Environmental stochasticity and extinction risk in a population of a small songbird, the great tit

Using a long-term demographic data set, we estimated the separate effects of demographic and environmental stochasticity in the growth rate of the great tit population in Wytham Wood, United Kingdom. Assuming logistic density regulation, both the demographic (sigma2d = 0.569) and environmental (sigma2e = 0.0793) variance, with interactions included, were significantly greater than zero. The estimates of the demographic variance seemed to be relatively insensitive to the length of the study period, whereas reliable estimates of the environmental variance required long time series (at least 15 yr of data). The demographic variance decreased significantly with increasing population density. These estimates are used in a quantitative analysis of the demographic factors affecting the risk of extinction of this population. The very long expected time to extinction of this population (approximately 10(19) yr) was related to a relatively large population size (>/=120 pairs during the study period). However, for a given population size, the expected time to extinction was sensitive to both variation in population growth rate and environmental stochasticity. Furthermore, the form of the density regulation strongly affected the expected time to extinction. Time to extinction decreased when the maximum density regulation approached K. This suggests that estimates of viability of small populations should be given both with and without inclusion of density dependence.     

Interactions between Operophtera brumata and Tortrix viridana on oak: new evidence from time-series analysis

1. Yearly population estimates of the green oak leaf roller moth, Tortrix viridana (Lepidoptera: Tortricidae), and the winter moth, Operophtera brumata (Lepidoptera: Geometridae), were taken from the pedunculate oak, Quercus robur, from 1951 to 1966 in Wytham Woods, Oxfordshire, U.K. Larval mortality from parasitoids was recorded for both species over the same period. 2. Operophtera brumata population density fluctuated around a constant mean, whereas T. viridana population density exhibited a linear decline over the sampling period. Population counts were subjected to time-series analysis after the linear decline was removed from the T. viridana data by detrending. Multiple regression models were built so that variation in the per capita rate of increase of each population could be partitioned among (a) current and previous population density, (b) current and previous population density of the second moth species, and (c) rates of larval parasitism. 3. Multiple regression analysis suggested that variation in the per capita rate of increase of O. brumata could be explained by negative feedback from O. brumata density at T–2 (32%), a negative relationship with T. viridana density at T–1 (18%), and a positive relationship with parasitism suffered by T. viridana at T–1 (24%). 4. The T. viridana population time-series was dominated by a rapid feedback process such that per capita rate of increase at time T was negatively related to population density at time T–1, explaining 53% of variance in population growth rate. Per capita rate of increase was unrelated to larval parasitism or densities of O. brumata. 5. In light of previous life-table and experimental studies on O. brumata and T. viridana, the current analyses suggest that pupal predation (data not presented) and interspecific competition are significant determinants of O. brumata population growth rates in Wytham Woods. In contrast, T. viridana population growth rates appear to be dominated by rapid negative feedback consistent with intraspecific competition.     

Rate of population change in voles from different phases of the population cycle

Factors involved in causing cyclic vole populations to decline, and in preventing populations from recovering during the subsequent low density phase have long remained unidentified. The traditional view of self-regulation assumes that an increase in population density is prevented by a change in the quality of individuals within the population itself, but this is still inadequately tested in the field. We compared the population growth of wild field voles ( Microtus agrestis ) from the low phase (conducted in 1998) with that of voles from the increase phase (conducted in 1999) in predator-proof enclosures (each 0.5 ha) in western Finland. Within a few months, enclosed vole populations increased to high density, and the realised per capita rate of change over the breeding season did not differ between the populations from different cycle phases. This implies that the recovery of populations from the low phase was not hindered by an impoverishment in quality of individual voles. Accordingly, we suggest that population intrinsic factors (irrespective of the mechanisms they are based on) are unlikely to play a significant role in the generation of cyclic density fluctuations of voles. Instead, we discovered direct density-dependent regulation in the vole populations. Accurate estimates of population growth and the observed density dependence provide important information for empirically based models on population dynamics of rodents.     

