Increased toxicity of Karenia brevis during phosphate limited growth: ecological and evolutionary implications

Karenia brevis is the dominant toxic red tide algal species in the Gulf of Mexico. It produces potent neurotoxins (brevetoxins [PbTxs]), which negatively impact human and animal health, local economies, and ecosystem function. Field measurements have shown that cellular brevetoxin contents vary from 1–68 pg/cell but the source of this variability is uncertain. Increases in cellular toxicity caused by nutrient-limitation and inter-strain differences have been observed in many algal species. This study examined the effect of P-limitation of growth rate on cellular toxin concentrations in five Karenia brevis strains from different geographic locations. Phosphorous was selected because of evidence for regional P-limitation of algal growth in the Gulf of Mexico. Depending on the isolate, P-limited cells had 2.3- to 7.3-fold higher PbTx per cell than P-replete cells. The percent of cellular carbon associated with brevetoxins (%C-PbTx) was ~ 0.7 to 2.1% in P-replete cells, but increased to 1.6–5% under P-limitation. Because PbTxs are potent anti-grazing compounds, this increased investment in PbTxs should enhance cellular survival during periods of nutrient-limited growth. The %C-PbTx was inversely related to the specific growth rate in both the nutrient-replete and P-limited cultures of all strains. This inverse relationship is consistent with an evolutionary tradeoff between carbon investment in PbTxs and other grazing defenses, and C investment in growth and reproduction. In aquatic environments where nutrient supply and grazing pressure often vary on different temporal and spatial scales, this tradeoff would be selectively advantageous as it would result in increased net population growth rates. The variation in PbTx/cell values observed in this study can account for the range of values observed in the field, including the highest values, which are not observed under N-limitation. These results suggest P-limitation is an important factor regulating cellular toxicity and adverse impacts during at least some K. brevis blooms.

Growth and reproductive performance of locally isolated brackish water rotifer (Brachionus plicatilis) feeding different micro algae

Population growth and reproductive capacity of brackishwater rotifer, Brachionus plicatilis, were evaluated, for a period of 8 days in a temperature controlled ( =25°C) microalgallaborarory, under three different algal feeding regimens. The algal species that were tested are: (i) Chlorella sp. (T1), Tetraselmis chui (T2), Nannochloropsis oculata (T 3). The feeding density of each algal species was maintained similar as of 4.5xW6 ceHs mi. The rotifer fed on T. chui showed the highest (p<0.05) population growth (131.5 ind./ml), compared to that fed on Chlorella sp (45.67 ind./ml) and N oculata (43.44 ind./ml). The abundance of egg bearing rotifers was also higher (35.77%) with T. chuithan with Chlorella sp (27.76%) and N oculara (24.60%). The results of the present study indicate that T. chui could be the most suitable algal food for the stock culture of locally isolated rotifer B. plicatilis.

The study of biological characteristic and population dynamic of Caspian vimba (Vimba vimba persa), in coastal waters of Caspian Sea (Mazandaran Province)

This study was conducted to determine reproduction characteristics, diet regime, age structure and population dynamics parameters of the vimba vimba persa (Pallas, 1811) in Mazandaran waters of the Caspian Sea, from October 2008 to September 2009. A total of 994 specimens were monthly collected by beach seine and cast net from six fish landings of Ramsar, Tonekabon, Chaloos, Mahmood Abad, Sari and Behshahr. Biometric characters were measured for each specimen at the laboratory. Scales were used for age determination. Sex determination and fecundity were determined. Population dynamic parameters as well as stock assessment including cohort analysis were estimated using FISAT software. The finding showed that the mean of fork length and body weight of the Caspian Vimba were 168.4±2.6 mm and 71.94±32.24 g respectively. Strong correlation was found between these two variables (a= 0.012; b = 3.047; r2 = 0.955). 92 specimens were studied from the fecundity point of view. This species was found to have more abundance in spring (esp. Apr-May). The samples composed of 397(42.6%) male, 537(57.4%) female; Overall sex ratio (M: F =1: 1.35) was significantly different from the expected 1:1 ratio (p ≤0.05). The advanced stages of maturity (4th & 5th) were found in April and May. The highest Gonadosomatic Index in female was in May and the lowest one was in July. This fish is therefore a spring spawner. The maximum absolute and relative fecundities were 34640 and 260.9, respectively; the minimum absolute and relative fecundities were 5400 and 94.5 respectively. The averages of absolute and relative fecundities were 17198±7710 and 171.85±48.8, respectively. Coefficient vacuity index was 59.2% which indicates that this fish is mesophagous. Among of living creature consumes by Caspian Vimba mollusks, 76 arthropods, worms, plants, detritus and fishes were found 32.9% , 26.7% , 13.4% , 17% , 4.4% and 1.6% respectively. The infinite fork lengths were 261 mm for females, 25mm for males and 261 mm for both sexes respectively. For population growth and mortality parameters; K ( 0.28 per year for both sexes, 0.3 per year for males, 0.33 per year for females); t0 ( -0.65 year for both sexes, -0.23 year in females, -0.51 year in males ); Φ' ( 2.28 ); Z ( 0.98 per year ); M ( 0.59 per year); F ( 0.39 per year) and Exploitation coefficient was 0.4. The analysis showed that total biomass and MSY were 1336 and 528.8 tonnes respectively.

