Is clutch size individually optimized?

Brood size manipulations were carried out to test whether clutchsize variation in individual great tits (Parus major) controlledfor laying date was tuned to their phenotypic quality and/orlocal food abundance (individual optimization hypothesis; IOH).Broods with different original clutch sizes, but equal hatchingdates, were manipulated to a common brood size. A third broodwas kept as a control. Under the IOH, we expected a positiveassociation between reproductive success and original clutchsize. Fledgling production varied in an inconclusive way aftermanipulation, with data from 1 out of 3 years favoring the IOH.The effect of manipulation on the probability of a second clutchwas consistent with the IOH in another 1 out of the 3 years.When fitness accrued to second broods was also taken into considerationin terms of annual fledgling production, results from 2 outof 3 years tended to support the IOH. There was no effect ofthe manipulation on fitness (estimated as the number of recruitsplus parents breeding in the next season). Both the clutch component(local recruitment) and the parental component (survival tillnext breeding season) varied inconclusively with respect tothe IOH. On the basis of fitness measurements, the IOH couldnot be confirmed as an explanation for clutch size variationin this population. In 2 out of 3 years one of the three fitnesscomponents measured varied in accordance with the IOH. Overallthe evidence for the IOH in this data set is therefore weak.     

Testing the geometric clutch hypothesis

The Geometric Clutch hypothesis is based on the premise that transverse forces (t-forces) acting on the outer doublets of the eukaryotic axoneme coordinate the action of the dynein motors to produce flagellar and ciliary beating. T-forces result from tension and compression on the outer doublets when a bend is present on the flagellum or cilium. The t-force acts to pry the doublets apart in an active bend, and push the doublets together when the flagellum is passively bent and thus could engage and disengage the dynein motors. Computed simulations of this working mechanism have reproduced the beating pattern of simple cilia and flagella, and of mammalian sperm. Cilia-like beating, with a clearly defined effective and recovery stroke, can be generated using one uniformly applied switching algorithm. When the mechanical properties and dimensions appropriate to a specific flagellum are incorporated into the model the same algorithm can simulate a sea urchin or bull sperm-like beat. The computed model reproduces many of the observed behaviors of real flagella and cilia. The model can duplicate the results of outer arm extraction experiments in cilia and predicted two types of arrest behavior that were verified experimentally in bull sperm. It also successfully predicted the experimentally determined nexin elasticity. Calculations based on live and reactivated sea urchin and bull sperm yielded a value of 0.5 nN/microm for the t-force at the switch-point. This is a force sufficient to overcome the shearing force generated by all the dyneins on one micron of outer doublet. A t-force of this magnitude should produce substantial distortion of the axoneme at the switch-point, especially in spoke or spoke-head deficient motile flagella. This concrete and verifiable prediction is within the grasp of recent advances in imaging technology, specifically cryoelectron microscopy and atomic force microscopy.     

Parent-offspring conflict over clutch size

Summary We present a model for sexually-reproducing diploids in which a female can produce a variable (generally large) clutch size, where the sibs then compete over some fixed resource, and where certain offspring use siblicide to reduce the primary clutch/brood size created by the mother. Where siblicide involves neither direct energy loss (e.g. fighting cost) nor gain (e.g. cannibalism) to an offspring, the optimal clutch size for an offspring can differ from the optimum for the mother, i.e. there can be parent-offspring conflict over clutch size. The magnitude of this evolutionary conflict (measured in terms of difference between clutch size optima) increases with multipaternity of the brood and with the steepness of the initial decline in offspring survivorship (through sib-competition as further offspring are added to the brood). However, the disparity in clutch size optima may not be great. Where the integer clutch size optima are the same, there will clearly be no conflict. Where this differs, resolution of the evolutionary conflict could involve much apparent behavioral conflict, commonly manifest as siblicidal aggression.The ESS (evolutionarily stable strategy) for such a game will depend upon the direct costs and benefits of siblicide, as well as on the indirect costs to sibs via relatedness. If the only costs of siblicide arise through relatedness, then offspring will win in the sense that the eventual clutch size will match the offspring optimum. Whether or not the mother will produce this clutch size depends on the mechanism controlling siblicide. A siblicidal ESS will occur when offspring are programmed to kill a fixed number/proportion of a brood (victim-based siblicide), but not if programmed to reduce the sibship to the offspring optimum (survivor-based siblicide). With survivor-based siblicide, the mother can do no better than to lay the offsprings' optimal clutch size.     

