Increased survival and breeding performance of double breeders in little penguins Eudyptula minor,New Zealand: evidence for individual bird quality?

The little penguin Eudyptula minor is unique among penguin species in being able to fledge chicks from two clutches in one breeding season. Pairs laying two clutches in a given season make a higher reproductive investment, and may be rewarded by a higher reproductive success as they may raise twice as many chicks as pairs laying one clutch. The higher effort made by pairs laying two clutches could correlate negatively with survival, future reproductive performance or offspring survival, indicating a cost of reproduction. Conversely, a positive relationship between the number of clutches produced in a given breeding season and survival, future reproductive performance or offspring survival would indicate that birds laying two clutches belonged to a category of birds with higher fitness, compared to birds laying only one clutch in the season. In this study we used a long‐term data set taken from an increasing population of little penguins in Otago, SE New Zealand. We modelled the relationship between the number of clutches laid in a breeding season and survival probability, reproductive performance in the next breeding season and first year survival of offspring using capture‐recapture modelling.
Birds laying two clutches produced 1.7 times more fledglings during a breeding season than pairs laying one clutch. We found that birds laying two clutches had a higher probability of breeding in the following breeding season, a higher probability of laying two clutches in the following breeding season and a higher survival probability. There was no overall difference in post‐fledging survival between the young of birds producing one clutch and the young of birds producing two clutches. However, the survival of young of single clutch breeders declined with laying date, whereas the young of double clutch breeders had the same survival rate irrespective of laying date. For a subset of data with birds of known age, we found evidence that the probability of laying two clutches increased with age. However, there were also indications for differences among birds in the tendency to lay two clutches that could not be attributed to age. We tentatively interpret our results as evidence of quality difference among little penguin breeders.     

Effects of pair migratory behavior on breeding phenology and success in a partially migratory shorebird population

In migratory systems, variation in individual phenology can arise through differences in individual migratory behaviors, and this may be particularly apparent in partial migrant systems, where migrant and resident individuals are present within the same population. Links between breeding phenology and migratory behavior or success are generally investigated at the individual level. However, for breeding phenology in particular, the migratory behaviors of each member of the pair may need to be considered simultaneously, as breeding phenology will likely be constrained by timing of the pair member that arrives last, and carryover effects on breeding success may vary depending on whether pair members share the same migratory behavior or not. We used tracking of marked individuals and monitoring of breeding success from a partially migrant population of Eurasian oystercatchers (Haematopus ostralegus) breeding in Iceland to test whether (a) breeding phenology varied with pair migratory behavior; (b) within‐pair consistency in timing of laying differed among pair migratory behaviors; and (c) reproductive performance varied with pair migratory behavior, timing of laying, and year. We found that annual variation in timing of laying differed among pair migratory behaviors, with resident pairs being more consistent than migrant and mixed pairs, and migrant/mixed pairs breeding earlier than residents in most years but later in one (unusually cold) year. Pairs that laid early were more likely to replace their clutch after nest loss, had higher productivity and higher fledging success, independent of pair migratory behavior. Our study suggests that the links between individual migratory behavior and reproductive success can vary over time and, to a much lesser extent, with mate migratory behavior and can be mediated by differences in laying dates. Understanding these cascading effects of pair phenology on breeding success is likely to be key to predicting the impact of changing environmental conditions on migratory species.     

Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Parental age and reproduction in the Marsh Tit Paras palustris

In the Marsh Tit Parus palustris , breeding performance improved with the age of both the male and the female parent—older birds produced larger clutches earlier in the season. The effect of male age, however, was mainly due to a positive association with the age of the female. When the effect of laying date was accounted for, there was no additional effect of female age on clutch-size. However, pairs with previous breeding experience laid larger clutches than newly formed pairs of the same age. There was some evidence that young females that laid late in the season were less likely to survive to breed again, leading to an age effect on reproductive performance. There was no evidence that individual birds laid earlier or produced larger clutches when they became older.     

