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The golden rule of reviewing

A major bottleneck in the time required to publish a scientific or scholarly paper is the speed with which reviews by peers are returned to journals. Peer review is a reciprocal altruistic system in which each individual may perform every task—editors, reviewers, and authors—at different times. Journals have no way to coerce reviewers to return their critiques faster. To greatly shorten the time to publication, all actors in this altruistic network should abide by the Golden Rule of Reviewing: review for others as you would have others review for you. Say yes to reviewing whenever your duties and schedule allow; provide a thorough, fair, and constructive critique of the work; and do it at your first opportunity regardless of the deadline.

American Naturalist↗

How vegetation and sediment transport feedbacks drive landscape change in the Everglades and wetlands worldwide

Mechanisms reported to promote landscape self‐organization cannot explain vegetation patterning oriented parallel to flow. Recent catastrophic shifts in Everglades landscape pattern and ecological function highlight the need to understand the feedbacks governing these ecosystems. We modeled feedback between vegetation, hydrology, and sediment transport on the basis of a decade of experimentation. Results from more than 100 simulations showed that flows just sufficient to redistribute sediment from sparsely vegetated sloughs to dense ridges were needed for an equilibrium patterned landscape oriented parallel to flow. Surprisingly, although vegetation heterogeneity typically conveys resilience, in wetlands governed by flow/sediment feedbacks it indicates metastability, whereby the landscape is prone to catastrophic shifts. Substantial increases or decreases in flow relative to the equilibrium condition caused an expansion of emergent vegetation and loss of open‐water areas that was unlikely to revert upon restoration of the equilibrium hydrology. Understanding these feedbacks is critical in forecasting wetland responses to changing conditions and designing management strategies that optimize ecosystem services, such as carbon sequestration or habitat provision. Our model and new sensitivity analysis techniques address these issues and make it newly apparent that simply returning flow to predrainage conditions in the Everglades may not be sufficient to restore historic landscape patterns and processes.

American Naturalist↗

Optimization of biomass composition explains microbial growth-stoichiometry relationships

Integrating microbial physiology and biomass stoichiometry opens far-reaching possibilities for linking microbial dynamics to ecosystem processes. For example, the growth-rate hypothesis (GRH) predicts positive correlations among growth rate, RNA content, and biomass phosphorus (P) content. Such relationships have been used to infer patterns of microbial activity, resource availability, and nutrient recycling in ecosystems. However, for microorganisms it is unclear under which resource conditions the GRH applies. We developed a model to test whether the response of microbial biomass stoichiometry to variable resource stoichiometry can be explained by a trade-off among cellular components that maximizes growth. The results show mechanistically why the GRH is valid under P limitation but not under N limitation. We also show why variability of growth rate-biomass stoichiometry relationships is lower under P limitation than under N or C limitation. These theoretical results are supported by experimental data on macromolecular composition (RNA, DNA, and protein) and biomass stoichiometry from two different bacteria. In addition, compared to a model with strictly homeostatic biomass, the optimization mechanism we suggest results in increased microbial N and P mineralization during organic-matter decomposition. Therefore, this mechanism may also have important implications for our understanding of nutrient cycling in ecosystems.

American Naturalist↗

Biogeographic and ecological regulation of disease: Prevalence of Sin Nombre virus in island mice is related to island area, precipitation, and predator richness

The relative roles of top-down and bottom-up forces in affecting disease prevalence in wild hosts is important for understanding disease dynamics and human disease risk. We found that the prevalence of Sin Nombre virus (SNV), the agent of a severe disease in humans (hantavirus pulmonary syndrome), in island deer mice from the eight California Channel Islands was greater with increased precipitation (a measure of productivity), greater island area, and fewer species of rodent predators. In finding a strong signal of the ecological forces affecting SNV prevalence, our work highlights the need for future work to understand the relative importance of average rodent density, population fluctuations, behavior, and specialist predators as they affect SNV prevalence. In addition to illustrating the importance of both bottom-up and top-down limitation of disease prevalence, our results suggest that predator richness may have important bearing on the risk of exposure to animal-borne diseases that affect humans.

