Between-clutch interactions affect a benefit of group feeding for pipevine swallowtail larvae

Abstract.  1. In California, early instar larvae of the pipevine swallowtail ( Battus philenor ) develop at an accelerated rate when feeding in large groups compared with small groups due to a plant-mediated response to feeding group size. Larvae benefit from accelerated growth because the time larvae remain in early stages, where mortality is highest, is reduced. Occasionally, multiple clutches are laid on the same plant stem. Clutch size modification by females ovipositing on plant with previously laid clutches and the effect of kinship and group size on larval behaviour was examined. The direct and indirect interactions between clutches were investigated to determine if group size and time between clutch establishment affects the performance of early instar larvae.
2. Larger groups consume the young foliage more quickly and develop at an accelerated rate compared with smaller groups. Older foliage available to later clutches is an inferior food resource compared with younger foliage.
3. There was no evidence that females adjust clutch size in response to the presence of conspecific clutches.
4. Second groups of larvae readily joined previously established feeding groups. There were no observed behavioural differences between sibling and mixed-family groups.
5. The effect of a second group on the growth of the initial group was dependent on the size of both groups and the time interval between the arrival of the two groups.
6. Accelerated growth associated with larger feeding aggregations was absent when these groups were introduced to plants with previously established groups.
7. It is beneficial for ovipositing females to avoid plants with previously laid clutches because direct and indirect interactions with established clutches compromises larval performance.     

Clutch size in frugivorous insects as a function of host firmness: the case of the tephritid fly Anastrepha ludens

Abstract. 1. Optimal clutch size theory predicts that individuals will oviposit the number of eggs that increases their fitness. In Anastrepha ludens Loew (Diptera: Tephritidae), females oviposit larger clutches in unripe (firm) fruits than in ripe (soft) fruits. The following hypotheses were tested: (1) Using fruit firmness as an indicator of fruit quality, A. ludens females vary the number of eggs per clutch every time they reach an oviposition decision. (2) Maximising offspring survival with respect to either unripe or ripe fruit requires placing large clutches in firm fruit and smaller clutches in soft fruit. 2. Agar spheres were used as artificial hosts. Three agar concentrations resulted in three degrees of firmness. Mango fruits Mangifera indica L. served as natural hosts. Ripe and unripe fruits were used to test soft and firm host conditions respectively. Females laid significantly larger clutches in the firmer artificial hosts than in the softer hosts. They also laid significantly more eggs in artificial hosts without sugar than in hosts with sugar. Firm (unripe) mangoes also received significantly larger clutches than soft (ripe) mangoes. 3. When an individual female was first presented with a firm artificial host, it laid a large clutch. If subsequently offered a soft host, the female laid a significantly smaller clutch. Finally, if again offered a firm host, clutch size was increased significantly. 4. Possible trade‐offs in offspring fitness were explored in ripe and unripe mangoes by measuring offspring egg‐to‐adult survival, pupal weight, mean adult longevity, and fecundity. Despite the fact that larval survival was greater in soft fruit than in firm fruit, parameters such as pupal weight, mean longevity, and fecundity of adults stemming from both fruit types did not differ significantly. 5. A probable trade‐off between high offspring mortality caused by host unsuitability and low offspring and adult mortality caused by parasitism and predation is discussed as the reason for the exploitation of sub‐optimal hosts.     

Consequences of infanticide for a gregarious ectoparasitoid of leafroller larvae

1. In this laboratory study, the clutch size and handling time of Goniozus jacintae were investigated, a comparison of its life‐history performance between primary and secondary (laid after infanticide events) broods was carried out, and the lipid and protein concentrations in the haemolymph of non‐parasitised and parasitised hosts were estimated. 2. It was found that G. jacintae temporarily paralysed its host larvae for 66 min and briefly guarded its brood for 66 min. The clutch size of G. jacintae increased from two to seven with increasing larval fresh weight of its host, and both ovicide and larvicide of primary clutches occurred in 81% of encounters. 3. Secondary clutches of G. jacintae were significantly larger than primary clutches in two of three ovicide treatments for the same host individuals. Secondary clutches also experienced greater brood survivorship than primary clutches. 4. Lipid concentrations were consistently higher in the haemolymph of parasitised hosts, and protein concentrations were initially higher (until egg hatch), but increased at a lower rate in parasitised hosts than in non‐parasitised hosts. 5. This study is the first to provide evidence that improved nutritional quality could be an important benefit of infanticide for an insect parasitoid, allowing for larger clutch size and improved brood survivorship among secondary broods.     

