A global analysis of avian island diversity-area relationships in the Anthropocene.

Research on island species-area relationships (ISAR) has expanded to incorporate functional (IFDAR) and phylogenetic (IPDAR) diversity. However, relative to the ISAR, we know little about IFDARs and IPDARs, and lack synthetic global analyses of variation in form of these three categories of island diversity-area relationship (IDAR). Here, we undertake the first comparative evaluation of IDARs at the global scale using 51 avian archipelagic data sets representing true and habitat islands. Using null models, we explore how richness-corrected functional and phylogenetic diversity scale with island area. We also provide the largest global assessment of the impacts of species introductions and extinctions on the IDAR. Results show that increasing richness with area is the primary driver of the (non-richness corrected) IPDAR and IFDAR for many data sets. However, for several archipelagos, richness-corrected functional and phylogenetic diversity changes linearly with island area, suggesting that the dominant community assembly processes shift along the island area gradient. We also find that archipelagos with the steepest ISARs exhibit the biggest differences in slope between IDARs, indicating increased functional and phylogenetic redundancy on larger islands in these archipelagos. In several cases introduced species seem to have 're-calibrated' the IDARs such that they resemble the historic period prior to recent extinctions.

Riskiness of Movement Lifestyle Varies Inversely with Adult Survival Probability among Species.

AbstractWhy do species differ in their movement lifestyles? Animals that spend more time sitting motionless and acquire food using less conspicuous movements can be more vigilant and less obvious to predators. More active animals that use food types and sites that require more conspicuous behaviors increase vulnerability to predators. Life history theory predicts that aversiveness to mortality risk evolves inversely to adult survival probability. Consequently, we postulated that long-lived species evolved inconspicuous movement lifestyles, whereas shorter-lived species use more conspicuous movement lifestyles. We tested this hypothesis by quantifying the movement lifestyles of nine tropical songbird species. Use of conspicuous movement and foraging behaviors, such as flying and hovering, was greatest in shorter-lived species and decreased with increasing adult survival probability across species. Similarly, foraging speed decreased with increasing adult survival based on a meta-analysis of 64 songbird species. Faster and conspicuous movement lifestyles of shorter-lived species likely increase food acquisition rates, which fits with faster life history strategies that include more feeding trips for young and faster growth. Similarly, slow movement lifestyles of long-lived species fit with the reduced food needs of slower life history strategies. Movement lifestyles may have evolved as an integrated component of the slow-fast life history continuum.

Contrasting seasonal effects of climate change influence density in a cold-adapted species.

Many ecological processes are profoundly influenced by abiotic factors, such as temperature and snow. However, despite strong evidence linking shifts in these ecological processes to corresponding shifts in abiotic factors driven by climate change, the mechanisms connecting population size to season-specific climate drivers are little understood. Using a 21-year dataset and a Bayesian state space model, we identified biologically informed seasonal climate covariates that influenced densities of snowshoe hares (Lepus americanus), a cold-adapted boreal herbivore. We found that snow and temperature had strong but conflicting season-dependent effects. Reduced snow duration in spring and fall and warmer summers were associated with lowered hare density, whereas warmer winters were associated with increased density. When modeled simultaneously and under two climate change scenarios, the negative effects of reduced fall and spring snow duration and warmer summers overwhelm the positive effect of warmer winters, producing projected population declines. Ultimately, the contrasting population-level impacts of climate change across seasons emphasize the critical need to examine the entire annual climate cycle to understand potential long-term population consequences of climate change.

Nest predation risk and deposition of yolk steroids in a cavity-nesting songbird: an experimental test.

Maternal hormones can shape offspring development and increase survival when predation risk is elevated. In songbirds, yolk androgens influence offspring growth and begging behaviors, which can help mitigate offspring predation risk in the nest. Other steroids may also be important for responding to nest predation risk, but non-androgen steroids have been poorly studied. We used a nest predator playback experiment and liquid chromatography with tandem mass spectrometry (LC-MS-MS) to assess whether nest predation risk influences deposition of 10 yolk steroids. We found no clear evidence that yolk androgen deposition changed when perception of nest predation risk was experimentally increased. However, elevated nest predation risk led to decreased yolk progesterone deposition. Overall, our results suggest yolk progesterone may be more important than yolk androgens in responses to offspring predation risk and highlight new avenues for research.

