A comparative study on the evolution of reversed size dimorphism in monogamous waders

Sexual dimorphism in size is common in birds. Males are usually larger than females, although in some taxa reversed size dimorphism (RSD) predominates. Whilst direct dimorphism is attributed to sexual selection in males giving greater reproductive access to females, the evolutionary causes of RSD are still unclear. Four different hypotheses could explain the evolution of RSD in monogamous birds: (1) The ‘energy storing’ hypothesis suggests that larger females could accumulate more reserves at wintering or refuelling areas to enable an earlier start to egg laying. (2) According to the ‘incubation ability’ hypothesis, RSD has evolved because large females can incubate more efficiently than small ones. (3) The ‘parental role division’ hypothesis suggests that RSD in monogamous waders has evolved in species with parental role division and uniparental male care of the chicks. It is based on the assumption that small male size facilitates food acquisition in terrestrial habitats where chick rearing takes place and that larger females can accumulate more reserves for egg laying in coastal sites. (3) The ‘display agility’ hypothesis suggests that small males perform better in acrobatic displays presumably involved in mate choice and so RSD may have evolved due to female preference for agile males. I tested these hypotheses in monogamous waders using several comparative methods. Given the current knowledge of the phylogeny of this group, the evolutionary history of waders seems only compatible with the hypothesis that RSD has evolved as an adaptation for increasing display performance in males. In addition, the analysis of wing shape showed that males of species with acrobatic flight displays had wings with higher aspect ratio (wing span/²wing area) than non-acrobatic species, which probably increases flight manoeuvrability during acrobatic displays. In species with acrobatic displays males also had a higher aspect ratio than females although no sexual difference was found in non-acrobatic species. These results suggest that acrobatic flight displays could have produced changes in the morphology of some species and suggest the existence of selection favouring higher manoeuvrability in species with acrobatic flight displays. This supports the validity of the mechanisms proposed by the ‘display agility’ hypothesis to explain the evolution of RSD in waders.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

THE EVOLUTION OF REVERSED SEXUAL DIMORPHISM IN SIZE IN MONOGAMOUS SPECIES OF BIRDS

Reversed sexual dimorphism in size (RSD) occurs in most species of several taxonomic groups of birds. The hypotheses proposed to explain this phenomenon are examined theoretically, using inequalities to state selection in the most rigorous possible terms. The most pertinent empirical evidence is also examined critically. Proponents of hypotheses on the evolution of RSD have failed to consider the genetic constraints on the evolution of dimorphism. Selection for dimorphism can act on only that small portion of the genetic determination of body size that is sex limited. In general, selection for body size is much more likely to lead to a similar change (e.g. larger) in both sexes than to dimorphism. The most popular hypotheses involve selection for size-related differences in foraging ability. It is unlikely that there is variation in size-related foraging differences available for selection in a monomorphic, ancestral population. Foraging differences between the sexes cannot lead to the evolution of RSD; evolution of large and small morphs of both sexes is a more likely outcome. Selection for sex-role differentiation factors (e.g. large females lay larger eggs, small males are more agile in flight) can lead to the evolution of RSD, but only if the magnitudes of opposing selection for small males and for large females are equal. Combining selection for size-related foraging differences with selection for sex-role differentiation factors hinders the evolution of RSD until the sexes differ in size by 3 s.d . Empirical evidence supports this assertion: statistically significant differences between the sexes in the size of prey taken are found only in highly dimorphic species. The sex-role differentiation factors that have been proposed appear unlikely to provide the equal selection necessary for the evolution of RSD. Several authors have proposed that small size in males is selected for foraging ability and large size in females for some sex-role differentiation factor. Males cannot be more efficient foragers without females being less efficient and efficiency cannot be a factor only when the male is feeding his family. RSD cannot evolve in monogamous species if large females survive less well than small males. RSD might evolve as the result of sexual selection for small size in males and constraints on the reduction of size in females because of some factor associated with reproduction. Examination of seven studies indicating a relationship between female size and reproductive success shows very little unequivocal evidence for small size in females allowing breeding earlier in the season. Large size in females allows females to breed at a younger age in the sparrowhawk and pairs to form more rapidly in three species of sandpipers. Both of these may be the result of sexual selection. There are fewer theoretical problems with sexual selection as a cause for the evolution of RSD than with the other hypotheses. Empirical evidence for sexual selection is scarce but better than that for the other hypotheses. Evidence is contradictory for the selection of small size in males for agility in aerial displays for courtship or defence of territory. Large size in females does not appear to be the result of selection for competitive ability to obtain mates. Facilitation of female dominance and hence of the formation and maintenance of a pair bond is the most viable explanation of the evolution of RSD. It is most likely that all dimorphism (normal or reversed) is the result of sexual selection. RSD is correlated with birds in the diet in the Falconiformes and this is a central theme in the foraging hypotheses. This correlation may be because birds are abundant and available in a continuum of sizes, thus permitting but not causing the evolution of RSD or because species that prey upon birds are better equipped physically (and perhaps more likely behaviourally) to inflict damaging attacks on conspecifics and the greater RSD increases female dominance and the ease of pair formation.     