Impact of simian immunodeficiency virus infection on chimpanzee population dynamics.

Like human immunodeficiency virus type 1 (HIV-1), simian immunodeficiency virus of chimpanzees (SIVcpz) can cause CD4+ T cell loss and premature death. Here, we used molecular surveillance tools and mathematical modeling to estimate the impact of SIVcpz infection on chimpanzee population dynamics. Habituated (Mitumba and Kasekela) and non-habituated (Kalande) chimpanzees were studied in Gombe National Park, Tanzania. Ape population sizes were determined from demographic records (Mitumba and Kasekela) or individual sightings and genotyping (Kalande), while SIVcpz prevalence rates were monitored using non-invasive methods. Between 2002-2009, the Mitumba and Kasekela communities experienced mean annual growth rates of 1.9% and 2.4%, respectively, while Kalande chimpanzees suffered a significant decline, with a mean growth rate of -6.5% to -7.4%, depending on population estimates. A rapid decline in Kalande was first noted in the 1990s and originally attributed to poaching and reduced food sources. However, between 2002-2009, we found a mean SIVcpz prevalence in Kalande of 46.1%, which was almost four times higher than the prevalence in Mitumba (12.7%) and Kasekela (12.1%). To explore whether SIVcpz contributed to the Kalande decline, we used empirically determined SIVcpz transmission probabilities as well as chimpanzee mortality, mating and migration data to model the effect of viral pathogenicity on chimpanzee population growth. Deterministic calculations indicated that a prevalence of greater than 3.4% would result in negative growth and eventual population extinction, even using conservative mortality estimates. However, stochastic models revealed that in representative populations, SIVcpz, and not its host species, frequently went extinct. High SIVcpz transmission probability and excess mortality reduced population persistence, while intercommunity migration often rescued infected communities, even when immigrating females had a chance of being SIVcpz infected. Together, these results suggest that the decline of the Kalande community was caused, at least in part, by high levels of SIVcpz infection. However, population extinction is not an inevitable consequence of SIVcpz infection, but depends on additional variables, such as migration, that promote survival. These findings are consistent with the uneven distribution of SIVcpz throughout central Africa and explain how chimpanzees in Gombe and elsewhere can be at equipoise with this pathogen.

Future requirements for and supply of ophthalmologists for an aging population in Singapore.

BACKGROUND: Singapore's population, as that of many other countries, is aging; this is likely to lead to an increase in eye diseases and the demand for eye care. Since ophthalmologist training is long and expensive, early planning is essential. This paper forecasts workforce and training requirements for Singapore up to the year 2040 under several plausible future scenarios. METHODS: The Singapore Eye Care Workforce Model was created as a continuous time compartment model with explicit workforce stocks using system dynamics. The model has three modules: prevalence of eye disease, demand, and workforce requirements. The model is used to simulate the prevalence of eye diseases, patient visits, and workforce requirements for the public sector under different scenarios in order to determine training requirements. RESULTS: Four scenarios were constructed. Under the baseline business-as-usual scenario, the required number of ophthalmologists is projected to increase by 117% from 2015 to 2040. Under the current policy scenario (assuming an increase of service uptake due to increased awareness, availability, and accessibility of eye care services), the increase will be 175%, while under the new model of care scenario (considering the additional effect of providing some services by non-ophthalmologists) the increase will only be 150%. The moderated workload scenario (assuming in addition a reduction of the clinical workload) projects an increase in the required number of ophthalmologists of 192% by 2040. Considering the uncertainties in the projected demand for eye care services, under the business-as-usual scenario, a residency intake of 8-22 residents per year is required, 17-21 under the current policy scenario, 14-18 under the new model of care scenario, and, under the moderated workload scenario, an intake of 18-23 residents per year is required. CONCLUSIONS: The results show that under all scenarios considered, Singapore's aging and growing population will result in an almost doubling of the number of Singaporeans with eye conditions, a significant increase in public sector eye care demand and, consequently, a greater requirement for ophthalmologists.