Adult survival probability and body size affect parental risk-taking across latitudes.

Parents faced with a predator must choose between their own safety versus taking care of their offspring. Each choice can have fitness costs. Life-history theory predicts that longer-lived species should be less willing than shorter-lived species to return to care for their offspring after a predator disturbance because they have more opportunities to reproduce in the future. We increased adult predation risk during incubation for 40 bird species in north temperate, tropical, and south temperate latitudes. We found that species with higher adult survival probabilities were more cautious, waiting longer before returning to the nest to provide care. Contrary to other studies, we also found that parents were more risk averse and waited longer to return in smaller than larger species, likely reflecting greater vulnerability of smaller species. Ultimately, the relative risk a predator poses to a species and the probability of future reproduction predict parental risk taking across the world.

Metabolic rate is negatively linked to adult survival but does not explain latitudinal differences in songbirds.

Survival rates vary dramatically among species and predictably across latitudes, but causes of this variation are unclear. The rate-of-living hypothesis posits that physiological damage from metabolism causes species with faster metabolic rates to exhibit lower survival rates. However, whether increased survival commonly observed in tropical and south temperate latitudes is associated with slower metabolic rate remains unclear. We compared metabolic rates and annual survival rates that we measured across 46 species, and from literature data across 147 species of birds in northern, southern and tropical latitudes. High metabolic rates were associated with lower survival but survival varied substantially among latitudinal regions independent of metabolism. The inability of metabolic rate to explain latitudinal variation in survival suggests (1) species may evolve physiological mechanisms that mitigate physiological damage from cellular metabolism and (2) extrinsic rather than intrinsic sources of mortality are the primary causes of latitudinal differences in survival.

Daily Nest Predation Rates Decrease with Body Size in Passerine Birds.

AbstractBody size evolution is generally framed by the benefits of being large, while costs are largely overlooked. An important putative cost of being large is the need to extend development periods, which should increase exposure to predation and potentially select against larger size. In birds, this selection pressure can be important because predation is the main source of offspring mortality and predators should more readily detect the larger nests associated with larger body sizes. Here, we show for diverse passerine birds across the world that counter to expectations, larger species suffer lower daily nest predation rates than smaller species. This pattern is consistent despite latitudinal variation in predation and does not seem to reflect a tendency of larger species to use more protected nests or less exposed nest locations. Evidence instead suggests that larger species attack a wider array of predator sizes, which could reduce predation rates in nests of large-bodied species. Regardless of the mechanism, the lower daily nest predation rates of larger species yield slightly lower predation rates over the entire development period compared with smaller species. These results highlight the importance of behavior as a mechanism to alter selection pressures and have implications for body size evolution.

Experimental Amelioration of Harsh Weather Speeds Growth and Development in a Tropical Montane Songbird.

AbstractOrganisms living at high elevations generally grow and develop more slowly than those at lower elevations. Slow montane ontogeny is thought to be an evolved adaptation to harsh environments that improves juvenile quality via physiological trade-offs. However, slower montane ontogeny may also reflect proximate influences of harsh weather on parental care and offspring development. We experimentally heated and protected nests from rain to ameliorate harsh montane weather conditions for mountain blackeyes (Chlorocharis emiliae), a montane songbird living at approximately 3,200 m asl in Malaysian Borneo. This experiment was designed to test whether cold and wet montane conditions contribute to parental care and postnatal growth and development rates at high elevations. We found that parents increased provisioning and reduced time spent warming offspring, which grew faster and departed the nest earlier compared with offspring from unmanipulated nests. Earlier departure reduces time-dependent predation risk, benefitting parents and offspring. These plastic responses highlight the importance of proximate weather contributions to broad patterns of montane ontogeny and parental care.

Nest predation and adult mortality relationships with post-natal metabolic rates and growth among songbird species.

Metabolism is thought to mediate the connection between environmental selection pressures and a broad array of life history tradeoffs, but tests are needed. High juvenile predation correlates with fast growth, which may be achieved via fast juvenile metabolism. Fast offspring metabolism and growth can create physiological costs later in life that should be minimized in species with low adult mortality. Yet, relationships between juvenile metabolism and mortality at offspring versus adult stages are unexplored. We found that post-natal metabolism was positively correlated with adult mortality but not nest predation rates among 43 songbird species on three continents. Nest predation, but not adult mortality, explained additional variation in growth rates beyond metabolism. Our results suggest that metabolism may not be the mechanism underlying the relationships between growth and mortality at different life stages.

Atom-by-atom analysis of sintering dynamics and stability of Pt nanoparticle catalysts in chemical reactions.

Supported Pt nanoparticles are used extensively in chemical processes, including for fuel cells, fuels, pollution control and hydrogenation reactions. Atomic-level deactivation mechanisms play a critical role in the loss of performance. In this original research paper, we introduce real-time in-situ visualization and quantitative analysis of dynamic atom-by-atom sintering and stability of model Pt nanoparticles on a carbon support, under controlled chemical reaction conditions of temperature and continuously flowing gas. We use a novel environmental scanning transmission electron microscope with single-atom resolution, to understand the mechanisms. Our results track the areal density of dynamic single atoms on the support between nanoparticles and attached to them; both as migrating species in performance degradation and as potential new independent active species. We demonstrate that the decay of smaller nanoparticles is initiated by a local lack of single atoms; while a post decay increase in single-atom density suggests anchoring sites on the substrate before aggregation to larger particles. The analyses reveal a relationship between the density and mobility of single atoms, particle sizes and their nature in the immediate neighbourhood. The results are combined with practical catalysts important in technological processes. The findings illustrate the complex nature of sintering and deactivation. They are used to generate new fundamental insights into nanoparticle sintering dynamics at the single-atom level, important in the development of efficient supported nanoparticle systems for improved chemical processes and novel single-atom catalysis. This article is part of a discussion meeting issue 'Dynamic in situ microscopy relating structure and function'.

Visualizing single atom dynamics in heterogeneous catalysis using analytical in situ environmental scanning transmission electron microscopy.

Progress is reported in analytical in situ environmental scanning transmission electron microscopy (ESTEM) for visualizing and analysing in real-time dynamic gas-solid catalyst reactions at the single-atom level under controlled reaction conditions of gas environment and temperature. The recent development of the ESTEM advances the capability of the established ETEM with the detection of fundamental single atoms, and the associated atomic structure of selected solid-state heterogeneous catalysts, in catalytic reactions in their working state. The new data provide improved understanding of dynamic atomic processes and reaction mechanisms, in activity and deactivation, at the fundamental level; and in the chemistry underpinning important technological processes. The benefits of atomic resolution-E(S)TEM to science and technology include new knowledge leading to improved technological processes, reductions in energy requirements and better management of environmental waste. This article is part of a discussion meeting issue 'Dynamic in situ microscopy relating structure and function'.

Higher Nest Predation Favors Rapid Fledging at the Cost of Plumage Quality in Nestling Birds.

High predation risk can favor rapid offspring development at the expense of offspring quality. Impacts of rapid development on phenotypic quality should be most readily expressed in traits that minimize fitness costs. We hypothesize that ephemeral traits that are replaced or repaired after a short period of life might express trade-offs in quality as a result of rapid development more strongly than traits used throughout life. We explored this idea for plumage quality in nestling body feathers, an ephemeral trait. We found a strong trade-off whereby nestlings that spend less time in the nest produced lower-quality plumage with less dense barbs relative to adults across 123 temperate and tropical species. For a subset of these species ( n=67 ), we found that variation in the risk of nest predation explained additional variation in plumage quality beyond development time. Ultimately, the fitness costs of a poor-quality ephemeral trait, such as nestling body feathers, may be outweighed by the fitness benefits of shorter development times that reduce predation risk. At the same time, reduced resource allocation to traits with small fitness costs, such as ephemeral traits, may ameliorate resource constraints from rapid development on traits with larger fitness impacts.

