Seasonal clutch size decline and individual variation in the timing of breeding are related to female body condition in a non-migratory species, the Upland Goose Chloephaga picta leucoptera

Several studies have shown that waterfowl females in good condition lay larger clutches and start egg-laying earlier in a breeding season. However, most of these studies lack corresponding data on individual condition and timing of breeding in different years. We analysed data on clutch size, egg size, hatching date and female body condition of a non-migratory species, the Upland Goose Chloephaga picta leucoptera, recorded in three seasons on New Island, Falkand Islands. We found a strong seasonal decline in both egg and clutch size. Egg size increased with clutch size. The mean egg volume and total clutch volume increased with female body condition and hatching date was earlier for females in higher body condition. Chicks hatched from nests with higher mean egg volume had higher early body condition. We also compared individual females between different study years and found that individual body condition showed a significant repeatability between years, whereas hatching date did not. While female aging did not systematically affect hatching date, females produced clutches that hatched earlier in years of higher body condition. We could thus show for the first time that the frequently reported negative relationship between female condition and hatching date also holds true on an individual basis. To our knowledge, this is the first study on seasonal clutch size decline in sheldgeese and one of a few in waterfowl comparing condition and timing of breeding within individuals between different years.     

Female gigantism in a New Guinea population of the spider Nephila maculata

In many arthropods, female clutch size is positively correlated with female size but the benefits of increased size are presumed to be counterbalanced by the mortality risks incurred through delayed maturation. The orb-weaving spiders of the genus Nephila are characterized in part by large female size, and among species female size declines with increasing latitude, suggesting that one important mortality risk associated with delayed female maturation is the end of the growing season. The best-studied species inhabit strongly seasonal habitats. To explore the hypothesis that female size is limited by season length, I collected data concerning size-dependent growth and development rates, prey capture success, predation risk, and female fecundity from a population of N. maculata inhabiting a nearly aseasonal habitat. Comparisons of these data with observations from another N. maculata population and from congeners inhabiting more seasonal sites support the hypothesis that female size in this genus may be limited primarily by seasons length. Combined with recent phylogenetic evidence that male size is an ancestral characteristic, these results indicate that sexual size dimorphism in this group has evolved due to fitness advantages of delayed female maturation.     

Timing of current reproduction directly affects future reproductive output in European coots

Life-history theory suggests that the variation in the seasonal timing of reproduction within populations may be explained on the basis of individual optimization. Optimal breeding times would vary between individuals as a result of trade-offs between fitness components. The existence of such trade-offs has seldom been tested empirically. We experimentally investigated the consequences of altered timing of current reproduction for future reproductive output in the European coot (Fulica atra). First clutches of different laying date were cross-fostered between nests, and parents thereby experienced a delay or an advance in the hatching date. The probability and success of a second brood, adult survival until and reproduction in the next season were then compared to the natural variation among control pairs. Among control pairs the probability of a second brood declined with the progress of season. Delayed pairs were less likely and advanced pairs were more likely to produce a second brood. These changes were quantitatively as predicted from the natural seasonal decline. The number of eggs in the second clutch was positively related to egg number in the first clutch and negatively related to laying date. Compared to the natural variation, delayed females had more and advanced females had fewer eggs in their second clutch. The size of the second brood declined with season, but there was no significant effect of delay or advance. Local adult survival was higher following a delay and reduced following an advance. The effect of the experiment on adult survival was independent of sex. Laying date and clutch size of females breeding in the next year were not affected by treatment. The study demonstrates the existence of a trade-off between increased probability of a second brood and decreased parental survival for early breeding. Timing-dependent effects of current reproduction on future reproductive output may thus play an important role in the evolution of the seasonal timing of reproduction.     

Field evidence challenges the often‐presumed relationship between early male maturation and female‐biased sexual size dimorphism