The problem of optimal clutch size in a tritrophic system: the oviposition strategy of the thistle gallfly Urophora cardui (Diptera,Tephritidae)

The problem of optimal clutch sizes is a central theme in life history theory. Optimal allocation of eggs is especially complicated for insects in tritrophic systems. In this study we analyze some of the processes determining clutch sizes of the thistle gallfly Urophora cardui, a monophagous tephritid fly associated with Cirsium arvense. U. cardui forms multilocular shoot galls, which vary broadly in their size and number of their gall cells. We investigate various fitness consequences of gall size. An analysis of the number of cells per gall (which is correlated with gall diameter and gall weight) showed that in U. cardui there is mutual facilitation rather than larval competition. Increasing numbers of larvae per gall led to a decreasing mortality and increasing larval weight. Larval weight in turn was positively correlated with the probability of survival to adulthood and with adult weight and fecundity. Thus, all fitness parameters measured favoured large galls. Clutch sizes in oviposition experiments were distinctly larger than the number of gall cells of field populations and in cage experiments, suggesting high mortality of eggs and/or early larval instars. There was a significant relationship between the internal structure (i.e., the size of the growing point) of the bud and clutch size, suggesting that U. cardui females are able to measure bud quality and adapt clutch sizes accordingly. Clutch size was positively correlated with the female's age at first oviposition and negatively with the number of previous ovipositions and previously laid eggs. Since the potential egg capacity per female is higher than the average number of larvae it is likely to produce during its short adult lifespan, U. cardui females tend to be time-limited rather than egglimited, which might favour large clutches once an appropriate oviposition site has been located. As the development of the gall and hence the fate of a clutch depends on a number of unpredictable factors, exclusive concentration of eggs in a few large clusters would involve risks which could be avoided by increasing the number of clutches. Therefore we interpret the high variation of clutch sizes in U. cardui as a mixed strategy of bet hedging and gambling.     

Models of clutch size in insect oviposition

Models for clutch size in species where a female deposits eggs into a larval resource of limited carrying capacity are developed. Previous models of clutch size related mainly to vertebrates (notably birds) where parental care limits clutch size. Our models cover cases where a single female “saturates” a larval food patch with larvae. The main predictions are that (1) extra eggs should be laid to compensate for larval moratility; (2) clutches should generally be smaller than the size that yields the maximum number of surviving larvae/clutch; (3) in species that gain resources for eggs in the adult stage, clutch size will be unaffected by age-independent parental mortality between clutches; (4) clutch size should reduce throughout life in species that gain resources for eggs before the adult stage; (5) similar species, but which are constrained to produce constant-sized clutches, should lay smaller clutches if their total potential egg production is low; (6) clutch size should increase with increasing search costs for oviposition sites. An ESS model of double-oviposition (where two females sometimes lay in the same larval food patch) indicates that the first female should generally lay more eggs than the second female; the difference in clutch size should decrease as the probability of double-oviposition increases, and should decrease as the search costs for larval food plants decreases. Many of the predictions have some support from data on insect oviposition.     

