A grazer's niche edge is associated with increasing diet diversity and poor population performance.

The core-periphery hypothesis predicts niche cores should be associated with greater survivorship, reproductive output and population performance rates than marginal habitats at niche edges. However, there is very little empirical evidence of whether niche centrality influences population trends in animals. Using the Cape mountain zebra (Equus zebra zebra) as a model system, we evaluated whether niche centrality is associated with population trends, resource availability and diet across a core-periphery gradient. Population growth rates and density progressively declined towards niche peripheries. Niche peripheries were resource-poor and Cape mountain zebra consumed more phylogenetically diverse diets dominated by non-grass families. In core habitats they consumed grass-rich diets and female reproductive success was higher. This combination of spatial niche modelling and functional ecology provides a novel evaluation of how bottom-up resource limitation can shape species distributions, population resilience and range change and can guide conservation management.

Recolonization following past persecution questions the importance of persistent snow cover as a range limiting factor for wolverines.

Globally, climate is changing rapidly, which causes shifts in many species' distributions, stressing the need to understand their response to changing environmental conditions to inform conservation and management. Northern latitudes are expected to experience strongest changes in climate, with milder winters and decreasing snow cover. The wolverine (Gulo gulo) is a circumpolar, threatened carnivore distributed in northern tundra, boreal, and subboreal habitats. Previous studies have suggested that wolverine distribution and reproduction are constrained by a strong association with persistent spring snow cover. We assess this hypothesis by relating spatial distribution of 1589 reproductive events, a fitness-related proxy for female reproduction and survival, to snow cover over two decades. Wolverine distribution has increased and number of reproductive events increased 20 times in areas lacking spring snow cover during our study period, despite low monitoring effort where snow is sparse. Thus, the relationship between reproductive events and persistent spring snow cover weakened during this period. These findings show that wolverine reproductive success and hence distribution are less dependent on spring snow cover than expected. This has important implications for projections of future habitat availability, and thus distribution, of this threatened species. Our study also illustrates how past persecution, or other factors, that have restricted species distribution to remote areas can mask actual effects of environmental parameters, whose importance reveals when populations expand beyond previously restricted ranges. Overwhelming evidence shows that climate change is affecting many species and ecological processes, but forecasting potential consequences on a given species requires longitudinal data to revisit hypotheses and reassess the direction and magnitude of climate effects with new data. This is especially important for conservation-oriented management of species inhabiting dynamic systems where environmental factors and human activities interact, a common scenario for many species in different ecosystems around the globe.

Host population dynamics in the face of an evolving pathogen.

Interactions between hosts and pathogens are dynamic at both ecological and evolutionary levels. In the resultant 'eco-evolutionary dynamics' ecological and evolutionary processes affect each other. For example, the house finch Haemorhous mexicanus and its recently emerged pathogen, the bacterium Mycoplasma gallisepticum, form a system in which evidence suggests that changes in bacterial virulence through time enhance levels of host immunity in ways that drive the evolution of virulence in an arms race. We use data from two associated citizen science projects in order to determine whether this arms race has had any detectable effect at the population level in the north-eastern United States. We used data from two citizen science projects, based on observations of birds at bird feeders, which provide information on the long-term changes in sizes of aggregations of house finches (host population density), and the probabilities that these house finches have observable disease (disease prevalence). The initial emergence of M. gallisepticum caused a rapid halving of house finch densities; this was then followed by house finch populations remaining stable or slowly declining. Disease prevalence also decreased sharply after the initial emergence and has remained low, although with fluctuations through time. Surprisingly, while initially higher local disease prevalence was found at sites with higher local densities of finches, this relationship has reversed over time. The ability of a vertebrate host species, with a generation time of at least 1 year, to maintain stable populations in the face of evolved higher virulence of a bacterium, with generation times measurable in minutes, suggests that genetic changes in the host are insufficient to explain the observed population-level patterns. We suggest that acquired immunity plays an important role in the observed interaction between house finches and M. gallisepticum.

The population density of an urban raptor is inextricably tied to human cultural practices.

Human socio-cultural factors are recognized as fundamental drivers of urban ecological processes, but their effect on wildlife is still poorly known. In particular, human cultural aspects may differ substantially between the extensively studied urban settings of temperate regions and the poorly studied cities of the tropics, which may offer profoundly different niches for urban wildlife. Here, we report how the population levels of a scavenging raptor which breeds in the megacity of Delhi, the black kite Milvus migrans, depend on spatial variation in human subsidies, mainly in the form of philanthropic offerings of meat given for religious purposes. This tight connection with human culture, which generated the largest raptor concentration in the world, was modulated further by breeding-site availability. The latter constrained the level of resource-tracking by the kites and their potential ecosystem service, and could be used as a density-management tool. Similar ties between animal population densities, key anthropogenic resources and human beliefs may occur in thousands of cities all over the globe and may fit poorly with our current understanding of urban ecosystem functioning. For many urban animals, key resources are inextricably linked with human culture, an aspect that has been largely overlooked.

