Optimal size and number of propagules: allowance for discrete stages and effects of maternal size on reproductive output and offspring fitness

Existing optimality models of propagule size and number are not appropriate for many organisms. First, existing models assume a monotonically increasing offspring fitness/propagule size relationship. However, offspring survival during certain stages may decrease with increasing propagule size, generating a peaked offspring fitness/propagule size function (e.g., egg size in oxygen-limited aquatic environments). Second, existing models typically do not consider maternal effects on total reproductive output and the expression of offspring survival/propagule size relationships. However, larger females often have greater total egg production and may provide better habitats for their offspring. We develop a specific optimality model that incorporates these effects and test its predictions using data from salmonid fishes. We then outline a general model without assuming specific functional forms and test its predictions using data from freshwater fishes. Our theoretical and empirical results illustrate that, when offspring survival is negatively correlated with propagule size, optimal propagule size is larger in better habitats. When larger females provide better habitats, their optimal propagule size is larger. Nevertheless, propagule number should increase more rapidly than propagule size for a given increase in maternal size. In the absence of density dependence, females with greater relative reproductive output (i.e., for a given body size) should produce more but not larger propagules.     

Mothers adjust egg size to helper number in a cooperatively breeding cichlid

Mothers should adjust the size of propagules to the selectiveforces to which these offspring will be exposed. Usually, alarger propagule size is favored when young are exposed to highmortality risk or conspecific competition. Here we test 2 predictionson how egg size should vary with these selective agents. Whenoffspring are cared for by parents and/or alloparents, protectionmay reduce the predation risk to young, which may allow mothersto invest less per single offspring. In the cooperatively breedingcichlid Neolamprologus pulcher, brood care helpers protect groupoffspring and reduce the latters' mortality rate. Therefore,females are expected to reduce their investment per egg whenmore helpers are present. In a first experiment, we tested thisprediction by manipulating the helper number. In N. pulcher,helpers compete for dispersal opportunities with similar-sizedindividuals of neighboring groups. If the expected future competitionpressure on young is high, females should increase their investmentper offspring to give them a head start. In a second experiment,we tested whether females produce larger eggs when perceivedneighbor density is high. Females indeed reduced egg size withincreasing helper number. However, we did not detect an effectof local density on egg size, although females took longer toproduce the next clutch when local density was high. We arguethat females can use the energy saved by adjusting egg sizeto reduced predation risk to enhance future reproductive output.Adaptive adjustment of offspring size to helper number may bean important, as yet unrecognized, strategy of cooperative breeders.     

Offspring fitness and the optimal propagule size in a fluctuating environment

Propagule size is an important maternal effect on offspring fitness and phenotype in birds and other oviparous animals. The performance of propagules often increases with size, but a fluctuating environment may introduce temporal variation in the optimal phenotype. Understanding these mechanisms will provide novel insights into the eco‐evolutionary dynamics of life history strategies in parental reproductive investment. We investigated the interaction between propagule size (measured as egg volume) and environmental conditions on offspring mortality and phenotype in a Norwegian house sparrow population. Increased propagule size reduced offspring mortality in early life, with more pronounced effects under heavy precipitation. However, the optimal propagule size for low offspring mortality until recruitment shifted from large to small as temperature increased. Propagule size had no significant effect on fledgling body mass and tarsus length. These results reveal a potential for eco‐evolutionary dynamics in propagule size, as populations adapt to fluctuating environmental conditions. The ultimate outcome of this dynamic process will also depend on variation in parental fitness and tradeoffs with other life‐history traits, particularly clutch size.     

