Body condition,migration, and timing of reproduction in snow geese: a test of the condition-dependent model of optimal clutch size

The seasonal decline of avian clutch size may result from the conflict between the advantage of early breeding (greater offspring value) and the advantage of a delay in lay date (improved body condition and hence clutch size). We tested predictions of a condition-dependent individual optimization model based on this trade-off (Rowe et al. 1994) in a long-distance migrant, the greater snow goose (Chen caerulescens atlantica), using data on condition, migration, and reproductive decisions of individuals. We closely tracked radio-marked females at their main spring staging area and on their breeding grounds. Our results were consistent with predictions of the model. Early-arriving females had a longer prelaying period and initiated their nests earlier than late arrivals. After controlling statistically for arrival date, we determined that females with high premigration condition had an earlier lay date than those in low condition. After controlling for the seasonal decline (i.e., lay date), we observed that clutch size was not related to premigration condition. Moreover, we took advantage of an unplanned manipulation of the prebreeding condition that occurred during our long-term study. We found that a reduction in condition caused a delay in lay date. However, after controlling for the seasonal decline, it did not affect clutch size. Our study indicates that geese simultaneously adjust their lay date and clutch size according to their premigration condition and migratory behavior as predicted by the condition-dependent optimization model.     

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.     

Seasonal declines in incubation periods of Brünnich's Guillemots Una lomvia:testing proximate causes

In many species of bird, eggs laid late in the laying period hatch after a shorter incubation period than do eggs laid early. These seasonal declines in incubation period are generally thought to confer evolutionary advantages, but the proximate mechanisms that underlie them are poorly understood. Seasonal declines in incubation period are usually attributed to: (1) seasonal increases in ambient air temperatures; (2) seasonal changes in the behaviour of incubating birds; and/or (3) seasonal declines in egg size. In a previous study, Common Guillemot Una aalge incubation periods declined with laying date at a low-Arctic colony. As there was no support for hypotheses 1 or 2, it was suggested that this occurred because egg size declined with laying date, but eggs were not measured in that study. We recorded similar seasonal declines in the incubation periods of the single eggs laid by Brünnich's Guillemots Una lomvia at two low-Arctic colonies in four years. Neither seasonal variation in ambient air temperatures, nor in the behaviour of incubating adults, appeared to cause the declines. As predicted for Common Guillemots, incubation period increased with egg size among Briinnich's Guillemots, in one of two years. However, incubation period declined with laying date in the absence of corresponding declines in egg size. We conclude that none of the three commonly proposed proximate mechanisms adequately explains the seasonal variation in guillemot incubation periods. Several testable, alternative mechanisms are explored.     

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.     

Seasonal clutch size variation of multi-brooded bird species: comparisons between breeding season and latitudes

Single- and multi-brooded species of birds differ in their seasonal patterns of clutch size. Single-brooded species start with a maximum clutch size that declines continuously as the season progresses, whereas the clutch sizes of multi-brooded species usually increase to a mid-season maximum peak and then decrease progressively until the end of the breeding season. Previous studies have shown that multi-brooded migrant species present seasonal patterns that are similar to single-brooded species at high latitudes but similar to multi-brooded non-migratory species at lower latitudes. We studied the Greenfinch Carduelis chloris and Goldfinch C. carduelis populations in eastern Spain (Sagunto, Valencia) between 1975 and 2002 to compare seasonal variations in clutch sizes between years with early and late starts to the breeding season. The period over which clutch sizes increase was longer when the breeding season started earlier. The Goldfinch population showed no pattern of initial increase in clutch size when there was a late start to the breeding season: a late start shortens the season giving them less time to breed, and may also coincide with maximum food availability. Thus, the pattern of single-brooded species was observed. In the Greenfinch population, a trend toward the seasonal pattern of single-brooded species was also observed when the following indices were compared: clutch size increase, modal timing, initial slope and timing of maximum clutch size. We have also compared the seasonal patterns of clutch size of both species in eastern Spain with the patterns observed in Britain. Our results show that for both Goldfinches and Greenfinches, the non-migrant southern populations of Sagunto in eastern Spain do not tend towards a more multi-brooded seasonal pattern of clutch size than the migrant Goldfinches of Britain.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