BODY SIZE AND HAREM SIZE IN MALE RED-WINGED BLACKBIRDS: MANIPULATING SELECTION WITH SEX-SPECIFIC FEEDERS

We experimentally manipulated the strength of selection in the field on red-winged blackbirds (Agelaius phoeniceus) to test hypotheses about contrasting selective forces that favor either large or small males in sexually size dimorphic birds. Selander (1972) argued that sexual selection favors larger males, while survival selection eventually stabilizes male size because larger males do not survive as well as smaller males during harsh winters. Searcy (1979a) proposed instead that sexual selection may be self limiting: male size might be stabilized not by overwinter mortality, but by breeding-season sexual selection that favors smaller males. Under conditions of energetic stress, smaller males should be able to display more and thus achieve higher reproductive success. Using feeders that provisioned males or females but not both, we produced conditions that mimicked the extremes of natural conditions. We found experimental support for the hypothesis that when food is abundant, sexual selection favors larger males. But even under conditions of severe energetic stress, smaller males did not gain larger harems, as the self-limiting hypothesis predicted. Larger males were more energetically stressed than smaller males, but in ways that affected their future reproductive output rather than their current reproductive performance. Stressed males that returned had smaller wings and tails than those that did not return; among returning stressed males, relative harem sizes were inversely related to wing and tail length. Thus, male body size may be stabilized not by survival costs during the non-breeding season, nor by energetic costs during the breeding season, but by costs of future reproduction that larger males pay for their increased breeding-season effort.     

Selection on body size in a raptor with pronounced reversed sexual size dimorphism: are bigger females better?

An overabundance of hypotheses have been proposed to accountfor reversed sexual size dimorphism (RSD; females the largersex) in raptors. Previous research principally focused on examininginterspecific patterns of RSD, rarely testing predictions ofvarious hypotheses within populations. To redress this, we useddata from both sexes of a large brown falcon, Falco berigora,population to evaluate the importance of size and body conditionindices on the hunting prowess of males and the reproductivesuccess, recruitment, and survival probabilities of both sexes.Female-female competition for territorial vacancies was likelyto be intense as the floating population was female-biased andintrasexual agonistic interactions were frequently observed.In this competitive population, larger adult females were morelikely to be recruited, indicating directional selection favoringincreased female body size. Furthermore, after recruitment largerfemales were more likely to successfully fledge offspring, providinga mechanism by which RSD is maintained in the population. Incontrast, male recruitment was unrelated to either body sizeor condition indices. Smaller immature males more often heldtheir territories (survived) over two breeding seasons thandid their larger counterparts; however, they also took smallprey more frequently, a diet related to poor reproductive success.We argue that, together, these results are indicative of selectionfavoring an increase in female body size and a reduction ormaintenance in male body size. Of all the hypotheses proposedto account for the maintenance and evolution of RSD in raptors,this scenario is consistent only with the predictions of theintrasexual competition hypothesis.     

