Social foraging and the associated benefits of group-living in Cliff Swallows decrease over 40 years
Animals that feed socially can sometimes better locate prey, often by transferring information about food that is patchy, dense, and temporally and spatially unpredictable. Information transfer is a potential benefit of living in breeding colonies where unsuccessful foragers can more readily locate successful ones and thereby improve feeding efficiency. Most studies on social foraging have been short-term, and how long-term environmental change affects both foraging strategies and the associated benefits of coloniality is generally unknown. In the colonial Cliff Swallow (), we examined how social foraging, information transfer, and feeding ecology changed over a 40-year period in western Nebraska. Relative to the 1980's, Cliff Swallows in 2016-2022 were more likely to forage solitarily or in smaller groups, spent less time foraging, were more successful as solitaries, fed in more variable locations, and engaged less in information transfer at the colony site. The total mass of insects brought back to nestlings per parental visit declined over the study. The diversity of insect families captured increased over time, and some insect taxa dropped out of the diet, although the three most common insect families remained the same among the decades. Nestling Cliff Swallow body mass at 10 days of age and the number of nestlings surviving per nest declined more sharply with colony size in 2015-2022 than in 1984-1991 at sites where the confounding effects of ectoparasites were removed. Adult body mass during provisioning of nestlings was lower in more recent years, but the change did not vary with colony size. The reason(s) for the reduction in social foraging and information transfer over time are unclear, but the consequence is that colonial nesting may no longer offer the same fitness advantages for Cliff Swallows as in the 1980's. The results illustrate flexibility of foraging behavior and dynamic shifts in the potential selective pressures for group-living.
Climate-mediated population dynamics of a migratory songbird differ between the trailing edge and range core
Understanding the demographic drivers of range contractions is important for predicting species' responses to climate change; however, few studies have examined the effects of climate change on survival and recruitment across species' ranges. We show that climate change can drive trailing edge range contractions through the effects on apparent survival, and potentially recruitment, in a migratory songbird. We assessed the demographic drivers of trailing edge range contractions using a long-term demography dataset for the black-throated blue warbler () collected across elevational climate gradients at the trailing edge and core of the breeding range. We used a Bayesian hierarchical model to estimate the effect of climate change on apparent survival and recruitment and to forecast population viability at study plots through 2040. The trailing edge population at the low-elevation plot became locally extinct by 2017. The local population at the mid-elevation plot at the trailing edge gradually declined and is predicted to become extirpated by 2040. Population declines were associated with warming temperatures at the mid-elevation plot, although results were more equivocal at the low-elevation plot where we had fewer years of data. Population density was stable or increasing at the range core, although warming temperatures are predicted to cause population declines by 2040 at the low-elevation plot. This result suggests that even populations within the geographic core of the range are vulnerable to climate change. The demographic drivers of local population declines varied between study plots, but warming temperatures were frequently associated with declining rates of population growth and apparent survival. Declining apparent survival in our study system is likely to be associated with increased adult emigration away from poor-quality habitats. Our results suggest that demographic responses to warming temperatures are complex and dependent on local conditions and geographic range position, but spatial variation in population declines is consistent with the climate-mediated range shift hypothesis. Local populations of black-throated blue warblers near the warm-edge range boundary at low latitudes and low elevations are likely to be the most vulnerable to climate change, potentially leading to local extirpation and range contractions.
Stable pollination service in a generalist high Arctic community despite the warming climate
Insects provide key pollination services in most terrestrial biomes, but this service depends on a multistep interaction between insect and plant. An insect needs to visit a flower, receive pollen from the anthers, move to another conspecific flower, and finally deposit the pollen on a receptive stigma. Each of these steps may be affected by climate change, and focusing on only one of them (e.g., flower visitation) may miss important signals of change in service provision. In this study, we combine data on visitation, pollen transport, and single-visit pollen deposition to estimate functional outcomes in the high Arctic plant-pollinator network of Zackenberg, Northeast Greenland, a model system for global warming-associated impacts in pollination services. Over two decades of rapid climate warming, we sampled the network repeatedly: in 1996, 1997, 2010, 2011, and 2016. Although the flowering plant and insect communities and their interactions varied substantially between years, as expected based on highly variable Arctic weather, there was no detectable directional change in either the structure of flower-visitor networks or estimated pollen deposition. For flower-visitor networks compiled over a single week, species phenologies caused major within-year variation in network structure despite consistency across years. Weekly networks for the middle of the flowering season emerged as especially important because most pollination service can be expected to be provided by these large, highly nested networks. Our findings suggest that pollination ecosystem service in the high Arctic is remarkably resilient. This resilience may reflect the plasticity of Arctic biota as an adaptation to extreme and unpredictable weather. However, most pollination service was contributed by relatively few fly taxa (Diptera: and [Muscidae] and species of [Empididae]). If these key pollinators are negatively affected by climate change, network structure and the pollination service that depends on it would be seriously compromised.