California↗

The effect of travel loss on evolutionarily stable distributions of populations in space

A key assumption of the ideal free distribution (IFD) is that there are no costs in moving between habitat patches. However, because many populations exhibit more or less continuous population movement between patches and traveling cost is a frequent factor, it is important to determine the effects of costs on expected population movement patterns and spatial distributions. We consider a food chain (tritrophic or bitrophic) in which one species moves between patches, with energy cost or mortality risk in movement. In the two-patch case, assuming forced movement in one direction, an evolutionarily stable strategy requires bidirectional movement, even if costs during movement are high. In the N -patch case, assuming that at least one patch is linked bidirectionally to all other patches, optimal movement rates can lead to source-sink dynamics where patches with negative growth rates are maintained by other patches with positive growth rates. As well, dispersal between patches is not balanced (even in the two-patch case), leading to a deviation from the IFD. Our results indicate that cost-associated forced movement can have important consequences for spatial metapopulation dynamics. Relevance to marine reserve design and the study of stream communities subject to drift is discussed.

American Naturalist↗

Carryover effects associated with winter location affect fitness, social status, and population dynamics in a long-distance migrant

We used observations of individually marked female black brant geese (Branta bernicla nigricans; brant) at three wintering lagoons on the Pacific coast of Baja California—Laguna San Ignacio (LSI), Laguna Ojo de Liebre (LOL), and Bahía San Quintín (BSQ)—and the Tutakoke River breeding colony in Alaska to assess hypotheses about carryover effects on breeding and distribution of individuals among wintering areas. We estimated transition probabilities from wintering locations to breeding and nonbreeding by using multistratum robust-design capture-mark-recapture models. We also examined the effect of breeding on migration to wintering areas to assess the hypothesis that individuals in family groups occupied higher-quality wintering locations. We used 4,538 unique female brant in our analysis of the relationship between winter location and breeding probability. All competitive models of breeding probability contained additive effects of wintering location and the 1997–1998 El Niño–Southern Oscillation (ENSO) event on probability of breeding. Probability of breeding in non-ENSO years was 0.98 ± 0.02, 0.68 ± 0.04, and 0.91 ± 0.11 for females wintering at BSQ, LOL, and LSI, respectively. After the 1997–1998 ENSO event, breeding probability was between 2% (BSQ) and 38% (LOL) lower than in other years. Individuals that bred had the highest probability of migrating the next fall to the wintering area producing the highest probability of breeding.

American Naturalist↗

Nest predation risk and growth strategies of passerine species: grow fast or develop traits to escape risk?

Different body components are thought to trade off in their growth and development rates, but the causes for relative prioritization of any trait remains a critical question. Offspring of species at higher risk of predation might prioritize development of locomotor traits that facilitate escaping risky environments over growth of mass. We tested this possibility in 12 altricial passerine species that differed in their risk of nest predation. We found that rates of growth and development of mass, wings, and endothermy increased with nest predation risk across species. In particular, species with higher nest predation risk exhibited relatively faster growth of wings than of mass, fledged with relatively larger wing sizes and smaller mass, and developed endothermy earlier at relatively smaller mass. This differential development can facilitate both escape from predators and survival outside of the nest environment. Tarsus growth was not differentially prioritized with respect to nest predation risk, and instead all species achieved adult tarsus size by age of fledging. We also tested whether different foraging modes (aerial, arboreal, and ground foragers) might explain the variation of differential growth of locomotor modules, but we found that little residual variation was explained. Our results suggest that differences in nest predation risk among species are associated with relative prioritization of body components to facilitate escape from the risky nest environment.

American Naturalist↗

Thermal sensitivity of immune function: evidence against a generalist-specialist trade-off among endothermic and ectothermic vertebrates

Animal body temperature (Tbody) varies over daily and annual cycles, affecting multiple aspects of biological performance in both endothermic and ectothermic animals. Yet a comprehensive comparison of thermal performance among animals varying in Tbody (mean and variance) and heat production is lacking. Thus, we examined the thermal sensitivity of immune function (a crucial fitness determinant) in Vertebrata, a group encompassing species of varying thermal biology. Specifically, we investigated temperature-related variation in two innate immune performance metrics, hemagglutination and hemolysis, for 13 species across all seven major vertebrate clades. Agglutination and lysis were temperature dependent and were more strongly related to the thermal biology of species (e.g., mean Tbody) than to the phylogenetic relatedness of species, although these relationships were complex and frequently surprising (e.g., heterotherms did not exhibit broader thermal performance curves than homeotherms). Agglutination and lysis performance were positively correlated within species, except in taxa that produce squalamine, a steroidal antibiotic that does not lyse red blood cells. Interestingly, we found the antithesis of a generalist-specialist trade-off: species with broader temperature ranges of immune performance also had higher peak performance levels. In sum, we have uncovered thermal sensitivity of immune performance in both endotherms and ectotherms, highlighting the role that temperature and life history play in immune function across Vertebrata.