Logistic Proliferation of Cells in Scratch Assays is Delayed

Scratch assays are used to study how a population of cells re-colonises a vacant region on a two-dimensional substrate after a cell monolayer is scratched. These experiments are used in many applications including drug design for the treatment of cancer and chronic wounds. To provide insights into the mechanisms that drive scratch assays, solutions of continuum reaction–diffusion models have been calibrated to data from scratch assays. These models typically include a logistic source term to describe carrying capacity-limited proliferation; however, the choice of using a logistic source term is often made without examining whether it is valid. Here we study the proliferation of PC-3 prostate cancer cells in a scratch assay. All experimental results for the scratch assay are compared with equivalent results from a proliferation assay where the cell monolayer is not scratched. Visual inspection of the time evolution of the cell density away from the location of the scratch reveals a series of sigmoid curves that could be naively calibrated to the solution of the logistic growth model. However, careful analysis of the per capita growth rate as a function of density reveals several key differences between the proliferation of cells in scratch and proliferation assays. Our findings suggest that the logistic growth model is valid for the entire duration of the proliferation assay. On the other hand, guided by data, we suggest that there are two phases of proliferation in a scratch assay; at short time, we have a disturbance phase where proliferation is not logistic, and this is followed by a growth phase where proliferation appears to be logistic. These two phases are observed across a large number of experiments performed at different initial cell densities. Overall our study shows that simply calibrating the solution of a continuum model to a scratch assay might produce misleading parameter estimates, and this issue can be resolved by making a distinction between the disturbance and growth phases. Repeating our procedure for other scratch assays will provide insight into the roles of the disturbance and growth phases for different cell lines and scratch assays performed on different substrates.     

Estimating the time to extinction in an island population of song sparrows

We estimated and modelled how uncertainties in stochastic population dynamics and biases in parameter estimates affect the accuracy of the projections of a small island population of song sparrows which was enumerated every spring for 24 years. The estimate of the density regulation in a theta-logistic model (theta = 1.09 suggests that the dynamics are nearly logistic, with specific growth rate r1 = 0.99 and carrying capacity K = 41.54. The song sparrow population was strongly influenced by demographic (ŝigma2(d) = 0.66) and environmental (ŝigma2(d) = 0.41) stochasticity. Bootstrap replicates of the different parameters revealed that the uncertainties in the estimates of the specific growth rate r1 and the density regulation theta were larger than the uncertainties in the environmental variance sigma2(e) and the carrying capacity K. We introduce the concept of the population prediction interval (PPI), which is a stochastic interval which includes the unknown population size with probability (1 - alpha). The width of the PPI increased rapidly with time because of uncertainties in the estimates of density regulation as well as demographic and environmental variance in the stochastic population dynamics. Accepting a 10% probability of extinction within 100 years, neglecting uncertainties in the parameters will lead to a 33% overestimation of the time it takes for the extinction barrier (population size X = 1) to be included into the PPI. This study shows that ignoring uncertainties in population dynamics produces a substantial underestimation of the extinction risk.     

The maximum yield problem: Distortion in the yield curve due to age structure

An unselective harvest is sustainable if the per capita removal rate balances the net per capita growth rate of the population. If the function relating the growth rate to population density is known, finding the density at which the maximal total harvest rate may be achieved is a simple exercise in parameter optimization. In age-structured populations the per capita growth rate is related in a complicated way to the age-specific vital rates upon which density feedbacks directly operate. Generally, the function relating the per capita growth rate to density will resemble a log transform of the function relating age-specific fecundities to density. Accordingly, maximum yield is attained at a population density that is closer to the saturation density than we would expect on the basis simply of substituting the functional form of the density dependence of fecundity into the parameter optimization model derived for the case without age structure. The amount of the discrepancy increases with the intensity of the density feedback and with the degree to which the reproduction of a cohort is dominated by reproduction taking place during a span of ages that is small relative to the generation time.     