Spatial and temporal drivers of avian population dynamics across the annual cycle.

Untangling the spatial and temporal processes that influence population dynamics of migratory species is challenging, because changes in abundance are shaped by variation in vital rates across heterogeneous habitats and throughout the annual cycle. We developed a full-annual-cycle, integrated, population model and used demographic data collected between 2011 and 2014 in southern Indiana and Belize to estimate stage-specific vital rates of a declining migratory songbird, the Wood Thrush (Hylocichla mustelina). Our primary objective was to understand how spatial and temporal variation in demography contributes to local and regional population growth. Our full-annual-cycle model allowed us to estimate (1) age-specific, seasonal survival probabilities, including latent survival during both spring and autumn migration, and (2) how the relative contribution of vital rates to population growth differed among habitats. Wood Thrushes in our study populations experienced the lowest apparent survival rates during migration and apparent survival was lower during spring migration than during fall migration. Both mortality and high dispersal likely contributed to low apparent survival during spring migration. Population growth in high-quality habitat was most sensitive to variation in fecundity and apparent survival of juveniles during spring migration, whereas population growth in low-quality sites was most sensitive to adult apparent breeding-season survival. These results elucidate how full-annual-cycle vital rates, particularly apparent survival during migration, interact with spatial variation in habitat quality to influence population dynamics in migratory species.

Quantifying drivers of population dynamics for a migratory bird throughout the annual cycle.

Worldwide, migratory species are undergoing rapid declines but understanding the factors driving these declines is hindered by missing information about migratory connectivity and the lack of data to quantify environmental processes across the annual cycle. Here, we combined range-wide information about migratory connectivity with global remote-sensing data to quantify the relative importance of breeding and non-breeding environmental processes to persistent long-term population declines of a migratory songbird, the wood thrush (Hylocichla mustelina). Consistent with theoretical predictions about population limitation of migratory birds, our results suggest that habitat loss and climate have contributed to the observed declines in wood thrush breeding abundance, yet the relative importance of breeding versus non-breeding factors is population-specific. For example, high-abundance core breeding populations appear to be more limited by habitat loss, whereas low-abundance, peripheral populations appear to be limited by climate-driven seasonal interactions. Further, our analysis indicates that the relative impact of breeding habitat loss is at least three to six times greater than the impact of equivalent non-breeding habitat loss and therefore the steepest regional declines have likely been driven by the loss of breeding habitat. These results underscore the need for population-specific conservation strategies implemented throughout the annual cycle to reverse long-term declines.

Food provisioning alters infection dynamics in populations of a wild rodent.

While pathogens are often assumed to limit the growth of wildlife populations, experimental evidence for their effects is rare. A lack of food resources has been suggested to enhance the negative effects of pathogen infection on host populations, but this theory has received little investigation. We conducted a replicated two-factor enclosure experiment, with introduction of the bacterium Bordetella bronchiseptica and food supplementation, to evaluate the individual and interactive effects of pathogen infection and food availability on vole populations during a boreal winter. We show that prior to bacteria introduction, vole populations were limited by food availability. Bordetella bronchiseptica introduction then reduced population growth and abundance, but contrary to predictions, primarily in food supplemented populations. Infection prevalence and pathological changes in vole lungs were most common in food supplemented populations, and are likely to have resulted from increased congregation and bacteria transmission around feeding stations. Bordetella bronchiseptica-infected lungs often showed protozoan co-infection (consistent with Hepatozoon erhardovae), together with more severe inflammatory changes. Using a multidisciplinary approach, this study demonstrates a complex picture of interactions and underlying mechanisms, leading to population-level effects. Our results highlight the potential for food provisioning to markedly influence disease processes in wildlife mammal populations.

Seasonal demography of a cyclic lemming population in the Canadian Arctic.

1. The causes of cyclical fluctuations in animal populations remain a controversial topic in ecology. Food limitation and predation are two leading hypotheses to explain small mammal population dynamics in northern environments. We documented the seasonal timing of the decline phases and demographic parameters (survival and reproduction) associated with population changes in lemmings, allowing us to evaluate some predictions from these two hypotheses. 2. We studied the demography of brown lemmings (Lemmus trimucronatus), a species showing 3- to 4-year population cycles in the Canadian Arctic, by combining capture-mark-recapture analysis of summer live-trapping with monitoring of winter nests over a 10-year period. We also examined the effects of some weather variables on survival. 3. We found that population declines after a peak occurred between the summer and winter period and not during the winter. During the summer, population growth was driven by change in survival, but not in fecundity or proportion of juveniles, whereas in winter population growth was driven by changes in late summer and winter reproduction. 4. We did not find evidence for direct density dependence on summer demographic parameters, though our analysis was constrained by the paucity of data during the low phase. Body mass, however, was highest in peak years. 5. Weather effects were detected only in early summer when lemming survival was positively related to snow depth at the onset of melt but negatively related to rainfall. 6. Our results show that high mortality causes population declines of lemmings during summer and fall, which suggests that predation is sufficient to cause population crashes, whereas high winter fecundity is the primary factor leading to population irruptions. The positive association between snow depth and early summer survival may be due to the protective cover offered by snow against predators. It is still unclear why reproduction remains low during the low phase.