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

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

The effect of parental age,experience and historical reproductive success on wandering albatross (Diomedea exulans) chick growth and survival

Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.     

The effect of mate change and new partner's age on reproductive success in the barnacle goose, Branta leucopsis

Mate retention frequencies and correlations between mate changeand reproductive performance were estimated in a populationof barnacle geese, Branta leucopsis, breeding on the islandof Gotland in the Baltic. About 90% of the pairs remained togetherfrom one breeding season to the next. Only 2.4% of the pairsdivorced, most mate changes being consequences of the deathof one partner. Divorces were not forecast by low reproductivesuccess, and seemed to be accidental. In the season before matechange, there was no difference in reproductive performance,measured as clutch size, hatching date, and number of fledgedyoung, between faithful pairs and pairs where one partner wassubsequently changed. However, in the first season with a newpartner, clutch size and number of fledged young decreased onaverage. Hence, because mate change led to a reduction in reproductivesuccess, it was concluded that mate retention is advantageous.Our results suggest that this reduction is more likely due tothe lower average age or breeding experience of new partnersthan to the benefits of breeding experience with one particularpartner. [Behav Ecol 1991 ;2:116–122]     

Is Divorce in Young Common Terns,Sterna hirundo,after Recruitment just a Question of Timing?

The probability of divorce in birds has been linked with age, breeding experience, reproductive output and synchrony in return. Here, we investigate the consequences of first breeding attempts in common terns for mating in the subsequent season. Nearly 20% of all first‐time breeders disappeared or skipped at least one season after recruitment. In 84 pairs, which consisted of at least one recruit and of which both partners returned to the colony, the divorce rate was 45%. We compared reproductive success, arrival dates, and asynchrony in arrival dates of pairs of the first breeding season against the second season, for both reunited and divorced pairs and males and females separately. First, in pairs of which both members came back to the colony, we found an increase of reproductive success most pronounced in males. In the second season reproductive success of divorced compared with reunited pairs was higher, as only divorced pairs significantly improved the number of fledglings, and again this relation was stronger in males. Secondly, females of reunited pairs arrived significantly earlier from the first to the second season and by far more days than their males. However, in divorced pairs former mates did not differ in the number of days they advance their arrivals. Finally, divorced males arrived on average 4 d earlier than their former mates, whereas divorced females arrived 5 d later compared with their former mates of the recruitment season. Contradictory to nearly all other divorce studies in birds so far, we found a clear fitness gain in divorced males. We suggest that the improvement in reproductive success of young males stems from a side‐effect of the birds’ quality and ability to reach the breeding site in appropriate time and earlier as potential competitors. In long‐lived bird species the heterogeneity among young individuals in the timing of arrival at the colony seems to explain why former recruit‐pairs reunite or split. For young males we suggest as best explanation of divorce that they profit from ‘pushing for an empty chair’, while females seem to profit from their choosiness and may actively decide between former and other mates.     

Differential maternal investment counteracts for late breeding in magpies Pica pica: an experimental study

Reproductive success in many avian populations declines throughout the breeding season. Two hypotheses have gained attention to explain such a decline: the "timing" hypothesis proposes that deteriorating food availability causes the decline in reproductive success (causal effect of breeding time), whereas the "quality" hypothesis proposes that individuals of lower phenotypic quality reproduce at the end of the breeding season, causing the correlation between breeding time and breeding success. We tested both of these hypotheses in a monogamous single breeder, the magpie Pica pica , by experimentally inducing some pairs to lay a replacement clutch, after removal of the first clutch. The first clutch was left in the nest of another magpie pair (matched by laying date and clutch size), and incubated and raised by these foster parents. In this way we obtained two clutches from the same magpie pair with full siblings raised in conditions of the first and second reproductive attempts. High quality pairs (with laying dates in the first half of the breeding season) reached similar breeding success in replacement clutches as compared to first clutches of the same female. In addition, experimental pairs reared significantly more offspring of similar quality in their replacement clutches as compared to late-season first clutches, thereby suggesting that late season first clutches were produced by pairs of lower phenotypic quality. Results indicate that high quality pairs trade-off clutch size for larger eggs in replacement clutches, which could help magpie pairs to partly compensate for poorer environmental conditions associated with a delayed breeding attempt.     