Female‐biased sexual size dimorphism (SSD) is often considered an epiphenomenon of selection for the increased mating opportunities provided by early male maturation (i.e., protandry). Empirical evidence of the adaptive significance of protandry remains nonetheless fairly scarce. We use field data collected throughout the reproductive season of an SSD crab spider, Mecaphesa celer, to test two hypotheses: Protandry provides fitness benefits to males, leading to female‐biased SSD, or protandry is an indirect consequence of selection for small male size/large female size. Using field‐collected data, we modeled the probability of mating success for females and males according to their timing of maturation. We found that males matured earlier than females and the proportion of virgin females decreased abruptly early in the season, but unexpectedly increased afterward. Timing of female maturation was not related to clutch size, but large females tended to have more offspring than small females. Timing of female and male maturation was inversely related to size at adulthood, as early‐maturing individuals were larger than late‐maturing ones, suggesting that both sexes exhibit some plasticity in their developmental trajectories. Such plasticity indicates that protandry could co‐occur with any degree and direction of SSD. Our calculation of the probability of mating success along the season shows multiple male maturation time points with similar predicted mating success. This suggests that males follow multiple strategies with equal success, trading‐off access to virgin females with intensity of male–male competition. Our results challenge classic hypotheses linking protandry and female‐biased SSD, and emphasize the importance of directly testing the often‐assumed relationships between co‐occurring animal traits.     

Seasonal clutch size variation of multi-brooded bird species: comparisons between breeding season and latitudes

Single- and multi-brooded species of birds differ in their seasonal patterns of clutch size. Single-brooded species start with a maximum clutch size that declines continuously as the season progresses, whereas the clutch sizes of multi-brooded species usually increase to a mid-season maximum peak and then decrease progressively until the end of the breeding season. Previous studies have shown that multi-brooded migrant species present seasonal patterns that are similar to single-brooded species at high latitudes but similar to multi-brooded non-migratory species at lower latitudes. We studied the Greenfinch Carduelis chloris and Goldfinch C. carduelis populations in eastern Spain (Sagunto, Valencia) between 1975 and 2002 to compare seasonal variations in clutch sizes between years with early and late starts to the breeding season. The period over which clutch sizes increase was longer when the breeding season started earlier. The Goldfinch population showed no pattern of initial increase in clutch size when there was a late start to the breeding season: a late start shortens the season giving them less time to breed, and may also coincide with maximum food availability. Thus, the pattern of single-brooded species was observed. In the Greenfinch population, a trend toward the seasonal pattern of single-brooded species was also observed when the following indices were compared: clutch size increase, modal timing, initial slope and timing of maximum clutch size. We have also compared the seasonal patterns of clutch size of both species in eastern Spain with the patterns observed in Britain. Our results show that for both Goldfinches and Greenfinches, the non-migrant southern populations of Sagunto in eastern Spain do not tend towards a more multi-brooded seasonal pattern of clutch size than the migrant Goldfinches of Britain.     

The effect of extreme northerly climatic conditions on the life history of the minnow, Phoxinus phoxinus (L.)

Over its exceptionally wide geographical range the minnow displays a remarkable variability in its life history. In southern England many fish mature after one year and few fish survive to their third birthday. Very few fish spawned for more than one season but a 65-mm female is estimated to spawn up to 3172 eggs over the extended spawning season. In the River Utsjoki in Finnish Lapland maturity was strongly size-dependent and delayed until the fish reached 5, 6 or even 7 years of age, with a maximum age of 13 years at a length of only 75 mm. Unlike some other small-sized species in unproductive environments, individual females continued to spawn successive clutches of eggs though over a much shorter spawning season. The maximum estimated egg production was 824 for a 65-mm fish. Temperature had a strong direct or indirect effect on growth, with significantly lower growth increments in cold summers. At a second Arctic site in eastern Lapland growth rates were higher and maturity earlier, yet clutch size was greatly reduced with a seasonal fecundity estimate of only 314. There were only small differences in size of eggs or larvae between the populations. Evidence from other species indicates that most of the observed changes over the range of the minnow are phenotypic responses to the contrasting environments. However, the different strategies displayed at the two Arctic sites could represent the results of selection for differences in pre- and post-reproductive survival.     

Melatonin delays clutch initiation in a wild songbird

The hormone melatonin is known to play an important role in regulating many seasonal changes in physiology, morphology and behaviour. In birds, unlike in mammals, melatonin has thus far been thought to play little role in timing seasonal reproductive processes. This view is mainly derived from laboratory experiments on male birds. This study tests whether melatonin is capable of influencing the timing of clutch initiation in wild female songbirds. Free-living female great tits (Parus major) treated with melatonin-filled implants prior to the breeding season initiated their first clutch of the season significantly later than females carrying an empty implant. Melatonin treatment did not affect clutch size. Further, melatonin treatment did not delay the onset of daily activity in the wild nor adversely affect body mass in captivity compared with controls. These data suggest a previously unknown role for this hormone in regulating the timing of clutch initiation in the wild.     

