Predicting fluctuations of reintroduced ibex populations: the importance of density dependence, environmental stochasticity and uncertain population estimates

1. Development of population projections requires estimates of observation error, parameters characterizing expected dynamics such as the specific population growth rate and the form of density regulation, the influence of stochastic factors on population dynamics, and quantification of the uncertainty in the parameter estimates. 2. Here we construct a Population Prediction Interval (PPI) based on Bayesian state space modelling of future population growth of 28 reintroduced ibex populations in Switzerland that have been censused for up to 68 years. Our aim is to examine whether the interpopulation variation in the precision of the population projections is related to differences in the parameters characterizing the expected dynamics, in the effects of environmental stochasticity, in the magnitude of uncertainty in the population parameters, or in the observation error. 3. The error in the population censuses was small. The median coefficient of variation in the estimates across populations was 5.1%. 4. Significant density regulation was present in 53.6% of the populations, but was in general weak. 5. The width of the PPI calculated for a period of 5 years showed large variation among populations, and was explained by differences in the impact of environmental stochasticity on population dynamics. 6. In spite of the high accuracy in population estimates, the uncertainty in the parameter estimates was still large. This uncertainty affected the precision in the population predictions, but it decreased with increasing length of study period, mainly due to higher precision in the estimates of the environmental variance in the longer time-series. 7. These analyses reveal that predictions of future population fluctuations of weakly density-regulated populations such as the ibex often become uncertain. Credible population predictions require that this uncertainty is properly quantified.     

Climate, food, density and wildlife population growth rate

1. The aim of this study was to derive and evaluate a priori models of the relationship between annual instantaneous population growth rate (r) and climate. These were derived from the numerical response of annual r and food, and the effect of climate on a parameter in the numerical response. The goodness of fit of a range of such deductive models to data on annual r of Soay sheep and red deer were evaluated using information-theoretic (AICc-based) analyses. 2. The analysis for sheep annual r showed negative effects of abundance and negative effects of the interaction of abundance and climate, measured as March rainfall (and winter NAO) in the best fitting models. The analysis for deer annual r showed a negative effect of deer abundance and a positive effect of climate measured as March rainfall (but a negative effect of winter NAO), but no interaction of abundance and climate in the best fitting models. 3. There was most support in the analysis of sheep dynamics for the ratio numerical response and the assumption that parameter J (equilibrium food per animal) was influenced by climate. In the analysis of deer dynamics there was most support for the numerical responses assuming effects of food and density (Ivlev and density, food and density, and additive responses) and slightly less support for the ratio numerical response. The evaluation of such models would be aided by the collection of and incorporation of food data into the analyses.     

宝天曼国家级自然保护区落叶阔叶林密度制约效应对树木存活的影响

密度制约对物种共存起着重要作用。随着密度制约效应研究的深入, 亲缘关系较近的物种间表现出的密度制约效应逐渐被人们认识。本研究基于2009和2014年对宝天曼1 ha落叶阔叶林样地的2次调查数据, 利用广义线性混合模型分析了重要值排名前11位的物种不同邻域尺度的密度制约效应对不同径级(小径级(1 cm ≤ DBH < 5 cm)、中径级(5 cm ≤ DBH < 10 cm)、大径级(DBH ≥ 10 cm))目标个体存活的影响。研究表明: (1) 5年间样地中DBH ≥ 1 cm的所有个体的年均死亡率和增员率分别为5.85%和0.27%; (2)有5个物种的个体存活率与同种邻体个体数及同种邻体胸高断面积显著负相关; (3)小径级个体的存活在5 m的邻域范围内受同种密度制约和谱系密度制约的影响都很显著; 中径级个体在3个尺度上受到的密度制约和谱系密度制约的影响都不大; 大径级个体在7.5 m、10 m邻域范围内受谱系密度制约影响显著。结果表明, 同种密度制约和谱系密度制约效应对宝天曼落叶阔叶林不同生长阶段的树木个体影响不同。     

Intraspecific variation in the strength of density dependence in aphid populations

Abstract.  1. Experimental evidence is presented for positive, negative, and no density dependence from 32 independent density manipulations of milkweed aphids ( Aphis nerii ) in laboratory and field experiments. This substantial variation in intraspecific density dependence is associated with temperature and host-plant species.
2. It is reported that as population growth rate increases, density dependence becomes more strongly negative, suggesting that the monotonic definition of density dependence used in many common population models is appropriate for these aphids, and that population growth rate and carrying capacity are not directly proportional.
3. For populations that conform to these assumptions, population growth rate may be widely applicable as a predictor of the strength of density dependence.     

Density dependence in cereal aphid populations

Long sequences of data on the incidence of cereal aphids from five European countries were analysed for evidence of density dependent processes occurring between years. Using a randomisation test (Pollard, Lakhani & Rotheray, 1987), density dependence was revealed in all (16) population censuses of Metopolo-phium dirhodum , 60% of Rhopalosiphum padi censuses (10 of 17) but only 25% of Sitobion avenae population censuses (4 of 16). Correcting for density independent effects of weather revealed the existence of significant direct density dependence in some populations censuses where it was previously undetected. The implications of density dependence in cereal aphid populations are considered.     