Egg size and offspring quality: a meta‐analysis in birds

Parents affect offspring fitness by propagule size and quality, selection of oviposition site, quality of incubation, feeding of dependent young, and their defence against predators and parasites. Despite many case studies on each of these topics, this knowledge has not been rigorously integrated into individual parental care traits for any taxon. Consequently, we lack a comprehensive, quantitative assessment of how parental care modifies offspring phenotypes. This meta‐analysis of 283 studies with 1805 correlations between egg size and offspring quality in birds is intended to fill this gap. The large sample size enabled testing of how the magnitude of the relationship between egg size and offspring quality depends on a number of variables. Egg size was positively related to nearly all studied offspring traits across all stages of the offspring life cycle. Not surprisingly, the relationship was strongest at hatching but persisted until the post‐fledging stage. Morphological traits were the most closely related to egg size but significant relationships were also found with hatching success, chick survival, and growth rate. Non‐significant effect sizes were found for egg fertility, chick immunity, behaviour, and life‐history or sexual traits. Effect size did not depend on whether chicks were raised by their natural parents or were cross‐fostered to other territories. Effect size did not depend on species‐specific traits such as developmental mode, clutch size, and relative size of the egg, but was larger if tested in captive compared to wild populations and between rather than within broods. In sum, published studies support the view that egg size affects juvenile survival. There are very few studies that tested the relationship between egg size and the fecundity component of offspring fitness, and no studies on offspring survival as adults or on global fitness. More data are also needed for the relationships between egg size and offspring behavioural and physiological traits. It remains to be established whether the relationship between egg size and offspring performance depends on the quality of the offspring environment. Positive effect sizes found in this study are likely to be driven by a causal effect of egg size on offspring quality. However, more studies that control for potential confounding effects of parental post‐hatching care, genes, and egg composition are needed to establish firmly this causal link.     

The evolution of offspring size and number: a test of the Smith-Fretwell model in three species of crickets

Most models of parental investment in offspring assume a trade-off between propagule size and number, and an increasing concave down function relating offspring fitness to propagule size. In this study, we test these two fundamental assumptions, using three closely related species of crickets, Gryllus firmus, G. veletis, and G. pennsylvanicus. Egg weight, 35-day fecundity and 35-day egg biomass were estimated in a population of each species, and the relationships between these reproductive traits and date of egg laying and body size were estimated. The relationships between egg weight and offspring survival were also sought for eggs buried at different depths, soil moistures, and soil types (G. firmus and G. veletis), as well as in the field (G. pennsylvanicus). A trade-off between egg weight and 35-day fecundity was revealed in a multivariate analysis taking into account among-species variation in egg weight and body size. Independent of the environmental conditions affecting the eggs, a positive correlation existed between the number of larvae that emerged from the soil and propagule weight in each species. Therefore, these findings provide partial support for the assumptions considered in the models mentioned above. A single optimal egg size was favored in two out of the three sets of conditions in which the functions relating egg weight to larval survival could be derived. The conditions encountered by the eggs, however, influenced the average survival of the larvae, as well as the shape of the relationship between egg weight and offspring survival. This suggests that cricket eggs frequently face heterogeneous environments with respect to egg and hatchling survival; the implication of habitat heterogeneity on the evolution of an optimal egg size is considered. The relationships between the reproductive components and female age and size, as well as between egg size and variation in cricket life-history, are discussed in an ecological and evolutionary context.     

Optimal offspring provisioning when egg size is "constrained": a case study with the painted turtle Chrysemys picta