Size-Related Advantages for Reproduction in a Slightly Dimorphic Raptor: Opposite Trends between the Sexes

Despite many comparative analyses and more than 20 proposed hypotheses, there is still little consensus over the factors promoting the evolution of reversed sexual dimorphism (RSD) in raptorial species. Furthermore, intrapopulation studies, which may elucidate how RSD is maintained once evolved, have been surprisingly scarce and only focused on a handful of species with medium to high dimorphism. We examined the reproductive advantages associated with body size and condition, measured in the pre‐laying period, in a diurnal raptor with low sexual dimorphism, the black kite (Milvus migrans). The study population was essentially monomorphic in size. For females, there was an evidence of reproductive benefits associated with larger size and/or with better body condition. Larger females had also access to higher quality partners and territories, consistent with the ‘intrasexual selection’ hypothesis, by which members of the larger sex enjoy size‐related advantages in intrasexual competition over a scarce resource, the smaller sex. Opposite trends emerged for males: smaller, leaner males had higher breeding output, consistent with the ‘small efficient male’ hypothesis. Overall, the fact that we observed in an essentially monomorphic population the same selection pressures previously found in species with marked dimorphism suggests that such reproductive advantages may be counterbalanced in our study model by opposite selection pressures during other stages of the life cycle. This casts some doubts on the evolutionary significance of studies focusing exclusively on reproduction and calls for the need of more comprehensive analyses incorporating trait‐mediated differentials in survival and recruitment.     

Female survival,lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value.     

Cross‐fostering eggs reveals that female collared flycatchers adjust clutch sex ratios according to parental ability to invest in offspring

Across animal taxa, reproductive success is generally more variable and more strongly dependent upon body condition for males than for females; in such cases, parents able to produce offspring in above‐average condition are predicted to produce sons, whereas parents unable to produce offspring in good condition should produce daughters. We tested this hypothesis in the collared flycatcher (Ficedula albicollis) by cross‐fostering eggs among nests and using the condition of foster young that parents raised to fledging as a functional measure of their ability to produce fit offspring. As predicted, females raising heavier‐than‐average foster fledglings with their social mate initially produced male‐biased primary sex ratios, whereas those raising lighter‐than‐average foster fledglings produced female‐biased primary sex ratios. Females also produced male‐biased clutches when mated to males with large secondary sexual characters (wing patches), and tended to produce male‐biased clutches earlier within breeding seasons relative to females breeding later. However, females did not adjust the sex of individuals within their clutches; sex was distributed randomly with respect to egg size, laying order and paternity. Future research investigating the proximate mechanisms linking ecological contexts and the quality of offspring parents are able to produce with primary sex‐ratio variation could provide fundamental insight into the evolution of context‐dependent sex‐ratio adjustment.     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Sex-Specific Recruitment and Brood Sex Ratios of Eurasian Kestrels in a Seasonally and Annually Fluctuating Northern Environment

Timing of birth and food availability may select for biased offspring sex ratios when they differentially affect the reproductive value of male and female young. Here we show that early hatching date enhances more the probability of male Eurasian kestrels (Falco tinnunculus) to breed as one-year-old than that of females in a Finnish population. This rarely documented phenomenon has been previously observed in a kestrel population in the Netherlands. As kestrels in the Finnish population are migratory, our results refute the hypothesis that early-fledged males would have an advantage for early breeding only in resident populations. Contrary to the predictions, the Finnish population showed no change in brood sex ratio during the breeding season in a long-term data from 8years. As far as we know, this is the first demonstration that biased sex allocation may not occur even when it would appear to be adaptive. This result is different from the Dutch kestrel population, in which the season began with a bias towards males and ended with a bias in favour of females. We suggest that high inter-annual variation in food abundance in Finland might reduce selection for a sex ratio trend.     