Population responses of small mammals to food supply and predators: a global meta-analysis.

1. The relative importance of food supply and predation as determinants of animal population density is a topic of enduring debate among ecologists. To address it, many studies have tested the potential effects of food on population density by experimentally supplementing natural populations, with much focus on terrestrial vertebrates, especially small mammals. 2. Here we perform a meta-analysis of such experiments, testing two complementary hypotheses: (i) small mammal populations are bottom-up limited and (ii) population increases in response to food supplementation are constrained by predation, a top-down limitation. 3. In the 148 experiments recorded, food supplementation had an overall positive and significant effect, increasing population densities by 1.5-fold. Larger population increases occurred when predation was reduced and populations were open to immigration. Predation appeared to be unimportant when populations were closed to immigration. Immigration was the major mechanism underlying increases in abundance by increasing local population density and crowding. Contributions of increased reproductive rate could be detected, but were minor compared to immigration, and no effects were detected from survival. 4. Our analyses support the view that animal population density is determined by both bottom-up and top-down forces. They also suggest the possibility that food supplementation experiments might unintentionally create ecological traps by aggregating both prey and predators in small areas of the landscape. We suggest an alternative experimental design to increase the contribution that food supplementation experiments can make in future.

Numbers and distribution of the Great Cormorant in Iceland: Limitation at the regional and metapopulation level.

We studied a metapopulation of great cormorant (Phalacrocorax carbo) in Iceland, using complete aerial censuses of nests in 25 years during 1975-2015. Age composition was estimated in 1998-2014 by ground surveys in September and February. Brood size was estimated from aerial photographs in 2007-2015.Weather, food, breeding habitat, and density were considered as explanatory variables when examining numerical and distributional changes in the cormorant metapopulation.In 1975-1990 total nest numbers changed little, very low numbers about 1992 were followed by an annual increase of 3.5% in 1994-2015. Total nest numbers were positively correlated with estimates of spawning stocks of cod and saithe and inversely related to the subpolar gyre index (SPG-I).During the increase in 1994-2015, average colony size at first increased and then declined. Habitat use also changed: the proportion of nests on small rocky islets (skerries) at first declined, from 69% to 44% in 1995-2003 and then increased again to about 58% in 2012-2014. Habitat changes were probably a response to changed patterns of human disturbance.Breeding density, as nests per km2 sea <20 m deep, was rather uniform among five defined regions in 1975-1996. Thereafter, densities became much higher in two sheltered regions with kelp forests and in one mostly exposed region. A second exposed region remained low and in the third nest numbers declined markedly. Thus, carrying capacity was higher in sheltered regions where cormorant breeding had historically been depressed by human disturbance.Brood size varied little among regions but declined with the years from about 2.5 to 1.8.The proportion of juveniles in September (fecundity) declined in 1998-2015 from over 0.4 to 0.3 and was inversely correlated with year and nest numbers, if outlier years were excluded, suggesting resource limitation. Survival of juvenile cormorants in September-February was estimated at 0.471 ± 0.066 SE. Commercial fish stocks and climate indices were not correlated with the proportion of juveniles.Annual survival of adults (breeding and nonbreeding) was estimated from nest counts and age composition 1999-2014, as 0.850 ± 0.026 SE and showed no trend in 1998-2014.We conclude that the metapopulation of cormorants in Iceland was resource-limited at two levels: fecundity at the regional and winter survival at the total level.

Resource limitation, competition and the influence of life history in a freshwater snail community.

Previous work on a snail community occurring throughout lakes in southwestern Michigan showed that predation by molluscivorous sunfish had large impacts on only the rarest snail species. Thus, competition might play a major role in population limitation because dominant members of the snail community are relatively immune to predation. The present experiments were conducted to determine the extent to which the snail community depleted the abundance of food resources (epiphytes) and the extent to which epiphyte abundances limited snail production. An experimental gradient in snail densities showed that removal of snails increased epiphyte biomass by approximately 3-fold relative to that observed at natural snail densities. Enrichment of the environment with phosphorus fertilizer increased epiphyte biomass by approximately 20-fold and provided a test of food limitation in the snail community. All snail taxa exhibited positive numerical or growth responses to enrichment. The observations that snails depleted resources and that resources limited snail production demonstrated that snails competed exploitatively for epiphytes. The response of each snail species to increased food abundance differed depending on the timing of fertilization relative to the snails' life histories. Snails hatched before the experiment began were larger in fertilized treatments, due to increased growth, but their densities were similar among treatments. On the other hand, densities of snails born during the experiment were up to 15-fold greater in fertilized treatments, due in part to increased survival of newborn snails. Comparison of the responses of snails to food addition and to predator removals (based on prior experiments) suggested that food availability limits snail production more than predators do. Additionally, the large responses by algae and snails to fertilization demonstrated that both the producers and herbivores in this simplified food chain were strongly resource limited.

Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses.

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection.