Sexual conflict over parental care in Penduline Tits Remiz pendulinus: the process of clutch desertion

Do the two parents at a nest make simultaneous decisions whether to care for their offspring or to desert? If a single parent is sufficient for rearing young, one parent (typically, the male) may desert and reproduce with a new mate within the same breeding season, leaving the other parent with the brunt of care. As each parent is expected to maximize its own reproductive success, the interests of the two parents do not necessarily coincide, and a sexual conflict over care may emerge. Here we investigate the process of clutch desertion in a small passerine bird, the Penduline Tit Remiz pendulinus. Among birds, this species has a remarkably variable breeding system, because a single parent (either the male or the female) may provide the full care of the young, whereas about 30% of clutches are abandoned by both parents. First, we show that biparental desertion occurs within a single day in 73.7% of the clutches (n = 14), whereas desertion decisions are sequential in 26.3% of the clutches (n = 5) (male first: 10.5% (n = 2); female first: 15.8% (n = 3); n = 19 clutches in total). Secondly, we observed the behaviour of both parents before desertion, and investigated whether desertion can be predicted from their behaviour. However, neither singing nor nest‐building behaviour predicted whether the male or the female would desert. We therefore suggest that biparental desertion may be simultaneous by male and female in our population of Penduline Tits. Furthermore, the parents do not appear to signal their intention to desert their mate. We argue that the parents’ interest may be actually to disguise their intention to desert.     

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.     

Reproductive parameters of a captive colony of capuchin monkeys (<Emphasis Type="Italic">Cebus apella</Emphasis>) from 1984 to 2006

This paper presents the results of a demographic analysis of 22 years of data recorded on a colony of tufted capuchin monkeys (Cebus apella) in captivity at the CNR Primate Centre (Rome, Italy). Information is provided on reproduction, sex ratio, inter-birth interval (IBI), seasonality, and body weight. From 1984 to 2006, 46 live births were recorded. There were births in almost all months of the year, but a higher frequency was observed during spring and summer (71.1%). The sex ratio was 1:1 M:F for newborns and 1:1.06 M:F for surviving offspring. At birth, infants’ average weight was 238.13 ± 37.51 g, i.e. 250 ± 56.79 g for males and 231 ± 26.08 g for females. Age at first birth for females ranged from 4.9 to 7 years (n = 9), while males achieved first paternity between the ages of 5 and 9.2 years (n = 6). Only one pair of twins was recorded during this period. For females, the mean IBI was 17.88 ± 1.84 months, when they reared infants, and 12.70 ± 1.73 months, when they did not rear offspring. Infant mortality within the first 2 months was 28.3%.     

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.     

Sex-biases in the hatching sequence of cooperatively breeding apostlebirds <Emphasis Type="Italic">Struthidea cinerea</Emphasis>

In the cooperatively breeding apostlebird (Struthidea cinerea, Corcoracidae) both sexes are philopatric and help to raise offspring. However, male helpers provision nestlings more often than females, an activity associated with reduced nestling starvation and enhanced fledgling production. Presuming that males are the more helpful sex, we examined the helper repayment hypothesis by testing the predictions that offspring sex ratio should be skewed toward the production of males (a) among breeding groups with relatively few helpers, and (b) in the population as a whole. The relationship between sex and hatching order was examined as a potential mechanism of biasing sex allocation. The sex ratio of all sexed offspring was male biased (57.9%; n = 171) as was the mean brood sex ratio (0.579; n = 70 broods). These biases were less pronounced in the subset of clutches/broods in which all offspring were sexed. This overall bias appeared to result from two distinct patterns of skew in the hatching order. First, mothers in small breeding groups produced significantly more males among the first-hatching pair. This is consistent with the helper repayment hypothesis given that later hatching chicks were less likely to survive, particularly in small groups. Second, almost all fourth-hatching chicks, usually the last in the brood, were male (91.7%, n = 12). This bias is difficult to interpret but demonstrates the value of examining hatching sequences when evaluating specific predictions of sex allocation theory in birds.     

Breeding success in Blue Tits: good territories or good parents?