Ashmole's hypothesis and the latitudinal gradient in clutch size

One enduring priority for ecologists has been to understand the cause(s) of variation in reproductive effort among species and localities. Avian clutch size generally increases with increasing latitude, both within and across species, but the mechanism(s) driving that pattern continue to generate hypotheses and debate. In 1961, a Ph.D. student at Oxford University, N. Philip Ashmole, proposed the influential hypothesis that clutch size varies in direct proportion to the seasonality of resources available to a population. Ashmole's hypothesis has been widely cited and discussed in the ecological literature. However, misinterpretation and confusion has been common regarding the mechanism that underlies Ashmole's hypothesis and the testable predictions it generates. We review the development of well-known hypotheses to explain clutch size variation with an emphasis on Ashmole's hypothesis. We then discuss and clarify sources of confusion about Ashmole's hypothesis in the literature, summarise existing evidence in support and refutation of the hypothesis, and suggest some under-utilised and novel approaches to test Ashmole's hypothesis and gain an improved understanding of the mechanisms responsible for variation in avian clutch size and fecundity, and life-history evolution in general.     

Capital energy drives production of multiple clutches whereas income energy fuels growth in female collared lizards Crotaphytus collaris

Animals invest energy in reproduction that is obtained at two distinct times relative to the reproductive cycle. Energy obtained during egg production is referred to as income energy whereas stored energy acquired prior to reproduction is capital energy. Similar to most ectotherms, squamate reptiles are generally hypothesized to be capital breeders. Nearly all squamates in which income/capital energy investment has been examined thus far produce only one clutch per reproductive season. Although it is likely that squamates producing multiple seasonal clutches fuel first clutches with capital energy, either capital or income energy may be used to produce later clutches. We first monitored female eastern collared lizards over 14 reproductive seasons to confirm that the number of clutches females produce seasonally is a plastic response to variable environmental parameters, and to examine the effects of female body condition at the beginning of the reproductive season on clutch production. Clutch production varied annually and both the size and number of clutches were positively correlated with body condition. We then tested the competing predictions of the income and capital hypotheses experimentally by supplementing the diets of female collared lizards in situ for one season. Diet‐supplementation had no effect on the number of clutches produced but increased growth rates of gravid females. We further tested the competing predictions of these two hypotheses by examining variation in maternal energy investment per clutch using preserved specimens collected near our primary field site. Clutch size was highly correlated with female body size. Together, our results suggest that variation in reproductive output by female collared lizards is linked to stored capital energy rather than income energy, similar to most ectotherms.     

Maternal investment in reproduction and its consequences in leatherback turtles

Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.     

Cyclic variation in seasonal recruitment and the evolution of the seasonal decline in Ural owl clutch size

Plastic life-history traits can be viewed as adaptive responses to environmental conditions, described by a reaction norm. In birds, the decline in clutch size with advancing laying date has been viewed as a reaction norm in response to the parent's own (somatic or local environmental) condition and the seasonal decline in its offspring's reproductive value. Theory predicts that differences in the seasonal recruitment are mirrored in the seasonal decrease in clutch size. We tested this prediction in the Ural owl. The owl's main prey, voles, show a cycle of low, increase and peak phases. Recruitment probability had a humped distribution in both increase and peak phases. Average recruitment probability was two to three times higher in the increase phase and declined faster in the latter part of the season when compared with the peak phase. Clutch size decreased twice as steep in the peak (0.1 eggs day-1) as in the increase phase (0.05 eggs day-1). This result appears to refute theoretical predictions of seasonal clutch size declines. However, a re-examination of current theory shows that the predictions of modelling are less robust to details of seasonal condition accumulation in birds than originally thought. The observed pattern can be predicted, assuming specifically shaped seasonal increases in condition across individuals.     

Seasonal clutch size decline and individual variation in the timing of breeding are related to female body condition in a non-migratory species, the Upland Goose Chloephaga picta leucoptera

Several studies have shown that waterfowl females in good condition lay larger clutches and start egg-laying earlier in a breeding season. However, most of these studies lack corresponding data on individual condition and timing of breeding in different years. We analysed data on clutch size, egg size, hatching date and female body condition of a non-migratory species, the Upland Goose Chloephaga picta leucoptera, recorded in three seasons on New Island, Falkand Islands. We found a strong seasonal decline in both egg and clutch size. Egg size increased with clutch size. The mean egg volume and total clutch volume increased with female body condition and hatching date was earlier for females in higher body condition. Chicks hatched from nests with higher mean egg volume had higher early body condition. We also compared individual females between different study years and found that individual body condition showed a significant repeatability between years, whereas hatching date did not. While female aging did not systematically affect hatching date, females produced clutches that hatched earlier in years of higher body condition. We could thus show for the first time that the frequently reported negative relationship between female condition and hatching date also holds true on an individual basis. To our knowledge, this is the first study on seasonal clutch size decline in sheldgeese and one of a few in waterfowl comparing condition and timing of breeding within individuals between different years.     

