Change in offspring sex ratio over a very short season in Lincoln's Sparrows: the potential role of bill development

ABSTRACT The sex ratios of offspring are targets of natural selection that can affect parental energy expenditure and fitness, adult sex ratios, and population dynamics. Parents may manipulate offspring sex ratios based on sex differences in their offsprings' potential for reproductive success. In Lincoln's Sparrows (Melospiza lincolnii), male bill shape is associated with the quality of songs, and song quality predicts female preferences in a reproductive context. Males and females that hatch later relative to brood mates or later in the breeding season tend to develop bill shapes that are, for males, associated with low‐quality song. Because females do not sing and do not experience this selection pressure, we predicted that the sex of offspring produced late relative to their brood mates or relative to the season should be biased toward females. Using a molecular technique to sex nestlings, we found no effects of hatching order or any interaction between date of clutch initiation (season) and hatching order on offspring sex. However, we found a seasonal decline in the proportion of male offspring, from approximately 0.8 at the beginning to 0.4 at the end of a clutch initiation season only 19 d in duration. To our knowledge, this is the shortest period over which the offspring sex ratio has been shown to change in a bird population. Moreover, these findings are consistent with the hypothesis that sex differences in the potential attractiveness of offspring ultimately influence offspring sex ratios.     

SEX-RATIO SELECTION IN A BIVOLTINE THRIPS. I. CONDITIONAL SEX-RATIO MANIPULATION AND FITNESS VARIATION

Females of the bivoltine thrips Elaphrothrips tuberculatus (Hood) (Insecta: Thysanoptera) produce broods of either all males (by viviparity) or all females (by oviparity). Measurements of the sex-allocation ratio, ecological and physiological conditions affecting male and female offspring body size, and correlates of the relative fitnesses of adult males and females in relation to size indicate that female parents tend to be viviparous (produce males) if their offspring will become relatively large adults, and that males gain more in fitness from large size than do females. However, the conditions that link sex allocation with offspring fitness differ between the spring and summer generations. In spring, when breeding is synchronous, 1) oviparous and viviparous females do not differ in body size, 2) females tend to be viviparous where the fungus upon which they feed is relatively dense and where their offspring will become relatively large adults, and 3) fungus density is highly correlated with male and female offspring size. In summer, when breeding is relatively asynchronous, 1) viviparous females are much larger than oviparous females early (but not late) in the season, 2) large viviparous females begin breeding earlier than smaller ones, 3) offspring developing earlier in the season become larger adults, and 4) a higher proportion of females are viviparous earlier than later. Field experiments and field collections show that the covariation among sex allocation, conditions, and fitness is not caused by differential mortality by size or sex. Differences between the spring and summer generations in the cues used by females to adjust offspring sex ratio may be caused by seasonal variation in the factors that affect offspring size. However, in both generations, females tend to produce sons only when their offspring will become relatively large adults, whereas daughters are produced regardless of offspring size. These data suggest that females of E. tuberculatus avoid production of males (the sex with higher variance in expected fitness) when the size of their offspring is relatively uncertain.     

Strong temporal consistency in the individual foraging behaviour of Imperial Shags Phalacrocorax atriceps

Individual consistency in foraging behaviour can generate behavioural variability within populations and may, ultimately, lead to species diversification. However, individual‐based long‐term behavioural studies are particularly scarce in seabird species. Between 2008 and 2011, breeding Imperial Shags Phalacrocorax atriceps at the Punta León colony, Argentina, were tracked with GPS devices to evaluate behavioural consistency during their foraging trips. Within a breeding season, individuals were highly consistent in the maximum distances they reached from the shore and the colony, as well as in the time invested in flight and diving across consecutive days during early chick rearing. In addition, each individual had its specific foraging area distinct from the foraging area of other individuals. Comparing between early and late chick rearing in the same season, individuals were consistent, to a lesser degree, in the maximum distance they reached from the colony and the shore, increasing in consistency later on in the season. Within the season, females were more consistent than males in the maximum distance they moved from the colony and the shore, the sexes segregated in their foraging areas and individual females were segregated from one another. Twenty‐eight individuals tracked in different breeding seasons were marginally consistent in their trip durations and maximum distance reached from shore across seasons. Among seasons, foraging locations differed between sexes and among individual females. Individuals from this colony exhibited consistency over time in several aspects of foraging behaviour, which may be due to a combination of individual characteristics such as learning abilities, breeding experience or health, as well as targeted prey type and stability of the environment at this location.     