Territorial quality and parental quality are usually assumed to be the main sources of variation in the reproductive success of passerine birds. To evaluate their relative importance for variation in breeding time (itself an important factor for breeding success), clutch size and offspring condition at fledging, we analysed six years of data from a Blue Tit Parus caeruleus population breeding on the island of Gotland, Sweden. Hatching dates and the condition of offspring were consistent for territories between years and accounted for 30% and 33% of the variation in these variables, respectively. After removing the effect of territory quality, none of the breeding parameters were significantly repeatable for individual females, but offspring condition was repeatable for males, accounting for 28% of variation. For females breeding on the same territory in subsequent seasons (combined effect of individual and territory quality) only hatching date was repeatable (45% of variation accounted for). In males, the combined effect of individual and territory quality was repeatable for offspring condition and accounted for 33% of variation, but this result was only marginally significant. Consistency of the peak frass-fall date for individual frass collectors over the study period suggests that repeatable hatching dates on territories may be related to the relationship between timing of breeding and timing of peak food availability on territories. Our results suggest that territory quality is more important than parental quality for breeding success in the Blue Tit, and that male (but not female) quality makes a considerable contribution to reproductive success.     

Ecological and phenological covariates of offspring sex ratio in barn swallows

Non-random sex allocation in relation to parental, ecological and phenological factors has been investigated in several correlational studies of birds, mostly based on few breeding seasons and relatively small sample sizes, which have led to different results. We investigated sex ratio of nestling barn swallows (Hirundo rustica) in relation to adult sex ratio, laying date, clutch size, colony size and meteorological conditions in a sample of 553 broods (>2200 nestlings) during 10 years. At the population level, nestling sex ratio varied among years and deviated from parity in two years. Sex ratio among adults did not predict offspring sex ratio in the current or the following year. At the within-family level, the proportion of sons increased with laying date in large clutches, did not vary among clutches of intermediate size, and tended to decline with laying date in small clutches. Large colonies harbored more sons. The proportion of males increased with temperature during laying whereas the effects of temperature during the pre- or post-laying periods and that of rainfall were non-significant. These patterns of variation of offspring sex ratio did not differ between years. Thus, we identified several potential causal sources of variation in barn swallow offspring sex ratio, including temporal, phenological and ecological factors. The observation of an association of offspring sex with temperature during laying is novel for birds and may be mediated by effects on maternal steroid hormones profile. The ecological and evolutionary implications of present findings are discussed in the light of adaptive sex allocation theory.     

Does the differential cost of sons and daughters lead to sex ratio adjustment in great frigatebirds Fregataminor?

Sex allocation theory predicts that if benefits of producing sons and daughters differ and outweigh the costs of sex ratio adjustment, parents should produce more of the offspring that provide them with greater fitness. Potential benefits may be more likely to outweigh costs where sexual size dimorphism and, in birds, single‐egg clutches exist. Great frigatebirds Fregataminor are seabirds in which females are larger than males and clutch size is one egg. In our study population, sexual size dimorphism develops primarily during the period of complete juvenile dependence on parental care, consistent with a higher cost of producing daughters than sons. Over the course of the 1998 breeding season there was a shift from early season prevalence of daughters to late‐season prevalence of sons. Variation in food availability at time of egg laying, as indexed by sea surface temperature (SST), was a strong predictor of offspring sex in 1998. In contrast, SST in 2003 was not a predictor of offspring sex, nor was there a seasonal shift in the hatching sex ratio, despite a seasonal shift in SST. Besides food availability, we tested two additional factors in 2003 that could explain sex ratio adjustment in relation to the cost of reproduction. Offspring sex in 2003 was not related to natural or experimentally induced variation in maternal body condition; pre‐laying food supplements raised the body condition of females at the time of egg laying but did not affect offspring sex or egg mass. In addition, offspring sex was not predicted by the length of maternal telomere restriction fragments (TRFs), an index of age and possibly of reproductive experience. Broad confidence intervals on effect size suggest that undetected effects of maternal condition on offspring sex ratio could easily exist, but confidence intervals were narrower on the non‐significant effects of SST and TRF length on offspring sex ratio. The cause of different seasonal patterns of hatching sex ratio and different SST effects in 1998 and 2003 is unclear.     

Adaptive seasonal trend in brood sex ratio: test in two sister species with contrasting breeding systems

Evolutionary theory predicts adaptive adjustment in offspring sex ratio by females. Seasonal change in sex ratio is one possibility, tested here in two sister species, the Common sandpiper and the Spotted sandpiper Actitis hypoleucos and A. macularia. In the monogamous Common sandpiper, males are the most competitive sex. In each of 3 years, there was a change from mainly sons in early clutches to mainly daughters in late clutches. This seasonal adjustment of clutch sex ratio took place within the female before the eggs were laid, not by differential egg or chick survival. The sex of all eggs laid in the clutches used here was determined molecularly from chick blood taken at the time of hatching. The Spotted sandpiper in contrast is polyandrous, with partly reversed sex roles. There was no seasonal trend from sons to daughters in this species. When tested together, the two species differed significantly as predicted by the hypothesis of adaptive sex ratio adjustment by females.     