Guiding the prioritization of the most endangered and evolutionary distinct birds for new zoo conservation programs.

Zoos have played a pivotal role in the successful reinforcement and reintroduction of species threatened with extinction, but prioritization is required in the face of increasing need and limited capacity. One means of prioritizing between species of equal threat status when establishing new breeding programs is the consideration of evolutionary distinctness (ED). More distinct species have fewer close relatives such that their extinction would result in a greater overall loss to the Tree of Life. Considering global ex situ holdings of birds (a group with a complete and well-detailed evolutionary tree), we investigate the representation of at-risk and highly evolutionarily distinct species in global zoo holdings. We identified a total of 2,236 bird species indicated by the Zoological Information Management System as being held in zoological institutions worldwide. As previously reported, imperiled species (defined as those possessing endangered or critically endangered threat status) in this database are less likely to be held in zoos than non-imperiled species. However, we find that species possessing ED scores within the top 10% of all bird species are more likely to be held in zoos than other species, possibly because they possess unique characteristics that have historically made them popular exhibits. To assist with the selection of high priority ED species for future zoo conservation programs, we provide a list of imperiled species currently not held in zoos, ranked by ED. This list highlights species representing particular priorities for ex situ conservation planners, and represents a practical tool for improving the conservation value of zoological collections.

Fitness Consequences of Interspecific Nesting Associations among Cavity-Nesting Birds.

Interspecific aggregations of prey may provide benefits by mitigating predation risk, but they can also create costs if they increase competition for resources or are more easily detectable by predators. Variation in predation risk and resource availability may influence the occurrence and fitness effects of aggregating in nature. Yet tests of such possibilities are lacking. Cavity-nesting birds provide an interesting test case. They compete aggressively for resources and experience low nest predation rates, which might predict dispersion, but across 19 years of study we found that they commonly aggregate by sharing nest trees. Tree sharing was more common when aspen were more abundant and was somewhat more common in years with higher nest predation risk. Nest success was higher in shared trees when nest predation risk was higher than average. Ultimately, the costs and benefits of aggregating (nest tree sharing) varied across years, and we outline hypotheses for future studies.

Seasonal fecundity and costs to λ are more strongly affected by direct than indirect predation effects across species.

Increased perceived predation risk can cause behavioral and physiological responses to reduce direct predation mortality, but these responses can also cause demographic costs through reduced reproductive output. Such indirect costs of predation risk have received increased attention in recent years, but the relative importance of direct vs. indirect predation costs to population growth (λ) across species remains unclear. We measured direct nest predation rates as well as indirect benefits (i.e., reduced predation rates) and costs (i.e., decreased reproductive output) arising from parental responses to perceived offspring predation risk for 10 songbird species breeding along natural gradients in nest predation risk. We show that reductions in seasonal fecundity from behavioral responses to perceived predation risk represent significant demographic costs for six of the 10 species. However, demographic costs from these indirect predation effects on seasonal fecundity comprised only 12% of cumulative predation costs averaged across species. In contrast, costs from direct predation mortality comprised 88% of cumulative predation costs averaged across species. Demographic costs from direct offspring predation were relatively more important for species with higher within-season residual-reproductive value (i.e., multiple-brooded species) than for species with lower residual-reproductive value (i.e., single-brooded species). Costs from indirect predation effects were significant across single- but not multiple-brooded species. Ultimately, demographic costs from behavioral responses to offspring predation risk differed among species as a function of their life-history strategies. Yet direct predation mortality generally wielded a stronger influence than indirect effects on seasonal fecundity and projected λ across species.

Costs of fear: behavioural and life-history responses to risk and their demographic consequences vary across species.