A Comparison of Grizzly Bear Demographic Parameters Estimated from Non-Spatial and Spatial Open Population Capture-Recapture Models

Capture-recapture studies are frequently used to monitor the status and trends of wildlife populations. Detection histories from individual animals are used to estimate probability of detection and abundance or density. The accuracy of abundance and density estimates depends on the ability to model factors affecting detection probability. Non-spatial capture-recapture models have recently evolved into spatial capture-recapture models that directly include the effect of distances between an animal’s home range centre and trap locations on detection probability. Most studies comparing non-spatial and spatial capture-recapture biases focussed on single year models and no studies have compared the accuracy of demographic parameter estimates from open population models. We applied open population non-spatial and spatial capture-recapture models to three years of grizzly bear DNA-based data from Banff National Park and simulated data sets. The two models produced similar estimates of grizzly bear apparent survival, per capita recruitment, and population growth rates but the spatial capture-recapture models had better fit. Simulations showed that spatial capture-recapture models produced more accurate parameter estimates with better credible interval coverage than non-spatial capture-recapture models. Non-spatial capture-recapture models produced negatively biased estimates of apparent survival and positively biased estimates of per capita recruitment. The spatial capture-recapture grizzly bear population growth rates and 95% highest posterior density averaged across the three years were 0.925 (0.786–1.071) for females, 0.844 (0.703–0.975) for males, and 0.882 (0.779–0.981) for females and males combined. The non-spatial capture-recapture population growth rates were 0.894 (0.758–1.024) for females, 0.825 (0.700–0.948) for males, and 0.863 (0.771–0.957) for both sexes. The combination of low densities, low reproductive rates, and predominantly negative population growth rates suggest that Banff National Park’s population of grizzly bears requires continued conservation-oriented management actions.     

Nutritional availability and larval density dependence in Aedes aegypti

1. Density dependence is the effect of density on population growth. Density dependence is an aggregate term for a suite of complex interactions between animals and their environment. 2. Mechanistic studies of density dependence in mosquito ecology are sparse, and the role of environmental factors is poorly understood. 3. Two empirical study designs were compared to consider the interaction between nutritional availability and density in Aedes aegypti. First, larvae were fed per capita. Second, larvae were fed a fixed amount of food unadjusted for the number of individuals; therefore, at higher densities, individuals received less per capita. 4. Survivorship, wing length, and development rate were lower at high densities when larvae were fed a fixed, unadjusted amount of food. The opposite was observed when food was adjusted per capita, suggesting that high densities may be beneficial for larval development when per capita nutrition is held constant 5. These results demonstrate that negative associations between Ae. aegypti larval density and larval development are a manifestation of decreased per capita nutrient uptake at high densities. 6. Population regulation is a proportional response to environmental variability in Ae. aegypti. Increased survivorship at high densities when larvae were fed per capita demonstrates that nutritional availability is not the only mechanism of density dependence in mosquitoes. Further studies should characterise density dependence in mosquitoes by using mechanistic study designs across diverse environmental conditions.     

Wealth as a source of density dependence in human population growth

In modern industrialized countries, human birth rates have been declining persistently for decades. In many cases they have now fallen below the replacement threshold. However, unlike in natural populations where population growth is constrained by limited resources, birth rates in modern industrialized countries are negatively correlated with resource availability. Here, declining birth rates in human populations are shown to be a manifestation of density‐dependent population growth brought on by socioeconomic development. This is demonstrated by combining empirical power law relations between population size, gross domestic product (GDP) per capita, and fertility in a simple theoretical model describing population dynamics in developed countries. For a closed population, the model exhibits growth to a globally stable equilibrium population size, for both national and city populations. A version of the model that is open with respect to immigration and the influence of foreign technology and capital exhibits a good fit to long‐term time series data on population size, GDP per capita, and birth rates for the United States, France and Japan.     

Competition and Population Time Lags in Bosmina (Cladocera,Crustacea)

The competition between Bosmina coregoni and B. longirostris was studied by culturing of mixed and isolated populations in lake water. While interaction was detectable between the two species, they were capable of coexisting for at least 124 days. The pattern of predominance was found to depend on initial frequencies. The ratio of B. coregoni to B. longirostris in the pelagic zone of a mesotrophic lake could be explained on the basis of competition between either species. Birth, death, and population growth rates proved to be delayed functions of stock density, as was borne out by cross-correlation analysis of experimental population dynamics. Time lag estimates for all three rates varied for different phases of population oscillation. For accurate assessment of time lags, the phases of monotonous changes in density should have been separated from one another. The time lag estimated for the rate of population growth in mixed cultures was significantly higher for the superior competitor (B. coregoni). This estimate may well be predicted as an arithmetic mean value cf time lag estimates for birth and death rates. The duration of population growth (or decline) was correlated to the time lag of the death rate, whereas the amplitude was not correlated to any time lag at all.     