Worldwide trends in diabetes since 1980: a pooled analysis of 751 population-based studies with 4.4 million participants.

One of the global targets for non-communicable diseases is to halt, by 2025, the rise in the age-standardised adult prevalence of diabetes at its 2010 levels. We aimed to estimate worldwide trends in diabetes, how likely it is for countries to achieve the global target, and how changes in prevalence, together with population growth and ageing, are affecting the number of adults with diabetes. We pooled data from population-based studies that had collected data on diabetes through measurement of its biomarkers. We used a Bayesian hierarchical model to estimate trends in diabetes prevalence-defined as fasting plasma glucose of 7.0 mmol/L or higher, or history of diagnosis with diabetes, or use of insulin or oral hypoglycaemic drugs-in 200 countries and territories in 21 regions, by sex and from 1980 to 2014. We also calculated the posterior probability of meeting the global diabetes target if post-2000 trends continue. We used data from 751 studies including 4,372,000 adults from 146 of the 200 countries we make estimates for. Global age-standardised diabetes prevalence increased from 4.3% (95% credible interval 2.4-7.0) in 1980 to 9.0% (7.2-11.1) in 2014 in men, and from 5.0% (2.9-7.9) to 7.9% (6.4-9.7) in women. The number of adults with diabetes in the world increased from 108 million in 1980 to 422 million in 2014 (28.5% due to the rise in prevalence, 39.7% due to population growth and ageing, and 31.8% due to interaction of these two factors). Age-standardised adult diabetes prevalence in 2014 was lowest in northwestern Europe, and highest in Polynesia and Micronesia, at nearly 25%, followed by Melanesia and the Middle East and north Africa. Between 1980 and 2014 there was little change in age-standardised diabetes prevalence in adult women in continental western Europe, although crude prevalence rose because of ageing of the population. By contrast, age-standardised adult prevalence rose by 15 percentage points in men and women in Polynesia and Micronesia. In 2014, American Samoa had the highest national prevalence of diabetes (>30% in both sexes), with age-standardised adult prevalence also higher than 25% in some other islands in Polynesia and Micronesia. If post-2000 trends continue, the probability of meeting the global target of halting the rise in the prevalence of diabetes by 2025 at the 2010 level worldwide is lower than 1% for men and is 1% for women. Only nine countries for men and 29 countries for women, mostly in western Europe, have a 50% or higher probability of meeting the global target. Since 1980, age-standardised diabetes prevalence in adults has increased, or at best remained unchanged, in every country. Together with population growth and ageing, this rise has led to a near quadrupling of the number of adults with diabetes worldwide. The burden of diabetes, both in terms of prevalence and number of adults affected, has increased faster in low-income and middle-income countries than in high-income countries. Wellcome Trust.

Labor supply, biased technological change and economic growth

Se utiliza un modelo de innovaciones sesgadas para estudiar los efectos de cambios exógenos en la oferta laboral. En un contexto de innovaciones sesgadas, a medida que las economías acumulan capital, el trabajo se hace relativamente más escaso y más caro, por este motivo, hay incentivos para adoptar tecnologías ahorradoras de trabajo. Del mismo modo un cambio en la oferta laboral afecta la abundancia de factores y sus precios relativos. En general, una reducción de la oferta laboral, hace que el trabajo sea más caro y genera incentivos para cambio tecnológico ahorrador de trabajo. Así, el efecto inicial que tiene el cambio en la oferta laboral sobre los precios de los factores es mitigado por el cambio tecnológico. Finalmente, los movimientos en la remuneración a los factores afectan las decisiones de ahorro y, por lo tanto, la dinámica del crecimiento. En este trabajo se exploran las consecuencias de una reducción de la oferta laboral en dos contextos teóricos diferentes: un modelo de agentes homogéneos y horizonte infinito y un modelo de generaciones traslapadas.