Temporal dynamics of range expander and congeneric native plant responses during and after extreme drought events
Climate change is causing range shifts of many species to higher latitudes and altitudes and increasing their exposure to extreme weather events. It has been shown that range-shifting plant species may perform differently in new soil than related natives; however, little is known about how extreme weather events affect range-expanding plants compared to related natives. In this study we used outdoor mesocosms to study how range-expanding plant species responded to extreme drought in live soil from a habitat in a new range with and without live soil from a habitat in the original range (Hungary). During summer drought, the shoot biomass of the range-expanding plant community declined. In spite of this, in the mixed community, range expanders produced more shoot biomass than congeneric natives. In mesocosms with a history of range expanders in the previous year, native plants produced less biomass. Plant legacy or soil origin effects did not change the response of natives or range expanders to summer drought. During rewetting, range expanders had less biomass than congeneric natives but higher drought resilience (survival) in soils from the new range where in the previous year native plant species had grown. The biomass patterns of the mixed plant communities were dominated by spp.; however, not all plant species within the groups of natives and of range expanders showed the general pattern. Drought reduced the litter decomposition, microbial biomass, and abundances of bacterivorous, fungivorous, and carnivorous nematodes. Their abundances recovered during rewetting. There was less microbial and fungal biomass, and there were fewer fungivorous nematodes in soils from the original range where range expanders had grown in the previous year. We concluded that in mixed plant communities of range expanders and congeneric natives, range expanders performed better, under both ambient and drought conditions, than congeneric natives. However, when considering the responses of individual species, we observed variations among pairs of congenerics, so that under the present mixed-community conditions there was no uniformity in responses to drought of range expanders versus congeneric natives. Range-expanding plant species reduced soil fungal biomass and the numbers of soil fungivorous nematodes, suggesting that the effects of range-expanding plant species can trickle up in the soil food web.
Causes and consequences of pair-bond disruption in a sex-skewed population of a long-lived monogamous seabird
Many animals form long-term monogamous pair bonds, and the disruption of a pair bond (through either divorce or widowhood) can have significant consequences for individual vital rates (survival, breeding, and breeding success probabilities) and life-history outcomes (lifetime reproductive success [LRS], life expectancy). Here, we investigated the causes and consequences of pair-bond disruption in wandering albatross (). State-of-the-art statistical and mathematical approaches were developed to estimate divorce and widowhood rates and their impacts on vital rates and life-history outcomes. In this population, females incur a higher mortality rate due to incidental fishery bycatch, so the population is male-skewed. Therefore, we first posited that males would show higher widowhood rates negatively correlated with fishing effort and females would have higher divorce rates because they have more mating opportunities. Furthermore, we expected that divorce could be an adaptive strategy, whereby individuals improved breeding success by breeding with a new partner of better quality. Finally, we posited that pair-bond disruptions could reduce survival and breeding probabilities owing to the cost of remating processes, with important consequences for life-history outcomes. As expected, we showed that males had higher widowhood rates than females and females had higher divorce rates in this male-skewed population. However, no correlation was found between fishing effort and male widowhood. Secondly, contrary to our expectation, we found that divorce was likely nonadaptive in this population. We propose that divorce in this population is caused by an intruder who outcompetes the original partner in line with the so-called forced divorce hypothesis. Furthermore, we found a 16.7% and 18.0% reduction in LRS only for divorced and widowed males, respectively, owing to missing breeding seasons after a pair-bond disruption. Finally, we found that divorced individuals were more likely to divorce again, but whether this is related to specific individual characteristics remains an important area of investigation.