Shifting trends: detecting environmentally mediated regulation in long-lived marine vertebrates using time-series data

Assessing the status and trends in animal populations is essential for effective species conservation and management practices. However, unless time-series abundance data demonstrate rapid and reliable fluctuations, objective appraisal of directionality of trends is problematic. We adopted a multiple-working hypotheses approach based on information-theoretic and Bayesian multi-model inference to examine the population trends and form of intrinsic regulation demonstrated by a long-lived species, the southern elephant seal. We also determined the evidence for density dependence in 11 other well-studied marine mammal species. (1) We tested the type of population regulation for elephant seals from Marion Island (1986–2004) and from 11 other marine mammal species, and (2) we described the trends and behavior of the 19-year population time series at Marion Island to identify changes in population trends. We contrasted five plausible trend models using information-theoretic and Bayesian-inference estimates of model parsimony. Our analyses identified two distinct phases of population growth for this population with the inflexion occurring in 1998. Thus, the population decreased between 1986 and 1997 (−3.7% per annum) and increased between 1997 and 2004 (1.9% per annum). An index of environmental stochasticity, the Southern Oscillation Index, explained some of the variance in r and N. We determined analytically that there was good evidence for density dependence in the Marion Island population and that density dependence was widespread among marine mammal species (67% of species showed evidence for population regulation). This approach demonstrates the potential functionality of a relatively simple technique that can be applied to short time series to identify the type of regulation, and the uncertainty associated with the phenomenon, operating in populations of large mammals.     

Density-dependent growth as a key mechanism in the regulation of fish populations: evidence from among-population comparisons.

It is generally assumed that fish populations are regulated primarily in the juvenile (pre-recruit) phase of the life cycle, although density dependence in growth and reproductive parameters within the recruited phase has been widely reported. Here we present evidence to suggest that density-dependent growth in the recruited phase is a key process in the regulation of many fish populations. We analyse 16 fish populations with long-term records of size-at-age and biomass data, and detect significant density-dependent growth in nine. Among-population comparisons show a close, inverse relationship between the estimated decline in asymptotic length per unit biomass density, and the long-term average biomass density of populations. A simple population model demonstrates that regulation by density-dependent growth alone is sufficient to generate the observed relationship. Density-dependent growth should be accounted for in fisheries' assessments, and the empirical relationship established here can provide indicative estimates of the density-dependent growth parameter where population-specific data are lacking.     

How Human Household Size Affects the Habitat of Black-and-White Snub-Nosed Monkeys (<Emphasis Type="Italic">Rhinopithecus bieti</Emphasis>) in Hongla Snow Mountain Nature Reserve in Tibet,China

Human impacts on the environment at local or regional scales largely depend on intrinsic characteristics of the population, such as household size, household number, and human population growth. These demographic factors can vary considerably among ethnic groups sharing similar ecological landscapes, yet the role of traditional cultural practices in shaping local environmental impacts is not well known for many parts of the world. We here quantify land-cover changes and their relation to the habitat of the endangered Rhinopithecus bieti in Tibet, in 2 areas populated by different ethnic groups (polyandrous Tibetans and monogamous Naxi) from 1986 to 2006. Results indicate that habitat of the monkey decreased greatly within our study area over the 20-yr period. Polyandrous and monogamous ethnic communities differed in household size, household number, population growth, and per capita and per household land use. The practice of polyandry by ethnic Tibetan appears to have reduced per capita resource consumption by reducing the growth of overall household number and increasing household size, which can mitigate the negative effects of higher human density and population growth on the environment. Ethnic Tibetan may also reduce land impacts by adhering to Buddhist customs and alternative, more sustainable means of livelihood. Accordingly, the protection of traditional cultural resources, such as polyandry and Buddhist beliefs, could be an effective way to aid biodiversity and environmental conservation efforts in this key ecosystem.     

