Female Sexual Interference in the Smooth Newt,Triturus vulgaris vulgaris

Competition between unmated females for males was observed in the smooth newt, T. vulgaris, during the early part of the breeding season. Females interfered at the spermatophore-transfer stage of courtship, preventing a rival female from picking up the sperm mass. Interference often resulted in the rival female abandoning the courtship encounter. In many of the encounters (70%), the interfering female was inseminated. However, interference between females was not observed after the commencement of ovulation. By contrast, the incidence of sexual interference between males was low during the preovulatory period and increased significantly post-ovulation. Sexual interference in newts and salamanders has previously only been observed between males. During the greater part of the breeding season, when females are ovipositing, eggs are the limiting resource and males compete for access to receptive females. However, this study suggests that early in the breeding season, prior to commencement of ovulation, the limiting resource may temporarily be sperm, due to the physiological constraint of spermatophore production. This may lead to female competition for mates.     

The Cloaca and Spermatheca of the Female Smooth Newt, Triturus vulgaris L. (Amphibia: Urodela: Salamandridae)

The anatomy and histology of the cloaca and spermatheca of 13 female smooth newts ( Triturus vulgaris ) were studied by light microscopy. The cloaca consists of a short, anterior tube (which receives the oviducts), which opens into a larger, more posterior chamber. The spermatheca, which is the only gland in the cloaca, consists of a mass of exocrine, acinar tubules which empty individually and dorsolaterally into the posterior part of the cloacal tube and all but the most posterior part of the cloacal chamber. Stored sperm are most abundant in the spermathecal tubules during the peak of the breeding season (in May and June); during their period of storage, the sperm do not appear to make intimate contact with the epithelial cells lining the spermathecal tubules. Longterm storage of viable sperm from one breeding season to the next is probably absent in the smooth newt. Sperm storage by and multiple insemination of females both occur in this species and are necessary preconditions for competition between the sperm of different males for the fertilization of eggs. However, sperm competition has not been demonstrated in the smooth newt.     

Forced Copulations and Intra-specific Parasitism: Two Costs of Social Living in the White-fronted Bee-eater

White-fronted bee-eaters are colonially breeding birds that exhibit highly developed helping-at-the-nest. Through long-term studies of an individually-marked population, we have documented two costs of social living: 1) harassment of mated females by extra-pair males, and 2) intra-specific parasitism by females who lay eggs in the nests of others. Breeding females are sexually chased and, occasionally, forceably mated by males other than their mates. Focal-sampling of females throughout their period of receptivity revealed that the average female is involved in 5 to 8 sexual chases and is forceably copulated 0.15 to 0.23 times per breeding season. This risk to females would be much greater were it not for the behavior of male mates who remain close to, and actively defend, their partners. Such mate-guarding is highly effective — females entering and leaving the colony in consort with their mates are sexually harassed only 1/10 as often as females travelling alone. Although sexual harassment of females is common at bee-eater colonies, the risk of paternity uncertainty arising from forced copulations is thought to be low. The reason is that females copulate repeatedly with their male mates on all days immediately prior to as well as during egg laying. This point has been overlooked in previous reports and has led to an exaggeration of the paternity risks associated with forced sexual chases. We conclude that sexual chasing of extra-pair females is a low yield reproductive tactic employed primarily by monogamously mated males whose presence at the colony is required to allofeed and mateguard their own egg-laying females. Female white-fronted bee-eaters lay eggs in nests other than their own. This intraspecific parasitism constitutes a greater threat to certainty of parentage than does forced copulation. Over four years of study, 16% of nests were parasitized and 7 % of all eggs were laid by a female other than the breeder (Table 2). Parasitizing females come primarily from two sources: (1) members of mated pairs whose own breeding attempt is disrupted at the time of egg laying, and (2) single females who opportunistically add an egg at the nest of their parents (or parent plus step-parent). In each case of kin-parasitism, the “parasitic” female remained socially integrated with the host group and helped in the rearing of the young. In contrast, 9 of 10 females that parasitized the nests of non-relatives had no other interactions with the hosts (Table 3). Parasitizing females exhibited two specialized behaviors that enhanced their reproductive effectiveness: (1) they spent many hours observing, investigating, and testing the defenses of potential host nests, and (2) they preferentially laid in hosts' nests at the appropriate chronological stage of development. Breeding females also exhibited counterbehaviors against being parasitized. These included: (1) remaining sequestered in their nest chambers for 64%-65% of the daylight hours and 94 % of the pre-roost hours during their days of egg laying, (2) aggressively defending their nest entrances against all investigating (potentially parasitic) females, and (3) actively removing any eggs laid in their nests prior to the initiation of their own clutch. These tactics and countertactics suggest a long evolutionary history of parasitic opportunities and risks among white-fronted bee-eaters.     

