Latitudinal and seasonal patterns in clutch size of some single‐brooded British birds

Capsule Daylength, rather than latitude, was found to be an important determinant of variation in clutch size.

Aims To describe the nature of spatial and temporal variation in clutch size, and explore the ecological correlates of these patterns.

Methods We tested the prediction that seasonal declines in clutch size will be greater at higher latitudes. The environmental variables focused on were the influence of daylength, plant productivity, seasonality (i.e. Ashmole's hypothesis) and physiological mechanisms that relate clutch size to ambient temperature. We used data from 1980 to 2003 on spatial variation in clutch size across Britain for single‐brooded species, in which clutch size can be taken as a measure of annual reproductive investment. We included all seven species, from five families, with sufficient data in the British Trust for Ornithology's Nest Record Scheme.

Results There are strong seasonal declines in clutch size but little evidence for latitudinal gradients in clutch size or in latitudinal gradients in the rate of seasonal clutch size decline. Of the environmental variables investigated, daylength had the most marked effect on clutch size; this was positive in diurnal species and negative in the one nocturnal species.

Conclusions Although this study was confined to a relatively small latitudinal range of 8°, we found marked latitudinal gradients in a number of factors thought to drive spatial patterns in clutch size. Moreover, such variation is of sufficient magnitude to generate spatial patterns in other ecological variables in Britain. There is thus no simple explanation for the lack of a latitudinal gradient in clutch size. The results concerning daylength indicate that the time available for foraging is an important determinant of variation in clutch size.     

Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited

There are two major competing hypotheses for variation in clutch size among cavity-nesting species. The nest site limitation hypothesis postulates that nesting opportunities are more limited for weak excavators, which consequently invest more in each breeding attempt by laying larger clutches. Alternatively, clutch size may be determined by diet; the clutch sizes of strong excavators may be smaller because they are able to specialize on a more seasonally stable prey. We built a conceptual model that integrated hypotheses for interspecific variation in clutch size and tested it with comparative data on life-history traits of woodpeckers (Picidae) and nuthatches (Sittidae). In most analyses, diet explained more variation in clutch size among species than did propensity to excavate. Migratory status was positively associated with clutch size but was difficult to distinguish from diet since resident species consumed more bark beetles (a prey available in winter) and had smaller clutches than migratory species. The literature suggests that cavities are not limited in natural, old-growth forests. Although our data do not rule out nest site limitation, we conclude that annual stability of food resources has a larger impact on the evolution of clutch sizes in excavators than does limitation of nest sites.     

Ecological and environmental factors related to variation in egg size of New World flycatchers

Geographical variation in egg size is well documented for several taxa, but remains insufficiently described for birds in spite of a well‐known latitudinal gradient in clutch size. Several hypotheses have been proposed to explain avian egg size variation; however, they were not tested on a continental scale. Egg size is a key component of reproductive investment that influences offspring fitness. It is thought to vary geographically as one of a set of correlated life‐history traits that are under selection from varying ecological conditions. We completed a comprehensive literature review and calculated egg sizes for the most widespread clade within tyrant flycatchers, describing for the first time the geographical variation in egg size on a continental scale. We examined the relative support for ecological and environmental variables in explaining egg size variation using multi‐model inference and linear mixed models controlled for phylogenetic autocorrelation among species. We tested five hypotheses and found that: larger eggs occur in colder sites, which is consistent with the embryonic temperature hypothesis; medium/long‐distance migrants had smaller eggs than resident species while short‐distance migrants had the largest eggs; neither species clutch size, nor species nest type, nor evapotranspiration seasonality influenced egg size. Avian egg size is larger in Austral and Neotropical America (ANA), where species are resident or short‐distance migrants, and smaller across the medium/long‐distance migrants of the Nearctic region. In addition, while clutch size increases towards higher northern latitudes and is almost invariable across ANA species, egg sizes vary largely across ANA sites, increasing with southern latitudes and higher elevations and being influenced by summer temperature. While the embryonic temperature hypothesis has been usually linked to parental nest attentiveness, we highlight that environmental temperatures also have strong effects in shaping investment in egg size.     

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.     