Food web structure and climate effects on the dynamics of small mammals and owls in semi-arid Chile

Population dynamics of small mammals and predators in semi-arid Chile is positively correlated with rainfall associated with incursions of El Niño (El Niño Southern Oscillation: ENSO). However, the causal relationships between small mammal fluctuations, predator oscillations, and climatic disturbances are poorly understood. Here, we report time series models for three species of small mammal prey and two species of owl predators. The large differences in population fluctuations between the three small mammal species are related to differences in their respective feedback structures. The analyses reveal that per capita growth rate of the leaf-eared mouse is a decreasing function of log density and of log barn owl abundance together with a positive rainfall effect. In turn, per capita population growth rate ( R -function) of the barn owl is a negative function of log barn owl abundance and a positive function of leaf-eared mouse abundance, suggesting a predator–prey interaction. The dramatic population fluctuations exhibited by leaf-eared mouse ( Phyllotis darwini ) are caused by climate effects coupled with a complex food web architecture.     

Pattern of variation in avian population growth rates

A central question in population ecology is to understand why population growth rates differ over time. Here, we describe how the long-term growth of populations is not only influenced by parameters affecting the expected dynamics, for example form of density dependence and specific population growth rate, but is also affected by environmental and demographic stochasticity. Using long-term studies of fluctuations of bird populations, we show an interaction between the stochastic and the deterministic components of the population dynamics: high specific growth rates at small densities r(1) are typically positively correlated with the environmental variance sigma(e)(2). Furthermore, theta, a single parameter describing the form of the density regulation in the theta-logistic density-regulation model, is negatively correlated with r(1). These patterns are in turn correlated with interspecific differences in life-history characteristics. Higher specific growth rates, larger stochastic effects on the population dynamics and stronger density regulation at small densities are found in species with large clutch sizes or high adult mortality rates than in long-lived species. Unfortunately, large uncertainties in parameter estimates, as well as strong stochastic effects on the population dynamics, will often make even short-term population projections unreliable. We illustrate that the concept of population prediction interval can be useful in evaluating the consequences of these uncertainties in the population projections for the choice of management actions.     

Allee effects and population regulation: a test for biotic resistance against an invasive leafroller by resident parasitoids

Resident natural enemies can impact invasive species by causing Allee effects, leading to a reduction in establishment success of small founder populations, or by regulating or merely suppressing the abundance of established populations. Epiphyas postvittana, the Light Brown Apple Moth, an invasive leafroller in California, has been found to be attacked by a large assemblage of resident parasitoids that cause relatively high rates of parasitism. Over a 4-year period, we measured the abundance and per capita growth rates of four E. postvittana populations in California and determined parasitism rates. We found that at two of the sites, parasitism caused a component Allee effect, a reduction in individual survivorship at lower E. postvittana population densities, although it did not translate into a demographic Allee effect, an impact on per capita population growth rates at low densities. Instead, E. postvittana populations at all four sites exhibited strong compensatory density feedback throughout the entire range of densities observed at each site. As we found no evidence for a negative relationship between per capita population growth rates and parasitism rates, we concluded that resident parasitoids were unable to regulate E. postvittana populations in California. Despite a lack of evidence for regulation or a demographic Allee effect, the impact of resident parasitoids on E. postvittana populations is substantial and demonstrates significant biotic resistance against this new invader.     