Seasonal shifts in sex ratios of fledgling American kestrels (Falco sparverius paulus): The Early Bird Hypothesis

We document a seasonal shift in the sex ratios of broods produced by resident southeastern American kestrels (Falco sparverius paulus) breeding in nest boxes in Florida. Early in the breeding season, most biased broods were biased towards males, whereas later in the season, most biased broods were biased towards females. Computer-simulated broods subjected to sex-biased egg and/or nestling mortality demonstrate that it is possible that differential mortality produced the pattern of bias that we observed. However, these simulations do not exclude the possibility that female kestrels were manipulating the primary sex ratio of the broods. We present evidence that this sex ratio shift is adaptive: for males we detected breeding as yearlings, all had fledged early the previous season. No such relationship between season and the probability of breeding as a yearling was found for females. We propose the Early Bird Hypothesis as the ecological basis for the advantage of fledg ing early in males. We hypothesize that pre-emptive competition among post-fledging, dispersing males for breeding sites confers an advantage to males fledged early in the season. This hypothesis may explain why a non-migratory population of the Eurasian kestrel (F. tinnunculus) and non-migratory American kestrels breeding in Florida (F. s. paulus) exhibit this seasonal shift in sex ratios, whereas migratory American kestrels (F. s. sparverius) breeding in Saskatchewan, Canada, do not. We discuss the relevance of the Early Bird Hypothesis for other animal species.     

Age-related change in breeding performance in early life is associated with an increase in competence in the migratory barn swallow Hirundo rustica

1. We investigated age-related changes in two reproductive traits (laying date and annual fecundity) in barn swallows Hirundo rustica L. using a mixed model approach to di-stinguish among between- and within-individual changes in breeding performance with age. 2. We tested predictions of age-related improvements of competence (i.e. constraint hypothesis) and age-related progressive disappearance of poor-quality breeders (i.e. selection hypothesis) to explain age-related increase in breeding performance in early life. 3. Reproductive success increased in early life, reaching a plateau at middle age (e.g. at 3 years of age) and decreasing at older age (> 4 years). Age-related changes in breeding success were due mainly to an effect of female age. 4. Age of both female and male affected timing of reproduction. Final linear mixed effect models (LME) for laying date included main and quadratic terms for female and male age, suggesting a deterioration in reproductive performance at older age for both males and females. 5. We found evidence supporting the constraints hypothesis that increases in competence within individuals, with ageing being the most probable cause of the observed increase in breeding performance with age in early life. Two mechanisms were implicated: (1) advance in male arrival date with age provided middle-aged males with better access to mates. Yearling males arrived later to the breeding grounds and therefore had limited access to high-quality mates. (2) Breeding pairs maintaining bonds for 2 consecutive years (experienced pairs) had higher fecundity than newly formed inexperienced breeding pairs. 6. There was no support for the selection hypothesis because breeding performance was not correlated with life span. 7. We found a within-individual deterioration in breeding and migratory performance (arrival date) in the oldest age-classes consistent with senescence in these reproductive and migratory traits.     

宁波滑蜥两性异形和雌性繁殖

蜥蜴的雌性繁殖特征对理解两性异形的进化原因起着重要作用。于2011年4月在安徽滁州采集宁波滑蜥(Scincella modesta),定量研究该种形态特征的两性异形和雌性繁殖特征,检验成体形态特征两性异形与雌性繁殖的相关性。研究共采集43条(17♀♀,26♂♂)宁波滑蜥,雄性和雌性个体的最大体长分别为47.4 mm和46.6 mm。雌雄两性在体长和头宽上没有差异,而在腹长和头长上差异显著,雄性有较大的头长,雌性有较大的腹长。宁波滑蜥年产单窝卵。窝卵数和窝卵重与雌体体长及腹长呈正相关,卵重与雌体体长无相关性。窝卵数及卵重的变异系数分别为0.20和0.12。卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数来增加繁殖输出。这些结果表明,宁波滑蜥是雌雄个体大小同形的两性异形模式,性选择使得雄性有着较大的头长,以具有较高的交配成功率,生育力选择使得雌性有着较大的腹长,以具有较大的生育力和繁殖输出。     