American Naturalist↗

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 paper 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.

American Naturalist↗

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.

American Naturalist↗

Evolution of pathogen virulence across space during an epidemic

We explore pathogen virulence evolution during the spatial expansion of an infectious disease epidemic in the presence of a novel host movement trade-off, using a simple, spatially explicit mathematical model. This work is motivated by empirical observations of the Mycoplasma gallisepticum invasion into North American house finch ( Haemorhous mexicanus ) populations; however, our results likely have important applications to other emerging infectious diseases in mobile hosts. We assume that infection reduces host movement and survival and that across pathogen strains the severity of these reductions increases with pathogen infectiousness. Assuming these trade-offs between pathogen virulence (host mortality), pathogen transmission, and host movement, we find that pathogen virulence levels near the epidemic front (that maximize wave speed) are lower than those that have a short-term growth rate advantage or that ultimately prevail (i.e., are evolutionarily stable) near the epicenter and where infection becomes endemic (i.e., that maximize the pathogen basic reproductive ratio). We predict that, under these trade-offs, less virulent pathogen strains will dominate the periphery of an epidemic and that more virulent strains will increase in frequency after invasion where disease is endemic. These results have important implications for observing and interpreting spatiotemporal epidemic data and may help explain transient virulence dynamics of emerging infectious diseases.

American Naturalist↗

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.

The American Naturalist↗

Reproductive allochrony in seasonally sympatric populations maintained by differential response to photoperiod: Implications for population divergence and response to climate change

Reproductive allochrony presents a potential barrier to gene flow and is common in seasonally sympatric migratory and sedentary birds. Mechanisms mediating reproductive allochrony can influence population divergence and the capacity of populations to respond to environmental change. We asked whether reproductive allochrony in seasonally sympatric birds results from a difference in response to supplementary or photoperiodic cues and whether the response varies in relation to the distance separating breeding and wintering locations as measured by stable isotopes. We held seasonally sympatric migratory and sedentary male dark-eyed juncos ( Junco hyemalis ) in a common garden in early spring under simulated natural changes in photoperiod and made measurements of reproductive and migratory physiology. On the same dates and photoperiods, sedentary juncos had higher testosterone (initial and gonadotropin-releasing hormone induced), more developed cloacal protuberances, and larger testes than migrants. In contrast, migratory juncos had larger fat reserves (fuel for migration). We found a negative relationship between testis mass and feather hydrogen isotope ratios, indicating that testis growth was more delayed in migrants making longer migrations. We conclude that reproductive allochrony in seasonally sympatric migratory and sedentary birds can result from a differential response to photoperiodic cues in a common garden, and as a result, gene flow between migrants and residents may be reduced by photoperiodic control of reproductive development. Further, earlier breeding in response to future climate change may currently be constrained by differential response to photoperiodic cues.

American Naturalist↗

Fatal attraction? Intraguild facilitation and suppression among predators

Competition and suppression are recognized as dominant forces that structure predator communities. Facilitation via carrion provisioning, however, is a ubiquitous interaction among predators that could offset the strength of suppression. Understanding the relative importance of these positive and negative interactions is necessary to anticipate community-wide responses to apex predator declines and recoveries worldwide. Using state-sponsored wolf ( Canis lupus ) control in Alaska as a quasi experiment, we conducted snow track surveys of apex, meso-, and small predators to test for evidence of carnivore cascades (e.g., mesopredator release). We analyzed survey data using an integrative occupancy and structural equation modeling framework to quantify the strengths of hypothesized interaction pathways, and we evaluated fine-scale spatiotemporal responses of nonapex predators to wolf activity clusters identified from radio-collar data. Contrary to the carnivore cascade hypothesis, both meso- and small predator occupancy patterns indicated guild-wide, negative responses of nonapex predators to wolf abundance variations at the landscape scale. At the local scale, however, we observed a near guild-wide, positive response of nonapex predators to localized wolf activity. Local-scale association with apex predators due to scavenging could lead to landscape patterns of mesopredator suppression, suggesting a key link between occupancy patterns and the structure of predator communities at different spatial scales.