Classic egg size theory predicts that, in a given environment, there is a level of maternal investment per offspring that will maximize maternal fitness. However, positive correlations among egg size and female body size are observed within populations in diverse animal taxa. A popular explanation for this phenomenon is that, in some populations, morphological constraints on egg size, such as ovipositor size (insects) or pelvic aperture width (lizards and turtles), limit egg size. Egg size may therefore increase with female body size due to body size‐specific constraints on investment per offspring, coupled with selection towards an optimal egg size. We use 17 years of data from a population of painted turtles Chrysemys picta to evaluate this hypothesis. In accordance with our predictions, we find that (1) morphological constraints on egg size are apparent only in relatively small females, similarly (2) egg mass exhibits a strong asymptotic relationship with female body size, suggesting egg mass is optimized only at large body sizes, (3) clutch size, not egg mass, varies with female condition, and (4) clutch size varies more than egg mass across years. Contrary to our predictions, we observe that (5) the egg mass‐clutch size tradeoff is not less pronounced at large body sizes. Our data do not fully support the traditional hypothesis, and recent models suggest that this hypothesis is indeed overly simplistic. When the selective environment of a female's offspring is influenced by her phenotype, optimal egg size may vary among maternal phenotypes. This concept can explain correlations among egg size and body size in many taxa, as well as the patterns observed in the present study. In this paradigm, a tight coupling of aperture width (or other ‘constraints’) and egg size may occur in small females, even when such morphological features are not causally related to variation in egg size. In this spirit, we question validity of invoking morphological constraints to explain covariation among egg size and female body size.     

The Particular Maternal Effect of Propagule Size, Especially Egg Size: Patterns, Models, Quality of Evidence and Interpretations

SYNOPSIS. Propagule size is perhaps the most widely recognizedand studied maternal effect in ecology, yet its evolution isnot well-understood. The large body of extant optimality theorytreats parental investment solely as an ecological problem,largely from the perspective of progeny. This approach has hadlimited success explaining the ubiquitous variation in propagulesize within and among natural populations at most temporal andspatial scales. This problem aside, an unassailable gap in propagulesize theory is that it pays little heed to the fact that offspringsize is a joint phenotype of two individuals- the offspringand its mother. Hence, the ecology of mothers is decidedly asimportant in shaping the evolution of propagule size phenotypes.There are two reasons to suspect that this gap may account forthe lack of success of optimality theory to explain variationin nature. The first is that optimality models of propagulesize make no allowance for, nor can they explain, widespread,multivariate correlations between maternal characters and clutchparameters, namely the positive phenotypic covariances of maternalage, size, fecundity, and per-propagule investment found inmany organisms. If per-propagule investment is optimized byselection based on the expectation of offspring fitness, thenwhy should that phenotype be a function of maternal age or sizewhen the ecological circumstances of progeny are not changingas a function of maternal age or size? The second gap in currenttheory is that, like all optimization theory, it is patentlynon-genetic in that it is assumed that the phenotypes optimizedare evolutionarily accessible. Recent maternal effects theoryindicates that traits subject to maternal influence behave inunanticipated ways. Specifically, there may be time lags inresponse to selection, and hence, selection away from the optimumphenotype. This paper explores a suite of issues pertainingto the evolution of propagule size from the broader perspectiveof propagule size as a maternal effect (PSME) with a goal ofwidening the lens through which propagule size is viewed byevolutionary ecologists. Two themes are developed. First, Isuggest that, to understand egg size variance and its implicationsfor both maternal and offspring fitness, it is necessary toconsider explicitly the ecological context in which a motheris producing eggs, not just that into which offspring will enter.I argue that some of the variables that have only been incorporatedin pairwise fashion (or not at all) into studies of propagulesize might account for the lack of agreement about how thisimportant life history feature evolves. Further, I suggest thatfailure to consider other sources of selection on maternal phenotypes,driven by a narrow adaptationist view that has historicallybeen taken of PSMEs, has obfuscated many interesting questionssurrounding their coevolution with maternal characters. Thus,the second theme is that it is necessary to consider other explanationsfor why prop-agule size varies apart from those pertaining tooffspring fitness per si. Based on a detailed review of theempirical literature, I conclude that the concept of an optimalpropagule size is not only an insufficient construct to explainthe evolution of propagule size, but that continued relianceon an optimization approach is likely to stifle developmentof more realistic and predictive theory for the evolution ofthis key life history trait. Novel theory should incorporaterealities from physiology, development and genetics and shouldaccommodate the dynamic nature of the selective environmentsin which propagule size evolves, all of which have been shownby empiricists to play a role in determining propagule sizephenotypes. A key feature of this theory should be the explicittreatment of propagule size as a maternal effect.     