Southern Lapwing Vanellus chilensis cooperative helpers at nests are older siblings

Cooperative breeding is rare in shorebirds, and when found it is thought to be due to polygamous mating (cooperative polygamy). Here we describe the social structure of cooperatively breeding groups in Southern Lapwing Vanellus chilensis and test the prediction that offspring sex ratio is skewed towards the sex that helps. The social groups consisted of a breeding pair with one or two young (mostly males, 68%) from the previous breeding season, and offspring sex ratio was not skewed. Cooperative breeding in the Southern Lapwing is not the consequence of cooperative polygamy, but rather groups were formed by a mated pair and some of their offspring from previous nests as helpers.     

Males paving the road to polyandry? Parental compensation in a monogamous nesting cuckoo and a classical polyandrous relative

Comparative studies have established the necessity for biparental care as an important factor for monogamy in freshwater fish and birds. However, whether two parents are really needed for offspring care remains an open question in many cases. I experimentally studied female and male contributions to offspring care in the white-browed coucal (Centropus superciliosus), a monogamous and biparental cuckoo with a balanced adult sex ratio, and contrasted it with the sympatric black coucal (C. grillii), a classically polyandrous species with a male-biased adult sex ratio and male-only care. To study the necessity for biparental care, I temporarily removed one partner for 2 days to see whether the remaining parent compensated for the absence of its partner. Both female and male white-browed coucals approximately doubled their feeding rates when their partner was absent, thus fully compensating the number of feeding visits to the nest. However, nestlings maintained their growth only, when males were present and females were removed. When males were removed and only females were present, nestling growth declined. Hence, only male white-browed coucals fully compensated for the temporary loss of the partner, suggesting that females could benefit most from nesting with additional males—if these should become available. Removing female black coucals had no consequence for nestling feeding rates of male black coucals. But male black coucals had to be returned to their territories within a few hours to avoid harming the brood because female black coucals typically would not commence feeding their offspring. In conclusion, the breeding system of white-browed coucals seems quite flexible and the relatively balanced adult sex ratio may stabilize monogamy in this species. Should ecological factors ever favour a stronger bias in the adult sex ratio towards males, female white-browed coucals may easily become polyandrous and relinquish parental care entirely to males.     

The dynamics of male harem dominance in southern elephant seals (Mirounga leonina) at the South Shetland Islands

Breeding chronology, harem structure and changes in male harem dominance were studied at Stranger Point, Isla 25 de Mayo/King George Island, principally by extensive field census work during the 2003 breeding season. Males were individually identified and their size estimated by using a photogrammetric method. Peak female haul out for the population occurred on 31 October, when a total of 276 females were observed along 7 km of coastline, distributed in ten harems with a median size of 16 females. Overall sex ratio and harem sex ratio for the breeding population were 1:6.7 and 1:10.6, respectively. A total of 33 males were identified associated with harems. Male size conferred an advantage in terms of dominance hierarchy, since dominant males (4.91±0.15 m) were significantly longer than subordinate males (4.63±0.19 m). Harems were dominated by an average of 4.5 (range 2–7) different males during the breeding season. Elephant seals at Stranger Point breed in very low density aggregations. The main breeding events in this population occurred later than at other breeding sites, which agrees with previous observations in the area. Male movement among harems suggests that differences in mating success among males could be achieved through their different behaviours.     

Mate choice and cannibalism in a natural population of a stream goby,Rhinogobius sp.

Breeding ecology of the stream goby,Rhinogobius sp. LD (Large Dark), was investigated under natural conditions. Males selectively courted females of similar size to lead them to the nests, whereas females followed courting males preferentially when the relative male size was greater. Male-male competition for a female was relatively infrequent and not severe. Developmental stages of eggs and egg numbers in one nest indicated that males receive 1–3 clutches during one breeding cycle. Males guarding multiple clutches frequently ate some of the eggs, but those guarding single clutches rarely did so. Gravid females in the nest also frequently cannibalized eggs laid by a previous female, probably in order to extend the area available for egg deposition. Mate choice in this species is discussed in relation to paternal ability, limitation of available spawning area and the female-biased sex ratio.     

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.     