Parasites in kelp-forest food webs increase food-chain length, complexity, and specialization, but reduce connectance
We explored whether parasites are important in kelp forests by examining their effects on a high-quality, high-resolution kelp-forest food web. After controlling for generic effects of network size, parasites affected kelp-forest food web structure in some ways consistent with other systems. Parasites increased the trophic span of the web, increasing top predator vulnerability and the longest chain length. Unique links associated with parasites, such as concomitant predation (consumption of parasites along with their hosts by predators) increased the frequency of network motifs involving mutual consumption and decreased niche contiguity of free-living species. However, parasites also affected kelp-forest food web structure in ways not seen in other systems. Kelp-forest parasites are richer and more specialized than other systems. As a result, parasites reduced diet generality and decreased connectance in the kelp forest. Although mutual consumption motifs increased in frequency, this motif type was still a small fraction of all possible motifs, so their increase in frequency was not enough to compensate for the decrease in connectance caused by adding many specialist parasite species.
Remotely detected aboveground plant function predicts belowground processes in two prairie diversity experiments
Imaging spectroscopy provides the opportunity to incorporate leaf and canopy optical data into ecological studies, but the extent to which remote sensing of vegetation can enhance the study of belowground processes is not well understood. In terrestrial systems, aboveground and belowground vegetation quantity and quality are coupled, and both influence belowground microbial processes and nutrient cycling. We hypothesized that ecosystem productivity, and the chemical, structural and phylogenetic-functional composition of plant communities would be detectable with remote sensing and could be used to predict belowground plant and soil processes in two grassland biodiversity experiments: the BioDIV experiment at Cedar Creek Ecosystem Science Reserve in Minnesota and the Wood River Nature Conservancy experiment in Nebraska. We tested whether aboveground vegetation chemistry and productivity, as detected from airborne sensors, predict soil properties, microbial processes and community composition. Imaging spectroscopy data were used to map aboveground biomass, green vegetation cover, functional traits and phylogenetic-functional community composition of vegetation. We examined the relationships between the image-derived variables and soil carbon and nitrogen concentration, microbial community composition, biomass and extracellular enzyme activity, and soil processes, including net nitrogen mineralization. In the BioDIV experiment-which has low overall diversity and productivity despite high variation in each-belowground processes were driven mainly by variation in the amount of organic matter inputs to soils. As a consequence, soil respiration, microbial biomass and enzyme activity, and fungal and bacterial composition and diversity were significantly predicted by remotely sensed vegetation cover and biomass. In contrast, at Wood River-where plant diversity and productivity were consistently higher-belowground processes were driven mainly by variation in the quality of aboveground inputs to soils. Consequently, remotely sensed functional, chemical and phylogenetic composition of vegetation predicted belowground extracellular enzyme activity, microbial biomass, and net nitrogen mineralization rates but aboveground biomass (or cover) did not. The contrasting associations between the quantity (productivity) and quality (composition) of aboveground inputs with belowground soil attributes provide a basis for using imaging spectroscopy to understand belowground processes across productivity gradients in grassland systems. However, a mechanistic understanding of how above and belowground components interact among different ecosystems remains critical to extending these results broadly.
Water and nitrogen shape winter annual plant diversity and community composition in near-urban Sonoran Desert preserves
Increased nitrogen (N) deposition threatens global biodiversity, but its effects in arid urban ecosystems are not well studied. In addition to altered N availability, urban environments also experience increases in other pollutants, decreased population connectivity, and altered biotic interactions, which can further impact biodiversity. In deserts, annual plant communities make up most of the plant diversity, support wildlife, and contribute to nutrient cycling and ecosystem processes. Functional tradeoffs allowing coexistence of a diversity of annual plant species are well established, but maintenance of diversity in urban conditions and with increased availability of limiting nutrients has not been explored. We conducted a 13-year N and phosphorus (P) addition experiment in Sonoran Desert preserves in and around Phoenix, AZ, to test how nutrient availability interacts with growing season precipitation, urban location, and microhabitat to affect winter annual plant diversity. Using structural equation modeling and generalized linear mixed modeling, we found that annual plant taxonomic diversity was significantly reduced in N-enriched and urban plots. Water availability in both current and previous growing seasons impacted annual plant diversity, with significant interaction effects showing increased diversity in wetter years and greater responsiveness of the community to water following a wet year. However, there were no significant interactions between N enrichment and water availability, urban location, or microhabitat. Lowered diversity in urban preserves may be partly attributable to increased urban N deposition. Changes in biodiversity of showy species like annual wildflowers in urban preserves can have important implications for connections between urban residents and nature, and reduced diversity and community restructuring with N enrichment represents a challenge for future preservation of aridland biodiversity.