Parent–offspring conflict and selection on egg size in turtles

The trade‐off between offspring size and number can present a conflict between parents and their offspring. Because egg size is constrained by clutch size, the optimal egg size for offspring fitness may not always be equivalent to that which maximizes parental fitness. We evaluated selection on egg size in three turtle species (Apalone mutica, Chelydra serpentina and Chrysemys picta) to determine if optimal egg sizes differ between offspring and their mothers. Although hatching success was generally greater for larger eggs, the strength and form of selection varied. In most cases, the egg size that maximized offspring fitness was greater than that which maximized maternal fitness. Consistent with optimality theory, mean egg sizes in the populations were more similar to the egg sizes that maximized maternal fitness, rather than offspring fitness. These results provide evidence that selection has maximized maternal fitness to achieve an optimal balance between egg size and number.     

The causes and consequences of variation in offspring size: a case study using Daphnia

Offspring size can have large and direct fitness implications, but we still do not have a complete understanding of what causes offspring size to vary. Daphnia (water fleas) generally produce fewer and larger offspring when food is limited. Here, we use a mathematical model to show that this could be explained by either: (1) an advantage of producing larger eggs when food is limited; or (2) a lower boundary on egg volume (below which eggs do not have sufficient resources to be viable), that is similar in volume to the evolutionarily stable egg volume predicted by standard clutch size models. We tested the first possibilities experimentally by placing offspring from mothers kept at two food treatments (high and low - leading to relatively small and large eggs respectively) into two food treatments (same as maternal treatments, in a fully factorial design) and measuring their fitness (reproduction, age at maturity, and size at maturity). We also tested survival under starvation conditions of offspring produced from mothers at low and high food treatments. We found that (larger) offspring produced by low-food mothers actually had lower fitness as they took longer to reproduce, regardless of their current food treatment. Additionally, we found no survival advantage to being born of a food-stressed mother. Consequently, our results do not support the hypothesis that there is an advantage to producing larger eggs when food is limited. In contrast, data from the literature support the importance of a lower boundary on egg size.     

Rates of parasitism,but not allocation of egg resources,vary among and within hosts of a generalist avian brood parasite

Brown-headed cowbirds (Molothrus ater) deposit their eggs into the nests of other birds, which then raise the cowbird chick. Female cowbirds thus have limited options for impacting their offspring’s development via maternal effects compared to most other passerines. Cowbirds can impact their offspring’s phenotype by choosing among potential host nests, and by adjusting egg resources based on host characteristics. To examine whether cowbirds exhibit either or both of these strategies, we investigated rates of cowbird parasitism and egg investment (egg size, yolk-to-albumen ratio, and yolk testosterone and androstenedione) among and within host species in a shrubland bird community. We found that the probability of being parasitized by cowbirds, controlling for host status as a cowbird egg accepter or rejecter and ordinal date, varied significantly among host species, indicating an apparent preference for some hosts. Parasitism rates did not differ with host size, however, and despite variation in cowbird egg size among host species, this variation was not related to host size or cowbird preference. Among host species with eggs that are larger than those of the cowbird, cowbirds were significantly more likely to parasitize nests with relatively smaller eggs, whereas parasitism rates did not vary with relative egg size in host species with smaller eggs. There was no evidence for variation in cowbird egg components among or within host species. Our data indicate that cowbirds discriminate among host nests, but do not appear to adjust the composition of their eggs based on inter- or intraspecific host variation.     