Seasonal sex role changes in the blenniid Petroscirtes breviceps, a nest brooder with paternal care

The plasticity of the sex roles in the blenniid fish Petroscirtes breviceps , a nest brooder with exclusive paternal care, was studied throughout an 8 month breeding season. Males performed most courtships early and late in the breeding season, whereas females performed most in the middle of the season. These results indicated that the sex of individuals initiating courtship changed seasonally, with courtship role reversal in the middle of the season. Intrasexual aggression in both sexes occurred much more frequently in mid-season than in the early and late seasons. Males frequently fought when available nest sites were limited, regardless of the presence of females, suggesting that males competed for nests in order to qualify to mate (resource competition). In contrast, courting females fought only in mid-season, when females' relative success in entering nests decreased, indicating that females competed for limited mating opportunities (mating competition). The reversed courtship roles and female mating competition in mid-season suggested that the sex roles in P. breviceps changed seasonally from the conventional roles to reversed roles and back again during one breeding season. This study provides the first empirical evidence of multiple changes in the sex roles of animals within a breeding season.     

Non-breeding behaviour of Magenta Petrels Pterodroma magentae at Chatham Island, New Zealand

Magenta Petrels Pterodroma magentae were caught at light-attraction stations on southwest Chatham Island, New Zealand, and most were fitted with transmitters. Of 52 captured since 1993, 71% were males, and all 36 tracked adequately proved to be non-breeders in the breeding season of capture. Our data indicated no sex bias in their probability of being captured at lights. Males provided 86% of trackings, and 87% of trackings of birds flying over the breeding area were males. Males landed 118 times; females 13 times. Only males were found on the ground, by night and day, apparently unassociated with burrows (three with and ten without transmitters), but subsequently digging burrows ( n  = 8). Of 19 birds banded as fledglings up to 2000, males were first recaptured nearing 4 years old (at lights and on the ground) and a female nearing 6 years old (in burrow). Among 37 fledglings, the sex ratio was even. Nine tracked males occupied burrows, as did two females, but the latter were older recaptures (10+ and 25+ years old). It appears that only males claimed existing, or dug new, burrows. They then attracted a mate to the burrow by means unknown, but from among females frequenting an inshore courtship area near the colony, or occasionally flying over the colony, at night. Females established in burrows, but then losing their mate, were able to re-mate there, by calling from near the burrow or by attracting a mate in flight or from the postulated inshore courtship area. Both sexes sometimes took years to pair or re-mate, possibly reflecting the dearth of available mates.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Body Size and Multiple Copulations in a Neotropical Grasshopper with an Extraordinary Mate-Guarding Duration

After copulation, male grasshoppers of Sphenarium purpurascens (Orthoptera: Pyrgomorphidae) remain in a postinsemination association with their mate. A male can spend as many as 17 days mounted on a female. Guarding duration is related to both male and female body size and the female's mating history. Longest guarding durations were recorded at the middle of the reproductive season, when the probability of encounter between the sexes (sex ratio and population density) was decreasing, at the beginning of the associated dry season. These guardings were associated with large individuals of both sexes and with females that had more previous partners. Moreover, a positive association was found among guarding duration, female and male body size and age, and number of copulations performed by the males. Maybe males invest time and sperm in females as a function of the probability of sperm competition. Nevertheless, guarding may provide benefits to both sexes. Males may reduce the possibility of sperm competition, and females may obtain nutritional benefit for themselves or their offspring as a result of multiple copulations. Changes in male investment in guarding duration and number of copulations may be the result of physiological constraints related to seminal and/or sperm production. Moreover, guarding duration could be constrained by ecological factors such as a reduction of food availability associated with the beginning of the dry season.     