Nitrogen deposition and climate change effects on tree species composition and ecosystem services for a forest cohort
The composition of forests in the northeastern United States and the ecosystem services they provide to future generations will depend on several factors. In this paper, we isolate the effects of two environmental drivers, nitrogen (N) deposition and climate (temperature and precipitation) change, through an analysis of a single cohort of 24 dominant tree species. We assembled a tree database using data from U.S. Forest Service Forest Inventory and Analysis monitoring plots. Applying observed species-specific growth and survival responses, we simulated how forest stands in a 19-state study area would change from 2005 to 2100 under 12 different future N deposition-climate scenarios. We then estimated implications for three selected forest ecosystem services: merchantable timber, aboveground carbon sequestration, and tree diversity. Total tree biomass (for 24 species combined) was positively associated with both increased N deposition and temperatures; however, due to differences in the direction and magnitude of species-specific responses, forest composition varied across scenarios. For example, red maple () trees gained biomass under scenarios with more N deposition and more climate change, whereas biomass of yellow birch () and red pine () was negatively affected. Projections for ecosystem services also varied across scenarios. Carbon sequestration, which is positively associated with biomass accumulation, increased with N deposition and increasing climate change. Total timber values also increased with overall biomass; however, scenarios with increasing climate change tended to favor species with lower merchantable value, whereas more N deposition favored species with higher merchantable value. Tree species diversity was projected to decrease with greater changes in climate (warmer temperatures), especially in the northwestern, central, and southeastern portions of the study area. In contrast, the effects of N deposition on diversity varied greatly in magnitude and direction across the study area. This study highlights species-specific and regional effects of N deposition and climate change in northeastern U.S. forests, which can inform management decision for air quality and forests in the region, as well as climate policy. It also provides a foundation for future studies that may incorporate other important factors such as multiple cohorts, sulfur deposition, insects, and diseases.
Age × stage-classified demographic analysis: a comprehensive approach
This paper presents a comprehensive theory for the demographic analysis of populations in which individuals are classified by both age and stage. The earliest demographic models were age classified. Ecologists adopted methods developed by human demographers and used life tables to quantify survivorship and fertility of cohorts and the growth rates and structures of populations. Later, motivated by studies of plants and insects, matrix population models structured by size or stage were developed. The theory of these models has been extended to cover all the aspects of age-classified demography and more. It is a natural development to consider populations classified by both age and stage. A steady trickle of results has appeared since the 1960s, analyzing one or another aspect of age × stage-classified populations, in both ecology and human demography. Here, we use the vec-permutation formulation of multistate matrix population models to incorporate age- and stage-specific vital rates into demographic analysis. We present cohort results for the life table functions (survivorship, mortality, and fertility), the dynamics of intra-cohort selection, the statistics of longevity, the joint distribution of age and stage at death, and the statistics of life disparity. Combining transitions and fertility yields a complete set of population dynamic results, including population growth rates and structures, net reproductive rate, the statistics of lifetime reproduction, and measures of generation time. We present a complete analysis of a hypothetical model species, inspired by poecilogonous marine invertebrates that produce two kinds of larval offspring. Given the joint effects of age and stage, many familiar demographic results become multidimensional, so calculations of marginal and mixture distributions are an important tool. From an age-classified point of view, stage structure is a form of unobserved heterogeneity. From a stage-classified point of view, age structure is unobserved heterogeneity. In an age × stage-classified model, variance in demographic outcomes can be partitioned into contributions from both sources. Because these models are formulated as matrices, they are amenable to a complete sensitivity analysis. As more detailed and longer longitudinal studies are developed, age × stage-classified demography will become more common and more important.