Population Assessment of an Endangered Shorebird: the Piping Plover (Charadrius melodus melodus) in Eastern Canada

Small, at-risk populations are those for which accurate demographic information is most crucial to conservation and recovery, but also where data collection is constrained by logistical challenges and small sample sizes. Migratory animals in particular may experience a wide range of threats to survival and reproduction throughout each annual cycle, and identification of life stages most critical to persistence may be especially difficult for these populations. The endangered eastern Canadian breeding population of Piping Plover (Charadrius melodus melodus) was estimated at only 444 adults in 2005, and extensive effort has been invested in conservation activities, reproductive monitoring, and marking of individual birds, providing a comprehensive data set on population dynamics since 1998. We used these data to build a matrix projection model for two Piping Plover population segments that nest in eastern Canada in order to estimate both deterministic and stochastic rates of population growth (λd and λs, respectively). Annual population censuses suggested moderate growth in abundance between 1998–2003, but vital rate estimates indicated that this temporary growth may be replaced by declines in the long term, both in southern Nova Scotia (λd = 1.0043, λs = 0.9263) and in the Gulf of St. Lawrence (λd = 0.9651, λs = 0.8214). Nonetheless, confidence intervals on λ estimates were relatively wide, highlighting remaining uncertainty in future population trajectories. Differences in projected growth between regions appear to be driven by low estimated juvenile post-fledging survival in the Gulf, but threats to juveniles of both population segments following departure from nesting beaches remain unidentified. Similarly, λ in both population segments was particularly sensitive to changes in adult survival as expected for most migratory birds, but very little is understood about the threats to Piping Plover survival during migration and overwintering. Consequently, we suggest that future recovery efforts for these and other vulnerable migrants should quantify and manage the largely unknown sources of both adult and juvenile mortality during non-breeding seasons while maintaining current levels of nesting habitat protection.

Lesser Scaup Population Dynamics: What Can Be Learned from Available Data?

Populations of Lesser Scaup (Aythya affinis) have declined markedly in North America since the early 1980s. When considering alternatives for achieving population recovery, it would be useful to understand how the rate of population growth is functionally related to the underlying vital rates and which vital rates affect population growth rate the most if changed (which need not be those that influenced historical population declines). To establish a more quantitative basis for learning about life history and population dynamics of Lesser Scaup, we summarized published and unpublished estimates of vital rates recorded between 1934 and 2005, and developed matrix life-cycle models with these data for females breeding in the boreal forest, prairie-parklands, and both regions combined. We then used perturbation analysis to evaluate the effect of changes in a variety of vital-rate statistics on finite population growth rate and abundance. Similar to Greater Scaup (Aythya marila), our modeled population growth rate for Lesser Scaup was most sensitive to unit and proportional change in adult female survival during the breeding and non-breeding seasons, but much less so to changes in fecundity parameters. Interestingly, population growth rate was also highly sensitive to unit and proportional changes in the mean of nesting success, duckling survival, and juvenile survival. Given the small samples of data for key aspects of the Lesser Scaup life cycle, we recommend additional research on vital rates that demonstrate a strong effect on population growth and size (e.g., adult survival probabilities). Our life-cycle models should be tested and regularly updated in the future to simultaneously guide science and management of Lesser Scaup populations in an adaptive context.

Nonbreeding-Season Drivers of Population Dynamics in Seasonal Migrants: Conservation Parallels Across Taxa

For seasonal migrants, logistical constraints have often limited conservation efforts to improving survival and reproduction during the breeding season only. Yet, mounting empirical evidence suggests that events occurring throughout the migratory life cycle can critically alter the demography of many migrant species. Herein, we build upon recent syntheses of avian migration research to review the role of non-breeding seasons in determining the population dynamics and fitness of diverse migratory taxa, including salmonid fishes, marine mammals, ungulates, sea turtles, butterflies, and numerous bird groups. We discuss several similarities across these varied migrants: (i) non-breeding survivorship tends to be a strong driver of population growth; (ii) non-breeding events can affect fitness in subsequent seasons through seasonal interactions at individual- and population-levels; (iii) broad-scale climatic influences often alter non-breeding resources and migration timing, and may amplify population impacts through covariation among seasonal vital rates; and (iv) changes to both stationary and migratory non-breeding habitats can have important consequences for abundance and population trends. Finally, we draw on these patterns to recommend that future conservation research for seasonal migrants will benefit from: (1) more explicit recognition of the important parallels among taxonomically diverse migratory animals; (2) an expanded research perspective focused on quantification of all seasonal vital rates and their interactions; and (3) the development of detailed population projection models that account for complexity and uncertainty in migrant population dynamics.

