Theory predicts shorter embryonic periods in species with greater embryo mortality risk and smaller body size. Field studies of 80 passerine species on three continents yielded data that largely conflicted with theory; incubation (embryonic) periods were longer rather than shorter in smaller species, and egg (embryo) mortality risk explained some variation within regions, but did not explain larger differences in incubation periods among geographic regions. Incubation behavior of parents seems to explain these discrepancies. Bird embryos are effectively ectothermic and depend on warmth provided by parents sitting on the eggs to attain proper temperatures for development. Parents of smaller species, plus tropical and southern hemisphere species, commonly exhibited lower nest attentiveness (percent of time spent on the nest incubating) than larger and northern hemisphere species.Lower nest attentiveness produced cooler minimum and average embryonic temperatures that were correlated with longer incubation periods independent of nest predation risk or body size. We experimentally tested this correlation by swapping eggs of species with cool incubation temperatures with eggs of species with warm incubation temperatures and similar egg mass. Incubation periods changed (shortened or lengthened) as expected and verified the importance of egg temperature on development rate. Slower development resulting from cooler temperatures may simply be a cost imposed on embryos by parents and may not enhance offspring quality. At the same time, incubation periods of transferred eggs did not match host species and reflect intrinsic differences among species that may result from nest predation and other selection pressures. Thus, geographic variation in embryonic development may reflect more complex interactions than previously recognized.

Causes of interspecific variation in growth rates within and among geographic regions remain poorly understood. Passerine birds represent an intriguing case because differing theories yield the possibility of an antagonistic interaction between nest predation risk and food delivery rates on evolution of growth rates. We test this possibility among 64 Passerine species studied on three continents, including tropical and north and south temperate latitudes. Growth rates increased strongly with nestling predation rates within, but not between, sites. The importance of nest predation was further emphasized by revealing hidden allometric scaling effects. Nestling predation risk also was associated with reduced total feeding rates and per-nestling feeding rates within each site. Consequently, faster growth rates were associated with decreased per-nestling food delivery rates across species, both within and among regions. These relationships suggest that Passerines can evolve growth strategies in response to predation risk whereby food resources are not the primary limit on growth rate differences among species. In contrast, reaction norms of growth 10 Present address:

Broad geographic patterns in egg and clutch mass are poorly described, and potential causes of variation remain largely unexamined. We describe interspecific variation in avian egg and clutch mass within and among diverse geographic regions and explore hypotheses related to allometry, clutch size, nest predation, adult mortality, and parental care as correlates and possible explanations of variation. We studied 74 species of Passeriformes at four latitudes on three continents: the north temperate United States, tropical Venezuela, subtropical Argentina, and south temperate South Africa. Egg and clutch mass increased with adult body mass in all locations, but differed among locations for the same body mass, demonstrating that egg and clutch mass have evolved to some extent independent of body mass among regions. A major portion of egg mass variation was explained by an inverse relationship with clutch size within and among regions, as predicted by life‐history theory. However, clutch size did not explain all geographic differences in egg mass; eggs were smallest in South Africa despite small clutch sizes. These small eggs might be explained by high nest predation rates in South Africa; life‐history theory predicts reduced reproductive effort under high risk of offspring mortality. This prediction was supported for clutch mass, which was inversely related to nest predation but not for egg mass. Nevertheless, clutch mass variation was not fully explained by nest predation, possibly reflecting interacting effects of adult mortality. Tests of the possible effects of nest predation on egg mass were compromised by limited power and by counterposing direct and indirect effects. Finally, components of parental investment, defined as effort per offspring, might be expected to positively coevolve. Indeed, egg mass, but not clutch mass, was greater in species that shared incubation by males and females compared with species in which only females incubate eggs. However, egg and clutch mass were not related to effort of parental care as measured by incubation attentiveness. Ecological and life‐history correlates of egg and clutch mass variation found here follow from theory, but possible evolutionary causes deserve further study.

Parental behavior and effort vary extensively among species. Life-history theory suggests that age-specific mortality could cause this interspecific variation, but past tests have focused on fecundity as the measure of parental effort. Fecundity can cause costs of reproduction that confuse whether mortality is the cause or the consequence of parental effort. We focus on a trait, parental allocation of time and effort in warming embryos, that varies widely among species of diverse taxa and is not tied to fecundity. We conducted studies on songbirds of four continents and show that time spent warming eggs varies widely among species and latitudes and is not correlated with clutch size. Adult and offspring (nest) mortality explained most of the interspecific variation in time and effort that parents spend warming eggs, measured by average egg temperatures. Parental effort in warming eggs is important because embryonic temperature can influence embryonic development period and hence exposure time to predation risk. We show through correlative evidence and experimental swapping of embryos between species that parentally induced egg temperatures cause interspecific variation in embryonic development period. The strong association of age-specific mortality with parental effort in warming eggs and the subsequent effects on embryonic development time are unique results that can advance understanding of broad geographic patterns of life-history variation.

