Japanese monkeys (Macaca fuscata) discriminate between pictures of conspecific males and females without specific training

We investigated whether Japanese monkeys can discriminate pictures of conspecific males and females using a visual paired comparison (VPC) task. Whole-body pictures of adult and nonadult monkeys were used as stimuli. The monkeys were first familiarized with pairs of pictures of different monkeys from one sex category (the familiarized sex). Pairs of novel pictures of a member of the familiarized sex and the opposite sex (novel) were then presented in test. The monkeys showed a preference for novel-sex pictures of both adult and nonadult individuals, indicating that they perceive the differences between familiarized- and novel-sex pictures. These results suggest that monkeys discriminate between pictures of males and females without specific training.     

The role of sex steroids in courtship, pairing and pairing behaviors in the socially monogamous zebra finch

The purpose of this study was to test whether sex steroid actions are necessary for courtship and pairing in socially monogamous birds. We examined the effects of an aromatase inhibitor, 1,4,6-androstatriene-3,17-dione (ATD), combined with an anti-androgen, flutamide (F), on the behavior and pairing status of initially unpaired male and female zebra finches (Taeniopygia guttata). In the first experiment, 24 adult males were implanted with either a combination of ATD and flutamide or empty implants. Two weeks after implantation, birds were housed in aviaries containing 3 ATD + F males, 3 control males, and 3 females and allowed 2 weeks to pair, with observations 7 times during the 2-week period. A second experiment tested the effects of these same treatments in females. During the first 4 days of testing, ATD + F males were less likely to attack conspecifics than were control males. ATD + F males were also less likely to "greet," or approach, females than were control males, but other courtship behaviors, including directed singing, were unaffected. ATD + F females did not differ from control females on any courtship behavior measured. Furthermore, these treatments did not affect pairing behaviors (time spent clumping or in a nest box together) or the likelihood of pairing with a partner of the opposite sex. ATD + F treatments in females did, however, increase the likelihood of same-sex pairing. This suggests that, although sex steroids may regulate some courtship behaviors in males, they do not regulate pairing behaviors and have little effect on the likelihood that a male or female will be chosen as a mate by a bird of the opposite sex.     

Evolution of male parental care and female multiple mating: game-theoretical and two-locus diploid models

Males gain a fitness benefit by mating with many females, whereas the number of progeny per female does not increase as a function of additional mates. Furthermore, males run the risk of investing in the offspring of other males if they provide parental care. Nevertheless, in various species, males provide parental care, and females mate with multiple males. We investigate a game-theoretical model in which females gain a direct benefit by multiple mating from the paternal care they elicit for their offspring. The parameters that directly favor male parental care, such as small cost of paternal care, have indirect positive effects on the evolution of female multiple mating, while they have negative effects in the opposite case. Both traits are more likely to evolve when the number of matings is smaller. The individual-based model of a diploid two-locus, two-allelic genetic model confirms the result.     

Extreme bias in sex allocation in Eclectus parrots

We investigated extraordinary patterns of sex allocation in captive eclectus parrots (Eclectus roratus). These birds are extremely unusual as they show reverse sexual dichromatism, they are the only cooperatively breeding parrot, and they are one of the few birds with nestlings that are easily sexed. They lay two eggs per clutch, but often only fledge one young, and the sex ratio of 209 fledglings did not differ significantly from parity. However, when two young are fledged together they are very likely to be of the same sex, and some females produce long unbroken runs of one sex (the maximum was 20 males) before switching to the other sex. Monte-Carlo simulations show that these runs of same-sex clutches defy expectation if we assume that the sex of chicks within each clutch is independent of the previous clutch. We use further simulations to show that the sex bias must occur at fertilization (i.e. the primary sex ratio), although the female may make further adjustments via infanticide. Control over sex allocation in eclectus parrots is one of the most extreme reported from birds.     