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.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Clutch size manipulation, hatching success and offspring phenotype in the ball python (Python regius)

In a diverse array of avian and mammalian species, experimental manipulations of clutch size have tested the hypothesis that natural selection should adjust numbers of neonates produced so as to maximize the number of viable offspring at the end of the period of parental care. Reptiles have not been studied in this respect, probably because they rarely display parental care. However, females of all python species brood their eggs until hatching, but they do not care for their neonates. This feature provides a straightforward way to experimentally increase or reduce clutch size to see whether the mean clutch size observed in nature does indeed maximize hatching success and/or optimize offspring phenotypes. Eggs were removed or added to newly laid clutches of Ball Pythons ( Python regius ) in tropical Africa (nine control clutches, eight with 50% more eggs added, six with 42% of eggs removed). All clutches were brooded by females throughout the 2-month incubation period. Experimental manipulation of clutch-size did not significantly affect the phenotypes (morphology, locomotor ability) of hatchlings, but eggs in 'enlarged' clutches hatched later, and embryos were more likely to die before hatching. This mortality was due to desiccation of the eggs, with females being unable to cover 'enlarged' clutches sufficiently to retard water loss. Our results support the notion of an optimal clutch size, driven by limitations on parental ability to care for the offspring. However, the proximate mechanisms that generate this optimum value differ from those previously described in other kinds of animals. © 2003 The Linnean Society of London . Biological Journal of the Linnean Society 2003, 78 , 263–272.     

SOME ASPECTS OF THE BEHAVIOUR OF THE WATTLED PLOVER AFRIBYX SENEGALLUS (LINNAEUS)

The question of how aridity might influence avian clutch size, through the influences of rainfall seasonality and environmental stochasticity (unpredictability), has received little attention. A marked east-west gradient in aridity across South Africa provides a unique opportunity to test for such influences. Using an extensive collection of nest records for 106 terrestrial bird species from the South African Nest Record Card Scheme, we tested three predictions related to rainfall seasonality and stochasticity. Analyses were conducted at two levels, the first examining each species independently, and the second grouping species into five dietary guilds. The first prediction, that clutch size should generally increase with higher rainfall seasonality (i.e. higher seasonal fluctuation of food availability), was supported, particularly in the most arid environments where food abundance is more closely linked to rainfall. Controlling for rainfall seasonality, the second prediction, that clutch size should generally decrease as a bet-hedging strategy in arid, stochastic environments, was also supported. Although the timing of the rainy season differs among regions in South Africa (winter, early summer, later summer, year-round), birds primarily nest during spring. The relative timing of rainfall and breeding is expected to have different consequences for seasonal variation in clutch size among rainfall regions. The third prediction, of different patterns of seasonal variation in clutch size between rainfall regions, was also supported. In the winter and early-summer rainfall regions, early-nesting birds (breeding with or soon after the rains) generally had a larger clutch size than late-nesting birds. In the late-summer rainfall region, early-nesting birds (breeding well before the rains) had a smaller clutch size than late-nesting birds.     

Allometry of reproduction in two species of gekkonid lizards (Gehyra): effects of body size miniaturization on clutch and egg sizes

In squamate reptiles there is an allometric pattern for small-bodied females to have smaller clutches and proportionally larger eggs than large-bodied females, and this pattern occurs both among and within species. The allometric patterns in two species of the gecko Gehyra were studied to see how evolutionary reductions in adult body size affect fecundity and offspring size among species, and how these changes affect allometric relationships within species. Gehyra dubia has two eggs per clutch (the typical clutch size for gekkonid lizards), whereas the smallerbodied G. variegata has a single egg per clutch. Within both species, egg size increased with female body size. The data are consistent with at least two mechanistic hypotheses: (1) that the width of the pelvis constrains egg size; and (2) in species with invariant clutch sizes, larger females can only allocate additional energy towards egg size and not number. More direct tests of these hypotheses are warranted. Miniaturization of body sizes in Gehyra is correlated with a clutch size reduction of 50% (from two to one), and a large (1.7-fold) compensatory increase in relative egg mass. However, the small-bodied G. variegata (one egg per clutch) had a lower relative clutch mass than did G. dubia. These findings have implications for understanding the influence of evolutionary reductions in body size on reproductive traits, and for allometric trends in squamate reptiles in general.     