A Demographic Model to Evaluate Population Declines in the Endangered Streaked Horned Lark

The Streaked Horned Lark (Eremophila alpestris strigata) is listed as endangered by the State of Washington, USA and by Canada under the Species at Risk Act and is also classified as a federal candidate for listing under the Endangered Species Act in the USA. A substantial portion of Streaked Horned Lark habitat has been lost or degraded, and range contraction has occurred in Oregon, Washington, and British Columbia. We estimate the vital rates (fecundity, adult and juvenile survival) and population growth rate (λ) for Streaked Horned Larks breeding in Washington, USA and conduct a Life-Stage Simulation Analysis (LSA) to evaluate which vital rate has the greatest influence on λ. We simulated changes in the three vital rates to examine how much they would need to be adjusted either independently or in concert to achieve a stable Streaked Horned Lark population (λ = 1). We also evaluated which fecundity component (the number of fledglings per egg laid or renesting interval) had the greatest impact on λ. The estimate of population growth suggests that Streaked Horned Larks in Washington are declining rapidly (λ = 0.62 ± 0.10) and that local breeding sites are not sustainable without immigration. The LSA results indicate that adult survival had the greatest influence on λ, followed by juvenile survival and fecundity. However, increases in vital rates led to λ = 1 only when adult survival was raised from 0.47 to 0.85, juvenile survival from 0.17 to 0.58, and fecundity from 0.91 to 3.09. Increases in breeding success and decreases in the renesting interval influenced λ similarly; however, λ did not reach 1 even when breeding success was raised to 100% or renesting intervals were reduced to 1 day. Only when all three vital rates were increased simultaneously did λ approach 1 without requiring highly unrealistic increases in each vital rate. We conclude that conservation activities need to target all or multiple vital rates to be successful. The baseline data presented here and subsequent efforts to manage Streaked Horned Larks will provide valuable information for management of other declining Horned Lark subspecies and other grassland songbirds across North America.

Population Change in a Marine Bird Colony is Driven By Changes in Recruitment

The population dynamics of long-lived birds are thought to be very sensitive to changes in adult survival. However, where natal philopatry is low, recruitment from the larger metapopulation may have the strongest effect on population growth rate even in long-lived species. Here, we illustrate such a situation where changes in a seabird colony size appeared to be the consequence of changes in recruitment. We studied the population dynamics of a declining colony of Ancient Murrelets (Synthliboramphus antiquus) at East Limestone Island, British Columbia. During 1990-2010, Ancient Murrelet chicks were trapped at East Limestone Island while departing to sea, using a standard trapping method carried on throughout the departure period. Adult murrelets were trapped while departing from the colony during 1990-2003. Numbers of chicks trapped declined during 1990-1995, probably because of raccoon predation, increased slightly from 1995-2000 and subsequently declined again. Reproductive success was 30% lower during 2000-2003 than in earlier years, mainly because of an increase in desertions. The proportion of nonbreeders among adult birds trapped at night also declined over the study period. Mortality of adult birds, thought to be mainly prebreeders, from predators more than doubled over the same period. Apparent adult survival of breeders remained constant during 1991-2002 once the first year after banding was excluded, but the apparent survival rates in the first year after banding fell and the survival of birds banded as chicks to age three halved over the same period. A matrix model of population dynamics suggested that even during the early part of the study immigration from other breeding areas must have been substantial, supporting earlier observations that natal philopatry in this species is low. The general colony decline after 2000 probably was related to diminished recruitment, as evidenced by the lower proportion of nonbreeders in the trapped sample. Hence the trend is determined by the recruitment decisions of externally reared birds, rather than demographic factors operating on the local breeding population, an unusual situation for a colonial marine bird. Because of the contraction in the colony it may now be subject to a level of predation pressure from which recovery will be impossible without some form of intervention.