Larval Competition, Adult Fitness, and Reproductive Strategies in the Acoustically Orienting Ormiine Homotrixa alleni (Diptera: Tachinidae)

Homotrixa alleni is a gregarious endoparasitoid fly that attacks adult male Sciarasaga quadrata (Orthoptera: Tettigoniidae) in southwestern Western Australia. Gravid female flies acoustically orient to their host's call and deposit live first-instar larvae upon or near their calling host. Up to 16 larvae may be found developing in the one host, and since only calling adult male S. quadrata are parasitized, host size and hence larval resources are essentially fixed at parasitism. This study examines parasitism by H. alleni in relation to intraspecific larval competition and adult fitness. The mean number of larvae emerging per host failed to increase significantly beyond a clutch size of four. Mean pupal weight and survival to the adult stage decreased linearly with increasing clutch size across the entire range of clutch sizes examined. Within a clutch, heavier pupae successfully completed pupal development significantly more often than lighter pupae. Pupal weight was directly related to adult size, with adult males being significantly larger than adult females at any given pupal weight. Female body size was positively correlated with fecundity. The size distribution of emerging females was normally distributed, while the distribution of searching gravid females collected at acoustic traps in the field was significantly skewed toward larger flies, suggesting yet another fitness benefit associated with large size. Using fecundity and survival to adulthood as our measure of fitness we calculated the optimal clutch size maximizing fitness per host to be seven, which exceeds the majority of observed clutch sizes in the field. Uncertainties associated with larvae successfully entering the host following larviposition are likely to reduce clutch sizes of H. alleni below this optimum in the field.     

Encountering competitors reduces clutch size and increases offspring size in a parasitoid with female-female fighting

Understanding the size of clutches produced by only one parent may require a game-theoretic approach: clutch size may affect offspring fitness in terms of future competitive ability. If larger clutches generate smaller offspring and larger adults are more successful in acquiring and retaining resources, clutch size optima should be reduced when the probability of future competitive encounters is higher. We test this using Goniozus nephantidis, a gregarious parasitoid wasp in which the assumption of size-dependent resource acquisition is met via female-female contests for hosts. As predicted, smaller clutches are produced by mothers experiencing competition, due to fewer eggs being matured and to a reduced proportion of matured eggs being laid. As assumed, smaller clutches generate fewer but larger offspring. We believe this is the first direct evidence for pre-ovipositional and game-theoretic clutch size adjustment in response to an intergenerational fitness effect when clutches are produced by a single individual.     

Resource-Dependent Clutch Size Decisions and Size-Fitness Relationships in a Gregarious Ectoparasitoid Wasp, <Emphasis Type="Italic">Bracon brevicornis</Emphasis>

Gregarious parasitic wasps, which lay more than one egg into or onto a host arthropod’s body, are usually assumed to lay an optimal number of eggs per host. If females would lay too few eggs, some resources may be wasted, but if females lay too many eggs, offspring may develop into substantially smaller-sized adults or may not develop successfully and die. The availability of hosts can further influence a female’s clutch size decision, as more eggs should be laid when hosts are scarce. Here, we analyzed clutch size decisions and the fitness consequences thereof in the ectoparasitic wasp Bracon brevicornis (Hymenoptera: Braconidae), a potential biocontrol agent against pest moth species. For experiments, larvae of the Mediterranean flower moth, Ephestia kuehniella (Lepidoptera: Pyralidae) were used. Using artificially created as well as naturally laid clutches of eggs, the effects of clutch size on fitness of first (F1) and second (F2) generation offspring were investigated. Our results revealed that the fitness consequences of large clutches included both increased mortality and smaller adult sizes of the emerging offspring (F1). Smaller F1 females matured fewer eggs during their lifetime and their offspring (F2) had reduced egg-to-adult survival probability. Naturally laid clutches varied with host size up to a maximum, which probably reflects egg limitation. Clutches remained smaller than the calculated optimal (Lack) clutch size and females responded to high host availability with a decreased number of eggs laid. We thus conclude that large clutches may result in significantly smaller offspring with reduced fitness, and that host size as well as host availability influence the clutch size decision made by B. brevicornis females.     

