Polygyny in the red-winged blackbird: do females prefer monogamy or polygamy?

Five hypotheses have been proposed to explain polygyny in the red-winged blackbird (Agelaius phoeniceus). We categorized the hypotheses into three groups based on female preference for unmated versus monogamously mated males: (1) the “polygyny threshold” model, “sexy son” hypothesis and the “asynchronous settlement” model, which assume that females prefer unmated males to mated males on breeding situations of homogeneous quality; (2) the “neutral mate choice” hypothesis, which assumes that females have no preference; and (3) the “cooperative female choice” model, which assumes that females prefer monogamously mated males to unmated males. We tested the direction of female preference in two field experiments. In both experiments, newly settling females were given a choice of two adjacent territories, one defended by an unmated male and the other by a monogamously mated male. Male mating status was randomized with respect to the variation in territory quality and male quality. Early in the breeding season, significantly more females settled with the unmated males than with the mated males. Although more females settled with the unmated males than with the mated males late in the breeding season, the difference was no longer significant. Female settlement late in the season appeared to be related to the tenure of the resident females: the new females avoided territories where the resident females were in early stages of their nesting, but settled on territories where the resident females were in late stages. The pattern of female settlement shows that females prefer unmated males to mated males. The preference is consistent with the polygyny threshold model, sexy son hypothesis and the asynchronous settlement model, and inconsistent with the neutral mate choice hypothesis and the cooperative female choice model. For this reason, the latter two hypotheses are unlikely to explain the occurrence of polygyny in our population of red-winged blackbirds. Received: 1 December 1994 / Accepted after revision: 28 October 1995     

Sex allocation in the solitary bee Osmia cornuta: do females behave in agreement with Fisher's theory?

Fisher's theory of sex allocation predicts that, in a panmictic population, parental investment will be equally distributed between male and female progeny. Most studies on parental investment in nesting solitary bees and wasps use offspring or provision weight as estimators of parental investment and do not corroborate Fisher's theory. The measurement of parental investment may be confounded by several factors. First, the use of offspring or provision size does not account for seasonal variation in foraging costs associated with aging of nesting females. Second, provision or offspring size do not reflect parental investment associated with nest construction. In this two-year study we measured parental investment in a solitary bee. We calculated sex allocation using both provision weight and foraging time as parental investment estimators. Investment in pollen-nectar provisions decreased, while investment in mud structures (nest construction) increased, as the nesting period progressed. Overall investment in provisions per nest was ∼25 times higher than investment in mud. Pollen-nectar foraging trips became longer as the season progressed, but mud trip duration did not vary. Due to weather differences between years, more offspring per female were produced in the first year, but progeny sex ratio and mean offspring size of both sexes were similar between years. Mortality did not differ between sexes. As predicted by Fisher's theory, production cost ratios did not differ from 1 in either year, irrespective of the currency used to estimate parental investment (provision weight or foraging time). Our results strongly support Fisher's theory.     

Extra-pair paternity in tree swallows: why do females mate with more than one male?

Recent studies of monogamous tree swallows (Tachycineta bicolor) suggest that females may receive some type of genetic benefit from extra-pair fertilizations. In this study we attempted to determine what type of genetic benefits might be gained by females. We compared numerous morphological and behavioral traits (Table 1) of every male nesting on one grid of nest-boxes (n = 23) to determine what male traits were correlated with male success at gaining extra-pair fertilizations. DNA fingerprinting revealed an increase in the level of extra-pair paternity from the previous year (50% of broods contained extra-pair young in 1990 vs. 87% of broods in 1991), but no significant correlates of paternity. We found six extra-pair fathers at seven nests (20 nests had extra-pair young). The traits of these extra-pair males did not differ from those of the males they cuckolded. We discuss several reasons for this lack of difference, but argue that our results are not inconsistent with females choosing extra-pair males to enhance the genotypic quality of their offspring. Despite a complete search of the nest-box grid for extra-pair fathers, we were able to explain the paternity of just 21% (13/63) of all extra-pair young. This suggests that extra-pair fathers were either residents off our study grid or non-territorial floaters. Tree swallows are quite mobile and spend only part of the day at their nest prior to laying. In addition, we rarely see swallows visiting other grids of nest-boxes. Therefore, we suggest that most extra-pair copulations occur at some unknown location, possibly at a feeding or roosting area where females may be able to choose from many more potential extra-pair fathers than at their nest-site.     

Mating in the afternoon: Time-saving in courtship and remating by females of a polyandrous butterfly Pieris napi L.

Summary Among butterflies where females are polygamous, interactions with males after the first mating entail time costs associated with both courtship and mating. In the green-veined white butterfly, Pieris napi, spermatophore counts on wild-caught females revealed that the average number of matings that females had performed increased gradually from 1.1 during the first week of the flight period to 2.4 during the 4th and final week. By dissecting pairs found in copula in the field, we found that female rematings occurred solely in the afternoon, whereas virgin females mated most frequently earlier in the day. We suggest that non-virgin females minimize the time cost of rematings by allocating them to an hour when the egglaying frequency has decreased to a low level. Virgin females were very attractive to males, judging by their courtship persistence (several minutes), but when mated, females became so unattractive that courting males left them within a few seconds. However, females' attractiveness increased again with time elapsed since the last mating. These observations support the idea that females emit repellent pheromones after mating, which may be male derived. Since newly mated females were not harassed by males or hindered in their egglaying activities, it is conceivable that the time cost of remating may be outweighed by the benefit of the regained ability to curtail male courtship. This advantage to newly remated females may in itself be an explanation for polyandry in P. napi and is alternative or complementary to the idea that females that remate receive a material benefit that can be used to increase their realized fecundity.     

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.     

Internal eutrophication: How it works and what to do about it—a review

In the 1980s and 1990s, it became increasingly clear that changes in external nutrient loads alone could not entirely explain the severe eutrophication of surface waters in the Netherlands. Nowadays, 'internal eutrophication' has become a widely accepted term in Dutch water management practice to describe the eutrophication of an ecosystem without additional external input of nutrients (N, P, K). This review surveys the principal mechanisms involved in this process. It also discusses possible remedies to combat internal eutrophication.