Dominance and Mating in a Communal Polygynandrous Bird: Cooperation or Indifference towards Mating Competitors?

The effect of dominance on the mating behaviour of both females and males was investigated for the communally breeding pukeko (Porphyrio porphyrio melanotus). Pukeko are polygynandrous with most groups consisting of three to 7 breeding males and one or two breeding females that lay eggs in a single nest. Linear dominance hierarchies are well established within breeding groups. Alpha females were involved in more copulations than beta females, but this difference did not appear to be a result of dominance status. There was also no evidence of egg tossing or egg destruction by breeding females. There was no positive correspondence between dominance rank of males and their frequency of sexual activity or copulations during the prelaying or laying periods. Dominants did interrupt matings by subordinates, but both qualitative and quantitative analyses of behaviour found no evidence of mate guarding. Furthermore dominants were no more likely to interrupt a copulation involving a subordinate male than they were to passively observe it or not react at all. This led us to suggest that males tended to behave indifferently toward mating competitors. As a result, multiple paternity is predicted to be high in pukeko groups. We also question the standard interpretation that males are behaving cooperatively in communal groups rather than coactively.     

Producing extra eggs does not deplete macronutrient reserves in European Starlings Sturnus vulgaris

Recent studies have suggested that the size of avian clutches may be determined in part by the costs of egg production, which are thought to involve the depletion of endogenous nutrient reserves. I tested the hypothesis that producing extra eggs would deplete protein and lipid reserves in a passerine species, the European Starling Sturnus vulgaris . The first three eggs of clutches were removed on the days they were laid to induce females to lay extra eggs. Females responded to this manipulation by laying, on average, 1.3 eggs more than controls. However, based on the distribution of clutch sizes, it appears that some females responded to the removal of three eggs by producing more than one extra egg, whereas others responded by laying no extra eggs or by deserting. In unmanipulated females, the lean dry mass of the flight muscles and total body lipid were found to decline throughout breeding, an observation which is often interpreted as evidence that egg production depletes endogenous reserves. However, females experimentally induced to produce an extra egg did not have reduced stores of protein or lipid at clutch completion. No effect was observed in starlings even though this study had substantial (80%) power to detect a change in protein reserves of the same magnitude as that observed in a previous study of gulls. The decrease in flight muscle mass that has frequently been observed during breeding in passerines may not be due simply to mobilization of endogenous protein reserves.     

Paternity in the Australian brush-turkey, Alectura lathami, a megapode bird with uniparental male care

Male Australian brush-turkeys, Alectura lathami, provide allparental care by building and tending large incubation mounds.Females visit and lay eggs in the mounds of several males sequentially,but they provide no parental care after laying. Because malesand females meet only briefly at mounds to copulate and lay,males have no obvious means of ensuring paternity. I used DNAfingerprinting techniques to determine paternity for 65 brush-turkeychicks. Eighteen chicks (27.7%) did not match the mound-tendingmale. Some of these paternity exclusions were evidently causedby females switching rapidly from one mound to another, butthe majority (23.1% of eggs) appeared to result from femalescopulating with males other than the one in whose mound theywere currently laying. However, the frequency of these copulations(43%) was much lower than the estimated frequency with whichthey fertilized eggs, perhaps because their timing during theovulatory cycle differed relative to most other copulations.The percentage of eggs excluded in paternity analyses rangedfrom 20.0% to 43.8% for individual males but did not appearto affect male parental care. Several factors favor male parentalcare regardless of paternity. Males can accommodate eggs fromseveral females in one mound, which increases the opportunitiesfor additional matings without increasing the cost of parentalcare. In addition, paternity appears to be unpredictable andhard to assess, and a facultative reduction in care would bedifficult without abandoning a mound entirely.     