Endogenous gonadal,LH and molt rhythms in tropical stonechats: effect of pair bond on period,amplitude, and pattern of circannual cycles

To investigate the effects of reproduction and associated stimuli on the circannual cycles of African stonechats Saxicola torquata axillaris birds were held for 29 months in aviaries under a constant equatorial (12.25 h) photoperiod, either singly (10 females and 10 males) or in 10 male/female pairs. The birds of all 3 groups went through circannual cycles in gonadal size, plasma LH and molt, but groups differed with regard to actual reproductive performance. During the second cycle, only one of the singly-held females laid eggs and incubated. In constrast, in the paired females egg-laying and incubation occurred in all but one bird. About 50% of the clutches from paired females contained fertilized eggs confirming the expectation of behavioral differences between the paired and unpaired birds. However, despite differences in reproductive performance there were no differences in either circannual period or duration of reproductive phases. Moreover, there was no correlation between number of broods produced per season and circannual parameters of the paired females. Therefore, the temporal course and, particularly, the period during which reproduction is possible is rigidly determined by an endogenous program that is not influenced by reproductive performance. A rigid program of this kind may be advantageous in the tropics because it prevents prolongation of the breeding season in years with favourable conditions which in turn could jeopardize optimal timing of breeding in the following year and thus reduce lifetime reproductive success.     

Delayed female reproduction in equilibrium and chaotic populations

Behavioural and life history polymorphisms are often observed in animal populations. We analyse the timing of maturation and reproduction in risky and resource-limited environments. Field and laboratory evidence suggests that female voles and mice, for example, can adjust their breeding according to the level of risk to their own survival and to survival probabilities and recruitment of young produced under different environmental conditions. Under risky or harsh conditions breeding can be postponed until later in the current breeding season or even to the next breeding season. We develop a population dynamics and life history model for polymorphism in reproduction (co-existence of breeding and non-breeding behaviours) of females in an age-structured population, with two temporally distinct mating events within the breeding season. We assume that, after overwintering, the females can breed in spring and again in summer or they can delay breeding in spring and breed in summer only. Young females born in spring can either mature and breed in summer or stay immature and postpone breeding over the winter to the next breeding season. We show that an evolutionarily stable breeding strategy is either an age-structured combination of pure breeding behaviours (old females breed and young delay maturity) or a mixed breeding behaviour within age-classes (a fraction of females breed and the rest of the age class postpones breeding). Co-occurrence of mixed reproductive behaviour in spring and summer within a single breeding season is observed in fluctuating populations only. The reproductive patterns depend on intraspecific, possibly interspecific, and ecological factors. The density dependence (e.g. social suppression) and predation risk are shown to be possible evolutionary mechanisms in adjusting the relative proportions of the different but co-existing reproductive behaviours.     

Seasonal decline in provisioning effort and nestling mass of Blue Tits Cyanistes caeruleus: experimental support for the parent quality hypothesis