Sex‐specific patterns of fuelling and pre‐breeding body moult of Little Stints Calidris minuta in South Africa

The timing and duration of each stage of the life of a long‐distance migrant bird are constrained by time and resources. If the parental roles of males and females differ, the timing of other life stages, such as moult or pre‐migratory fuelling, may also differ between the sexes. Little is known about sexual differences for species with weak sexual dimorphism, but DNA‐sexing enables fresh insights. The Little Stint Calidris minuta is a monomorphic long‐distance migrant wader breeding in the Arctic tundra. Males compete for territories and perform elaborate aerial displays. Females produce two clutches a season. Each sex may be a bigamist and incubate one nest a season, each with a different partner. We expect that these differences in breeding behaviour entail different preparations for breeding by males and females, so we aimed to determine whether Little Stints showed any sex differences in their strategies for pre‐breeding moult and pre‐migratory fuelling at their non‐breeding grounds in South Africa. We used body moult records, wing length and body mass of 241 DNA‐sexed Little Stints that we caught and ringed between 27 January and 29 April in 2008–2018 at two neighbouring wetlands in North West Province, South Africa. For each individual we assessed the percentage of breeding plumage on its upperparts and took blood samples for DNA‐sexing. We calculated an adjusted Body Moult Index and an adjusted Wing Coverts Moult Index, then used the Underhill–Zucchini moult model to estimate the start dates and the rate of body moult in males and females. We estimated the changes in the sex ratio of the local population during their stay in South Africa, and also estimated the timing and rate of pre‐migratory fuelling and the potential flight ranges for males and females. The males started body moult on average on 7 February and the females on 12 February, but the sexes did not differ in their timing of wing covert moult, which started on average on 10 February. In January to mid‐February, males constituted c. 57% of the population, but their proportion declined afterwards, indicating an earlier departure than females. We estimated that both sexes began pre‐migratory fuelling on average on 15 March. The sexes did not differ in fuelling rate, but most females stayed at the non‐breeding site longer than the males, and thus accumulated more fuel and had longer potential flight ranges. These patterns of moult and fuelling suggest sex differences in preparations for breeding. We suggest that the males depart from South Africa earlier but with smaller fuel loads than the females to establish breeding territories before the females arrive. We conclude that for each sex the observed trade‐offs between fuelling and moult at the non‐breeding grounds are precursors to different migration strategies, which in turn are adaptations for their different roles in reproductive behaviour.     

Cuckoo Hosts in Southern Africa

There is currently no reliable, affordable method of sexing Mauritius Fodies in their first non-breeding season. Ringed immature fodies from a released population on an offshore island were caught in April and May 2005 and sexed in later breeding seasons. Males had longer wing lengths and tarsi than females, with no overlap in wing length between sexes. Males in their first breeding season could usually be differentiated from older males by the paleness and completeness of their breeding plumage. Two adult females grew red feathers characteristic of breeding males in the winter of 2006. It is possible to sex Mauritius Fodies using wing length and separate immature males from adults using the darkness of the bill in the non-breeding season. It is not possible to separate unringed adult and immature females following the postjuvenile and postbreeding moults using current knowledge.     

Sexual Selection on a Coloured Ornament in King Penguins

Species in which the sexes equally exhibit colourful ornaments are an issue for evolutionary theory. Among several hypotheses, sexual selection for mutual mate choice and social selection for signals of behavioural dominance are most commonly supported. We examined the previously documented sex‐similar size of yellow‐orange ear patches in the king penguin, Aptenodytes patagonicus. This species is monogamous and pairs just before reproduction. Raising a chick requires considerable effort by both parents, as they alternate care of their single offspring with foraging at sea. The size of the ear patches appears to signal aggressive territoriality in the breeding colony for both sexes. However, experiments suggest that females prefer large patch size during mate choice, and males do not prefer this trait. We tested whether the size of the coloured ear patch was influenced by sexual selection for couples that had recently paired. We used analyses of covariance to compare the size of the ear patch to a measure of body size and then tested for the difference between males and females. Males were 6.2% larger in ear patch width and 7.7% larger in ear patch area than females, and the distance between the ear patches over the head was 7.5% smaller in males, with all differences highly significant. Consequently, sexual selection appears to favour larger ear patches in males, possibly because of an excess of males that promotes female choice. Social selection also appears to favour the evolutionary maintenance of ear patches of males, and thus both types of selection may contribute to enlarged ear patches.     