Patterns of density dependence in growth,reproduction and survival in the invasive freshwater snail Pomacea canaliculata in Japanese rice fields

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Cycles, stochasticity and density dependence in pink salmon population dynamics

Complex dynamics of animal populations often involve deterministic and stochastic components. A fascinating example is the variation in magnitude of 2-year cycles in abundances of pink salmon (Oncorhynchus gorbuscha) stocks along the North Pacific rim. Pink salmon have a 2-year anadromous and semelparous life cycle, resulting in odd- and even-year lineages that occupy the same habitats but are reproductively isolated in time. One lineage is often much more abundant than the other in a given river, and there are phase switches in dominance between odd- and even-year lines. In some regions, the weak line is absent and in others both lines are abundant. Our analysis of 33 stocks indicates that these patterns probably result from stochastic perturbations of damped oscillations owing to density-dependent mortality caused by interactions between lineages. Possible mechanisms are cannibalism, disease transmission, food depletion and habitat degradation by which one lineage affects the other, although no mechanism has been well-studied. Our results provide comprehensive empirical estimates of lagged density-dependent mortality in salmon populations and suggest that a combination of stochasticity and density dependence drives cyclical dynamics of pink salmon stocks.     

Long-term variability and density dependence in Hudson River Dreissena populations

1. We used a 27-year record of Dreissena populations in the freshwater tidal Hudson River to describe interannual variation in population density, body size, and body condition; estimate long-term variation in recruitment, survivorship, and shell growth; and assess possible controls on the populations.
2. Dreissena populations in the Hudson have been highly variable, with interannual ranges of c. 100-fold in abundance and biomass, and 7-fold in mean body mass. This large interannual variation arises from both long-term trends and 2–5-year cycles.
3. Long-term trends include the 2008 appearance of the quagga mussel (Dreissena rostriformis), which still forms a small part (<10%) of the dreissenid community, and a decline in zebra mussel body size. The decline in body size was caused by a long-term decline in adult survivorship rather than a decline in rates of shell growth. We could detect no long-term trends in adult abundance or spread of Dreissena onto soft sediments in the Hudson.
4. We observed persistent, strong cycles in adult abundance and body size. These were driven by the appearance and decay of eight dominant year classes over the 27 years of our study, and were a result of temporal variation in recruitment rather than temporal variation in survivorship. The observed strongly irregular recruitment appears to arise from strong adult–larval interactions, and is consistent with previous simulation model results showing that interactions between adults and larvae can drive persistent cycling.
5. We found evidence that negative density dependence affects recruitment, somatic growth, and body condition of Dreissena in the Hudson. Warm summers may also cause high adult mortality.
6. We put our results into the context of a conceptual model of Dreissena population dynamics, and argue that neither the dynamics nor the controls of populations of these important invaders is known satisfactorily.

     

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.     

Mechanisms of density dependence in fluctuating vole populations: deducing annual density dependence from seasonal processes

Based on recent advances in time-series analyses of ecological dynamics using statistical and mathematical models, we summarise our recent results on the seasonal processes in the annual population dynamics of the grey-sided vole Clethrionomys rufocanus (Sundevall, 1846) in Hokkaido, Japan, and report additional analyses on annual and seasonal density dependence. Annual direct density dependence was strong in almost all populations. In contrast, delayed density dependence was generally weak, although clear delayed density dependence was detected in some of the studied populations. Although seasonal density dependence was observed both in winter and summer, direct density dependence was much more profound during winter; thus, winter density dependence contributed most to the overall annual direct density dependence. We found no correlation between the seasonal components of annual direct density dependence; however, the corresponding seasonal components for annual delayed density dependence were positively correlated. We conclude that winter conditions influence the strength of annual direct density dependence most profoundly. Moreover, we conclude that direct density dependence during summer and winter may be generated by different mechanisms, whereas delayed density dependence seems to be generated by a common mechanism. Candidate mechanisms are discussed in relation to general knowledge of northern rodent populations and to specific insights provided by earlier studies of grey-sided voles in Hokkaido.     

Native insect herbivory overwhelms context dependence to limit complex invasion dynamics of exotic weeds

Understanding the role of consumers in density‐dependent plant population dynamics is a long‐standing goal in ecology. However, the generality of herbivory effects across heterogeneous landscapes is poorly understood due to the pervasive influence of context‐dependence. We tested effects of native insect herbivory on the population dynamics of an exotic thistle, Cirsium vulgare, in a field experiment replicated across eight sites in eastern Nebraska. Using hierarchical Bayesian analysis and density‐dependent population models, we found potential for explosive low‐density population growth (λ > 5) and complex density fluctuations under herbivore exclusion. However, herbivore access drove population decline (λ < 1), suppressing complex fluctuations. While plant–herbivore interaction outcomes are famously context‐dependent, we demonstrated that herbivores suppress potentially invasive populations throughout our study region, and this qualitative outcome is insensitive to environmental context. Our novel use of Bayesian demographic modelling shows that native insect herbivores consistently prevent hard‐to‐predict fluctuations of weeds in environments otherwise susceptible to invasion.     