Aggregative Behavior is Not Explained by an Allee Effect in the Walnut Infesting Fly, Rhagoletis juglandis

Component Allee effects are considered to be a driving force in the origin and maintenance of aggregative behavior. In this study, we examine whether a pattern of active host reuse by the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), involves an Allee effect. We examined how the density of clutches deposited within a fruit, the temporal pattern in which successive clutches are deposited and the spatial distribution of clutches over a fruit’s surface influences survival to pupation and pupal size. Within the density range used in this experiment (1 to 7 clutches), increases in larval density strongly reduced pupal weight but not larval survival to pupation. The temporal staggering of clutches into a host strongly reduced offspring survival and, probably owing to competitive release, increased the pupal weight of survivors. Offspring survival and pupal weight were affected relatively little by whether two clutches were deposited within the same oviposition punctures or were evenly spaced. In contrast, in three-clutch treatments offspring survival was higher when clutches were placed within the same oviposition cavity. However, pupal weights did not significantly increase when clutches were placed together and this relatively higher survival rate was not greater than that associated with hosts that contained fewer clutches. The results of the study failed to provide evidence of an Allee effect. We put forward a scenario under which females appear to reuse larval hosts to maximize their own reproductive success, albeit at the expense of the per capita fitness of their offspring.     

Clutch size manipulations in the chestnut weevil, Curculio elephas: fitness of oviposition strategies

We test the adaptive value of clutch size observed in a natural population of the chestnut weevil Curculio elephas. Clutch size is defined as the number of immatures per infested chestnut. In natural conditions, clutch size averages 1.7 eggs. By manipulating clutch size in the field, we demonstrate that deviations from the theoretical ”Lack clutch size”, estimated as eight immatures, are mainly due to proximate and delayed effects of clutch size on offspring performance. We show the existence of a trade-off between clutch size and larval weight. The latter, a key life-history trait, is highly correlated with fitness because it is a strong determinant of larval survival and potential fecundity of offspring females. The fitness of different potential oviposition strategies characterized by their clutch sizes, ranging from one to nine immatures, was calculated from field- estimated parameters. Chestnut weevil females obtain an evolutionary advantage by laying their eggs singly, since, for instance, fitness of single-egg clutches exceeds fitness of two-egg clutches and four-egg clutches by 8.0% and 15.1% respectively. Received 4 August 1999 / Accepted: 7 October 1999     

The cost of incubation in relation to clutch-size in the Collared Flycatcher Ficedula albicollis

This study examines the importance of avian incubation costs as determinants of clutch-size variation by performing clutch-size and brood-size manipulations in the same population of Collared Flycatchers Ficedula albicollis during the same breeding season. In 2 5 cases when three or more clutches of the same size were completed on the same day, we moved two eggs on the day after the last egg had been laid from one randomly selected clutch (C) to another (C) and moved two other eggs from this to a third clutch (C⁺). In 20 other cases of simultaneously completed clutches of the same size, we moved two randomly selected young from one brood to a second and from that moved two other young to a third (B, B and B⁺groups). Most females were weighed the day after completion of the clutch and 1–4 days before hatching of the young, and some of them also 10–14 days after hatching of the young. We measured the daily energy expenditure of females incubating manipulated clutches of 4, 6 and 8 eggs by means of the doubly-labelled water (D₂¹⁸O) technique and also recorded their nest attendance. Hatching success of fertilized eggs was reduced in the enlarged clutches compared with control and reduced clutches. Females expired on average 3142.6 ml CO₂ and expended 78.6 kJ per day while incubating, which corresponds to a metabolic intensity of 3.3 times BMR. Daily energy expenditure increased with clutch-size due to higher costs while incubating, and not because of changed activity patterns. There were no significant differences in length of incubation, female mass or mass changes between phases for the C, C and C⁺groups. In both the C and B groups, enlarged broods produced significantly more fledged young than control broods, and those significantly more than reduced broods. Fledgling tarsus-length and mass did not differ significantly between treatments in either the C or B groups. There was no significant difference in breeding success between clutch and brood manipulations. In this season, incubation costs did not entail significant fitness losses, expressed either as fledgling production or female condition. Also, control females could have raised more young to fledging age than they did with no apparent costs.     