Sex and environmental sensitivity in blue tit nestlings

In birds and mammals with sexual size dimorphism (SSD), the larger sex is typically more sensitive to adverse environmental conditions, such as food shortage, during ontogeny. However, some recent studies of altricial birds have found that the larger sex is less sensitive, apparently because large size renders an advantage in sibling competition. Still, this effect is not an inevitable outcome of sibling competition, because several studies of other species of altricial birds have found the traditional pattern. We investigated if the sexes differ in environmental sensitivity during ontogeny in the blue tit, a small altricial bird with c. 6% SSD in body mass (males larger than females). We performed a cross-fostering and brood size manipulation experiment during 2 years to investigate if the sexes were differently affected as regards body size (body mass, tarsus and wing length on day 14 after hatching) and pre-fledging survival. We also investigated if the relationship between body size and post-fledging survival differed between the sexes. Pre-fledging mortality was higher in enlarged than in reduced broods, representing poor and good environments, respectively, but the brood size manipulation did not affect the mortality rate of males and females differently. In both years, both males and females were smaller on day 14 after hatching in enlarged as compared to reduced broods. In one of the years, we also found significant Sex × Experiment interactions for body size, such that females were more affected by poor environmental conditions than that of males. Body size was positively correlated with post-fledging survival, but we found no interactive effects of sex and morphological traits on survival. We conclude that in the blue tit, females (the smaller sex) are more sensitive to adverse environmental conditions which, in our study, was manifest in terms of fledgling size. A review of published studies of sex differences in environmental sensitivity in sexually size-dimorphic altricial birds suggests that the smaller sex is more sensitive than the larger sex in species with large brood size and vice versa.     

Does breeding population trajectory and age of nesting females influence disparate nestling sex ratios in two populations of Cooper's hawks?

Offspring sex ratios at the termination of parental care should theoretically be skewed toward the less expensive sex, which in most avian species would be females, the smaller gender. Among birds, however, raptors offer an unusual dynamic because they exhibit reversed size dimorphism with females being larger than males. And thus theory would predict a preponderance of male offspring. Results for raptors and birds in general have been varied although population‐level estimates of sex ratios in avian offspring are generally at unity. Adaptive adjustment of sex ratios in avian offspring is difficult to predict perhaps in part due to a lack of life‐history details and short‐term investigations that cannot account for precision or repeatability of sex ratios across time. We conducted a novel comparative study of sex ratios in nestling Cooper's hawks (Accipiter cooperii) in two study populations across breeding generations during 11 years in Wisconsin, 2001–2011. One breeding population recently colonized metropolitan Milwaukee and exhibited rapidly increasing population growth, while the ex‐Milwaukee breeding population was stable. Following life‐history trade‐off theory and our prediction regarding this socially monogamous species in which reversed sexual size dimorphism is extreme, first‐time breeding one‐year‐old, second‐year females in both study populations produced a preponderance of the smaller and cheaper sex, males, whereas ASY (after‐second‐year), ≥2‐year‐old females in Milwaukee produced a nestling sex ratio near unity and predictably therefore a greater proportion of females compared to ASY females in ex‐Milwaukee who produced a preponderance of males. Adjustment of sex ratios in both study populations occurred at conception. Life histories and selective pressures related to breeding population trajectory in two age cohorts of nesting female Cooper's hawk likely vary, and it is possible that these differences influenced the sex ratios we documented for two age cohorts of female Cooper's hawks in Wisconsin.     

Family-based winter territoriality in western bluebirds, Sialia mexicana: the structure and dynamics of winter groups