FACTORS DETERMINING A CLUTCH SIZE REDUCTION IN CALIFORNIA GULLS (LARUS CALIFORNICUS): A MULTI-HYPOTHESIS APPROACH

In the thirty-five years since David Lack first highlighted the importance of clutch size, a large number of hypotheses have been proposed relating clutch size variation to various environmental and demographic factors. Despite a great deal of both empirical and theoretical work on clutch size, there has been very little effort to test many of the competing hypotheses in explaining a clutch size difference between two populations of the same species. I have taken the latter approach in an effort to explain a clutch size reduction in the California Gull (Larus californicus) population at Mono Lake, California. I compared the breeding biologies of the gulls at Mono Lake and at Great Salt Lake, Utah, collecting data for three breeding seasons at Mono Lake and one breeding season at Great Salt Lake. These data included measurements of the conditions of 60 adults, growth and mortality measurements for approximately 900 chicks, 4450 nest-hours of parental care observations, and the results of egg-removal experiments on 40 females. I tested seven hypotheses to explain the clutch size reduction: age structure, egg predation, bet-hedging, effort reallocation, most productive brood size, parental mortality, and pre-egg food limitation. Each of these hypotheses is described in detail in the introduction. The pre-egg food limitation hypothesis is best able to explain the clutch size reduction at Mono Lake, although the egg-removal experiments show that the resource limitation is relative and not absolute. Clutch size variation at each site need not be viewed as the result of fine-scaled evolutionary adjustment, although the general clutch size decision machinery is presumably molded by selection. Future research must focus on the details of this clutch size decision machinery and its application to the concept of reproductive effort.     

A comparative test of adaptive hypotheses for sexual size dimorphism in lizards

It is commonly argued that sexual size dimorphism (SSD) in lizards has evolved in response to two primary, nonexclusive processes: (1) sexual selection for large male size, which confers an advantage in intrasexual mate competition (intrasexual selection hypothesis), and (2) natural selection for large female size, which confers a fecundity advantage (fecundity advantage hypothesis). However, outside of several well-studied lizard genera, the empirical support for these hypotheses has not been examined with appropriate phylogenetic control. We conducted a comparative phylogenetic analysis to test these hypotheses using literature data from 497 lizard populations representing 302 species and 18 families. As predicted by the intrasexual selection hypothesis, male aggression and territoriality are correlated with SSD, but evolutionary shifts in these categorical variables each explain less than 2% of the inferred evolutionary change in SSD. We found stronger correlations between SSD and continuous estimates of intrasexual selection such as male to female home range ratio and female home range size. These results are consistent with the criticism that categorical variables may obscure much of the actual variation in intrasexual selection intensity needed to explain patterns in SSD. In accordance with the fecundity advantage hypothesis, SSD is correlated with clutch size, reproductive frequency, and reproductive mode (but not fecundity slope, reduced major axis estimator of fecundity slope, length of reproductive season, or latitude). However, evolutionary shifts in clutch size explain less than 8% of the associated change in SSD, which also varies significantly in the absence of evolutionary shifts in reproductive frequency and mode. A multiple regression model retained territoriality and clutch size as significant predictors of SSD, but only 16% of the variation in SSD is explained using these variables. Intrasexual selection for large male size and fecundity selection for large female size have undoubtedly helped to shape patterns of SSD across lizards, but the comparative data at present provide only weak support for these hypotheses as general explanations for SSD in this group. Future work would benefit from the consideration of alternatives to these traditional evolutionary hypotheses, and the elucidation of proximate mechanisms influencing growth and SSD within populations.     

Colony size and foraging range in seabirds

The reasons for variation in group size among animal species remain poorly understood. Using ‘Ashmole's halo’ hypothesis of food depletion around colonies, we predict that foraging range imposes a ceiling on the maximum colony size of seabird species. We tested this with a phylogenetic comparative study of 43 species of seabirds (28 262 colonies), and investigated the interspecific correlation between colony size and foraging ranges. Foraging range showed weak relationships with the low percentiles of colony size of species, but the strength of the association increased for larger percentiles, peaking at the maximum colony sizes. To model constraints on the functional relationship between the focal traits, we applied a quantile regression based on maximum colony size. This showed that foraging range imposes a constraint to species’ maximum colony sizes with a slope around 2. This second‐order relationship is expected from the equation of the area of a circle. Thus, our large dataset and innovative statistical approach shows that foraging range imposes a ceiling on seabird colony sizes, providing strong support to the hypothesis that food availability is an important regulator of seabird populations.     