Density-dependent population dynamics in clonal organisms: a modelling approach

1. Density dependence may act at several stages in an organisms life-cycle (e.g. on mortality, fecundity, etc.), but not all density-dependent processes necessarily regulate population size. In this paper I use a density manipulation experiment to determine the effects of density on the transition rates between different size classes of the clonal zoanthid Palythoa caesia Dana 1846. I then formulate a density-dependent matrix model of population dynamics of Palythoa , and perform a series of sensitivity analyses on the model to determine at what stage in the life-cycle regulation acts.
2. Seven of the 16 transition probabilities decreased with density, most of them being shrinkage (due to loss of tissue or fission) and stasis (the self–self transition) of medium and large colonies. The only probability to increase was for the stasis of large colonies. Recruitment was quadratically dependent on density, peaking at intermediate densities.
3. Equilibrium cover in the model was 84% and was reached in ≈40 years. To determine which density-dependent transitions were involved in population regulation, the strength of density dependence was varied in each independently. This sensitivity analysis showed that only changes in the probabilities of large colonies remaining large and producing medium colonies, were regulating.
4. These results suggest that regulation is primarily acting on fission of large colonies to produce intermediate-sized colonies, in combination with size specific growth rates. Fission rates decrease greatly with density, resulting in a greater proportion of large colonies at high densities and large colonies grow more slowly than small. Overall, this behaviour is very similar to that of clonal plants which have a phalanx type life history.     

Demographic buffering and compensatory recruitment promotes the persistence of disease in a wildlife population

Demographic buffering allows populations to persist by compensating for fluctuations in vital rates, including disease‐induced mortality. Using long‐term data on a badger (Meles meles Linnaeus, 1758) population naturally infected with Mycobacterium bovis, we built an integrated population model to quantify impacts of disease, density and environmental drivers on survival and recruitment. Badgers exhibit a slow life‐history strategy, having high rates of adult survival with low variance, and low but variable rates of recruitment. Recruitment exhibited strong negative density‐dependence, but was not influenced by disease, while adult survival was density independent but declined with increasing prevalence of diseased individuals. Given that reproductive success is not depressed by disease prevalence, density‐dependent recruitment of cubs is likely to compensate for disease‐induced mortality. This combination of slow life history and compensatory recruitment promotes the persistence of a naturally infected badger population and helps to explain the badger's role as a persistent reservoir of M. bovis.     

Cowpox virus infection in natural field vole Microtus agrestis populations: delayed density dependence and individual risk

1. Little is known about the dynamics of pathogen (microparasite) infection in wildlife populations, and less still about sources of variation in the risk of infection. Here we present the first detailed analysis of such variation. 2. Cowpox virus is an endemic sublethal pathogen circulating in populations of wild rodents. Cowpox prevalence was monitored longitudinally for 2 years, in populations of field voles exhibiting multiannual cycles of density in Kielder Forest, UK. 3. The probability that available susceptible animals seroconverted in a given trap session was significantly positively related to host density with a 3-month time lag. 4. Males were significantly more likely to seroconvert than females. 5. Despite most infection being found in young animals (because transmission rates were generally high) mature individuals were more likely to seroconvert than immature ones, suggesting that behavioural or physiological changes associated with maturity contribute to variation in infection risk. 6. Hence, these analyses confirm that there is a delayed numerical response of cowpox infection to vole density, supporting the hypothesis that endemic pathogens may play some part in shaping vole cycles.     

Multiple sources of evidence for density dependence in the endangered Hawaiian stilt (Himantopus mexicanus knudseni)

Hawaiian stilts (Himantopus mexicanus knudseni) are an endangered subspecies of the Black-necked stilt endemic to the Hawaiian Islands. Despite long-term study, the main drivers of Hawaiian stilt population dynamics are poorly understood. We tested for density dependence using two sources of evidence: a 30-year time series of annual estimated range-wide abundance, and two 15+ year time series of reproductive success. Using separate methods with independent data, sources allowed us to make up for the potentially positive bias of one approach with the more conservative nature of the second. We compared nonlinear density-dependent and density-independent population model fits to our time-series data, using both frequentist and Bayesian state-space approaches. Across both approaches, density-dependent models best fit observed population dynamics, with lower AICc and cross-validation statistics compared to density-independent models. Among density-dependent models, a conditional model in which density-independent dynamics occur below a population size threshold (~850–1,000 birds), and then density-dependent dynamics occur above that threshold, performed best across Bayesian and frequentist model comparisons, with the Ricker model ranked next or equivalently. Our analysis of reproduction data revealed a strong negative effect of local adult density on nest success (proportion of nests hatching at least one chick) at Kealia National Wildlife Refuge on Maui, where few alternative breeding habitats are available, but no such effect at another site where many nearby alternative wetlands are available. These congruent results across independent datasets and analytical approaches support the hypothesis that Hawaiian stilts exhibit density dependence across their range.