Density dependent population growth of the two-spotted spider mite, Tetranychus urticae, on the host plant Leonurus cardiaca

We observed Tetranychus urticae (Koch), a polyphagous spider mite herbivore, on Leonurus cardiaca (L.) at several sites in eastern North America at variable density, ranging from extremely dense to sparse. To understand the nature of T. urticae 's population dynamics we experimentally manipulated population densities on L. cardiaca and assessed per capita growth after 1 to 2 generations in laboratory and field experiments. In particular, we took a 'bottom-up' approach, manipulating both plant size and quality to examine effects on mite dynamics. Per capita growth was strongly dependent on the initial density of the mite population. Spider mite populations grew (1) in a negatively density dependent manner on small plants and (2) unhindered by density dependence on large plants. Mean per capita growth was 59% higher on small plants compared to large plants, irrespective of mite density. We also found evidence for density dependent induced susceptibility to spider mites in small plants and density dependent induced resistance in large plants. Hence, spider mite populations grew at a relatively fast rate on small plants, and this was associated with negative density dependence due to factors that depress population growth, such as food deterioration or limitation. On large plants, spider mite populations grew at a relatively slow rate, apparently resulting in herbivore densities that may not have been high enough to cause intraspecific competition or other forms of negative density dependence.     

Subnational greenhouse gas and land‐based biodiversity footprints in the European Union

Insights into subnational environmental impacts and the underlying drivers are scarce, especially from a consumption‐based perspective. Here, we quantified greenhouse gas (GHG) emissions and land‐based biodiversity losses associated with final consumption in 162 regions in the European Union in 2010. For this purpose, we developed an environmentally extended multi‐regional input–output (MRIO) model with subnational European information on demand, production, and trade structures subdivided into 18 major economic sectors, while accounting for trade outside Europe. We employed subnational data on land use and national data on GHG emissions. Our results revealed within‐country differences in per capita GHG and land‐based biodiversity footprints up to factors of 3.0 and 3.5, respectively, indicating that national footprints may mask considerable subnational variability. The per capita GHG footprint increased with per capita income and income equality, whereas we did not find such responses for the per capita land‐based biodiversity footprint, reflecting that extra income is primarily spent on energy‐intensive activities. Yet, we found a shift from the domestic to the foreign part of the biodiversity footprints with rising population density and income. Because our analysis showed that most regions are already net importers of GHG emissions and biodiversity losses, we conclude that it is increasingly important to address the role of trade in national and regional policies on mitigating GHG emissions and averting further biodiversity losses, both within and outside the region itself. To further increase the policy relevance of subnational footprint analyses, we also recommend the compilation of more detailed subnational MRIO databases including harmonized environmental data.     

Non‐native plants have greater impacts because of differing per‐capita effects and nonlinear abundance–impact curves

Invasive, non‐native species can have tremendous impacts on biotic communities, where they reduce the abundance and diversity of local species. However, it remains unclear whether impacts of non‐native species arise from their high abundance or whether each non‐native individual has a disproportionate impact – that is, a higher per‐capita effect – on co‐occurring species compared to impacts by native species. Using a long‐term study of wetlands, we asked how temporal variation in dominant native and non‐native plants impacted the abundance and richness of other plants in the recipient community. Non‐native plants reached higher abundances than natives and had greater per‐capita effects. The abundance–impact relationship between plant abundance and richness was nonlinear. Compared with increasing native abundance, increasing non‐native abundance was associated with steeper declines in richness because of greater per‐capita effects and nonlinearities in the abundance–impact relationship. Our study supports eco‐evolutionary novelty of non‐natives as a driver of their outsized impacts on communities.     

Experimental evidence that livestock grazing intensity affects cyclic vole population regulation processes

Grazing by domestic ungulates may limit the densities of small herbivorous mammals that act as key prey in ecosystems. Whether this also influences density dependence and the regulation of small herbivore populations, hence their propensity to exhibit multi-annual population cycles, is unknown. Here, we combine time series analysis with a large-scale grazing experiment on upland grasslands to examine the effects of livestock grazing intensity on the population dynamics of field voles (Microtus agrestis). Using log-linear modelling of replicated time series under different grazing treatments, we show that increased sheep densities weaken delayed density dependent regulation of vole population growth, hence reducing the cyclicity in vole population dynamics. While population regulation is commonly attributed to both top-down and bottom up processes, our results suggest that regulation of cyclic vole populations can be disrupted by the influence of another grazer in the same trophic level. These results support the view that ongoing changes in domestic grazing intensity, by affecting small mammal dynamics, can potentially have cascading impacts on higher trophic levels, and strongly influence the dynamics of upland grassland systems.