The evolution of different maternal investment strategies in two closely related desert vertebrates

We compared egg size phenotypes and tested several predictions from the optimal egg size (OES) and bet‐hedging theories in two North American desert‐dwelling sister tortoise taxa, Gopherus agassizii and G. morafkai, that inhabit different climate spaces: relatively unpredictable and more predictable climate spaces, respectively. Observed patterns in both species differed from the predictions of OES in several ways. Mean egg size increased with maternal body size in both species. Mean egg size was inversely related to clutch order in G. agassizii, a strategy more consistent with the within‐generation hypothesis arising out of bet‐hedging theory or a constraint in egg investment due to resource availability, and contrary to theories of density dependence, which posit that increasing hatchling competition from later season clutches should drive selection for larger eggs. We provide empirical evidence that one species, G. agassizii, employs a bet‐hedging strategy that is a combination of two different bet‐hedging hypotheses. Additionally, we found some evidence for G. morafkai employing a conservative bet‐hedging strategy. (e.g., lack of intra‐ and interclutch variation in egg size relative to body size). Our novel adaptive hypothesis suggests the possibility that natural selection favors smaller offspring in late‐season clutches because they experience a more benign environment or less energetically challenging environmental conditions (i.e., winter) than early clutch progeny, that emerge under harsher and more energetically challenging environmental conditions (i.e., summer). We also discuss alternative hypotheses of sexually antagonistic selection, which arise from the trade‐offs of son versus daughter production that might have different optima depending on clutch order and variation in temperature‐dependent sex determination (TSD) among clutches. Resolution of these hypotheses will require long‐term data on fitness of sons versus daughters as a function of incubation environment, data as yet unavailable for any species with TSD.     

Sensory control of clutch size in the Common Swift Apus apus

Clutch size varies among individuals in most bird species. A widespread assumption is that such variation results from variable timing in the disruption of ovarian follicular growth that brings, with a few days' lag, egg‐laying to an end. Currently, there is empirical evidence that this is the case in Blue Tits Cyanistes caeruleus but not in Zebra Finches Taeniopygia guttata, in which the timing of follicular disruption has been shown to be invariant. Here, I investigate clutch size regulation of Common Swifts Apus apus. Using experimental egg removal, I show that, assuming the female gets enough egg contact, the determination of clutch size occurs at noon solar time on the day the first egg of a clutch is laid. In spite of individual variation in clutch size, there was no hint of any variability in the timing of clutch‐size determination. In this species therefore, the timing of the signal disrupting follicular growth appears invariant. The physiological mechanism that controls clutch size is discussed, including the existence of an endogenous circadian clock and potential zeitgeber, the developmental range of clutch size for the species, the sensory nature of the input that triggers follicular disruption, and the stress of laying extra eggs.     

Origin of evolutionary change in avian clutch size

Why different bird species lay different numbers of eggs is a question that has long been associated with factors external to the organism, that is, factors which operate on inherited variation in clutch size through the action of natural selection. Yet, while external factors are important, the extent of what is evolutionarily possible rests with the mechanisms developed by birds for clutch‐size control. Hitherto neglected, these mechanisms generate factors internal to the organism that are central to the origin of evolutionary change. They are related to the fact that a species‐specific range of clutch size arises from the differential survival of pre‐ovulatory follicles undergoing growth when the signal causing egg laying to end reaches the ovary. Herein, I examine three internal factors that, together with external factors, could impact the evolution of avian clutch size. Each factor acts by changing either the number of pre‐ovulatory follicles present in the ovary at the time of follicular disruption or the timing of this event. These changes to clutch size can be explained by the concept of heterochrony. In light of this, the role of phenotypic plasticity and genes determining clutch size is discussed. Finally, to account for the origin of evolutionary change in clutch size, I detail an hypothesis involving a process similar to Waddington's theory of genetic assimilation.     

Litter size and latitude in a large mammal: the wild boar Sus scrofa

• 1 A positive relationship between clutch size or litter size and latitude exists in birds and many species of small mammal. Hitherto, however, analyses for large mammals have failed to provide evidence that litter sizes increase with latitude.
• 2 We collated data from published studies of wild boar in Europe, to analyse the relationship between litter size and latitude in this widely distributed terrestrial mammal.
• 3 Depending on the specific data set (whether only the most reliable data or all available data were included), latitude explained 58% to 72% of the variation in mean litter sizes across studies. On average, litter size increases by approximately 0.15 piglets per degree of latitude.
• 4 A strong correlation between litter size and latitude for wild boar in Europe provides a starting point for demographic modelling of this species of both ecological and economic importance.
• 5 The pattern for wild boar is consistent with Ashmole's explanation for the effects of latitude on reproduction. The contrast between our results and those generated for other large mammals may result from our focus on an herbivore in contrast to previous work which was focused on carnivores. Further work could usefully examine the extent of seasonality in the availability of resources for species of different dietary types.