Photoperiodic control of reproduction and molt in the kestrel, Falco tinnunculus

An adaptive decline in average clutch size with progressive date of laying is characteristic of most bird species with a single clutch of variable size per year. The effect of photoperiod on timing of laying, clutch size, and subsequent molt was investigated in kestrel pairs breeding and raising their young in captivity. In natural daylight (nLD), clutch size, under ad libitum feeding, showed the same decline with date as in nature. Birds breeding later also started molt later and molted faster (more feathers simultaneously), so that all birds completed molt more or less at the same time. Constant long days (LD 17.5:6.5 and LD 13:11) from December 1 onward advanced both reproduction and molt. The LD 17.5:6.5 group developed the reproductive system faster, had a shorter courtship period, and laid eggs earlier than the LD 13:11 group. In both photoperiods there was a decrease in clutch size with progressive laying date, similar to that in nLD. Molt started in both groups at about the end of the laying period and slowed down in the longer photoperiod, especially in males. Plasma luteinizing hormone (LH) changes in the two photoperiods were different for males and females. Males showed the expected slower LH response in the shorter photoperiod, but the initial LH response by females was the same in both photoperiods. Data on LH, reproductive behavior, and body mass suggest that females have a wider annual reproductive window than males. Data on time of laying and number of eggs suggest that clutch size in the kestrel is determined by laying date itself, following an endogenous rhythm that is phase-locked to the reproductive cycle.     

Intraspecific nest parasitism in Starlings Sturnus vulgaris: effects of disturbance on laying females

Female Starlings Sturnus vulgaris that laid distinctive eggs at Worplesdon, Surrey, in 1988 and 1989 were made to desert their partially completed first clutches by removing their eggs part way through the laying sequence. Most of these Starlings continued to lay, but in other Starlings' nests, and most of them subsequently laid a replacement clutch that they reared themselves. Using estimates of the success of nests in the colony over 14 breeding seasons, it is concluded that intraspecific nest parasitism offers a means of maximizing annual chick production in the event of disturbance causing the loss of first-clutch eggs.     

Some aspects of adaptation in a polymorphism for breeding colours in the Threespine stickleback (Gasterosteus aculeatus))

Polymorphism in the nuptial colouration of male Threespine sticklebacks ( Garterosteus aculeatus L.) was studied. In Lake Wapato in Eastern Washington, approximately 14% of the breeding males develop typical red nuptial colouration while others develop various colours which were grouped into a non-red category. The hypothesis was tested that red males are adaptive in Lake Wapato as a result of sexual selection and differential egg predation.
In a series of mate preference tests, females demonstrated a significant preference for red males over non-red males. In a second experiment, using artiticially coloured non-red males, it was shown that the females are responding to the red colouration alone. This evidence suggests that one of the selective pressures maintaining the red morph is sexual selection.
In a stomach analysis of male and female sticklebacks collected from a breeding colony it was found that 51 % of the fish examined had stickleback eggs in their stomachs. Both male and female sticklebacks eat a large number of eggs; however, males eat significantly more eggs than females. Eggs were found to be most susceptible to predation during their first four days of development. Given that red ventral colouration has threat value, red males may also be adaptive because of a reduction in egg predation within their territories in contrast to non-red males.
One of the selective forces against red males is probably differential predation by trout ( Salmo gairdneri ). This trout predation has been shown to be extensive in Lake Wapato. The opposing selective forces discussed in this paper cannot maintain the polymorphism; they are only components of fitness of the colour morphs.     