Load Lightening in Southern Lapwings: Group‐Living Mothers Lay Smaller Eggs than Pair‐Living Mothers

Females of some cooperative‐breeding species can decrease their egg investment without costs for their offspring because helpers‐at‐the‐nest compensate for this reduction either by feeding more or by better protecting offspring from predation. We used the southern lapwing (Vanellus chilensis) to evaluate the effects of the presence of helpers on maternal investment. Southern lapwings are cooperative (some breeding pairs are aided by helpers), chick development is precocial, thus adults do not feed the chicks, and adults offer protection from predators through mobbing behaviors. We tested whether southern lapwing females reduced their reproductive investment (i.e. load‐lightening [LL] hypothesis) or increased their investment (i.e. differential allocation hypothesis) when breeding in groups when compared with females that bred in pairs. We found that increased group size was associated with lower egg volume. A significant negative association between the combined egg nutritional investment (yolk, protein, and lipid mass) and group size was observed. Chicks that hatched from eggs laid in nests of groups were also smaller than chicks hatched in nests of pairs. However, there was no relationship between the body mass index of chicks, or clutch size and group size, which suggests that such eggs are, simply, proportionally smaller. Our results support the LL hypothesis even in a situation where adults do not feed the chicks, allowing females to reduce investment in eggs without incurring a cost to their offspring.     

Allocation of offspring size and sex by female black ratsnakes

How females allocate resources to each offspring and how they allocate the sex of their offspring are two powerful potential avenues by which mothers can affect offspring fitness. Previous research has focussed extensively on mean offspring size, with much less attention given to variance in offspring size. Here we focussed on variation in offspring size in black ratsnakes, Elaphe obsoleta . We collected and hatched 105 clutches (1283 eggs) over 9 years. We predicted that females should lay larger eggs, or more variable eggs, when the environment is less predictable. We also predicted that females laying early or laying larger eggs should produce mostly sons because adult males are larger than adult female ratsnakes. The largest hatchling was more than twice the length and almost four times the mass of the smallest hatchling. Variation in offspring size was itself highly variable, with CVs in offspring mass among clutches ranging from 1% to 25%. With one exception, the variables we expected should influence variation in offspring size had little effect. We found that clutch size increased with maternal size and that egg size decreased with clutch size, but we found no evidence that variance in egg size among clutches increased as the season progressed or that females increased the mean size of their offspring the later in the season they laid their eggs. Females in better condition after they finish laying their eggs did produce larger eggs. There was no relationship between within-clutch variation in egg size and laying date or mean egg size. Finally, sex ratio did not vary with mean egg size or hatching date. Given evidence that offspring size in snakes affects survival, selection should reduce variation in offspring size unless that variance enhances maternal fitness and yet we found little support for hypothesized advantages of varying offspring size.     

Egg size plasticity corresponding to maternal food conditions in an annual fish,Ayu <Emphasis Type="Italic">Plecoglossus altivelis</Emphasis>

The classic model of Smith and Fretwell predicts that the optimal egg size will vary according to the shape of the relationship between offspring size and offspring fitness, which may vary among environments. Adaptive significance of intrapopulation egg size variation was examined using Ayu (Plecoglossus altivelis). The species has an annual and migratory life history. Fish under controlled rearing conditions become sexually mature with a trend that smaller females produced larger eggs later in the season. Observed egg size variation was explained by the maternal specific growth rate, which was composed of maternal body size and growing period. Hatchlings from larger eggs had a larger notochord length, larger yolk-sac and grew faster. Such offspring traits provide general advantages of increased larval size, which confer competitive ability for assuring early survivorship. In conclusion, egg size plasticity in Ayu suggests higher offspring fitness through enhancement of their accessibility to food.     