Temperature-dependent development of the predatory mite <Emphasis Type="Italic">Cheyletus malaccensis</Emphasis> (Acari: Cheyletidae)

The effect of temperature on the development of immature stages of the predator Cheyletus malaccensis Oudemans, produced by either fertilized or virgin females, was studied at 17.5, 20, 25, 30, 32.5, and 35°C, 80 ± 5% relative humidity, in complete darkness, and fed Tyrophagus putrescentiae (Schrank). The population maintained at 15°C failed to reproduce. The thermal data obtained were used for the estimation of the thermal requirements (developmental thresholds, thermal constant, optimum temperature) of this predator by a linear and nonlinear model (Logan type I model). Upper and lower developmental thresholds ranged between 37.4–37.8 and 11.6–12.0°C, respectively. Optimum temperature for development was estimated at between 33.1 and 33.5°C. The thermal constant ranged between 238.1 and 312.5 degree-days. Based on the data of the total pre-imaginal period, immatures’ survival peaked at 25°C. Arrhenotokous parthenogenesis (haplodiploidy) is confirmed in the species: virgin females always produce males, whereas fertilized females give rise to offspring of both sexes. Survival of the immature stages and offspring sex ratio were not significantly influenced by temperature.     

Correlations between ultraviolet coloration, overwinter survival and offspring sex ratio in the blue tit

We studied the correlations between offspring sex ratio, UV coloration and overwinter survival in a population of blue tits, breeding in Gotland, Sweden, over three consecutive breeding seasons. In 2 of 3 years, we found that females paired to males with relatively brighter UV-coloration produced a greater proportion of sons in their broods, and that this effect was significant with all 3 years combined, despite a significant year by male UV interaction. In addition, we found other correlates of sex ratio (breeding time, female age and clutch size) in some, but not all years, and some of these showed significantly different relationships with sex ratio between years. In both years for which data were available, there were indications that males with relatively brighter UV coloration, and that paired with females that produced male-biased clutches, were more likely to survive to the next year. In addition, we also found that in both males and females, individuals produced similar sex ratios in consecutive years. Because correlations with the sex ratio may be expected to be weak, variation in results between years within the same population may be explained by low statistical power or genuine biological differences. Our results suggest that conclusions about sex ratio variation in birds should be based on multiple years. The correlations that we found in some years of this study are consistent with models of adaptive sex ratio adjustment in response to mate quality. However, careful experimental work is required to provide tests of the assumptions of these models, and should be a priority for future work.     

Prolonged and flexible primary moult overlaps extensively with breeding in beach‐nesting Hooded Plovers Thinornis rubricollis

We present the first report of complete overlap of breeding and moult in a shorebird. In southeastern Australia, Hooded Plovers Thinornis rubricollis spend their entire lives on oceanic beaches, where they exhibit biparental care. Population moult encompassed the 6‐month breeding season. Moult timing was estimated using the Underhill–Zucchini method for Type 2 data with a power transformation to accommodate sexual differences in rates of moult progression in the early and late stages of moult. Average moult durations were long in females (170.3 ± 14.2 days), and even longer in males (210.3 ± 13.5 days). Breeding status was known for most birds in our samples, and many active breeders (especially males) were also growing primaries. Females delayed the onset of primary moult but were able to increase the speed of moult and continue breeding, completing moult at about the same time as males. The mechanism by which this was achieved appeared to be flexibility in moult sequence. All moult formulae fell on one of two linked moult sequences, one faster than the other. The slower sequence had fewer feathers growing concurrently and also had formulae indicating suspended moults. Switching between sequences via common formulae is possible at many points during the moult cycle, and three of 12 recaptures were confirmed to have switched sequences in the same moult season. Hooded Plovers thus have a prolonged primary moult with the flexibility to change their rate of moult; this may facilitate high levels of replacement clutches that are associated with passive nest defence and low reproductive success.     

Causes and consequences of adaptive seasonal sex ratio variation in house sparrows

1. Here we examine how sex ratio variation in house sparrow broods interacts with other demographic traits and parental characteristics to improve the understanding of adaptive significance and demographic effects on variation in sex ratio. 2. The sex ratio in complete broods did not deviate significantly from parity (54.9% males). 3. There was sex-specific seasonal variation in the probability of recruitment. Male nestlings that hatched late in the breeding season had larger probability of surviving than early hatched males. 4. An adaptive adjustment of sex ratio should favour production of an excess of males late in the breeding season. Accordingly, the proportion of male offspring increased throughout the breeding season. 5. A significant nonlinear relationship was present between sex ratio and age of the female. However, there was no relationship between parental phenotype and standardized hatch day that could explain the observed seasonal change in sex ratio. 6. The sex-specific number of offspring recruited by a pair to subsequent generations was closely related to the brood sex ratio. 7. These results indicate an adaptive adjustment of sex ratio to seasonal variation in environmental conditions that affects the offspring fitness of the two sexes differently. Our results also suggest that such a sex ratio variation can strongly influence the demography and structural composition of small passerine populations.