     

Clutch size variation in the Nazca booby: a test of the egg quality hypothesis

In obligately siblicidal bird species, aggressive behavior bya dominant chick results in a fixed brood size of one, yetthese species usually show clutch size variation between individuals.Simmons proposed that variation in clutch size in obligatelysiblicidal species is related to a trade-off between egg qualityand egg quantity: some individuals produce a single highly hatchable egg, while others produce two small, lower qualityeggs. We tested the egg quality hypothesis as an explanationfor observed clutch size variation in the Nazca booby (Sulagranti), an obligately siblicidal seabird. We tested the assumptionthat egg volume is positively correlated with hatchabilityand the prediction that eggs from one-egg clutches are largerthan eggs from two-egg clutches. We did not find a positive relationship between egg volume and hatchability in this species.Eggs from two-egg clutches were either equivalent in volumeor larger than eggs from one-egg clutches. Thus, the egg qualityhypothesis was rejected as an explanation for clutch size variationin the Nazca booby. Instead, two-egg clutches appear to befavored because of the insurance value of the second-laid egg,while one-egg clutches result from food limitation.     

Do Rapoport's Rule,Mid-Domain Effect or Environmental Factors Predict Latitudinal Range Size Patterns of Terrestrial Mammals in China?

Background

Explaining species range size pattern is a central issue in biogeography and macroecology. Although several hypotheses have been proposed, the causes and processes underlying range size patterns are still not clearly understood. In this study, we documented the latitudinal mean range size patterns of terrestrial mammals in China, and evaluated whether that pattern conformed to the predictions of the Rapoport''s rule several analytical methods. We also assessed the influence of the mid-domain effect (MDE) and environmental factors on the documented range size gradient.

Methodology/Principal Findings

Distributions of 515 terrestrial mammals and data on nine environmental variables were compiled. We calculated mean range size of the species in each 5° latitudinal band, and created a range size map on a 100 km×100 km quadrat system. We evaluated Rapoport''s rule according to Steven''s, mid-point, Pagel''s and cross-species methods. The effect of the MDE was tested based on a Monte Carlo simulation and linear regression. We used stepwise generalized linear models and correlation analyses to detect the impacts of mean climate condition, climate variability, ambient energy and topography on range size. The results of the Steven''s, Pagel''s and cross-species methods supported Rapoport''s rule, whereas the mid-point method resulted in a hump-shaped pattern. Our range size map showed that larger mean latitudinal extents emerged in the mid-latitudes. We found that the MDE explained 80.2% of the range size variation, whereas, environmental factors accounted for <30% of that variation.

Conclusions/Significance

Latitudinal range size pattern of terrestrial mammals in China supported Rapoport''s rule, though the extent of that support was strongly influenced by methodology. The critical factor underlying the observed gradient was the MDE, and the effects of climate, energy and topography were limited. The mean climate condition hypothesis, climate variability hypothesis, ambient energy hypotheses and topographical heterogeneity hypotheses were not supported.     

Global patterns of body size evolution are driven by precipitation in legless amphibians

Body size shapes ecological interactions across and within species, ultimately influencing the evolution of large‐scale biodiversity patterns. Therefore, macroecological studies of body size provide a link between spatial variation in selection regimes and the evolution of animal assemblages through space. Multiple hypotheses have been formulated to explain the evolution of spatial gradients of animal body size, predominantly driven by thermal (Bergmann's rule), humidity (‘water conservation hypothesis’) and resource constraints (‘resource rule’, ‘seasonality rule’) on physiological homeostasis. However, while integrative tests of all four hypotheses combined are needed, the focus of such empirical efforts needs to move beyond the traditional endotherm–ectotherm dichotomy, to instead interrogate the role that variation in lifestyles within major lineages (e.g. classes) play in creating neglected scenarios of selection via analyses of largely overlooked environment–body size interactions. Here, we test all four rules above using a global database spanning 99% of modern species of an entire Order of legless, predominantly underground‐dwelling amphibians (Gymnophiona, or caecilians). We found a consistent effect of increasing precipitation (and resource abundance) on body size reductions (supporting the water conservation hypothesis), while Bergmann's, the seasonality and resource rules are rejected. We argue that subterranean lifestyles minimize the effects of aboveground selection agents, making humidity a dominant selection pressure – aridity promotes larger body sizes that reduce risk of evaporative dehydration, while smaller sizes occur in wetter environments where dehydration constraints are relaxed. We discuss the links between these principles with the physiological constraints that may have influenced the tropically‐restricted global radiation of caecilians.     

Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Ecological predictors of reduced avian reproductive investment in the southern hemisphere

Despite intensive research, the factors driving spatial patterns in life‐history traits remain poorly understood. One of the most frequently documented, and paradoxically, least understood patterns, is the latitudinal gradient of increasing avian clutch size at higher latitudes. These gradients are less marked in the southern hemisphere, thus clutch sizes tend to be smaller at southern latitudes than at equivalent northern ones. We exploited a natural experiment provided by the introduction of European passerines to New Zealand (NZ) to test three widely proposed ecological drivers of this pattern, i.e. the nest predation, Ashmole’s seasonality, and the breeding density hypotheses. We focus on the blackbird Turdus merula and the song thrush T. philomelos as founder effects do not have a major influence on the reproductive traits of their introduced populations. Both species laid smaller clutches in NZ than in Europe. These reductions had stabilised within one hundred years and were not associated with a compensatory increase in investment in individual offspring by laying larger eggs. In contrast to the nest predation hypothesis, daily nest predation rates were lower in NZ than in Europe. Smaller southern hemisphere clutches were associated with higher conspecific population densities and a relaxation of seasonal clutch size trends. These findings thus provide some support for both Ashmole’s seasonality and the breeding density hypotheses. Analyses across 11 European passerines introduced to NZ suggest, however, that neither of these hypotheses provide general explanations of smaller clutches in the southern hemisphere. We suggest that reduced seasonality and lower nest predation promote increased breeding densities and adult survival in the southern hemisphere. The later may drive smaller southern clutch sizes by generating spatial variation in the outcome of the trade‐off between reproductive investment and longevity.     

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.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

A comparative study of body size and clutch size across the parasitoid Hymenoptera

Across animal species, body size and clutch size often form part of a suite of associated life history traits, exemplified by the "fast-slow continuum" in mammals. Across the parasitoid Hymenoptera however, a major axis of life history variation is the development mode of the larva (koinobiosis versus idiobiosis), and body size and clutch size do not seem to form clear associations with this major axis. Here we use a large comparative data set and the latest phylogenetic information to explore hypotheses that might explain the variation in body size and clutch size across species in parasitoids. We find evidence for three novel evolutionary correlations: changes in the stage of host attacked by the parasitoid (i.e. egg, larva, pupa) significantly predict changes in both body size and clutch size, whilst in gregarious species changes to higher latitudes are associated with reduced clutch size. We also find a number of hypothesized cross-species (phenotypic) associations that, however, we cannot demonstrate are the result of evolutionary correlations: large bodied species in our data tend to lay small clutches; koinobionts are larger than idiobionts attacking the same host stage; tropical species are smaller than temperate species (Bergmann's rule). Our results provide support for theoretical models of trait evolution in parasitoids, whilst the associations between latitude and life history may help explain why species richness in the family Ichneumonidae peaks at intermediate latitudes. Our results also show the continuing value of phylogenetically-based comparative analyses and demonstrate that recent work on parasitoid phylogenetics has produced significant benefits for our understanding of life history evolution.     

MATERNAL INHERITANCE OF CONDITION AND CLUTCH SIZE IN THE COLLARED FLYCATCHER

Maternal effects may strongly influence evolutionary response to natural selection but they have been little studied in the wild. We use a novel combination of experimental and statistical methods to estimate maternal effects on condition and clutch size in the collared flycatcher, where we define “condition” to be the nongenetic component of clutch size. We found evidence of two maternal effects. The first (m) was the negative effect of mother's clutch size on daughter's condition, when mother's condition was held constant. The second (M) was the positive effect of mother's condition on daughter's condition, when mother clutch size was held constant. These two effects oppose one another because mothers in good condition also lay many eggs. The maternal effects were large: Experimentally adding an egg to a mother's nest reduced clutch sizes of her daughters by 1/4 egg (i.e., m = -0.25). Measured degree of resemblance between mother and daughter clutch sizes yielded M = 0.43. The results weakly support the presence of heritable genetic variation in clutch size: additive genetic variance/total phenotypic variance = 0.33. This estimate was highly variable probably because, as we show, mother-daughter resemblance may depend hardly at all on the amount of genetic variance when maternal effects are present. Daughter-mother regression (a standard method for estimating heritability) is consequently a poor guide to the amount of genetic variance in clutch size. Our results emphasize the value of combining field experiments with observations for studying inheritance.