Correlates of egg size variation in a population of house sparrow Passer domesticus

Propagule size represents an important life-history trait under maternal control. Despite a positive relationship between propagule size and components of fitness, propagule size displays tremendous amounts of variation which causes are poorly understood within natural populations. With a study of a house sparrows Passer domesticus, we investigate maternal and environmental correlates of egg size, quantify variation in egg size within and between females and broods, and estimate heritability. Egg size had a curvilinear relationship with clutch size and decreased significantly in subsequent broods within seasons. Furthermore, egg size increased with maternal body mass, was positively affected by spring temperatures and curvilinearly related to temperature during the 2 weeks prior to egg laying. Some 46.4 % of variation in egg size was due to differences between females, and 21.9 % was explained by variation between broods by the same female. The heritability of egg size was low (h ² = 0.26) compared to estimates from other studies (h ² > 0.6). The present study challenges the recent idea that egg size is an inflexible maternal characteristic with very high additive genetic variance, and suggests that females are subject to both intrinsic and extrinsic constraints prior to and during egg formation, leading to the observed plasticity in egg size. In a general sense, propagule size could be expected to be both limited by and adaptively adjusted in accordance to prevailing environmental conditions.     

The effect of egg size and habitat on starling nestling growth and survival

In spite of the fact that hatchling size and energy reserves in birds are affected by egg size, many studies have failed to find an effect of egg size on offspring fitness. One possibility is that this is because they have been performed in areas with high food availability and that effects of egg size on offspring fitness are most apparent in areas of low food availability. To investigate this, egg size,␣offspring mass and survival of European starlings (Sturnus vulgaris) were measured in an agricultural landscape with a low but variable amount of pasture, the preferred foraging habitat of parent starlings. Offspring mass was related to egg size, but egg size explained a declining proportion of the variation in nestling mean mass as nestlings grew older. Offspring survival during the early, but not during the late nestling period was related to egg size. Throughout the nestling␣period, survival was related to the mass of the nestlings. It is suggested that the effect of egg size on␣offspring survival is through the effect of egg size on offspring mass, this effect declining as offspring grow older. Offspring survival during the early part of the nestling period was related to egg size when availability of pasture was low, but not when it was high. However, the interaction was not significant. Selection for␣larger egg size is discussed in relation to the structuring␣of starling populations into sources and sinks. Received: 22 September 1997 / Accepted: 22 January 1998     

INTERACTIVE EFFECTS OF OFFSPRING SIZE AND TIMING OF REPRODUCTION ON OFFSPRING REPRODUCTION: EXPERIMENTAL,MATERNAL, AND QUANTITATIVE GENETIC ASPECTS

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season—a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect—experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.     

Egg load is a cue for offspring sex ratio adjustment in a fig‐pollinating wasp with male‐eggs‐first sex allocation

Fig‐pollinating wasps (Agaonidae) only reproduce within fig tree inflorescences (figs). Agaonid offspring sex ratios are usually female‐biased and often concur with local mate competition theory (LMC). LMC predicts less female‐bias when several foundresses reproduce in a fig due to reduced relatedness among intra‐sexually competing male offspring. Clutch size, the offspring produced by each foundress, is a strong predictor of agaonid sex ratios and correlates negatively with foundress number. However, clutch size variation can result from several processes including egg load (eggs within a foundress), competition among foundresses and oviposition site limitation, each of which can be used as a sex allocation cue. We introduced into individual Ficus racemosa figs single Ceratosolen fusciceps foundresses and allowed each to oviposit from zero to five hours thus variably reducing their eggs‐loads and then introduced each wasp individually into a second fig. Offspring sex ratio (proportion males) in second figs correlated negatively with clutch size, with males produced even in very small clutches. Ceratosolen fusciceps lay mainly male eggs first and then female eggs. Our results demonstrate that foundresses do not generally lay or attempt to lay a ‘fixed’ number of males, but do ‘reset to zero’ their sex allocation strategy on entering a second fig. With decreasing clutch size, gall failure increased, probably due to reduced pollen. We conclude that C. fusciceps foundresses can use their own egg loads as a cue to facultatively adjust their offspring sex ratios and that foundresses may also produce more ‘insurance’ males when they can predict increasing rates of offspring mortality.     