Colony attendance and at‐sea distribution of thin‐billed prions during the early breeding season

Procellariiform seabirds have extreme life histories; they are very long‐lived, first breed when relatively old, lay single egg clutches, both incubation and chick‐rearing are prolonged and chicks exhibit slow growth. The early part of the breeding season is crucial, when pair bonds are re‐established and partners coordinate their breeding duties, but is a difficult period to study in burrow‐nesting species. Miniature geolocators (Global Location Sensors or GLS loggers) now offer a way to collect data on burrow attendance, as well as determine at‐sea movements. We studied the early breeding season in thin‐billed prions Pachyptila belcheri breeding at New Island, Falkland Islands. Males and females arrived back at the colony at similar times, with peak arrival in the last days of September. However, males spent more time on land during the pre‐laying period, presumably defending and maintaining the burrow and maximising mating opportunities. Males departed later than females, and carried out a significantly shorter pre‐laying exodus. Males took on the first long incubation shift, whereas females returned to sea soon after egg laying. During the pre‐laying exodus and incubation, males and females travelled at similar speeds (> 250 km d^?1) and were widely distributed over large areas of the Patagonian Shelf. Inter‐annual differences in oceanographic conditions were stronger during the incubation than during the pre‐laying exodus and were matched by stronger differences in distribution. The study thus suggests that extended trips and flexible distribution enable thin‐billed prions to meet the high energy demands of egg production and incubation despite low productivity in waters around the colony during the early summer.     

Male-female interactions of sika deer (cervus nippon) in nara park through allogrooming during breeding and rutting seasons

The frequency and pattern of interactions between males and females of sika deerCervus nippon were surveyed in 3 sites of Nara Park, central Japan, mainly via observation of allogrooming frequency and aggressive behavior throughout the breeding and rutting seasons. In the breeding season, the Daibutsuden group contained several adult males while Ukimido and Hakuba groups contained only a few. Most allogroomers were adult and subadult females. Females groomed the same sex more than the opposite sex at Daibutsuden, but at the other 2 sites, there was no such difference. In general, male-female interactions through allogrooming frequencies in the breeding season were not significantly fewer than intra-sex interactions. Males that attacked females tended to be groomed by females in the breeding season. In the rutting season, all adult males identified in the breeding season were absent at the usual observation sites, and newly arrived males showed defensive and/or mating behaviors. Females groomed adult males in the rutting season regardless of group status or display of sexual interactions. This suggests that male-female allogrooming in the breeding season does not relate to mating in the following rutting season, but may reduce tension in the group during the current breeding season.     

Colony kin structure and breeding patterns in the social wasp, <Emphasis Type="Italic">Polistes biglumis</Emphasis>

We used DNA microsatellites to study colony kin structure and breeding patterns in the primitively eusocial wasp Polistes biglumis. P. biglumis inhabits cool areas at high altitudes and, as a consequence, has a reduced colony cycle compared to more temperate Polistes. P. biglumis colonies are always founded and controlled by a single foundress, but nest failure is common and foundresses losing their nests do not have time to start new ones due to the short season. Instead, nests are characterized by frequent female turnover, in the form of females taking over (usurpation) other con-specific nests. Our results showed that most nests had offspring from multiple unrelated females, including some where multiple females were not observed in monitoring. Reconstruction of behavioural events from the genetic data revealed three types of multiple matriline nests: (a) nests that were usurped by another female, where the original nest owner disappeared following the usurpation event, (b) nests that were joined by another female, where the original nest owner stayed following the joining event, (c) nests that were both usurped and joined by other females. We also found, for the first time, a clear indication of multiple mating by Polistes females. Moreover, males mating with the same female were related, which may be explained by the lek mating system of P. biglumis. Finally, we analysed the nest sex ratios and how it changed during the season and found that sexes were produced sequentially, males before females.     