Interactive effects of food and age on breeding in the Montagu's Harrier Circus pygargus

We evaluated the direct and interactive effects of food and age on reproduction in the Montagu's Harrier Circus pygargus , to test whether variation in food supply was likely to affect age-specific breeding probability or success. Younger females were more frequently non-breeders than older females. When breeding, older females laid earlier, produced larger clutches, failed less often and had higher number of fledglings than younger females. Probability of breeding was higher, laying was earlier, and clutch size and number of fledglings per pair increased with increasing food abundance. A significant interaction between food and age was observed in both breeding probability and breeding performance: older females were more likely to breed than younger females when food abundance was low, and younger females performed less well in good food conditions than older females. Overall, differences between age groups were most marked in extreme food conditions, regardless of the quality of the conditions.     

The apparent rôle of temperature in breeding initiation and winter population structure in Hyale nilssoni Rathke (Amphipoda): Field observations 1972–1983

Data are presented on the population structure of Hyale nilssoni Rathke (an amphipod inhabitant of high littoral seaweeds) over the period of breeding initiation (late February) of 11 yr. By sampling, as near as possible, the same date annually, photoperiodic effects are controlled. The percentage of ovigerous females on this date was significantly correlated with post-winter solstice mean sea temperature. February sex ratios (M : F) did not correlate with either preceding August or September mean sea temperature such as would have been expected was sex determination temperature sensitive. Sex ratio, however, was significantly positively related to mean winter sea temperatures, with warmer winters having male-dominated Hyale populations in late February. Behavioural hypotheses involving amphipod migration and/or size specific elimination of larger males (c.f. females) during harsher winters are offered in explanation. Population size-frequency distributions over 11 yr were remarkably consistent. Male size was more heterogeneous between years than female size. Mean male size (late February) proved to be significantly negatively correlated with mean winter sea temperatures. Mean female size showed no significant relationship. Given that large males are necessary for precopulatory carrying behaviour it is important that, as shown, they are available to initiate spring breeding, even after harsh winters.     

Body size and determinants of laying date variation in the Snow Petrel Pagodroma nivea

We studied several determinants of laying date variation and the relationship between laying date and reproductive success in the Snow Petrel Pagodroma nivea . The effects of female body size and condition, year, individual laying period, colony size, mate fidelity, previous reproductive success, and duration of the pre-laying exodus on laying date, were investigated during a 3-year study. The average laying date was 4 December. The laying period was compressed into 10–16 days and was very constant from year to year, both for the population as a whole and for individual females. The laying period of individual females varied from year to year on average by less than one day. Body size explained 24% of the variation in laying dates, with large females laying their egg later than small ones. Laying in large colonies occurred c. 2 days later than in small colonies, mainly because a higher proportion of large females bred in large colonies. There was no effect of mate fidelity, age, body condition and previous reproductive success on laying date, but the duration of the pre-laying exodus was strongly correlated with laying date. Smaller females had shorter pre-laying exodus (c. 17.7 days) than larger ones (c. 20.4 days). During the three years of the study reproductive success either increased, decreased or did not vary with laying date. Although body size is not maintained by selection on laying date alone, the genetic body size component of this species suggests that balancing selection on body size may act through laying date.     