Behavioural responses to reduce predation risk might cause demographic 'costs of fear'. Costs differ among species, but a conceptual framework to understand this variation is lacking. We use a life-history framework to tie together diverse traits and life stages to better understand interspecific variation in responses and costs. We used natural and experimental variation in predation risk to test phenotypic responses and associated demographic costs for 10 songbird species. Responses such as increased parental attentiveness yielded reduced development time and created benefits such as reduced predation probability. Yet, responses to increased risk also created demographic costs by reducing offspring production in the absence of direct predation. This cost of fear varied widely across species, but predictably with the probability of repeat breeding. Use of a life-history framework can aid our understanding of potential demographic costs from predation, both from responses to perceived risk and from direct predation mortality.

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.

Postnatal growth rates covary weakly with embryonic development rates and do not explain adult mortality probability among songbirds on four continents.

Growth and development rates may result from genetic programming of intrinsic processes that yield correlated rates between life stages. These intrinsic rates are thought to affect adult mortality probability and longevity. However, if proximate extrinsic factors (e.g., temperature, food) influence development rates differently between stages and yield low covariance between stages, then development rates may not explain adult mortality probability. We examined these issues based on study of 90 songbird species on four continents to capture the diverse life-history strategies observed across geographic space. The length of the embryonic period explained little variation (ca. 13%) in nestling periods and growth rates among species. This low covariance suggests that the relative importance of intrinsic and extrinsic influences on growth and development rates differs between stages. Consequently, nestling period durations and nestling growth rates were not related to annual adult mortality probability among diverse songbird species within or among sites. The absence of a clear effect of faster growth on adult mortality when examined in an evolutionary framework across species may indicate that species that evolve faster growth also evolve physiological mechanisms for ameliorating costs on adult mortality. Instead, adult mortality rates of species in the wild may be determined more strongly by extrinsic environmental causes.

A conceptual framework for clutch-size evolution in songbirds.

Causes of evolved differences in clutch size among songbird species remain debated. I propose a new conceptual framework that integrates aspects of traditional life-history theory while including novel elements to explain evolution of clutch size among songbirds. I review evidence that selection by nest predation on length of time that offspring develop in the nest creates a gradient in offspring characteristics at nest leaving (fledging), including flight mobility, spatial dispersion, and self-feeding rate. I postulate that this gradient has consequences for offspring mortality rates and parental energy expenditure per offspring. These consequences then determine how reproductive effort is partitioned among offspring, while reproductive effort evolves from age-specific mortality effects. Using data from a long-term site in Arizona, as well as from the literature, I provide support for hypothesized relationships. Nestling development period consistently explains fledgling mortality, energy expenditure per offspring, and clutch size while accounting for reproductive effort (i.e., total energy expenditure) to thereby support the framework. Tests in this article are not definitive, but they document previously unrecognized relationships and address diverse traits (developmental strategies, parental care strategies, energy requirements per offspring, evolution of reproductive effort, clutch size) that justify further investigations of hypotheses proposed here.

Comparative reproductive ecology of Old and New World Trogons, an order in decline across the world.

Many tropical species show declining populations. The pantropical order Trogoniformes has 76% of its species ranked as declining, reflecting a worldwide problem. Here, we report on the reproductive ecology and life history traits of the declining and near-threatened old world Whitehead's Trogon (Harpactes whiteheadi), the declining new world Collared Trogon (Trogon collaris), and the stable Masked Trogon (T. personatus). We also reviewed the literature on reproductive ecology and life history traits of trogons to assess possible commonalities that might help explain population declines. We found that the declining Whitehead's and Collared Trogons had reasonable nest success (32% and 25%, respectively), while the stable Masked Trogon had poor reproductive success (9%), all contrary to population trends. However, the limited literature data suggested that poor reproductive success may be common among trogons, which may contribute to population declines. Parents fed young at a low rate and had long on-bouts for incubation and nestling warming that reduced activity at the nest, as favored by high nest predation risk over evolutionary time. We found that young fledged from the nest with poorly developed wings, as also favored by high nest predation risk. Evolved nestling periods among trogon species suggests that poor wing development is likely common. Wing development has been shown to affect juvenile survival after leaving the nest. The poor wing development may be an important contributor to population declines that deserves more attention. Evolved life history traits are important to recognize as creating population vulnerabilities in a changing world.