Anna D. Chalfoun

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.

SONGBIRD NEST PREDATORS IN FOREST–PASTURE EDGE AND FOREST INTERIOR IN A FRAGMENTED LANDSCAPE

Many studies have compared songbird nesting success between forest edge and interior, but few have addressed potential factors underlying variation in nest predation pressure in relation to edge. We examined the relative abundance and species richness of songbird nest predators and the abundance of Brown-headed Cowbirds (Molothrus ater) in forest edge and interior within a fragmented, agricultural landscape in central Missouri, USA. Avian predators and cowbirds were more abundant in forest edges. There were no differences in small- or medium-sized mammalian predator abundance between edge and interior. Almost twice as many snakes were captured in edge as in interior. Predator species richness was significantly higher in forest edge. Forest vegetation structure was very similar between edge and interior, suggesting that differences in predator abundance and species richness were not driven by variation in habitat structure. Nest predator distribution in relation to habitat edge may therefore depend on factors at larger spatial scales, such as landscape context. We suggest that in areas fragmented by agriculture, nest predator assemblages in forest edges may differ from those in forest interior. Edges may attract a greater number of predator species, and some nest predators may be more abundant near the edges of forest patches, although the trend does not apply across all predator taxa. Generalizations about nest predators and edges should thus be made with caution, and conservation plans should consider the composition of local nest predator assemblages in order to predict potential impacts on nesting birds in edge habitat.

Habitat structure mediates predation risk for sedentary prey: Experimental tests of alternative hypotheses

1. Predation is an important and ubiquitous selective force that can shape habitat preferences of prey species, but tests of alternative mechanistic hypotheses of habitat influences on predation risk are lacking. 2. We studied predation risk at nest sites of a passerine bird and tested two hypotheses based on theories of predator foraging behaviour. The total-foliage hypothesis predicts that predation will decline in areas of greater overall vegetation density by impeding cues for detection by predators. The potential-prey-site hypothesis predicts that predation decreases where predators must search more unoccupied potential nest sites. 3. Both observational data and results from a habitat manipulation provided clear support for the potential-prey-site hypothesis and rejection of the total-foliage hypothesis. Birds chose nest patches containing both greater total foliage and potential nest site density (which were correlated in their abundance) than at random sites, yet only potential nest site density significantly influenced nest predation risk. 4. Our results therefore provided a clear and rare example of adaptive nest site selection that would have been missed had structural complexity or total vegetation density been considered alone. 5. Our results also demonstrated that interactions between predator foraging success and habitat structure can be more complex than simple impedance or occlusion by vegetation.

Energy Development Affects Populations of Sagebrush Songbirds in Wyoming

ABSTRACT Oil and natural gas development in the Intermountain West region of North America has expanded over the last 2 decades, primarily within sagebrush dominated landscapes. Although the effects of energy development on high-profile game species such as the greater sage-grouse (Centrocercus urophasianus) have been documented, studies examining responses of non-game birds are lacking. Simultaneously, many songbirds that breed within sagebrush steppe habitats have shown range-wide population declines that are likely due to widespread habitat loss and alteration. We evaluated songbird abundance and species richness across gradients of oil and natural gas development intensity, as indexed by well density, at 3 energy fields (2 natural gas and 1 oil) in the Upper Green River Basin, Wyoming, USA during 2008–2009. While simultaneously accounting for important habitat attributes, increased well density was associated with significant decreases in Brewers sparrow (Spizella breweri) and sage sparrow (Amphispiza belli) abundance, particularly in the Jonah natural gas field. Vesper sparrows (Pooecetes gramineus) were also negatively influenced by increased well density. Horned larks (Eremophila alpestris) increased with well density in the Pinedale Anticline natural gas field, and sage thrashers (Oreoscoptes montanus) showed no response to energy development. Species richness was not significantly affected by well density. Results suggest that regional declines of some songbird species, especially sagebrush-obligates, may be exacerbated by increased energy development. Understanding the specific mechanisms underlying responses to energy development is an important next step and will aid land managers in the development of effective mitigation and management strategies for the maintenance of stable bird communities in sagebrush habitat.

Facultative nest patch shifts in response to nest predation risk in the Brewer’s sparrow: a “win-stay, lose-switch” strategy?