Winter residency is characteristic of the majority of cooperatively breeding birds, but the composition and dynamics of winter groups have been examined in relatively few. In 1996-1998, we examined winter territoriality in the western bluebird, a year-round resident that shows a limited degree of helping behaviour in central coastal California, U.S.A. In spring, most western bluebirds breed as socially monogamous pairs, but a small proportion of pairs (3-16%) have additional breeding-age males helping at the nest, usually assisting parents or brothers. We found that year-round residents commonly wintered in family groups that defended territories similar to those used in spring. Winter groups had an even sex ratio and formed early in the autumn, when hatch-year birds dispersed. More females than males left their natal groups to be replaced by an influx of immigrant hatch-year birds. Winter groups typically consisted of breeders and one or two sons from the prior breeding season along with one or more immigrant females. A second period of dispersal occurred in spring when winter groups broke up and most birds other than the breeding pair left the winter territory. When they bred, yearling males and females often bred with unrelated individuals from their winter groups. Sons were more likely to remain on the study area as yearlings when they wintered with both parents than when they wintered with just one parent. We suggest that young males stay the winter due to benefits of remaining in family groups on mistletoe-based winter territories. Subsequent localized dispersal of sons then leads to opportunistic kin-based interactions later in life. Copyright 2001 The Association for the Study of Animal Behaviour.     

Young and female‐biased irruptions in pygmy owls Glaucidium passerinum in southern Finland

Irruptive migrants often show biased sex and age ratio, and typically young females are the most abundant class participating in irruptions. Several hypotheses have been proposed to explain differential migration in birds. We examined these hypotheses in a sexually size‐dimorphic species (females being larger than males), the pygmy owl Glaucidium passerinum, whose migration behaviour has also been poorly documented. Migration data were collected at the Hanko Bird Observatory, SW Finland during 1979–2010. Pygmy owls showed large fluctuations in migration numbers between years. Most of the migrants were young birds and females migrated in larger numbers than males. Results support the ‘territory defence hypothesis’ since males showed weaker tendency to migrate despite the smaller body size, but the ‘dominance hypothesis’ was also supported (subordinate young outnumber dominant adults in migration numbers). Findings support the idea that in owl species, which are dependent on nesting cavities, males show lower migration behaviour than females, whereas the pattern is opposite in species which normally breed outside cavities. In addition, adults migrated later than young, likely because they need to moult remiges before departure. Early hatched young also migrated earlier than late hatched young, suggesting that more dominant young migrate first.     

WHY SHOULD LEK-BREEDERS BE MONOMORPHIC?

Approximately one-quarter of all lek-breeding bird species are sexually monomorphic. Understanding the significance, if any, of this exception to the usual correlation between sexual selection and dimorphism requires detailed data on the mating systems of both monomorphic and dimorphic species. The capuchinbird (Perissocephalus tricolor) is a sexually monomorphic, lek-breeding member of the cotinga family. I studied the social and sexual behavior of this species, and compared it with the Guianan cock-of-the-rock (Rupicola rupicola), a dimorphic, lekking member of the same family. Male–male competition in capuchinbirds involved direct contests for dominance, rather than territorial displays as in classic lek species. In each year, one dominant individual was able to control the most desired display site on the 8-male lek, and was the only male that copulated. In contrast to dimorphic lek birds, female as well as male capuchinbirds engaged in frequent and intense aggression at the lek, and both males and females engaged in sexual mimicry. I suggest that plumage monomorphism in lek birds has evolved as a result of social competition affecting both sexes. This hypothesis accounts for the exaggerated plumage characters shared by males and females in capuchinbirds and a number of other monomorphic lek birds. The evolution of plumage can best be analyzed as an arms race, in which the balance of selective forces acting on each sex can produce a variety of equilibrium states, ranging from sexual indistinguishability to extreme dimorphism.     