A common feature of many birds breeding in seasonal environments is that fitness‐related parameters such as nestling mass or survival decline as the breeding season progresses. Consequently, there is a tendency for early breeders to have better reproductive performance than individuals breeding later in the season. This variation could be caused by factors associated with the date of laying, such as changing environmental conditions (date hypothesis), or by differences in parental quality between early and late breeders (parent quality hypothesis). To evaluate the relative importance of both hypotheses, we manipulated hatch dates of Blue Tits Cyanistes caeruleus by exchanging clutches with different incubation stages and assessed the impact on nestling mass, nestling diet and provisioning rates. Mean nestling mass declined significantly over the season. This was the combined result of differences in parental quality, which dominated in the early part of the season, and the influence of hatching date (date effect per se), which prevailed later in the season. Nestling diet composition was apparently uninfluenced by the manipulation, suggesting that deteriorating food supplies are the primary reason for the seasonal variation in the nestling diet. Counter to the date hypothesis, delayed parents did not feed their young less than control pairs did, but in fact exhibited higher provisioning rates. Our results suggest that in this population, parental quality seems to constrain post‐hatching reproductive performance and such intrinsic limitations may help to explain why certain individuals breed later.     

The influence of age on reproductive performance in the Brown Thornbill

I examined age effects on reproduction in the Brown Thornbill Acanthiza pusilla in Canberra, Australia. I found that the reproductive performance of both males and females improved with age, although only age-related improvement in male performance had a significant effect on annual reproductive success. Reproductive success improved with male age as a result of improved performance during two stages of the breeding cycle: first-year males were less likely to fledge young than those aged two or more, while both first and second-year males were less successful at raising fledglings to independence than males of three or more. Male performance appears to improve over three years as they gain experience at provisioning nestlings and caring for fledglings without attracting predators, rather than as a direct result of improved foraging skills. In contrast, reproductive success only improved slightly with female age, although females of two or more years initiated their first clutch earlier in the season than one-year-old females, and tended to be more likely to re-nest if a breeding attempt failed. The poor performance of young females appears unlikely to be related to their foraging ability but may be associated with costs imposed by dispersing to a breeding vacancy earlier in the year. Although the reproductive performance of Brown Thornbills improves considerably with age I found no evidence that performance improved as a result of repeated breeding attempts with the same partner.     

Seasonal declines in reproductive success of the common tern Sterna hirundo: timing or parental quality?

Reproductive success declines over the course of the breeding season in many bird species. Two categories of hypothesis have been evoked to explain this decline. The “timing” hypothesis suggests that seasonal declines in breeding success are attributable to the date of laying. The “parental quality” hypothesis suggests that seasonal declines result from the fact that young, inexperienced, or low quality birds breed later in the season. To evaluate the relative importance of timing and parental quality, egg exchanges and removals were used to manipulate hatching dates of common terns Sterna hirundo. Indices of quality, attendance, provisioning rates, and reproductive success of birds in three experimental groups (delayed hatch pairs, advanced hatch pairs, and pairs induced to relay) were compared to those of date‐matched controls. Pairs that hatched chicks early raised more chicks than pairs hatching chicks late in the season, regardless of initial laying date. This suggests that hatching chicks early is advantageous in itself. Our results, however, also support the parental quality hypothesis. There was a significant negative relationship between natural laying date and fledging success, independent of hatching date. Differences in chick growth and survival suggest that higher quality adults may be able to compensate for the disadvantages of late hatching dates and achieve similar reproductive success to that of pairs hatching chicks early. We found that pairs hatching chicks late in the season were subject to more incidents of kleptoparasitism than those hatching chicks early. This may be a proximate factor contributing to seasonal declines in reproductive success for common terns, although such a mechanism would not be likely in non‐colonial species. Failure to control for egg quality may have biased the results of some prior egg exchange experiments. Additionally, altered cost of incubation may be an unavoidable confounding factor in studies designed to manipulate timing of breeding.     