Geographic variation of multiple paternity in the American lobster, Homarus americanus

We studied the frequency of multiple paternity for American lobster (Homarus americanus) at three Canadian sites differing in exploitation rate and mean adult size. The probability of detecting multiple paternity using four microsatellite loci and 100 eggs per female was in excess of 99% under various scenarios of paternal contribution. Overall, 13% of the 108 examined females carried a clutch sired by two or three males. Multiple paternity was observed at the two most exploited sites (11% at Magdalen Islands and 28% at Grand Manan Island), whereas single paternity only was observed at the least exploited site (Anticosti Island). Within populations females with a clutch sired by more than one male tended to be smaller than females with a clutch sired by a single male. Based on these and other findings, we postulate a link between female promiscuity and sperm limitation in the American lobster.     

Offspring size and timing of hatching determine survival and reproductive output in a lizard

Selection on offspring size and timing of birth or hatching could have important consequences for maternal investment strategies. Here we show consistent viability selection on hatchling body length across 2 consecutive years in a lizard that lays several clutches per season. There was no effect of hatching date on survival to maturity. However, both early hatching and large hatchling size increased adult size, which has a positive effect on total reproductive output. Earlier hatching also led to an earlier onset of reproduction. Overall, increased survival probability for large hatchlings and a positive effect of clutch size on recruitment suggest consistent directional selection on both egg size and clutch size within and across years. Because offspring size and timing of hatching are strongly affected by environmental and maternal effects, there should be potential for strong transgenerational effects on reproductive output in this species. We briefly discuss the implications of these results for the evolutionary ecology of maternal investment and population fluctuations in short-lived lizards.     

Age and size-dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed,Lepomis gibbosus

Synopsis In this study, we used pumpkinseed,Lepomis gibbosus, populations in two east-central Ontario lakes to test for age and size-dependent effects on the probability and timing of reproduction of mature females. Pumpkinseed body size characteristics differed in the two lakes; Little Round Lake harbours a stunted population and Beloporine Lake does not. Age-class distribution of mature females was determined by a maturity assessment on fish collected just prior to spawning and at the mid-spawning period, combined with an early-season capture-mark-recapture survey. Spawning females were collected throughout the breeding season to assess age and size-related temporal trends, and to compare their age and size distribution with that of mature females at large. The proportion of age 2 females in Little Round Lake that actually spawned was considerably lower than the proportion of mature age 2 females at large. Furthermore, age 2 females that spawned in this lake did not do so until late in the breeding season. In contrast, the proportion of young/small females spawning in Beloporine Lake was comparable to the proportion of young/small mature females at large, and both small and large females spawned throughout the breeding season. Small mature females in Little Round Lake may have had no other option but to spawn late in the season because of their poor body condition. In Beloporine Lake, condition factor early in the breeding season in age 2–4 females was higher than that of Little Round Lake females, suggesting that limited energy reserves in the spring may have prevented young Little Round Lake females from early spawning. Our results show that the likelihood and timing of reproduction are both age and size-dependent in some populations. Small individuals that delay seasonal maturation and spawn late in the summer probably contribute little to the population due to the restricted growth and reduced overwinter survival of their progeny.To whom correspondence should be addressed     

Maturation increases early reproductive investment in Adélie Penguins Pygoscelis adeliae

Correlations between female investment in egg production and age, breeding experience and laying date have been reported in several seabird species. In general, clutch and egg sizes increase with female age and breeding experience but decrease with laying date. Positive correlations of clutch and egg size with age and breeding experience can be caused by an increase in reproductive investment with maturation or they may be an artefact of lower survival rates for individuals with poor-quality phenotypes. Negative correlations of clutch and egg size with laying date might signal an adaptive reduction in egg production or be due in part to variation among individuals. We examined the interactions of female age, breeding experience, laying date and clutch and egg size in Adélie Penguins Pygoscelis adeliae . Breeding experience strongly affected clutch size with 87.3% of all one-egg clutches laid by first-time breeders. In addition, increasing age had a positive influence on egg size and was associated with earlier laying dates. However, there was little evidence to suggest that either clutch or egg sizes are influenced by laying date. Laying dates and clutch and egg sizes did not affect a female's probability of returning to breed in the following year, indicating that increased investment is a product of maturation and not of the loss of poor-quality breeders from the population. Our results suggest that as female Adélie Penguins gain foraging and breeding experience they are able to initiate breeding earlier, to lay complete clutches of two eggs and to lay larger eggs.     