Facultative shifts in nesting habitat selection in response to perceived predation risk may allow animals to increase the survival probability of sessile offspring. Previous studies on this behavioral strategy have primarily focused on single attributes, such as the distance moved or changes in nesting substrate. However, nest site choice often encompasses multiple habitat elements at both the nest site and nest patch scales. We studied the within-season re-nesting strategy of a multi-brooded songbird, the Brewer’s sparrow (Spizella breweri), to determine whether pairs utilized a “win-stay, lose-switch” decision rule with respect to inter-nest distance, nest substrate and/or nest patch characteristics in response to previous nest fate. Pairs moved sequential nest sites slightly farther following nest predation versus success. When inter-nest distance was controlled, however, pairs changed nest patch attributes (shrub height, potential nest shrub density) associated with probability of nest predation to a greater extent following nest predation than success. The strategy appeared to be adaptive; daily nest survival probability for previously depredated pairs increased with greater Euclidian habitat distances between attempts, whereas previously successful pairs were more likely to fledge second attempts when nest sites were similar to those of previous attempts. Our results suggest that nesting birds can use prior information and within-season plasticity in response to nest predation to increase re-nesting success, which may be a critical behavioral strategy within complex nest predator environments. Re-nesting site selection strategies also appeared to integrate multiple habitat components and inter-nest distances. The consideration of such proximate, facultative responses to predation risk may clarify often unexplained variation in habitat preferences and requirements.

Nest predation research: Recent findings and future perspectives

Nest predation is a key source of selection for birds that has attracted increasing attention from ornithologists. The inclusion of new concepts applicable to nest predation that stem from social information, eavesdropping or physiology has expanded our knowledge considerably. Recent methodological advancements now allow focus on all three players within nest predation interactions: adults, offspring and predators. Indeed, the study of nest predation now forms a vital part of avian research in several fields, including animal behaviour, population ecology, evolution and conservation biology. However, within nest predation research there are important aspects that require further development, such as the comparison between ecological and evolutionary antipredator responses, and the role of anthropogenic change. We hope this review of recent findings and the presentation of new research avenues will encourage researchers to study this important and interesting selective pressure, and ultimately will help us to better understand the biology of birds.

Parental investment decisions in response to ambient nest-predation risk versus actual predation on the prior nest.

Abstract. Theory predicts that parents should invest less in dependent offspring with lower reproductive value, such as those with a high risk of predation. Moreover, high predation risk can favor reduced parental activity when such activity attracts nest predators. Yet, the ability of parents to assess ambient nest-predation risk and respond adaptively remains unclear, especially where nest-predator assemblages are diverse and potentially difficult to assess. We tested whether variation in parental investment by a multi-brooded songbird (Brewers Sparrow, Spizella breweri) in an environment (sagebrush steppe) with diverse predators was predicted by ambient nest-predation risk or direct experience with nest predation. Variation among eight sites in ambient nest-predation risk, assayed by daily probabilities of nest predation, was largely uncorrelated across four years. In this system risk may therefore be unpredictable, and aspects of parental investment (clutch size, egg mass, incubation rhythms, nestling-feeding rates) were not related to ambient risk. Moreover, investment at first nests that were successful did not differ from that at nests that were depredated, suggesting parents could not assess and respond to territory-level nest-predation risk. However, parents whose nests were depredated reduced clutch sizes and activity at nests attempted later in the season by increasing the length of incubation shifts (on-bouts) and recesses (off-bouts) and decreasing trips to feed nestlings. In this unpredictable environment parent birds may therefore lack sufficient cues of ambient risk on which to base their investment decisions and instead rely on direct experience with nest predation to inform at least some of their decisions.

Microrefuges and the occurrence of thermal specialists: implications for wildlife persistence amidst changing temperatures