Stability of a Seabird Population in the Presence of an Introduced Predator

We hypothesized that although large populations may appear able to withstand predation and disturbance, added stochasticity in population growth rate (λ) increases the risk of dramatic population declines. Approximately half of the Aleutian Islands' population of Least Auklets (Aethia pusilla) breed at one large colony at Kiska Island in the presence of introduced Norway rats (Rattus norvegicus) whose population erupts periodically. We evaluated two management plans, do nothing or eradicate rats, for this colony, and performed stochastic elasticity analysis to focus future research and management. Our results indicated that Least Auklets breeding at Kiska Island had the lowest absolute value of growth rate and more variable λ's (neither statistically significant) during 2001-2010, when compared with rat-free colonies at Buldir and Kasatochi islands. We found variability in the annual proportional change in population size among islands with Kiska Island having the fastest rate of decline, 78% over 20 years. Under the assumption that the eradication of rats would result in vital rates similar to those observed at rat-free Buldir and Kasatochi islands, we found the projected population decline decreased from 78% to 24% over 20 years. Overall, eradicating rats at Kiska Island is not likely to increase Least Auklet vital rates, but will decrease the amount of variation in λ, resulting in a significantly slower rate of population decline. We recommend the eradication of rats from Kiska Island to decrease the probability of dramatic population declines and ensure the future persistence of this important colony.

Managing threatened species – the ecological toolbox, evolutionary theory and the declining-population paradigm

1. The management of threatened species is an important practical way in which conservationists can intervene in the extinction process and reduce the loss of biodiversity. Understanding the causes of population declines (past, present and future) is pivotal to designing effective practical management. This is the declining-population paradigm identified by Caughley. 2. There are three broad classes of ecological tool used by conservationists to guide management decisions for threatened species: statistical models of habitat use, demographic models and behaviour-based models. Each of these is described here, illustrated with a case study and evaluated critically in terms of its practical application. 3. These tools are fundamentally different. Statistical models of habitat use and demographic models both use descriptions of patterns in abundance and demography, in relation to a range of factors, to inform management decisions. In contrast, behaviourbased models describe the evolutionary processes underlying these patterns, and derive such patterns from the strategies employed by individuals when competing for resources under a specific set of environmental conditions. 4. Statistical models of habitat use and demographic models have been used successfully to make management recommendations for declining populations. To do this, assumptions are made about population growth or vital rates that will apply when environmental conditions are restored, based on either past data collected under favourable environmental conditions or estimates of these parameters when the agent of decline is removed. As a result, they can only be used to make reliable quantitative predictions about future environments when a comparable environment has been experienced by the population of interest in the past. 5. Many future changes in the environment driven by management will not have been experienced by a population in the past. Under these circumstances, vital rates and their relationship with population density will change in the future in a way that is not predictable from past patterns. Reliable quantitative predictions about population-level responses then need to be based on an explicit consideration of the evolutionary processes operating at the individual level. 6. Synthesis and applications. It is argued that evolutionary theory underpins Caughley’s declining-population paradigm, and that it needs to become much more widely used within mainstream conservation biology. This will help conservationists examine critically the reliability of the tools they have traditionally used to aid management decision-making. It will also give them access to alternative tools, particularly when predictions are required for changes in the environment that have not been experienced by a population in the past.

The effects of spatial and temporal heterogeneity on the population dynamics of four animal species in a Danish landscape

Background: Variation in carrying capacity and population return rates is generally ignored in traditional studies of population dynamics. Variation is hard to study in the field because of difficulties controlling the environment in order to obtain statistical replicates, and because of the scale and expense of experimenting on populations. There may also be ethical issues. To circumvent these problems we used detailed simulations of the simultaneous behaviours of interacting animals in an accurate facsimile of a real Danish landscape. The models incorporate as much as possible of the behaviour and ecology of skylarks Alauda arvensis, voles Microtus agrestis, a ground beetle Bembidion lampros and a linyphiid spider Erigone atra. This allows us to quantify and evaluate the importance of spatial and temporal heterogeneity on the population dynamics of the four species. Results: Both spatial and temporal heterogeneity affected the relationship between population growth rate and population density in all four species. Spatial heterogeneity accounted for 23–30% of the variance in population growth rate after accounting for the effects of density, reflecting big differences in local carrying capacity associated with the landscape features important to individual species. Temporal heterogeneity accounted for 3–13% of the variance in vole, skylark and spider, but 43% in beetles. The associated temporal variation in carrying capacity would be problematic in traditional analyses of density dependence. Return rates were less than one in all species and essentially invariant in skylarks, spiders and beetles. Return rates varied over the landscape in voles, being slower where there were larger fluctuations in local population sizes. Conclusion: Our analyses estimated the traditional parameters of carrying capacities and return rates, but these are now seen as varying continuously over the landscape depending on habitat quality and the mechanisms of density dependence. The importance of our results lies in our demonstration that the effects of spatial and temporal heterogeneity must be accounted for if we are to have accurate predictive models for use in management and conservation. This is an area which until now has lacked an adequate theoretical framework and methodology.