Egg size and clutch size in two Daphnia species grown at different food levels

The effect of three different Scenedesmus food concentrations(0.04, 0.2 and 1 mg C l^–1) on maternal investment wasstudied in two cladoceran species of similar size, Daphnia pulicariaand D.hyalina. It was observed that as food concentration decreased(between 1 and 0.2 mg C l^–1), there was an increase insize, protein content, lipid content, carbon and mass of theegg, while, at the same time, the clutch size of the femalesbecame smaller. Such an increase in ‘per offspring investment’was reflected in an increase in body length, body carbon andbody mass of neonates as the food available for females decreased.However, in D.pulicaria this tendency was not maintained downat 0.04 mg C l^–1 in which there was a decrease of theegg characteristics mentioned above. Although, there are notavailable all the egg and neonate parameters of D.hyalina at0.04 mg C l^–1, the body length of the neonates was largerthan at 0.2 and 1 mg C l^–1. These results show that, asfood diminishes, these two cladoceran species are able to respondby decreasing clutch size, but increasing the size of egg, therebyincreasing the probability of neonate survival. This tendencyis probably maintained until the food concentration is too lowand the females have to reduce the energy allocated for reproduction.     

Recurrent violations of invariant rules for offspring size: evidence from turtles and the implications for small clutch size models

Smith and Fretwell’s classic model predicts that parents can maximize fitness by dividing the energy available for reproduction into offspring of an optimal size. However, this model breaks down when clutch size is small (~1–10 offspring). Invariant rules are an extension of the Smith–Fretwell model, and these rules predict how offspring size will vary among and within individuals that produce small clutch sizes. Here, we provide a narrow test of invariant rules using three turtle species, then we synthesize and re-analyze existing data from 18 different species (comprising five Orders) to evaluate whether invariant rules are followed across broad taxa. We do not find support for most invariant rules in turtles, and our re-analysis demonstrates a general mismatch between observed and expected values across all taxa evaluated, suggesting that invariant rules fail to predict reproductive patterns in nature. Morphological constraints on offspring size and reproductive effort may be important reasons for disparities between theory and observation both in turtles and other taxa. Paradoxically, morphological constraints are most common in small-bodied species and individuals, but these same candidates are also those which produce the small clutch sizes that are necessary to test invariant rules, such that a fair test of invariant rules will often be challenging. Mismatches between theory and observation might also occur because theory assumes that mothers exert control over resource allocation to offspring. In fact, there is evidence of widespread genetic correlations among investment per offspring and reproductive effort, such that these traits are not independent.     

Testing demographic models of effective population size

Due to practical difficulties in obtaining direct genetic estimates of effective sizes, conservation biologists have to rely on so-called 'demographic models' which combine life-history and mating-system parameters with F-statistics in order to produce indirect estimates of effective sizes. However, for the same practical reasons that prevent direct genetic estimates, the accuracy of demographic models is difficult to evaluate. Here we use individual-based, genetically explicit computer simulations in order to investigate the accuracy of two such demographic models aimed at investigating the hierarchical structure of populations. We show that, by and large, these models provide good estimates under a wide range of mating systems and dispersal patterns. However, one of the models should be avoided whenever the focal species' breeding system approaches monogamy with no sex bias in dispersal or when a substructure within social groups is suspected because effective sizes may then be strongly overestimated. The timing during the life cycle at which F-statistics are evaluated is also of crucial importance and attention should be paid to it when designing field sampling since different demographic models assume different timings. Our study shows that individual-based, genetically explicit models provide a promising way of evaluating the accuracy of demographic models of effective size and delineate their field of applicability.     

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 the relationship between clutch size and brood size in the Coot (Fulica atra)

The time between egg laying and chick fledging is of crucial importance for the survival of young birds. I analyzed breeding output at consecutive phases of growth of young Coots (Fulica atra) relative to the clutch size and laying date. Considering the specific breeding biology of the Coot, I tested whether chick survival reveals clutch size-dependent variability. Clutch size did not affect hatching success; it only affected brood size, and that merely temporarily. During the first 20 days after hatching, i.e. during the time of the highest chick mortality, birds with larger clutches lost chicks at a higher rate. As a result, the number of fledged chicks was independent of the initial number of chicks, and pairs with different clutch sizes had a similar number of fledglings. The laying date had no effect. This pattern of age-related chick survival points to the greater role of the type of chick growth (semi-precocial) and behavior in their survival.     

Evolution of clutch size in insects. I. A review of static optimality models

Abstract This paper reviews the importance of constraint assumptions to the predictions of static optimality models of insect clutch size. This allows us to identify predictions that distinguish between models embodying different constraints on female oviposition behaviour and hence to determine which resources or other factors limit clutch size evolutionarily. We conclude that while some models may be distinguished using qualitative criteria, others require the testing of quantitative predictions. In a companion paper (Wilson 1994) these models are tested using the bruchid beetle Callosobruchus maculatus.     

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.     

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.     

