Penn Lloyd

GEOGRAPHIC VARIATION IN AVIAN INCUBATION PERIODS AND PARENTAL INFLUENCES ON EMBRYONIC TEMPERATURE

Abstract 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.

LIFE-HISTORY AND ECOLOGICAL CORRELATES OF GEOGRAPHIC VARIATION IN EGG AND CLUTCH MASS AMONG PASSERINE SPECIES

Abstract 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.

LINKING DEMOGRAPHIC EFFECTS OF HABITAT FRAGMENTATION ACROSS LANDSCAPES TO CONTINENTAL SOURCE–SINK DYNAMICS

Forest fragmentation may cause increased brood parasitism and nest pre- dation 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 (Mol- othrus 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.

GROWTH RATE VARIATION AMONG PASSERINE SPECIES IN TROPICAL AND TEMPERATE SITES: AN ANTAGONISTIC INTERACTION BETWEEN PARENTAL FOOD PROVISIONING AND NEST PREDATION RISK

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 rate relative to brood size suggest that food may limit growth rates within species in temperate, but not tropical, regions. Results here provide new insight into evolution of growth strategies relative to predation risk and food within and among species.

A TIGHT BALANCE BETWEEN NATURAL SELECTION AND GENE FLOW IN A SOUTHERN AFRICAN ARID-ZONE ENDEMIC BIRD

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‐robins 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.

Adult Mortality Probability and Nest Predation Rates Explain Parental Effort in Warming Eggs with Consequences for Embryonic Development Time.

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.

INVESTIGATOR EFFECTS ON THE NESTING SUCCESS OF ARID-ZONE BIRDS

Abstract We examined whether regular researcher visits affected egg hatchability or nest predation for three ground-nesting bird species that incur high levels of nest predation, primarily by small mammals. Frequently visited finch-lark (Eremopterix verticalis and E. australis) nests suffered similar predation to nests visited infrequently, suggesting that regular visits had no additive effects on nest survival. A comparison of finch-lark nests visited for the second time either one or two days after the first visit found that predation during the first 24 h (7.4%) was lower than predation during the second 24-h period (9.9%), suggesting that the act of visiting a nest did not increase the risk of predation. Daily predation rates on Namaqua Sandgrouse (Pterocles namaqua) and finch-lark nests showed no observable trend with an increasing number of visits over time, indicating that frequent visits had no cumulative effect on predation probabilities. Nests of both Grey-backed Finch-lark (E. verticalis) and Black-eared Finch-lark (E. australis) discovered at the egg stage did not fledge significantly fewer young than nests discovered at the nestling stage, suggesting that investigator disturbance had no effect on egg hatchability. These results from the southern hemisphere subtropics support the findings of limited north-temperate studies that largely mammalian nest predation does not increase after researcher disturbance.

Microgeographic socio‐genetic structure of an African cooperative breeding passerine revealed: integrating behavioural and genetic data

Dispersal can be motivated by multiple factors including sociality. Dispersal behaviour affects population genetic structure that in turn reinforces social organization. We combined observational information with individual‐based genetic data in the Karoo scrub‐robin, a facultative cooperatively breeding bird, to understand how social bonds within familial groups affect mating patterns, cause sex asymmetry in dispersal behaviour and ultimately influence the evolution of dispersal. Our results revealed that males and females do not have symmetrical roles in structuring the population. Males are extremely philopatric and tend to delay dispersal until they gain a breeding position within a radius of two territories around the natal site. By contrast, females dispersed over larger distances, as soon as they reach independence. This resulted in male neighbourhoods characterized by high genetic relatedness. The long‐distance dispersal strategy of females ensured that Karoo scrub‐robins do not pair with relatives thereby compensating for male philopatry caused by cooperation. The observed female‐biased strategy seems to be the most prominent mechanism to reduce the risk of inbreeding that characterizes social breeding system. This study demonstrates that tying together ecological data, such as breeding status, determining social relationships with genetic data, such as kinship, provides valuable insights into the proximate causes of dispersal, which are central to any evolutionary interpretation.

The influence of nest-site characteristics on the nesting success of the Karoo Prinia (Prinia maculosa)

Choice of nest site has important consequences for nest survival. We examined nest-site characteristics relative to nest success in Karoo Prinias breeding in coastal dwarf shrubland, where high nest predation is the main cause of nest failure. Initially, we compared nests that failed during the building, laying, incubation and nestling stages and those from which young were successfully raised, to test whether nests that survived to progressive stages in the nesting cycle differed in their nest-site characteristics. Subsequently, we compared the characteristics of successful nests with those of unsuccessful nests. The nest-site characteristics considered included nest height, nest-plant height, nest-plant species, distance from lateral foliage edge, nest concealment, nest-patch heterogeneity and vegetation cover at four different heights. We were unable to distinguish between the nest-site characteristics of nests that failed during the various stages of the nesting cycle. Concealment was the main nest-site characteristic that differentiated successful nests from unsuccessful nests, with successful nests being located in more concealed sites. The other variables that contributed to the discrimination between successful and unsuccessful nests by discriminant function analysis included nest-plant type and distance from edge, which are also directly related to concealment. This suggests that nest concealment is the most important variable influencing nesting success at this site, which has a preponderance of visually-oriented predators.

Avian clutch size in relation to rainfall seasonality and stochasticity along an aridity gradient across South Africa

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.