Breeding biology of a non-diadromous galaxiid, Galaxias vulgaris Stokell, in a New Zealand river

In the Glentui River, Canterbury, New Zealand, Galaxias vulgaris Stokell spawned in late winter and early spring (from July to September), the onset of spawning being temperature-dependent. No females matured in their first year, but 95 and 68 % of age 0+ males were mature in consecutive years. With the exception of one female, all fish aged 1 + and older spawned. In mature fish, overies reached nearly 23%, and testes 11%, of somatic weight. Maturity in males depended on size, whereas in females, age appeared to be the major factor affecting maturity. Recruitment eggs for next season's spawning were distinct from primary oocytes soon after the ripe eggs were shed. Egg numbers varied between 284 and 1911 per fish, and weight was the best indicator of fecundity. Sex ratios of whole samples were not significantly different from 1 : 1, but ratios of potential spawners revealed an excess of males, due mainly to the presence of age 0 + fish. Age0+ males probably play an important role in adverse conditions and act as a form of insurance by significantly increasing the proportion of ripe males, thereby ensuring that all eggs are fertilized. The nest, egg mass and behaviour during the spawning period are described. More than one female may lay eggs in each nest and it is probable that each egg mass receives sperm from more than one male. It is suggested that the presence of fewer and larger eggs in G. vulgaris , compared with the diadromous galaxiids, is the result of a more precise fertilization process.     

The evolution of sexual size dimorphism in the house finch. V. Maternal effects

The phenotype of a mother and the environment that she provides might differentially affect the phenotypes of her sons and daughters, leading to change in sexual size dimorphism. Whereas these maternal effects should evolve to accommodate the adaptations of both the maternal and offspring generations, the mechanisms by which this is accomplished are rarely known. In birds, females adjust the onset of incubation (coincident with the first egg or after all eggs are laid) in response to the environment during breeding, and thus, indirectly, determine the duration of offspring growth. In the two house finch (Carpodacus mexicanus) populations that breed at the extremes of the species' distribution (Montana and Alabama), females experience highly distinct climatic conditions during nesting. We show that in close association with these conditions, females adjusted jointly the onset of incubation and the sequence in which they produced male and female eggs and consequently modified the growth of sons and daughters. The onset of incubation in newly breeding females closely tracked ambient temperature in a pattern consistent with the maintenance of egg viability. Because of the very different climates in Montana and Alabama, females in these populations showed the opposite patterns of seasonal change in incubation onset and the opposite sex bias in egg-laying order. In females with breeding experience, incubation onset and sex bias in laying order were closely linked regardless of the climatic variation. In nests in which incubation began with the onset of egg laying, the first-laid eggs were mostly females in Montana, but mostly males in Alabama. Because in both populations, male, but not female, embryos grew faster when exposed to longer incubation, the sex-bias produced highly divergent sizes of male and female juveniles between the populations. Overall, the compensatory interaction between the onset of incubation and the sex-biased laying order achieved a compromise between maternal and offspring adaptations and contributed to rapid morphological divergence in sexual dimorphism between populations of the house finch breeding at the climatic extremes of the species range.     

Immunoglobulin plasma concentration in relation to egg laying and mate ornamentation of female barn swallows (Hirundo rustica)

In vertebrates, offspring have a relatively inefficient immune system soon after birth. Female birds transmit immunoglobulins to the egg, which can confer protection against parasites to their offspring after hatching, but allocation of immune factors can depend on the reproductive value of the offspring as affected, for example, by the quality of their father. We analyse the variation in immunoglobulin levels of female barn swallows (Hirundo rustica) during the breeding cycle in relation to the expression of a secondary sexual character of their mates. Circulating immunoglobulins peaked on approximately the day before that of laying of their first egg, but postlaying concentration was similar to the concentration well before laying. Immunoglobulin levels per unit volume of plasma were lower for females breeding late compared with those breeding early. Haematocrit of females reached an absolute minimum on the day of laying of the third egg. In males, concentration of immunoglobulins relative to other plasma proteins did not change in relation to the breeding stage. Smaller relative concentrations of immunoglobulins and haematocrit were observed in males breeding late in the season. Immunoglobulin concentration of females was positively correlated with the level of ornamentation of their mates. These results suggest that females alter their immune profile to transmit humoral factors providing immune defence against pathogens to their offspring after hatching. This enhancement of immune protection is larger when the offsprings are of relatively large reproductive value as when sired by high quality males. Alternatively, females with relatively large immunoglobulin concentration preferentially mate with the most ornamented males.     