OBSERVATIONS OF GROUP BEHAVIOUR AND BREEDING BIOLOGY OF THE YELLOW-BILLED SHRIKE CORVINELLA CORVINA

Yellow-billed Shrikes were found to live in groups throughout the year. Within the group, each member helped to defend the group's territory, warn against predators and feed the breeding female, nestlings and fledglings.
During the study there was little change in the location of the boundaries and in the areas of the territories occupied by the majority of the groups. The densities of the larger groups were in general two to three times that of smaller groups. Numbers within one group varied by ±24% of the average (12) during a period of three years.
Progeny remained in a group for some years before dispersing, sometimes in parties of the same sex. Both sexes exchanged groups, the females moving on average further than males. During successive periods in the history of a group the representation of the sexes varied from a surplus of females to a surplus of males. In the population as a whole the sex ratio was probably parity.
Only one female bred in a group at a particular time and she alone incubated. Eggs were laid on consecutive days. Breeding started at the height of the dry season; the first peak in egg laying occurred at the beginning of the rains; laying continued through the wet season and ceased usually in August. The most frequent clutch size was four, and varied little within a breeding season or between seasons. The incubation period ranged from 15 to 18 days, the most frequently recorded being 17 days. The nestling period was 19 days. The percentage of total eggs laid that produced fledglings was 25% and yearlings 11%.
Young shrikes were independent in the seventh week, participated in group displays in their tenth week and fed fledglings in their fourteenth week.
The age of first breeding was not discovered. Two females in their sixth year were still helpers in a group at the end of the study.     

THE EVOLUTION OF FEMALE BODY SIZE IN RED-WINGED BLACKBIRDS: THE EFFECTS OF TIMING OF BREEDING,SOCIAL COMPETITION,AND REPRODUCTIVE ENERGETICS

We examined opposing selective forces on female body size in the sexually dimorphic red-winged blackbird: social competition favoring larger females, and energetic advantages favoring smaller females. Downhower proposed that selection might drive female birds to be smaller than the optimum for survival, if smaller females were able to exceed their energetic requirements for self-maintenance earlier in the season and therefore breed earlier. Since in most birds the earliest breeders fledge the most young, this could favor the evolution of smaller female size, and therefore contribute to the magnitude of sexual size dimorphism in these birds. We tested this hypothesis in 1987 and 1988 by comparing the size and breeding date of female red-winged blackbirds. Consistent with our preditions, early-nesting females had much higher nesting success, but contrary to prediction, larger females bred earlier. We then examined the effects of female size on competition. If large females have an advantage in social competition, and if competition influences breeding date and reproductive success, then larger females might breed earlier. Primary females, the first females to arrive and nest on a territory, were more aggressive than lower ranked females; more aggressive females settled on better territories and laid earlier than less aggressive females; and larger females were more aggressive. Social competition between females may therefore favor large females. Finally, we tested the prediction that selection favoring large females might be limited by energetic constraints on large females. We found that large females had less fat than small females during breeding, and that the levels of fat that females of a given size carried affected breeding date and egg size. Therefore, social competition may favor large females, but reproductive energetics favoring smaller females may constrain selection for large female body size.     

Variation in selection pressure acting on body size by age and sex in a reverse sexual size dimorphic raptor

Body size strongly influences fitness, with larger individuals benefiting in terms of both greater productivity and survivorship; for reverse sexual size dimorphic (RSD) species, this relationship may be more complex. We examined the selection pressures acting on body size in male and female Merlins Falco columbarius to assess whether larger or smaller individuals of this RSD species were favoured in terms of survival and breeding performance. For males and females there were clear links between body size and survival but the exact relationship varied by sex. Among males, birds that survived each year class were larger than those that died and yearlings were on average smaller than older birds, but there were no measurable differences among adult males (age 2+). Among females, larger individuals aged 1 and 2 years were more likely to survive, but this size‐based pattern was not apparent in older age classes. Size early in life predicted the lifespan in male Merlins but not as strongly as for females and not for the largest individuals. Reproductive performance based on brood size was not associated with body size in either males or females, but there was a weak positive relationship between female body size and lifetime reproductive success. Selection appears to favour larger males and females but there is no indication that the population is evolving towards bigger individuals, perhaps in part due to selection against the largest birds. Increased survival may allow larger and higher quality individuals to occupy higher quality territories as they age and thereby to accrue greater lifetime reproductive success in the process.