Sex allocation in black-capped chickadees Poecile atricapilla

Optimal sex allocation for individuals can be predicted from a number of different hypotheses. Fisherian models of sex allocation predict equal investment in males and females up to the end of parental care and predict brood compositions based on the relative costs of producing males and females. The Trivers-Willard hypothesis predicts that individual females should alter the sex ratio of their broods based on their own condition if it has a differential impact on the lifetime reproductive success of their sons and daughters. The Charnov model of sex allocation predicts that females should alter sex allocation based on paternal attributes that may differentially benefit sons versus daughters. Because females are the heterogametic sex in birds, many recent studies have focussed on primary sex ratio biases. In black-capped chickadees Poecile atricapilla , males are larger than females suggesting they may be more costly to raise than females. Female condition affects competitive ability in contests for mates, and thus may be related to variance in fecundity. Females prefer high-ranking males as both social and extrapair partners. These observations suggest that females might vary the sex ratio of their broods based on the predictions of any of the above models. Here, we report on the results of PCR based sex determination of 1093 nestlings in 175 broods sampled from 1992 to 2001. Population-wide, we found a mean brood sex ratio of 0.525±0.016, with no significant deviation from a predicted binomial distribution. We found no effect of clutch size, female condition, hatch date, parental rank or paternity. Our results reject the idea that female black-capped chickadees systematically vary sex allocation in their broods.     

Priority versus brute force: when should males begin guarding resources?

When should males begin guarding a resource when both resources and guarders vary in quality? This general problem applies, for example, to migrant birds occupying territories in the spring and to precopula in crustaceans where males grab females before they molt and become receptive. Previous work has produced conflicting predictions. Theory on migrant birds predicts that the strongest competitors should often arrive first, whereas some models of mate guarding have predicted that the strongest competitors wait and then simply usurp a female from a weaker competitor. We build a general model of resource guarding that allows varying the ease with which takeovers occur. The model is phrased in terms of mate-guarding crustaceans, but the same logic can be applied to other forms of resource acquisition where priority plays a role but takeovers might be possible too. The race to secure breeding positions can lead to strong competitors (large males) taking females earliest, even though this means accepting a lower-quality female. Paradoxically, this means that small males, which have fewer breeding opportunities, are more choosy than larger ones. Such solutions are found when takeovers are impossible. The easier the takeovers and the higher the rate of finding guarded resources, the more likely are solutions where guarding durations are short, where strong competitors begin guarding only just before breeding, and where they do this by usurping the resource. The relationship between an individual's competitive ability and its timing of resource acquisition can also be nonlinear if takeovers are moderately common; if this is the case, then males of intermediate size guard the longest.     

Female mate preference to maximize paternal care. II. Female competition leads to monogamy

ABSTRACT A two-step game model of female mate preference and paternal care is examined, with a particular focus on the case of two females and two males. In a mating season, females choose their mates, and in the following breeding season males invest in paternal care, knowing the likelihood of their paternity in chicks. If parental ability is the same between individuals of each sex, the evolutionarily stable mating pattern is always monogamy. If females differ in fecundity and males differ in paternal care capacity, monogamy with assortative mating is likely to be evolutionarily stable. If the male cost function increases at a strongly accelerating rate, however, polyandry is evolutionarily stable when the difference of female fecundity is very large, but the game may have no evolutionarily stable state when the difference of female fecundity is small. The care graph (in which females are connected to males giving paternal care to their chicks) is often much simpler than the mating graph (in which females are connected to males they accepted). To be exact, no "loop" should be included in the evolutionarily stable care graph for the general case of n females and m males. This prediction is in accord with the observed prevalence of social monogamy in spite of genetic promiscuity among altricial birds.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

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.     

Sex ratio, survival and territorial behaviour of polygynous Hen Harriers Circus c. cyaneus in Orkney

Polygyny is widespread among Orkney's Hen Harriers Circus cyaneus . From 1975–81, it was associated with a sex ratio estimated from resightings of colour-marked birds up to six years of age to be 29 females:ten males. The sex ratio of fledglings changed significantly from a greater proportion of females in the 1950s and early 1960s to a greater proportion of males subsequently. The mean estimated 'survival' rates (birds colour-marked in Orkney which were seen there in later years) of males and of females 0–2 years old were 14 and 29%. The mean estimated annual survival rates of males and females from 2–6 years old were 72 and 90%. It is suggested that the uneven sex ratio resulted in more frequent intra-sexual encounters and displays by females. The results of temporary removals of two females (immediately replaced) and of two males (not replaced) in spring indicates that there was a shortage of males in the population.     