Influence of nest-site and individual quality on breeding performance in Merlins Falco columbarius

We examined the effects of nest-site quality and bird quality on breeding performance in male and female Merlins Falco columbarius from a long-term study in Saskatoon, Saskatchewan. In addition, we tested whether nest-site quality was associated with survival, as well as lifetime reproductive success (LRS). For females, nest-site quality had little influence on any of the measures of breeding performance or survival. Even so, when females switched nest-sites, they tended to move to better ones. Hatch date was repeatable for the same females occupying different nest-sites but not for the same sites occupied by different females. Among males, birds surviving past each age category tended to occupy nest-sites of higher quality, and LRS was positively correlated with nest-site quality. The relationship between nest-site quality and LRS was heavily influenced by the poorest nest-sites. When males switched nest-sites, they too tended to move to ones of higher quality. In addition, chick hatch date was repeatable neither for the same males occupying different sites nor for the same sites occupied by different males. As with most other raptors, male Merlins provide most of the food for the pair and their young during the breeding season, and differences in nest-site quality may have affected the effort needed by males to secure food. Female Merlins, however, appear still to have considerable control over the timing of breeding.     

Environmental and social cues can be used in combination to develop sustainable breeding techniques for goat reproduction in the subtropics

Goat breeds from subtropical latitudes show different annual reproductive cycles. Some of them display large seasonal variations in their annual breeding season, while others display a moderate seasonality or sexual activity all year round. This reproductive seasonality causes seasonality of milk, cheese and meat productions and, as a consequence, induces wide variation in producer incomes. To solve this problem and provide methods allowing producers to breed animals during the anestrous period and stabilize production all year round, it is necessary to have a deep knowledge of their annual sexual activity and to identify the environmental factors controlling the timing of the annual reproductive cycle. Then, it is possible to build on these knowledge sustainable breeding techniques adapted to the environmental, economic and social characteristics of the local breeding system. In this review, I will illustrate this strategy through the example of our experiments in subtropical goats. First, we determined the characteristics of the annual breeding season in both male and female goats. Second, we identified the photoperiod as the major environmental factor controlling the timing of this annual breeding season. Third, we used the photoperiod to stimulate indirectly the sexual behavior of does. Indeed, we used photoperiodic treatments to stimulate the sexual activity of bucks during the non-breeding season. These sexually active male goats were then used to induce and synchronize the estrous behavior and ovulatory activity of anestrous females in confined or grazing conditions by using the 'male effect'. Under subtropical conditions, these results constitute an original manner to control the reproductive activity of local goats using the photoperiod combined with the 'male effect.'     

Timing of arrival at breeding grounds determines spatial patterns of productivity within the population of white stork (Ciconia ciconia)

Early arrival at breeding grounds have important fitness consequences for migratory birds, both at individual and population level. The aim of this study was to investigate how the timing of arrival at the breeding territories affects the spatial patterns of reproductive success within a population of white storks (Ciconia ciconia). Data were gathered annually for ca. 200 pairs of storks breeding in central Poland between 1994 and 2011. Geostatistical analysis of data indicated that in years of delayed arrival of the population (measured by the first quartile arrival date), the reproductive output of storks was negatively autocorrelated, which indicated that there was a tendency for pairs of high breeding success to neighbour with pairs of low success. By contrast, in years when first storks returned in early dates to the breeding grounds, their reproductive success did not show any kind of spatial autocorrelation. These results suggest that delayed return of the first-arriving storks of the population may increase intensity of intra-specific competition to the level at which high-quality breeding pairs monopolize most of available resources at the expense of neighbouring low-quality pairs, which have lower reproductive success as a consequence. Such hypothesis was further supported with the analysis of nesting densities, showing that the late-arriving breeding pairs incurred greater fitness costs (or derived lower fitness benefits) while breeding in high densities comparatively to the early-arriving conspecifics.     

Variation in the timing of the reproductive season among breeds of sheep in relation to differences in photoperiodic synchronization of an endogenous rhythm.