The costs of reproduction to males and females of a paternal mouthbrooding cardinalfish Apogon notatus

In the cardinalfish Apogon notatus , starving males mouthbrood the eggs, and females are more active than males in courtship and in attacks against conspecific intruders upon their breeding territory, but sexual ornaments are developed in males. Condition factor, hepatosomatic index and fat body-somatic index declined in both sexes during the breeding season. The deterioration of somatic condition was more severe in females, suggesting that the overall energetic costs were larger for females which produced more than twice as many clutches as males brooded in a season. Selective mortality of females could be due primarily to the depletion of their energy reserves. The increased mortality in females is a possible mechanism for the male-biased operational sex ratio which may provide the potential for sexual selection on males.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

Effect of male-to-male aggressiveness and feed-restriction during rearing on sexual behaviour and aggressiveness towards females by male domestic fowl

This experiment was one part of a larger study investigating problems of aggression towards females by male broiler breeder fowl. To investigate causal mechanisms, we were interested in determining (1) if feed-restriction during rearing affects behaviour towards females at sexual maturity and (2) if aggressiveness towards females is correlated with general levels of aggressiveness. We compared broiler breeder males with commercial laying strain males, which were either fed ad libitum or were feed-restricted during the rearing phase, and with game strain males, bred for fighting. Differences in behaviour were determined by observing males during interactions with small groups of females.Laying strain males did not behave aggressively towards females, whether feed-restricted or fed ad libitum during rearing. Despite genetic selection for fighting ability, game strain males also were not aggressive towards females. Conversely, broiler breeder males displayed significantly higher levels of aggression towards females than did feed-restricted laying strain males (P<0.02). Broiler breeder males were rough with females during mating, whereas laying strain and game strain males were not. Females struggled more frequently during mating attempts by broiler breeder males (P<0.002) and interfered frequently when these males attempted to mate with other females.From our results, we conclude that (1) feed-restriction during rearing has little effect on the sexual and aggressive behaviour of laying strain males at maturity and (2) selection for aggressiveness has not resulted in males which are more aggressive to females. Aggression towards females appears to be a unique problem occurring in broiler breeder male strains and not a function of feed-restriction.     

Egg burial in penduline tits, Remiz pendulinus: its role in mate desertion and female polyandry

One important component in the mating strategy of an already-matedindividual is the decision to remain with the partner and carefor the offspring or to desert. Almost all research on nestdesertion has focused on the costs and benefits of continuedparental care versus desertion of both parents. However, ifit pays both parents to desert, the timing of desertion is mostimportant. In birds, where all eggs are fertilized well beforethe last egg is laid, males should be the first to desert. Evenif females try to hide their fertile period, it is likely thatthe appearance of eggs acts as a cue that males can use to calculatethe timing of their desertion. Here we examine egg burial behaviorin penduline tits and its possible effects on parental behaviorand desertion. Penduline tits perform uniparental care fromthe earliest point of breeding, and both sexes try to becomepolygamous. We found that 36% of investigated males were polygynousand deserted as soon as the first egg appeared in their nest,and 12.5% of females became polyandrous. About 27% of the nestswere deserted by both sexes, which means high costs for femalesin terms of wasted energy in eggs and for males in terms ofwasted energy and time in building elaborate nests. Femalescover the eggs, and several facts indicate that egg coveringis a deceptive behavior of females: (1) females cover the eggsin the morning before leaving the nest for the first time, (2)females are more aggressive toward their mates during the layingperiod than before laying, (3) females try to prevent malesfrom entering the nest when eggs have been experimentally uncovered,and (4) females uncover the eggs as soon as males are experimentallyremoved. Finally, we found that a female can only desert thenest before the male deserts when she covers the eggs. We concludethat the higher the proportion of eggs a female can hide, thegreater her chance of becoming polyandrous     

Alterations in serum steroid concentrations in the clearnose skate, Raja eglanteria: correlations with season and reproductive status.