Reconnaissance for future breeding sites by spotted sandpipers

We studied the sex-role-reversed, polyandrous spotted sandpiper(Actitis macularia) from 1974 to 1990 in northern Minnesota,USA. After peak arrival of breeding birds and before peak departureat the end of the breeding season, there were many short-termvisitors (transients) to the study site. Stepwise discriminantfunction analysis (DFA) was used to determine the importanceof absolute sex ratio (males/female), sex of the transient bird,number of nests, and number of breeding males and females duringthe week of visit in predicting whether a visiting bird wouldreturn the following year. In addition, multiple regressionwas used to determine how much variability in the number oftransient birds returning in subsequent yearscould be explainedby annual values during the year of transience for numbers ofbreeding males and females, numbers of eggs laid and hatched,and absolute sex ratio. Annual recruitment of foreign adultsranged from 1 to 20 birds, of which 0–56% were seen visitingin previous years. Female recruits were more likely than malesto have been observed previously as transients. Twenty-two chickshatched at our study site returned and bred for the first timemore than 1 year after hatching. Of these, 9 (41%) were seenas transients between the year of hatch and breeding. The DFAshowed that transient females returned more often than transientmales and that the number of transients returning in subsequentyears was positively associated with the absolute sex ratioduring the week visited. When the sexes were analyzed separately,none of the weekly variables significantly discriminated femalereturn, but sex ratio was positively associated with male return.Regression showed that the number of transient birds returningin subsequent years was positively associated with the numberof male breeders at our study site during the year a bird visited.Percentage return the year after transience was positively associatedwith the number of eggs laid at our study site during the yeara bird visited. When sexes were analyzed separately, the higherthe number of female breeders during the year a bird visited,the greater the number of males returning in subsequent years,and greater numbers of breeding males were positively associatedwith transient female return. Based on our results, we suggestthat transient birds were searching for better breeding areasfor future breeding and that intrasexual competition made thisinformation more important to females than to males.     

Turnover, age and sex ratios of kestrels (Falco tinnunculus) in south Scotland

This paper examines the composition and turnover of a population of European kestrels Falco tinnunculus L. in the Southern Uplands of Scotland. Turnover was generally high, with most birds staying in the area for only one summer or winter. The rate at which birds arrived or left was highest in autumn and spring, and was independent of population size. During winter, breeding males were more likely to stay on their territories than females, and males that did so were more likely to retain their partners the following year. Breeding success was associated with a greater likelihood of return the following year, except in first-year females which were unlikely to return anyway. The sex ratio was biased toward males in winter, especially in poor vole years when there were also few first-year birds present. The study confirmed the high rate of turnover reported elsewhere in small raptors, and further indicated that the composition and turnover of the population were related to food supply and seasonal patterns of migration.     

Sexual Imprinting on a Novel Adornment Influences Mate Preferences in the Javanese Mannikin Lonchura leucogastroides

We investigated whether sexual imprinting on an artificial novel adornment in the Javanese Mannikin Lonchura leucogastroides , a monomorphic estrildid finch, can occur and might provide a mechanism for the evolution of novel traits. We introduced a red feather on the forehead as a novel adornment. Young were raised by parents which were both adorned, which were both unadorned, or only one of which was adorned with the red feather. We tested the female and male offspring of those parents in mate choice tests with an adorned and unadorned conspecific of the opposite sex. Males raised by an adorned mother or adorned parents preferred adorned females significantly more often than males raised by unadorned parents. We conclude that males were sexually imprinted on the red feather. Females raised by an adorned mother or raised by adorned parents significantly preferred adorned males, whereas females raised by unadorned parents showed no preference for adorned males. Thus, females also became imprinted on the red feather. Males might learn the novel adornment in combination with the parent's sex or learn just the most conspicuous sex, whereas females showed a preference for the adornment independent of which sex bore the feather. Our study shows that sexual imprinting might be an effective mechanism for the evolution of a novel trait and that males and females might become imprinted on a novel trait in different ways.