Factors influencing double brooding in Eurasian Hoopoes Upupa epops

Double brooding may be a good strategy for short‐lived species to maximize annual and lifetime reproductive success (ARS and LRS, respectively). Nevertheless, there is typically individual variation in the probability of producing a second clutch. Here we evaluate factors that influence the decision to double brood in the Eurasian Hoopoe Upupa epops. Analyses of an 11‐year dataset showed that 36% of the females and 21% of the males produced a second clutch after successfully raising a first clutch. Double‐brooded females had higher ARS (9.1 ± 1.9 fledglings; mean ± se) and LRS (0.93 ± 0.08 recruits) than single‐brooded females (ARS: 4.5 ± 2.1 fledglings; LRS: 0.36 ± 0.03 recruits). This suggests that double brooding is adaptive in Hoopoes, and raises the question of why most individuals only produce one clutch per season. The probability of double brooding varied only slightly between years, suggesting that it is influenced by individual characteristics rather than by external, population‐level environmental factors. In both sexes, the probability of double brooding increased with earlier timing of the first clutch, and the timing of reproduction was the most important factor influencing reproductive success. The latter is likely to be mediated by changes in resources during the season. The probability of double brooding also increased slightly with female age, due to differences in intrinsic quality among females rather than to a gain in experience. In contrast to many other studies, the probability of double brooding increased with an increasing number of fledglings from the first clutch, suggesting that it is a strategy of individuals of high quality. Taken together, we show that the individual quality of the breeder and the timing of their first clutches are key factors influencing the decision to double brood, and thereby that they are important determinants of reproductive performance in Eurasian Hoopoes.     

Intra-seasonal decline of clutch size in Lesser Snow Geese

Summary Different mechanisms proposed to explain the intra-seasonal decline in clutch size of Lesser Snow Geese (Anser caerulescens caerulescens) were tested at the La Perouse Bay colony, Manitoba, Canada. Ovary examination of females collected after laying revealed that the actual number of eggs produced per female decreased over the laying period. This finding eliminates nest-parasitism, partial clutch predation and renesting attempts as sufficient explanations for seasonal clutch size decline. Follicular atresia induced by reserve depletion was also rejected since its occurrence was similar among early and late nesters. The decline in clutch size was observed within age-classes, and therefore age effects on clutch size and laying date per se do not account for the observed relationship. Clutch size and laying date were respectively positively and negatively correlated with the amount of nutrient reserves in females at the onset of laying, and also covaried within individuals observed breeding in several seasons. Laying date repeatability was estimated at 0.22. It is postulated that the seasonal decline in clutch size results mainly from a positive feed-back of female's nutrient reserves on the hormones controlling ovary development. Hatching synchrony or shortness of the breeding season may be the ultimate factors responsible for the intra-seasonal clutch size decline in Lesser Snow Geese.     

Timing of maturation and the mating system of the spider,Stegodyphus lineatus (Eresidae): how important is body size?

Selection on life history traits such as the timing of maturation and the size at maturity strongly depends on the mating system. In spiders, the mating system hypothesized to Be determined by spermathecal morphology and the related sperm precedence pattern. In a natural population of the eresid spider Stegodyphus lineatus , predictions concerning the timing of maturation, male mating behaviour and success were tested. Eresid spiders are supposed to show protandry, prematuration mate guarding and strong male-male competition resulting in selection for large body size and early maturation. In contrast to these predictions, male and female maturation overlapped largely. Males did not guard premature females nor was there evidence for male-male competition. Among mating pairs, male did not relate to female size, nor to duration of cohabitation. Evidence for an advantage of first over second or large over small males is weak. In males, body size at maturity and the time of maturation were negatively correlated although a trade off between timing of maturation and the body size reached by then should result in a positive correlation. Possible causes are discussed.     