Serum steroid hormones in the peripheral circulation of the clearnose skate, Raja eglanteria, were measured at the time of capture and at various times throughout the year while the animals were maintained as a captive breeding population. These analyses demonstrate interesting correlations between changes in hormone concentrations and annual reproductive events. Animals were sampled once (78 females, 20 males) or multiple times (15 females). For both groups of females, 17beta-estradiol was detected throughout the year with significant elevations occurring during October and November when ovarian follicles begin to mature (as determined through necropsy examinations), and January and February when maximum mating activity is observed and egg laying begins. Testosterone and dihydrotestosterone concentrations were significantly elevated in females only during January and February. Testosterone elevations were synchronous with longer-term elevations in 17beta-estradiol in females sampled either once or repetitively. Testosterone concentrations in males were significantly elevated during times of maximum breeding activity compared to periods of sexual inactivity. Data from females sampled during five stages of the egg laying process, as defined by the position of palpable egg capsules within the reproductive tract, revealed that 17beta-estradiol was highest when egg capsules were forming in the nidamental gland (stage 2) or uterus (stage 3); testosterone and dihydrotestosterone were maximal when eggs were in the uterus (stage 3) or cloaca (stage 4); and progesterone was significantly elevated immediately after oviposition (stage 5), suggesting a possible role for progesterone in the regulation of sequential laying of egg pairs. J. Exp. Zool. 284:575-585, 1999.     

Egg variability and hatching success in the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica

Eggs of the Cape petrel Daption capense at Nelson Island, South Shetland Islands, Antarctica, exhibited high variability in volume between females (up to 48%), while for each individual female, both volume and shape of eggs were highly correlated over two seasons. Both female size and body-condition at laying were unrelated to egg-volume in the 'good'season 1991, but larger females produced larger eggs in the 'bad'season 1990, when eggs were smaller on average. In 1991, females in better body-condition, but not larger, started laying earlier. In spite of synchronized laying (75% of eggs laid within five days), egg-volume decreased with laying date. Egg-volume differed significantly between the two years, which could not be explained by changes in the breeding population or timing of breeding. The laying of relatively small eggs and low overall breeding success in 1990 most probably reflected changes in food availability. Overall hatchability was 88% and did not differ significantly between disturbed and control colonies in 1991. Larger and more rounded eggs showed better hatching success, but hatchability was related more to egg-volume whereas hatching rate was related more to egg-shape. As the majority of egg losses were attributable to predation, the hatching rate was influenced by parental performance relatively more than by hatchability. Hence the relationship between hatching rate and egg-shape probably reflects improvement in hatching success with female age/experience, whereas the relationship between egg-size and hatchability suggests effect of egg-size independently of parental traits.     

Sexual dimorphism of egg size in the European Blackbird Turdus merula

Recently, a number of studies have shown that female birds are able to control the sex of their progeny at the stage of the gamete. There is also some evidence that females adjust their investment in offspring depending on the sex of the embryo during egg formation. Differential maternal investment to the eggs depending on their sex is usually interpreted as an adaptive strategy, by which females can increase competitive abilities of the smaller sex, or preferentially invest towards the sex with the potentially higher fitness returns. Here, we studied variation in egg size in relation to embryo sex and laying order in the European Blackbird Turdus merula. We found male and female eggs to differ in size, with larger eggs containing male embryos, as well as a significant interaction between embryo sex and laying order. This interaction resulted from the fact that egg size increased with the laying sequence among eggs bearing females but did not change with laying order among eggs bearing males. There was no relationship between offspring sex and the laying sequence within a clutch. We suggest that sexual dimorphism in egg size recorded in the European Blackbird may reflect favouritism of the sex which may give higher fitness returns.     

Factors affecting the cost of polygynous breeding for female Great Reed Warblers Acrocephalus arundinaceus

The male's role in feeding young and his effect on the mortality of eggs and nestlings of the Great Reed Warbler Acrocephalus arundinaceus were studied in Kahokugata, central Japan, during eight breeding seasons. Three potential costs of polygynous breeding for females were examined:(i) a reduction of male parental care per brood, (ii) an attraction of predators due to concentration of nests, and (iii) an increase of unfertilized eggs due to the reduced frequency of copulation per female. Only the first cost was found to occur:polygynous males fed their young, especially of their second broods, less frequently than monogamous ones. The lowered paternal feeding frequency, however, did not increase the mortality by starvation in second-hatched broods. This was because the warm climate emancipated females from brooding and enabled them to compensate for the deficiency of feeding by males. The paper discusses possible roles of the staggering of the time of laying among harem mates and the shift of breeding status of females caused by nesting failure, in reducing potential costs of polygynous breeding for females.