Testing small clutch size models with Daphnia

Life-history theory predicts that for small clutches, variance in egg size (between individuals) should decrease in a predictable invariant manner as clutch size increases. To test this, we studied Daphnia magna at 350 different food treatments and recorded the number of eggs and the volume of each egg for their first clutch. As predicted, we found that the relationship between clutch size and resources devoted to reproduction was linear, variance in egg volume decreased with increasing clutch size, and resources were shared relatively equally between the eggs in a clutch. However, we found that the rate at which the range of egg volumes decreased with clutch size was slower than predicted. We discuss possible explanations for this discrepancy, including a lower limit on the volume of eggs that are produced and selection for smaller eggs when food is abundant. Consistent with this, we found that mean egg volume decreased with increasing clutch size.     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

The demands of incubation and avian clutch size

We reviewed information on the demands of incubation to examine whether these could influence the optimal clutch size of birds. The results indicate that appreciable metabolic costs of incubation commonly exist, and that the incubation of enlarged clutches can impose penalties on birds. In 23 studies on 19 species, incubation metabolic rate (IMR) was not elevated above the metabolic rate of resting non-incubating birds (RMR), but contrary to the physiological predictions of King and others, IMR was greater than RMR in 15 studies on 15 species. Across species, IMR was substantially above basal metabolic rate (BMR), averaging 1.606 × BMR. Of six studies on three species performed under thermo-neutral conditions, none found IMR to be in excess of RMR. IMRs measured exclusively within the thermo-neutral zone averaged only 1.08 × BMR contrasting with the significantly higher figure of 1.72 × BMR under wider conditions. 16 of 17 studies on procellariiforms found IMR below RMR, indicating a significant difference between this and other orders. We could find no other taxonomic, or ecological factors which had clear effects on IMR. Where clutch size was adjusted experimentally during incubation, larger clutches were associated with: significantly lower percentage hatching success in 11 of 19 studies; longer incubation periods in eight of ten studies; greater loss of adult body condition in two of five studies; and higher adult energy expenditure in eight of nine studies. Given that incubation does involve metabolic costs and given that the demands of incubation increase sufficiently with clutch size to affect breeding performance, we propose that the optimal clutch size of birds may in part by shaped by the number of eggs the parents can afford to incubate.     

Offspring fitness and individual optimization of clutch size

Within-year variation in clutch size has been claimed to be an adaptation to variation in the individual capacity to raise offspring. We tested this hypothesis by manipulating brood size to one common size, and predicted that if clutch size is individually optimized, then birds with originally large clutches have a higher fitness than birds with originally small clutches. No evidence was found that fitness was related to the original clutch size, and in this population clutch size is thus not related to the parental capacity to raise offspring. However, offspring from larger original clutches recruited better than their nest mates that came from smaller original clutches. This suggests that early maternal or genetic variation in viability is related to clutch size.     

Maternal clutch reduction in the pied flycatcher Ficedula hypoleuca: an undescribed clutch size adjustment mechanism

During eight years of study of a population of pied flycatchers Ficedula hypoleuca breeding in central Spain, we observed throughout incubation 22 cases of intact eggs being placed on the nest rim and clearly outside of the nest bowl and remaining there, with consequent embryo mortality. We assume that the removal of eggs had been performed by the incubating female. There is no evidence that pied flycatcher parents can detect embryo mortality and eject unhatchable eggs. Maternal egg ejection constitutes a direct mechanism of clutch size adjustment during incubation and before hatching of any young of the brood. This adjustment of clutch size happened when the mean and minimum temperature of the day before ejection were especially low. Also, clutch-reducing females were significantly more prone to show moult-breeding overlap than other females, suggesting a lower disposition to invest in reproduction. Clutch-reducing females were also frequently older than four years and had indications of a lower immunocompetence than females not ejecting eggs. Our results support the hypothesis that adverse conditions can elicit parental family size adjustment at every stage of reproduction without the need to depend on mechanisms of sibling competition.     

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.     