中国石龙子雌体繁殖特征和卵孵化的地理变异

浙江丽水和广东韶关中国石龙子均年产单窝卵,窝卵数,窝卵重和卵重均与雌体SVL呈正相关,雌体头部形态,繁殖特征,产卵起始时间和孵孵化的热依赖性等有显著的地理变异;韶关石龙子产卵起始时间为5月中旬,比丽水经子约早两周,韶关石龙子窝卵数较大,卵较小,窝卵重与丽水石龙子无显著差异。韶关石龙子特定SVL的窝卵数比丽水石龙子多2．8枚卵,中国经子卵数量和大小之间有种群间权衡,无种数内权衡,同一种群内卵数量与卵大小无关,孵化温度影响石龙子孵出幼体的一些特征,24℃孵出细幼体比32℃孵出幼体大,躯干发育好,剩余卵黄少,韶关24℃孵出幼体的体重,躯干干重小于丽水幼体,韶关32℃孵出幼体的SVL小于丽水幼体,剩余卵黄大于丽水幼体,表明适宜卵孵化温度范围有地理变异。丽水石龙子卵对极端高温和低温的耐受性较强,适宜卵孵化温度范围较宽。     

The effects of rainfall on different components of seasonal fecundity in a tropical forest passerine

Seasonal fecundity is a composite metric that is determined by component parameters such as clutch size, nest survival and re‐nesting probability. Many of these component parameters are known to vary with environmental conditions, in particular rainfall prior to or during the breeding season. In some species, seasonal fecundity is positively related to rainfall, but little is known about which component parameters of seasonal fecundity respond most strongly to rainfall. We used intensive nest monitoring of a multi‐brooded tropical forest passerine, the Montserrat Oriole Icterus oberi, to examine the effects of rainfall during the pre‐breeding season on component parameters of annual fecundity. We monitored all nests of a total of 42 pairs over 5 years in which rainfall varied substantially. We then related clutch size, nest survival, onset and length of the breeding season, re‐nesting probability and re‐nesting interval to pre‐breeding season rainfall using generalized linear mixed models that accounted for random variation across sites and individual pairs, and incorporated other variables known to affect the response. Higher pre‐breeding season rainfall led to an increase in clutch size and a decrease in re‐nesting interval, but nest survival, re‐nesting probability and length of the breeding season were not affected by variation in rainfall. The onset of the breeding season was delayed in very dry years. We conclude that higher rainfall is likely to increase food availability and thus body condition of female Montserrat Orioles, leading to an increase in fecundity due to larger clutch sizes.     

Maturation of walleye by age, size and surplus energy

The probability of annual sexual maturation by male and female walleye Stizostedion vitreum was related to age, size and an index of condition, I _VF=[arcsine(visceral fat)^0·5(body mass)^−0·5]. Most males first matured at ages 2 and 3 years; size explained first maturation, but condition explained later maturation. In contrast, most females first matured at ages 4 and 5 years; maturity of females was more dependent upon condition. Maturity of females at ages 4 and 5 years was significantly correlated with average I _VF of the population ( PI _VF). The size reached by age 2 years (early growth) was correlated with the PI _VF. Growing degree-days, Secchi depth, latitude and lake morphology were not correlated with the PI _VF. Annual variations in female spawning stock size were related to the condition of the females, presumably reflecting the net acquisition of energy in the preceding growing season. Annual variations within lakes in the net acquisition of energy may exceed the variations in energy availability between lakes, dictated by lake morphology and geography. Thus, assessment of condition could be used to predict annual potential spawning stock size and egg production.     

Maternal investment in reproduction and its consequences in leatherback turtles

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.     

Female mating success and risk of pre-reproductive death in a protandrous grasshopper

Numerous studies have assessed the adaptive value of protandry for males in several insect species, considering that male emergence is determined by female availability. However, the possible advantage of the time of emergence for females on their mating success in protandrous insect species has only been explored theoretically. By studying the grasshopper Sphenarium purpurascens we evaluated the hypothesis that late emergence could be adaptive for females. If female maturation occurs when the population density is higher and the sex ratio (males/females) is biased to males, their probability of mating increases. Thus, in this study we estimated (1) the opportunity for mating in females as a function of their sexual maturation time, population density, and sex ratio at the moment they reached sexual maturity. In addition, (2) an analysis incorporating female body size and the total number of female matings was performed. Both analyses support the occurrence of protandry in the studied population. Under the first approach, females with intermediate maturation time had a higher probability of being mated than earlier and late matured females. Thus, it suggests that stabilising selection is acting on female maturation time and this may affect selection on male maturation time. Furthermore, the proportion of mated females increased when the sex ratio was biased to males, and stabilising selection on maturation time was detected also. However, the number of matings of a female depended on her body size. Females with larger body size had more matings than smaller ones at the beginning of the reproductive season. Because selection acts differently on maturation time in males and females of S. purpurascens this result is consistent with a condition for the maintenance of protandry in the population. The present results are discussed in the light of the models for the evolution of protandry.