Forest fragmentation may cause increased brood parasitism and nest predation of breeding birds. In North America, nest parasitism and predation are expected to increase closer to forest edges because the brood‐parasitic Brown‐headed Cowbird (Molothrus ater) and generalist nest predators often enter the forest from adjoining developed (largely agricultural) habitats. Yet the abundance of brood parasites and nest predators at the patch scale may be strongly constrained by the total area of developed habitat at landscape scales. The scale and extent of landscape effects are unclear, however, because past studies were mostly conducted within local landscapes rather than across independent landscapes. We report replicated studies from 30 independent landscapes across 17 states of the United States that show that nest parasitism is strongly affected by fragmentation at a 20 km radius scale, equivalent to the maximum foraging range of cowbirds. Nest predation is influenced by both edge and landscape effects, and increases with fragmentation at a 10 km radius scale. Predation is additive to parasitism mortality, and the two together yield decreased population growth potential with increasing forest fragmentation at a 10 km radius scale for 20 of 22 bird species. Mapping of population growth potential across continental landscapes displays broad impacts of fragmentation on population viability and allows geographic prioritization for conservation.

Rainfall is known t o b e a stimulus for initiating breeding in arid-zone birds, but its influence on clutch size is unknown for most southern African birds. The breeding activity of nine out of' 11 species studied in the arid Nama Karoo biome of South Africa increased markedly after appreciable rainfall. Some resident insectivores did breed in the absence of rainfall, even under drought conditions, but also laid eggs within one week of small rain showers. Nomadic granivores required more substantial rainfall to stimulate what was usually a synchronized, population-wide breeding response. The timing and length of the breeding season was dependent on the integrated effect that rainfall and temperature had on the growing season of the vegetation. Over 50%) of the 11 species studied showed evidence of an increase in clutch size following large rainfall events. Greybacked Finchlark Eremopterix verticalis began laying larger clutches within one week of a rainfall stimulus, illustrating how rapid this response can be.

Gene flow is traditionally thought to be antagonistic to population differentiation and local adaptation. However, recent studies have demonstrated that local adaptation can proceed provided that selection is greater than the homogenizing effects of gene flow. We extend these initial studies by combining ecology (climate), phenotype (body size), physiological genetics (oxidative phosphorylation genes), and neutral loci (nuclear microsatellites and introns) to test whether selection can counter-balance gene flow and hence promote local adaptation in a bird whose distribution spans an aridity gradient. Our results show that the Karoo scrub-robin's climatic niche is spatially structured, providing the potential for local adaptation to develop. We found remarkably discordant patterns of divergence among mtDNA, morphology, and neutral loci. For the mitochondrial genes, two amino acid replacements, strong population structure and reduced gene flow were associated with the environmental gradient separating western coastal sites from the interior of southern Africa. In contrast, morphology and the neutral loci exhibited variation independent of environmental variables, and revealed extensive levels of gene flow across the aridity gradient, 50 times larger than the estimates for mitochondrial genes. Together, our results suggest that selective pressures on physiology, mediated by the mitochondrial genome, may well be a common mechanism for facilitating local adaptation to new climatic conditions. K E Y W O R D S : climatic envelope, morphology, mtDNA OXPHOS genes, neutral loci, Passeriformes.

Disturbance by humans is widely expected to reduce the reproductive fitness of nesting birds if disturbance reduces nest attentiveness, and unattended eggs experience increased risk of predation or exposure to potentially lethal temperature extremes. Yet, relatively few studies have examined the physiological or behavioural mechanisms whereby disturbance influences reproductive fitness, or the extent to which the costs of disturbance may be reduced through habituation. We compared the behavioural responses, egg temperatures and reproductive success of shore-nesting white-fronted plovers Charadrius marginatus to disturbance at two breeding sites experiencing low versus high human recreational activity, respectively. Daytime nest attentiveness decreased with increasing experimental disturbance at both sites, but this relationship differed between sites; for any given level of disturbance, incubating birds at the more disturbed site had greater nest attentiveness. They achieved this through habituation, allowing a closer human approach before leaving the nest, and returning to the nest faster after a disturbance event. Despite lower average daytime nest attentiveness at the more disturbed site, incubation temperatures did not differ significantly between sites. Nest mortality, mostly by natural mammalian and corvid predators, was significantly lower at the site experiencing high recreational activity. However, chick mortality was significantly greater at the more disturbed site, most likely because of predation by domestic dogs. Chick mortality may have been increased by the habitation of chicks, whose escape responses were much reduced at the more disturbed site. Nonetheless, annual fecundity was substantially higher at the more disturbed site, showing that the overall reproductive fitness of wild birds is not always compromised by human disturbance and urbanization.