The American Naturalist↗

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.

The American Naturalist↗

Brood size affects future reproduction in a long-lived bird with precocial young

Estimation of trade-offs between current reproduction and future survival and fecundity of long-lived vertebrates is essential to understanding factors that shape optimal reproductive investment. Black brant geese (Branta bernicla nigricans) fledge more goslings, on average, when their broods are experimentally enlarged to be greater than the most common clutch size of four eggs. Thus, we hypothesized that the lesser frequency of brant clutches exceeding four eggs results, at least partially, from a future reduction in survival, breeding probability, or clutch size for females tending larger broods. We used an eight-year mark-recapture dataset (Barker robust design) with five years of clutch and brood manipulations to estimate long-term consequences of reproductive decisions in brant. We did not find evidence of a trade-off between reproductive effort and true survival or future initiation date and clutch size. Rather, future breeding probability was maximized (0.92 ± 0.03 [se]) for manipulated females tending broods of four goslings and lower for females tending smaller (one gosling; 0.63 ± 0.09 [se]) or larger broods (seven goslings; 0.52 ± 0.15 [se]). Our results suggest that demographic trade-offs for female brant tending large broods may reduce the fitness value of clutches larger than four and, therefore, contribute to the paucity of larger clutches. The lack of a trade-off between reproductive effort and survival provides evidence that survival, to which fitness is most sensitive in long-lived animals, is buffered against temporal variation in brant.

American Naturalist↗

Higher nest predation favors rapid fledging at the cost of plumage quality in nestling birds

Life-history theory predicts that rapid growth comes at a cost to offspring quality and adult longevity. However, trade-offs have been examined primarily based on proximate variation within species rather than evolved differences across species. Evolved differences are important to examine because species may co-evolve mechanisms to reduce long-term costs of rapid growth. For example, trade-offs associated with fast growth and short development time might be expressed in ephemeral traits because costs associated with these traits are paid for only a short duration. We explored this idea using nestling body feathers, an ephemeral trait, and examined whether shorter development time yielded lower quality feathers. We found a strong trade-off whereby nestlings that spend more time in the nest produced higher quality plumage across 123 temperate and tropical species. Using a subset of these species (n=67), we found that plumage quality was lower in species with higher rates of nest predation, which favors rapid nestling growth and short development times. Our results suggest that evolution of more rapid growth may co-evolve with other mechanisms whereby long-term costs are reduced by shifting costs of allocational trade-offs to ephemeral traits that are short-lived.

The American Naturalist↗

Life-history plasticity and water-use trade-offs associated with drought resistance in a clade of California jewelflowers

Water limitation is a primary driver of plant geographic distributions and individual plant fitness. Drought resistance is the ability to survive and reproduce despite limited water, and numerous studies have explored its physiological basis in plants. However, it is unclear how drought resistance and trade-offs associated with drought resistance evolve within plant clades. We quantified the relationship between water availability and fitness for 13 short-lived plant taxa in the Streptanthus clade that vary in their phenology and the availability of water in the environments where they occur. We derived two parameters from these relationships: plant fitness when water is not limiting and the water inflection point (WIF), the watering level at which additional water is most efficiently turned into fitness. We used phylogenetic comparative methods to explore trade-offs related to drought resistance and trait plasticity and the degree to which water relationship parameters are conserved. Taxa from drier climates produced fruits at the lowest water levels, had a lower WIF, flowered earlier, had shorter life spans, had greater plastic water-use efficiency (WUE), and had lower fitness at nonlimiting water. In contrast, later-flowering Streptanthus taxa from less xeric climates experienced high fitness at nonlimiting water but had no fitness at the lowest water levels. Across the clade, we found a trade-off between drought resistance and fitness at high water, though a single ruderal species was an outlier in this relationship. Our results suggest that drought escape trades off with maximal fitness under nonlimiting water, and both are tied to phenology. We also found that variation in trait plasticity determines how different plant species produce fitness over a water gradient.

California↗