Seasonal and diurnal variation in vocal behaviour of the South Island robin

Abstract

The proportion of time spent by South Island robins (Petroica australis australis) of different sex, age, and status giving full song, subsong, and downscales, on a monthly and diurnal basis, is described. Females never gave full song. Males did so throughout the year, but least while moulting. Since bachelors sang significantly more than paired males, song probably functioned largely to attract females. Breeding males sang most during the pre-laying stage. From January to April adult males sang significantly more than immature males, but from May to July the reverse was true. Robins sang mainly in the early morning. Robins generally spent less than 0.5% of time during any stage of the breeding cycle giving sub-song. Outside the breeding season, males spent significantly more time giving sub-song than females, and adult males sang more than immature males. Robins in all categories gave more sub-song during the moult than afterwards. The diurnal pattern of sub-song production was bimodal. About half the downscales were followed by no obvious action from the caller or response from other robins, nor were the calls preceded by an obvious event. About 25% of the downscales were associated with interactions between members of a pair; the remainder were associated with territorial defence. Downscales probably served to maintain contact between members of a pair and to advertise possession of territory. These calls were heard rarely in the breeding season, and most frequently in April. Males gave downscales more than females, but use of downscales by both sexes was evenly distributed through the day.     

Patterns of territory settlement and consequences for breeding success in the Northern Wheatear Oenanthe oenanthe

We examined the pattern of territory settlement and its consequences for breeding success in the Northern Wheatear Oenanthe oenanthe on Bardsey Island, Wales, during the breeding seasons of 1991-93. Males returned earlier than females, and older males returned earlier than first-year breeders. Although their boundaries shifted between years, the general location of territories was consistent during the three-year study. There was a high degree of fidelity to area and territory between years for both sexes. The order of territory settlement, from which a territory rank was calculated, was highly consistent for males between years irrespective of individual settlement patterns and territory fidelity. Patterns of territory settlement were less predictable for females, although there was a significant correlation between the mean territory ranks of paired males and females. There was a male-biased sex ratio in each year, and between 5% and 26% of males remained unpaired throughout the breeding season. Male mating status and breeding success were dependent on arrival date, territory rank and breeding density. Early-arriving, usually older, males were able to settle on territories first and were more likely to pair, while later-arriving individuals were more likely to remain unmated. These effects were consistent between years, and consequently territories could be classified as either preferred (accounting for proportionately more breeding attempts) or non-preferred. Territory quality as opposed to individual quality appeared to explain much of the variation in breeding success, and both sexes benefited by breeding on preferred territories through enhanced breeding success and an increased probability that their offspring would be recruited to the population.     

Factors Influencing the Breeding Activity in Male Masu Salmon, Oncorhynchus masou: The Relationship with Body Size

Synopsis Male body size has been emphasized as an important factor contributing to the breeding success of individuals. However, the operational sex ratio (OSR: ratio of mature females to males) during the breeding season significantly change due to differences in the breeding timing and period within and between males and females and may influence the selective advantage of the male large body size for mates. We examined the reproductive ecology of masu salmon Oncorhynchus masou, inhabiting Lake Toya, Hokkaido, northern Japan. Precipitation triggered the upstream migration, although males migrated into the river earlier than females. As a result, the OSR in the river changed markedly during the breeding season, as did the size structure of males in the river. Large males migrated into the river earlier than the smaller males. Differential male survival resulted in the different population size structure between the early and late periods. Under these conditions, we analyzed which males were more successful in releasing the more sperm throughout the breeding season by estimating the decrease in the sperm content in male carcasses. The body size and sperm-releasing success of the males were not related. With a weak male-biased OSR and synchronous timing of reproduction in females, males that had entered the river succeeded in releasing the sperm regardless of body size. Such a fluctuating advantage for different body sizes likely contributes to the maintenance of the size variation in the male salmon.     

