to existing adult females. These high adult mortality rates result in direct advantages to helpers in the form of colony inheritance, and indirect advantages via life insurance benefits.Received 23 October 2002; revised 13 March 2003; accepted 23 April 2003.     

Paternal genotype reconstruction reveals multiple paternity and sex ratios in a breeding population of leatherback turtles (<Emphasis Type="Italic">Dermochelys</Emphasis><Emphasis Type="Italic">coriacea</Emphasis>)

When animals are difficult to observe while breeding, insights into the mating system may be gained by using molecular techniques. Patterns of extra-pair copulation, multiple paternity and parental genotype analysis may elucidate population characteristics that help improve knowledge of life history while informing management decisions. During the course of a long-term study of leatherback turtles, we assessed the level of multiple paternity in successive clutches for 12 known females nesting at Sandy Point National Wildlife Refuge (St. Croix, U.S. Virgin Islands). We used seven polymorphic microsatellite markers to genotype the females and 1,019 hatchlings representing 38 nests (3–4 clutches from each female). Using deductive genotype reconstruction and GERUD1.0, we identified the 12 mothers and 17 different fathers that were responsible for 38 nests. We found that seven females (58.3%) showed no evidence of multiple paternity in their clutches, while five females (41.7%) had mated with two males each. There was evidence of two fathers (polyandry) in successive clutches for these five females. Multiple fathers didn’t contribute to clutches equally. For clutches laid by an individual female, the primary father was responsible for 53.7 to 85.9% of the hatchlings. We demonstrate the feasibility of using male genotype reconstruction to characterize the male component of this breeding population and to assess operational sex ratios for breeding sea turtles.     

Coevolution of female fidelity and male help in populations with alternative reproductive tactics

In socially monogamous species, pair-bonded males often continue to provide care to all offspring in their nests despite some degree of paternity loss due to female extra-pair copulation. Previous theoretical models suggested that females can use their within-pair offspring as ‘hostages'' to blackmail their social mates, so that they continue to provide care to the brood at low levels of cuckoldry. These models, however, rely on the assumption of sufficiently accurate male detection of cuckoldry and the reduction of parental effort in case of suspicion. Therefore, they cannot explain the abundant cases where cuckolded males continue to provide extensive care to the brood. Here we use an analytical population genetics model and an individual-based simulation model to explore the coevolution of female fidelity and male help in populations with two genetically determined alternative reproductive tactics (ARTs): sneakers that achieve paternity solely via extra-pair copulations and bourgeois that form a mating pair and spend some efforts in brood care. We show that when the efficiency of mate guarding is intermediate, the bourgeois males can evolve to ‘specialize'' in providing care by spending more than 90% of time in helping their females while guarding them as much as possible, despite frequent cuckoldry by the sneakers. We also show that when sneakers have tactic-specific adaptations and thus are more competitive than the bourgeois in gaining extra-pair fertilizations, the frequency of sneakers and the degrees of female fidelity and male help can fluctuate in evolutionary cycles. Our theoretical predictions highlight the need for further empirical tests in species with ARTs.