Photoperiod may regulate seasonal reproduction either by providing the primary driving force for the reproductive transitions or by synchronizing an endogenous reproductive rhythm. This study evaluated whether breed differences in timing of the reproductive seasons of Finnish Landrace (Finn) and Galway ewes are due to differences in photoperiodic drive of the reproductive transitions or to differences in photoperiodic synchronization of the endogenous rhythm of reproductive activity. The importance of decreasing photoperiod after the summer solstice in determining the onset and duration of the breeding season was tested by housing ewes from the summer solstice in either a simulated natural photoperiod or a fixed summer-solstice photoperiod (18 h light:6 h dark; summer-solstice hold). Onset of the breeding season within each breed did not differ between these photoperiodic treatments, but Galway ewes began and ended their breeding season earlier than Finn ewes. The duration of the breeding season was shorter in Galway ewes on summer-solstice hold than on simulated natural photoperiod; duration did not differ between photoperiodic treatments in Finn ewes. The requirement for increasing photoperiod after the winter solstice for initiation of anoestrus was tested by exposing ewes from the winter solstice to either a simulated natural photoperiod or a winter-solstice hold photoperiod (8.5 h light:15.5 h dark). Onset of anoestrus within each breed did not differ between these photoperiodic treatments, but the time of this transition differed between breeds. These observations suggest that genetic differences in timing of the breeding season in Galway and Finn ewes do not reflect differences in the extent to which photoperiod drives the reproductive transitions, because neither breed requires shortening days to enter the breeding season or lengthening days to end it at appropriate times. These findings are consistent with the hypothesis that photoperiod synchronizes an endogenous rhythm of reproductive activity in both breeds and that genetic differences in timing of the breeding season reflect differences in photoperiodic synchronization of this rhythm.     

Is food availability a circannual zeitgeber in tropical birds? A field experiment on stonechats in tropical Africa

Equatorial stonechats (Saxicola torquata axillaris) in Africa are seasonal breeders like their temperate-zone conspecifics (S.t. rubicola). Their annual cycle in gonadal size and function is controlled by an endogenous circannual rhythmicity that has been shown to run for up to 10 years in a constant equatorial photoperiod under laboratory conditions, with a period deviating from 12 months. In nature, however, this rhythm is synchronized with the actual year. Because photoperiod is essentially constant at the equator, it is likely that other environmental factors act as zeitgebers. The authors test whether food availability affects reproductive cycles of free-living East African stonechats. The authors offered supplemental food to the birds 2 months before the regular onset of the breeding season. Supplementally fed males started to sing and display earlier than males of control pairs that did not receive extra food. Although the supplemented food advanced the onset of the breeding season in the pairs that were fed, the onset of the postnuptial molt following the breeding season was not correspondingly shifted. Furthermore, in the year following the experiment, all pairs initiated breeding at the same time. The authors conclude that food availability does not act as a zeitgeber, but rather as a factor that modifies the timing of reproduction without affecting the underlying rhythmicity. The authors propose that this is adaptive under environmental conditions that are relatively constant within a given year but may vary considerably between years. The zeitgeber synchronizing the endogenous rhythmicity remains to be identified.     

Factors affecting variation in the reproductive investment of female treefrogs,Hyla intermedia

In anurans, fecundity (clutch size) is the most important determinant of female reproductive success. We investigated three possible causes responsible for fecundity variation in female Italian treefrogs, Hyla intermedia, during four breeding seasons: (i) variation in morphological (body size and condition) and life-history (age) traits; (ii) variation in the tradeoff between the number and the size of eggs; (iii) seasonal effects and within-season differences in the timing of deposition. At the population level, we found no evidence for a tradeoff between the number and the size of eggs, because they both correlated positively with females’ body size. Conversely, neither age nor post-spawning body condition showed any effect on female reproductive investment. Independent of body size, we found no evidence for variation in reproductive effort among different breeding seasons, but strong evidence for a decrease of clutch size and an increase of egg size with the advancing of a breeding season. To test for the functional significance of the observed temporal variation in allocation strategy, we carried out a rearing experiment in semi-natural conditions on a random sample of ten clutches. The experiment showed a negative effect of clutch size and a positive effect of egg size on both tadpole growth and developmental rates, suggesting that reproductive investment, although constrained by body size, can be adjusted by females to the time of deposition to increase the chances of offspring survival.