Termination of parental care due to small clutch size in the temperate damselfish,Chromis notata

Synopsis Reproductive behavior of the temperate damselfish, Chromis notata, was investigated on the island of Mukaishima, Japan, almost daily during the breeding season in 1982. Both males and females repeated reproductive cycles many times during the breeding season. Females had a strong tendency to spawn a whole clutch on one nest during a few hours. The average number of eggs which a male gained per reproductive cycle was estimated at 38560 (480–131100 eggs). Males ordinarily cared for eggs until just prior to hatching, but abandoned more than half of the nests with the eggs numbering less than 11568. Contribution 207 from the Mukaishima Marine Biological Station.     

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.     

Testing models of parental investment strategy and offspring size in ants

Parental investment strategies can be fixed or flexible. A fixed strategy predicts making all offspring a single ‘optimal’ size. Dynamic models predict flexible strategies with more than one optimal size of offspring. Patterns in the distribution of offspring sizes may thus reveal the investment strategy. Static strategies should produce normal distributions. Dynamic strategies should often result in non-normal distributions. Furthermore, variance in morphological traits should be positively correlated with the length of developmental time the traits are exposed to environmental influences. Finally, the type of deviation from normality (i.e., skewed left or right, or platykurtic) should be correlated with the average offspring size. To test the latter prediction, we used simulations to detect significant departures from normality and categorize distribution types. Data from three species of ants strongly support the predicted patterns for dynamic parental investment. Offspring size distributions are often significantly non-normal. Traits fixed earlier in development, such as head width, are less variable than final body weight. The type of distribution observed correlates with mean female dry weight. The overall support for a dynamic parental investment model has implications for life history theory. Predicted conflicts over parental effort, sex investment ratios, and reproductive skew in cooperative breeders follow from assumptions of static parental investment strategies and omnipresent resource limitations. By contrast, with flexible investment strategies such conflicts can be either absent or maladaptive. Electronic Supplementary Material Supplementary material is available for this article at     

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.     

Opposition site selection and clutch size in insects

Oviposition site selection and clutch size in parasitic insects can be viewed as problems in foraging theory. In this paper, a number of models for site selection and clutch size are developed, based on a dynamic state variable approach to optimal oviposition strategies. The models lead to predictions that are consistent with existing experimental data and suggest future experiments. Using these models shows the importance of constraints and state variables in the analysis of behavioral problems.     

A model of clutch size determination in birds

Summary This paper tests the hypothesis that the timing and the size of bird clutches are determined by (1) a tendency to delay reproduction so that the female can store enough internal resources for laying a large clutch; (2) a tendency to breed early because survival of offspring late in the breeding season might be lower than early. These factors are approximated by linear functions in a model. The model predicts that there will be a correlation between early breeding females and those that have large clutches. This agrees with published data for several bird species. Predictions from the model are also made of clutch size and the impact on final clutch-size if one egg is removed or added during the laying. These predictions were compared with data from a crow (Corvus cornix) population and were supported if it is assumed that the formation of an egg is a relatively costly process. More precisely, the model is supported if the formation of an extra egg requires a delay of about four or five days before laying can begin.     

Egg size and clutch size in the reproductive investment of American Kestrels

We studied causes and consequences of egg-size variation among clutches of American kestrels ( Falco sparverius ). Egg. from 275 clutches were measured 1990 to 1992. To test the hypothesis that the size of eggs was contrained by food availability in the pre-laying period, we censused small mammal populations in the three years and performed a food supplementation experiment in 1990 and 1991. Kestrels did not advance the date they laid their first egg but did lay significantly larger eggs in response to extra food. The size of eggs was correlated with small mammal abundance on the territoty, and females in good body condition tended to lay large eggs. Body size did not affect egg size, and there were no relationship between agg size and laying date except in 1900, the poorest food year. Clutches with a large mean egg volume had better hatching success than clutches containing small eggs. We argue that there is a phonetypic component to egg size in kestrels, and that kestrels use egg size to fine-tune reproductive investiment to available resources.     

On the evolution of clutch size and nest size in passerine birds

Summary I examined the hypothesis that the clutch size of some altricial birds may be limited by over-crowding of the nestlings in the nest, by comparing data on different species of European passerines. Large-sized birds build, relative to the body, larger nests than small-sized birds, both as regards the inner and the outer nest widths and as regards edge breadth; only inner nestcup depth did not change relatively to body size. Nest size also varied in relation to nesting place. Birds with open nests built off the ground had a rather narrow nestcup, whereas those with a domed nest, or which nest in a cavity, had a wide nestcup. When only open-nesters were compared, birds nesting on, or close to, the ground tended to have a wider nestcup than birds nesting above the ground. Inner nestcup width was correlated with the amount of mosses and lichens used in building the nest; the more of such materials the narrower the nestcup. The three variables: standardised body size, nesting place, and type of nesting material used accounted for 92% of the overall variation observed in inner nestcup width. When controlling for adult body size, clutch size was positively correlated with the size of the nestcup. A multiple regression analysis showed that relative nestcup depth, nest site, and type of nesting materials used, accounted for 64% of the overall variation in clutch size.