Patterns of life-history traits in open-nesting palearctic passerines as a function of the climatic variation of their ranges

Latitude, a surrogate of climatic conditions, is commonly used in the examination of life-history variation. However, the climatic mechanisms underlying latitudinal life-history variation have only rarely been tested. Here, we test whether the number of climates to which species are subjected in their ranges predicts geographical life-history variation. In particular, we examine whether eurytopic species, the range of which covers more climates, show different reproductive effort to stenotopic species, which are distributed over climatically more homogeneous environments. We examined female body mass, egg mass controlled for female body mass, clutch size and the number of breeding attempts per season for 34 sedentary and short-distance migratory passerine species of the Western Palearctic. For each species, we assessed how many climate zones extend over the species' wintering and breeding ranges. We found that avian body mass, and also clutch size, significantly increases with the number of climatic zones extended over the species' wintering range. In turn, species whose breeding ranges span more climates show more breeding attempts per season. Whereas the mass of a single egg declines, clutch size increases with increasing climatic variation in breeding ranges. Our study suggests that the level of climatic variation over species' ranges during and outside the breeding season might be responsible for variation in life-history traits in open-nesting Western Palearctic passerines.     

Life‐history tradeoffs revealed by seasonal declines in reproductive traits of Arctic‐breeding shorebirds

Seasonal declines in breeding performance are widespread in wild animals, resulting from temporal changes in environmental conditions or from individual variation. Seasonal declines might drive selection for early breeding, with implications for other stages of the annual cycle. Alternatively, selection on the phenology of nonbreeding stages could constrain timing of the breeding season and lead to seasonal changes in reproductive performance. We studied 25 taxa of migratory shorebirds (including five subspecies) at 16 arctic sites in Russia, Alaska, and Canada. We investigated seasonal changes in four reproductive traits, and developed a novel Bayesian risk‐partitioning model of daily nest survival to examine seasonal trends in two causes of nest failure. We found strong seasonal declines in reproductive traits for a subset of species. The probability of laying a full four‐egg clutch declined by 8–78% in 12 of 25 taxa tested, daily nest survival rates declined by 1–12% in eight of 22 taxa, incubation duration declined by 2.0–2.5% in two of seven taxa, and mean egg volume declined by 5% in one of 15 taxa. Temporal changes were not fully explained by individual variation. Across all species, the proportion of failed nests that were depredated declined over the season from 0.98 to 0.60, while the proportion abandoned increased from 0.01 to 0.35 and drove the seasonal declines in nest survival. An increase in abandonment of late nests is consistent with a life‐history tradeoff whereby either adult mortality increased or adults deserted the breeding attempt to maximize adult survival. In turn, seasonal declines in clutch size and incubation duration might be adaptive to hasten hatching of later nests. In other species of shorebirds, we found no seasonal patterns in breeding performance, suggesting that some species are not subject to selective pressure for early breeding.     

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.     

Hormone profiles of obligate avian brood parasites during the breeding season

The ultimate explanations for avian brood parasitism have been studied intensively as a model system for coevolution, but little is known about the proximate mechanisms, for example hormonal regulation, underlying brood parasitic behaviour. In this study, we explored seasonal hormone profiles in two brood parasitic Cuculus species breeding in the Republic of Korea. As brood parasites have relatively simple breeding stages without incubation and provisioning, we predicted that during the breeding season individuals would exhibit similar levels of testosterone (T) and stress‐induced corticosterone (CORT), hormones that are known to be closely related to the transition of breeding stages. We also assessed how these hormone profiles were associated with traits such as body size and sex. Overall, male cuckoos showed similarly high T levels throughout the breeding season, as predicted, but individual variation became greater as the season progressed. Individual CORT levels tended to decrease as the season progressed, although the decrease was significant only in Common Cuckoos Cuculus canorus. We also found that male Lesser Cuckoos Cuculus poliocephalus showed a much higher level of T than females, as expected, but this sexual difference was not observed in Common Cuckoos. Our results suggest that the seasonal hormone profiles of avian brood parasites are likely to be similar to typical hormone profiles expected for non‐brood parasites during the breeding season. This may suggest that not only the breeding cycle but also other factors such as social interaction may be affected by hormonal changes. Further studies are needed to fully understand the proximate mechanism of avian brood parasitism.