Parasitoid clutch size and irreversible evolution

Previously, theoretical and empirical studies suggested that parasitoids developing in small multiple-egg broods would evolve siblicidal behaviour, making such brood sizes rare and single-egg broods an evolutionary absorbing state. Recent evidence, however, suggests that small gregarious broods are relatively stable in many parasitoid taxa, and that gregarious development has evolved many times from solitary development. This suggests that new research is needed to assess how nonsiblicidal behaviour can spread and become stable. We discuss some potentially rewarding possibilities.     

Ecological patterns of relative clutch mass in snakes

Summary Data on the relative clutch mass of snakes are summarized for over 100 populations. RCM was significantly lower in live bearing versus egg laying forms. We suggest that the longer reproductive season of viviparous snakes results in higher overall mortality compared to oviparous species; by reducing RCM, viviparous snakes may reduce this risk of mortality. Unlike lizards, no differences in RCM were found between categories of either escape behavior or foraging mode, possibly because detailed information on these behaviors are lacking for most snakes. In four populations examined, RCM did not vary among years. When compared to lizards, snakes demonstrate significantly higher RCM, perhaps owing to a more energetically efficient means of locomotion. Our data support the contention that RCM should be considered a separate and distinctive life-history characteristic of reptiles.     

Geographic variation in calcium and clutch size

For over a half century numerous hypotheses have surfaced aimed at explaining a key life history trait, the evolution of clutch size in birds. A principal goal has been to explain why clutch size generally increases with latitude both within species and among closely related species. Most hypotheses have stressed food limitation, predation, or seasonality. I present a novel hypothesis to explain geographic variation: a limitation of calcium resulting from broad scale variation in this element. Because the storage capacity of avian medullary bone is limited or nonexistent, during egg formation, female birds must intake supplemental calcium. Yet calcium and other exchangeable bases are much rarer in tropical soils. I briefly review the abundant experimental and observational evidence supporting how calcium limitation affects clutch size and other life history traits, and I present a series of predictions (and apparent support for them), stemming from the calcium variation and limitation hypothesis. The balance of evidence suggests that variability in availability of environmental calcium plays a proximate and ultimate role in the evolution of clutch size. Although this hypothesis is not mutually exclusive with others, it highlights another factor that needs to be considered in studies of the geographic variation in clutch size.     

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.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

Experiments on clutch size and nest size in passerine birds

Summary Results of experiments on three passerine species suggest that brood size may be constrained by nest size, since the breeding success of pairs provided with large nestcups was greater than that of those provided with small artificial nestcups. These results may have important implications, e.g. to the design of experiments involving manipulation of clutch and brood size. A small nestcup is requisite for successful hatching during the incubation period, but a large one for successful rearing during the nestling period. In nature this difference may select for types of nesting materials that are elastic, such as mosses and lichens. However, experiments showed that such materials rapidly absorb rainwater but only slowly dry out. In addition, because large nests dry out more slowly than small nests, selection will favour small nests among those open-nesting species that have exposed nests. A further possible nest size constraint on open-nesters is nest predation. However, no difference in the predation rate was found in experiments with small and large artificial nests.     

Incubation prior to clutch completion accelerates embryonic development and so hatch date for eggs laid earlier in a clutch in the great tit Parus major

The ability of parents to respond to changes in food supply within a season will have a large effect on fitness through the number and quality of chicks fledged. Great tits, Parus major, attempt to synchronise their production of chicks with a seasonal food peak, but when food supply fails, hatching asynchrony of chicks provides a mechanism by which some young can be fledged because more developed chicks outcompete their less developed siblings for the reduced parental food supply. We tested whether female great tits can potentially control the degree of hatching asynchrony by using incubation before clutch completion, so that early laid eggs develop faster and hatch sooner. The temperature of an artificial egg placed in 29 nests during the laying period was measured with data loggers, and nocturnal incubation of eggs similar to incubation post clutch completion was recorded in all nests. We then demonstrated that eggs removed from the nest for 72 hour periods prior to clutch completion hatched later than eggs remaining in the nest for the entirety of the laying period. Our results show that variable pre clutch completion incubation (which was mostly nocturnal) can lead to faster embryo development and earlier hatching, so potentially providing a mechanism for adaptive female control of degree of hatching asynchrony.