From early life to senescence: individual heterogeneity in a long-lived seabird
Although population studies have long assumed that all individuals of a given sex and age are identical, ignoring among-individual differences may strongly bias our perception of eco-evolutionary processes. Individual heterogeneity, often referred to as individual quality, has received increasing research attention in the last decades. However, there are still substantial gaps in our current knowledge. For example, there is little information on how individual heterogeneity influences various life-history traits simultaneously, and studies describing individual heterogeneity in wild populations are generally not able to jointly identify possible sources of this variation. Here, based on a mark-recapture data set of 9,685 known-aged Wandering Albatrosses (), we investigated the existence of individual quality over the entire life cycle of this species, from early life to senescence. Using finite mixture models, we investigated the expression of individual heterogeneity in various demographic traits, and examined the origin of these among-individual differences by considering the natal environmental conditions. We found that some individuals consistently outperformed others during most of their life. In old age, however, the senescence rate was stronger in males that showed high demographic performance at younger ages. Variation in individual quality seemed strongly affected by extrinsic factors experienced during the ontogenetic period. We found that individuals born in years with high population density tended to have lower performances during their lifespan, suggesting delayed density dependence effects through individual quality. Our study showed that among-individual differences could be important in structuring individual life history trajectories, with substantial consequences at higher ecological levels such as population dynamics.
The impact of future forest dynamics on climate: interactive effects of changing vegetation and disturbance regimes
Currently, the temperate forest biome cools the earth's climate and dampens anthropogenic climate change. However, climate change will substantially alter forest dynamics in the future, affecting the climate regulation function of forests. Increasing natural disturbances can reduce carbon uptake and evaporative cooling, but at the same time increase the albedo of a landscape. Simultaneous changes in vegetation composition can mitigate disturbance impacts, but also influence climate regulation directly (e.g., via albedo changes). As a result of a number of interactive drivers (changes in climate, vegetation, and disturbance) and their simultaneous effects on climate-relevant processes (carbon exchange, albedo, latent heat flux) the future climate regulation function of forests remains highly uncertain. Here we address these complex interactions to assess the effect of future forest dynamics on the climate system. Our specific objectives were (1) to investigate the long-term interactions between changing vegetation composition and disturbance regimes under climate change, (2) to quantify the response of climate regulation to changes in forest dynamics, and (3) to identify the main drivers of the future influence of forests on the climate system. We investigated these issues using the individual-based forest landscape and disturbance model (iLand). Simulations were run over 200 yr for Kalkalpen National Park (Austria), assuming different future climate projections, and incorporating dynamically responding wind and bark beetle disturbances. To consistently assess the net effect on climate the simulated responses of carbon exchange, albedo, and latent heat flux were expressed as contributions to radiative forcing. We found that climate change increased disturbances (+27.7% over 200 yr) and specifically bark beetle activity during the 21st century. However, negative feedbacks from a simultaneously changing tree species composition (+28.0% broadleaved species) decreased disturbance activity in the long run (-10.1%), mainly by reducing the host trees available for bark beetles. Climate change and the resulting future forest dynamics significantly reduced the climate regulation function of the landscape, increasing radiative forcing by up to +10.2% on average over 200 yr. Overall, radiative forcing was most strongly driven by carbon exchange. We conclude that future changes in forest dynamics can cause amplifying climate feedbacks from temperate forest ecosystems.
Obtaining birth and mortality patterns from structured population trajectories
"A method is presented for unravelling the demographic equation for structured populations. A solution to the McKendrick-von Foerster equation is constructed using spline functions and this is fitted to stage-structured population data in such a way that the solution is smooth, positive, and does not imply negative death rates. The smoothness of the surface, and hence the complexity of the population model, is determined in a statistically optimum manner using cross validation. Time- and age-dependent death rates can be obtained as well as time-dependent birth rates. Confidence intervals are obtained for population size and death rates that give a 95% probability that the true population dynamics are within the intervals. Practical application of the method is demonstrated, and comparison made with three alternative methods."