BackgroundContemporary climate change is affecting nearly all biomes, causing shifts in animal distributions, phenology, and persistence. Favorable microclimates may buffer organisms against rapid changes in climate, thereby allowing time for populations to adapt. The degree to which microclimates facilitate the local persistence of climate-sensitive species, however, is largely an open question. We addressed the importance of microrefuges in mammalian thermal specialists, using the American pika (Ochotona princeps) as a model organism. Pikas are sensitive to ambient temperatures, and are active year-round in the alpine where conditions are highly variable. We tested four hypotheses about the relationship between microrefuges and pika occurrence: 1) Local-habitat Hypothesis (local-habitat conditions are paramount, regardless of microrefuge); 2) Surface-temperature Hypothesis (surrounding temperatures, unmoderated by microrefuge, best predict occurrence); 3) Interstitial-temperature Hypothesis (temperatures within microrefuges best predict occurrence), and 4) Microrefuge Hypothesis (the degree to which microrefuges moderate the surrounding temperature facilitates occurrence, regardless of other habitat characteristics). We examined pika occurrence at 146 sites across an elevational gradient. We quantified pika presence, physiographic habitat characteristics and forage availability at each site, and deployed paired temperature loggers at a subset of sites to measure surface and subterranean temperatures.ResultsWe found strong support for the Microrefuge Hypothesis. Pikas were more likely to occur at sites where the subsurface environment substantially moderated surface temperatures, especially during the warm season. Microrefugium was the strongest predictor of pika occurrence, independent of other critical habitat characteristics, such as forage availability.ConclusionsBy modulating surface temperatures, microrefuges may strongly influence where temperature-limited animals persist in rapidly warming environments. As climate change continues to manifest, efforts to understand the changing dynamics of animal-habitat relationships will be enhanced by considering the quality of microrefuges.

Towards a mechanistic understanding of human- induced rapid environmental change: a case study linking energy development, nest predation and predators

Summary 1. Demographic consequences of human-induced rapid environmental change (HIREC) have been widely documented for many populations. The mechanisms underlying such patterns, however, are rarely investigated and yet are critical to understand for effective conservation and management. 2. We investigated the mechanisms underlying reduced avian nest survival with intensification of natural gas development, an increasing source of human-induced rapid environmental change globally. We tested the hypothesis that energy development increased the local activity of important nest predator species, thereby elevating nest predation rates. During 2011–2012, we surveyed predators and monitored 668 nests of Brewer’s sparrows Spizella breweri (BRSP), sagebrush sparrows Artemisiospiza nevadensis (SASPs) and sage thrashers Oreoscoptes montanus (SATHs) breeding at twelve sites spanning a gradient of habitat loss from energy development in western Wyoming, USA. 3. Nine species, representing four mammalian and three avian families, were video-recorded depredating eggs and nestlings. Important nest predator species differed across songbird species, despite similar nesting habitats. Approximately 75% of depredation events were by rodents. 4. Consistent with our predictions, detections of most rodent nest predators increased with surrounding habitat loss due to natural gas development, which was associated with increased probability of nest predation for our three focal bird species. 5. An altered nest predator assemblage was therefore at least partly responsible for elevated avian nest predation risk in areas with more surrounding energy development. 6. Synthesis and applications. We demonstrate one mechanism, that is the local augmentation of predators, by which human-induced rapid environmental change can influence the demography of local populations. Given the accelerating trajectory of global energy demands, an important next step will be to understand why the activity and/or abundance of rodent predators increased with surrounding habitat loss from energy development activities.

Reproductive success of Horned Lark and McCown's Longspur in relation to wind energy infrastructure

ABSTRACT Wind energy is a rapidly expanding industry with potential indirect effects to wildlife populations that are largely unexplored. In 2011 and 2012, we monitored 211 nests of 2 grassland songbirds, Horned Lark (Eremophila alpestris) and McCowns Longspur (Rhynchophanes mccownii), at 3 wind farms and 2 undeveloped reference sites in Wyoming, USA. We evaluated several indices of reproductive investment and success: clutch size, size-adjusted nestling mass, daily nest survival rate, and number of fledglings. We compared reproductive success between wind farms and undeveloped sites and modeled reproductive success within wind farms as a function of wind energy infrastructure and habitat. Size-adjusted nestling mass of Horned Lark was weakly negatively related to turbine density. In 2011, nest survival of Horned Lark decreased 55% as turbine density increased from 10 to 39 within 2 km of the nest. In 2012, however, nest survival of Horned Lark was best predicted by the combination of vegetation height, distance to shrub edge, and turbine density, with survival increasing weakly with increasing vegetation height. McCowns Longspur nest survival was weakly positively related to vegetation density at the nest site when considered with the amount of grassland habitat in the neighborhood and turbine density within 1 km of the nest. Habitat and distance to infrastructure did not explain clutch size or number of fledglings for either species, or size-adjusted nestling mass for McCowns Longspur. Our results suggest that the influence of wind energy infrastructure varies temporally and by species, even among species using similar habitats. Turbine density was repeatedly the most informative measure of wind energy development. Turbine density could influence wildlife responses to wind energy production and may become increasingly important to consider as development continues in areas with high-quality wind resources.