Paternal effects associated with melanism in Harmonia axyridis (Coleoptera: Coccinellidae): mating sequence asymmetries and interactions with age‐specific maternal effects

1. For this study a cohort of melanic Harmonia axyridis males homozygous for the spectabilis allele was produced. These were used to produce four kinds of twice‐mated females, comprising all four permutations of melanic and non‐melanic (succinic) males. A series of 12 larvae were then reared from the first and 10th clutches of each female to compare progeny developmental phenotypes. 2. There were no effects of mating treatment on overall female reproductive performance (preoviposition period, 20‐day fecundity, or egg fertility). 3. Age‐specific maternal effects were evident in progeny developmental phenotypes; larvae of 10th clutches developed more slowly, pupation was shorter, and adults emerged at heavier weights. 4. Paternal effects were superimposed on maternal effects and affected progeny independent of their paternity; melanic males induced slower larval development and slower pupation, but only in 10th clutches and only when they mated second. 5. There was a significant three‐way interaction between male mating treatment, clutch number and progeny phenotype, indicating that progeny developmental responses to (mixed) paternal effects varied depending on their own phenotype and the time elapsed since their mother's last mating. 6. Melanic males mating second obtained a P2 advantage over succinic males, which increased from first to 10th clutch, but the reverse was not true when succinic males mated second. Thus, polyandry in H. axyridis facilitates both genetic and epigenetic competition among males while simultaneously enabling the sharing of predominant paternal effects among the progeny of different fathers.     

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.     

Egg size variation and its relationship with larval performance in the Lepidoptera: the case of the European grapevine moth Lobesia botrana

Life history theory attempts to define the pattern of resource distribution among offspring and predicts that egg size should be positively correlated with offspring fitness. In this paper we investigated the effect of an array of ecological and reproductive factors on the size of eggs laid by Lobesia botrana, and the ecological significance of egg size by experimentally testing whether or not egg size increased larval performance in this moth. Egg size was significantly affected by female age at mating, water availability, pupal weight loss, the phenological stage of the vine in which larvae developed, female body weight, and oviposition day, but was unaffected by the size of the spermatophore received. The greater the size of the first eggs produced, the smaller the size of the last eggs laid, as predicted by the resource depletion hypothesis. As expected, the larger the size of eggs the larger the size of larvae emerging from them. Large larvae displayed a better ability to endure starvation than small ones. When small and large larvae were allowed to develop on grape clusters, an adverse environment, large larvae performed better, settling and survival being significantly enhanced, almost tripled. Instead, when the same experiment was carried out on an artificial diet in the laboratory, a much more favourable milieu, survival in both sized-larval groups was not significantly different. We discuss our results in the context of current ideas relating egg size, larval performance and fitness in the Lepidoptera.     

The influence of competition between foragers on clutch size decisions in an insect parasitoid with scramble larval competition

The effect of competition between ovipositing females on theirclutch size decisions is studied in animals that lay their eggsin discrete units of larval food (hosts). In such species theeffect of competition depends on the form of the larval competitionwithin such units. In insect parasitoids, there might eitherbe contest (solitary parasitoids) or scramble competition (gregariousparasitoids) between larvae within a host For gregarious parasitoids,a decreasing clutch size with increasing competition betweenforagers is predicted. This prediction is tested in experimentsusing the parasitoid Aphaertta minuta. Parasitoids were eitherkept alone or in groups of four before the experiment, in whichthey were introduced singly in a patch containing unparasitizedhosts. Animals kept together laid on average clutches of 0.74eggs smaller than females kept alone (average clutch is 5.3),thereby confirming the prediction. Clutch size decreased withencounter number, which might be due to the adjustment of thefemale's estimate of the encounter rate with hosts. Finally,the results are compared with those reported for solitary parasitoids(that have scramble larval competition), for which it is predictedthat the clutch size will increase with increasing levels ofcompetition between females.     