DIFFERENCES IN BIOMASS ALLOCATION TO REPRODUCTIVE AND VEGETATIVE STRUCTURES OF MALE AND FEMALE PLANTS OF A DIOECIOUS,PERENNIAL HERB,SILENE ALBA (MILLER) KRAUSE

The pattern of biomass allocation of males and females and the sex ratio and growth characteristics of plants from three seed-size classes in Silene alba were investigated in a greenhouse study. Seed size significantly affected adult plant size and flower production of both male and female plants, but there was no significant difference in the proportion of males and females emerging in three seed-size categories. Male and female plants differed in the proportion of total biomass allocated to vegetative and reproductive structures and these differences were consistent across all seed-size categories. Males allocated a greater proportion of their biomass to flowers than did females. Female reproductive effort was dependent upon the percentage of flowers producing mature capsules. Only females with greater than 20% fruit set have a higher reproductive expenditure than males. Consequently, female expenditure is potentially greater than males, but is spread out over a longer portion of the growing season. This difference in the timing of reproductive expenditures by males and females allows females to allocate more biomass to growth during the early flowering period and may therefore account for the common pattern in herbaceous perennial dioecious species in which adult females are larger than adult males.     

Do females adjust brood sex ratio according to males’ genetic structures in a gregarious passerine,the vinous‐throated parrotbill Paradoxornis webbianus?

A growing number of bird species are known to have fine‐scale genetic structure during the breeding season, with relatives breeding in close vicinity. Such genetic structure often has fitness consequences for parents, and sex ratio theory predicts that females should respond adaptively when they determine offspring sex. We examined whether or not females allocate offspring sex adaptively in response to the local genetic structures as well as other biotic and abiotic factors in a population of the vinous‐throated parrotbill Paradoxornis webbianus, a small passerine with strong flocking habit and various genetic structures among neighbouring males during the breeding season. The average brood sex ratio of hatchlings (secondary sex ratio) did not deviate from parity. In addition, the observed brood sex ratio was independent of the fine‐scale genetic structure and other factors including breeding density, clutch size, laying date, parents’ quality, and the presence of extrapair paternity. Accordingly, we reject the hypothesis of adaptive sex allocation by female parrotbills in association with local genetic structure and other factors. Instead we conclude that despite the plausible benefits of biased sex allocation, this species determines brood sex ratio via random sex allocation with equal probability of male and female offspring.     

Late‐breeding Great Cormorants Phalacrocorax carbo sinensis produce fewer young of the more vulnerable sex

We examined the brood sex ratio and offspring body mass in relation to the timing of breeding and brood size in the Great Cormorant Phalacrocorax carbo sinensis. The brood sex ratio was not related to brood size but it was significantly related to the hatching date, with a decreasing proportion of males in the brood in the course of the season. Male chicks had significantly lower body mass if they hatched later in the season, whereas there was no such relationship for female offspring. Assuming that environmental conditions deteriorate with progress of the breeding season, and male offspring may be more vulnerable to poor environmental conditions, the observed decline in the proportion of male offspring late in the season may be adaptive.     

Reproductive strategies of the larval parasitoid Microplitis croceipes

Mate choice may have important consequences for offspring sex ratio and fitness of haplodiploid insects. Mate preference of females of the solitary larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) for virgin and mated males, and vice versa, and the reproductive consequences (i.e., the sex ratio expressed as the proportion of male offspring) were examined in choice and non‐choice experiments. In addition, the effect of repeated rapid and daily copulation of an individual male on the sex ratio of offspring of the female mates was assessed. Males preferred virgins over mated females, whereas females copulated with a male irrespective of his mating status. In both the rapid and daily copulation assay, females copulating with a male that had copulated five times or more produced a higher sex ratio than females that had copulated with a virgin male. Females that copulated with virgin males once or twice produced a significantly and considerably lower sex ratio than females that first copulated with a sperm‐depleted male followed by a virgin male. This indicates that copulating with a sperm‐depleted male has costs and limits acquisition by the female of sperm from virgin males.