Experimental (antiestrogen-mediated) reduction in egg size negatively affects offspring growth and survival

The relationship between egg size and offspring phenotype is critical to our understanding of the selective pressures acting on the key reproductive life-history traits of egg size and number. Yet there is surprisingly little empirical evidence to support a strong, positive relationship between egg size and offspring quality (i.e., offspring growth, condition, and survival) in birds, in part because of confounding effects of parental quality and the lack of experimental techniques for directly manipulating avian egg size independently of maternal condition. Previously, we showed that treatment of laying female zebra finches (Taeniopygia guttata) with the antiestrogen tamoxifen can decrease egg size by ca. 8% but that this reduction in egg size had few effects on offspring mass and size at fledging. Here, we extend the use of this technique to induce larger decreases in egg size (up to 50% in individual females) and show that a reduction in egg size of ca. 18% is associated with decreased embryo viability, increased hatchling mortality, and lower posthatching offspring survival. Furthermore, we show that although hatchlings from eggs reduced in size by ca. 9% can survive to fledging, these chicks show slower initial growth during the linear growth phase (5-10 d of age), fledge at lower masses than chicks from control eggs, and show postfledging compensatory growth. Our results provide empirical support for significant effects of egg size on offspring quality and further suggest that among individual females there is a minimum egg size required to maintain embryo viability and offspring quality.     

Of chickens and eggs: diverging propagule size of iteroparous and semelparous organisms

Interactive effects of two or more life-history traits on fitness have the potential to create suites of coadapted traits. Propagule (egg or seed) size is one such trait that is believed to have undergone coadaptation with other traits. Phylogenetic analyses of salmonid fishes have revealed an association between large eggs and semelparity, leading to the question of which came first. It has been hypothesized that an increased egg size would have increased juvenile relative to adult survival, favoring a subsequent increase in reproductive effort and eventually semelparity. Others have suggested that this is insufficient to cause a shift in parity, implying to the contrary that semelparity gave rise to larger eggs. In a previous study we showed that environmental unpredictability might select for production of larger propagules. Here we use simulations to directly model how propagule size evolves in response to environmental unpredictability with varying degrees of iteroparity. Our results demonstrate that environmental unpredictability causes pronounced propagule size divergence between iteroparous and purely semelparous species in taxa with a fixed age at maturity (e.g., pure annual species). However, even rare incidents of repeat breeding are sufficient to reduce selection for larger propagules substantially and thus divergence. Furthermore, introducing variation in age at maturity within propagule size genotypes has evolutionary effects similar to that of repeat breeding. Environmental unpredictability is thus unlikely to provide a general alternative explanation for the observed egg size divergence between iteroparous and semelparous salmonids.     

Climate change can reduce the risk of biological invasion by reducing propagule size

Climate change has been conclusively linked to species extinctions, and to expansion and contractions and shifts of species ranges. Climate change is exerting similarly profound pressures on the individual stages of biological invasion which can significantly impact the biodiversity and ecology of invaded areas. Propagule pressure is perhaps the single most important determinant of invasion success, but the effects of climate change on propagule pressure are still largely uncertain because we have few observations of introduction events (or their size) that can be analyzed together with climate records. The common surrogate variables for propagule pressure do not logically respond to climate. Here I use a process-based simulation model to examine the potential effects of climate change (specifically temperature) on propagule size of a common invading insect species by estimating in-transit survivorship rate of propagules using historical and future (projected) temperatures and two common trade routes between a donor and a recipient location (Yokohama, Japan and Sydney, Australia). Propagule size (=the number of individuals in an introduction event) was lower under climate change temperatures than under historical temperatures in both routes. The route had significant effects on propagule size through its influence on the duration (and also the timing) of exposure to temperature conditions that are of time-sensitive importance to the development of the invasive species. Under historical temperatures propagule size was higher and less variable in the direct than the indirect route in 20 independent iterations. Under the future temperatures propagule size was also higher in the direct route but it was more variable than in the indirect route. Increased trade is increasing the opportunities for introductions, but the results reported here suggest that climate change will have inconsistent effects on biological invasion because of the complex relationship between temperature and insect ontogeny.