Time-dependent reproductive decisions in the blue tit

Many breeding attempts in birds do not result in any fledged young due to predation on eggs or young. Consequently, the influence of time constraints on reproductive decisions are integrated parts of the reproductive behaviour of birds breeding within short, seasonal climate zones. In this study, I mimicked nest predation by removing blue tit (Parus caeruleus) clutches shortly after completion. Around 75% of the removed clutches were followed by a repeat clutch. Females producing their first clutch early in the season and females with an early onset of incubation in the laying sequence (an indication of high parental or territory quality) were most likely to initiate a repeat clutch. A trade‐off between the benefits of a repeat clutch and survival likely stopped late females in bad condition from investing more in the current reproductive season. Females producing a repeat clutch laid fewer eggs, had an earlier onset of incubation in the laying sequence and produced larger eggs than they did when producing their original first clutch. Eggs produced after the onset of incubation were especially large in the repeat clutches. Since food availability was presumably higher when the female produced her repeat clutch compared with her first clutch, females made a strategical decision when reducing clutch size, whereas onset of incubation and egg size may have been energetically constrained when producing the first clutch. Females that produced a relatively large clutch, had a relatively early onset of incubation, and laid relatively large eggs in their first clutch also did so when producing a repeat clutch, indicating that some of the variation in breeding parameters are due to differences in parental or territory quality. Differences between years in the temperature‐dependent development rate of caterpillars seem to affect the time constraints on breeding. A year with a predicted early seasonal decline in caterpillars resulted in short intervals between removal and relaying, small clutches and an early onset of incubation.     

The moult of British thrushes and chats

Incubated clutches of the Mandarin Duck were larger earlier in the breeding season. At all nests there were days on which a single egg, and days on which no egg was added to a clutch; days on which more than one egg was added to a clutch were less frequent. Larger clutches had proportionately more days on which two or more eggs were added; the proportion of days on which no eggs were added was unrelated to clutch size. Larger clutches had a relatively shorter laying period than expected, but a longer incubation period than smaller clutches. Unhatched eggs resulted mainly from embryonic mortality, which was concentrated in the period just prior to pipping. Similar proportions of unhatched eggs resulted from eggs laid into clutches before and after the start of incubation. Larger clutches did not have proportionately more unhatched eggs. As all clutches had at least one day on which no eggs were laid, it is proposed that the proportion of nests parasitized was greater than that estimated from nests with more than one egg laid on any one day.     

Clutch size in a larval-pupal endoparasitoid

Clutch size decisions by Aphaereta minuta (Nees) (Hymenoptera: Braconidae), a polyphagous, gregarious, larval-pupal endoparasitoid, were studied under laboratory conditions. This parasitoid attacks larvae of Diptera inhabiting ephemeral microhabitats such as decaying plant and animal material. Females oviposit in young larval stages, but the eventual size of the host pupa determines host food availability for competing offspring. The size of the pupa can differ greatly between host species. We questioned how A. minuta females deal with this delay between the moment of oviposition and eventual host food availability, and whether they make clutch size decisions that benefit their fitness. It was shown that females indeed vary their clutch size considerably and in an adaptive way: (1) females lay larger clutches in larvae of host species that produce larger pupae, even when the larvae are the same size at the moment of oviposition, and (2) females lay larger clutches in larger larvae than in smaller larvae of the same host species. The latter seems functional as larvae parasitized at an older stage indeed developed into larger pupae compared to larvae parasitized at a younger stage. Furthermore, mortality of parasitized young host larvae was greater than that of both unparasitized larvae and parasitized older larvae. Under field conditions the risk of mortality of young host larvae is expected to be even higher due to the limited period of microhabitat (host food) availability, strong scramble type competition between the host larvae, and the longer period of being exposed to predation.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

Brood Parasitism and Nest Takeover in Common Eiders

Conspecific brood parasitism (CBP) is an alternative breeding tactic that occurs in many brood‐tending animals and can have important fitness effects for both host and parasite. We use protein fingerprinting of egg albumen to distinguish the eggs from different females and to estimate the frequency, pattern and tactics of CBP and other forms of mixed maternity in a Hudson Bay population of common eiders (Somateria mollissima sedentaria). Mixed clutches, containing eggs from more than one female, occurred in 31% of the 86 nests studied that progressed to clutch completion. Other females than the host laid 8% of the eggs. In 11 (41%) of the mixed clutches another female laid before the host started laying, corroborating the hypothesis that takeover of nests started by other females accounts for many of the mixed clutches in this population. Our results also indicate that traditional non‐molecular methods of identifying foreign eggs may considerably underestimate the frequency of mixed clutches.