Understanding contributions of cohort effects to growth rates of fluctuating populations

1. Understanding contributions of cohort effects to variation in population growth of fluctuating populations is of great interest in evolutionary biology and may be critical in contributing towards wildlife and conservation management. Cohort-specific contributions to population growth can be evaluated using age-specific matrix models and associated elasticity analyses. 2. We developed age-specific matrix models for naturally fluctuating populations of stoats Mustela erminea in New Zealand beech forests. Dynamics and productivity of stoat populations in this environment are related to the 3-5 year masting cycle of beech trees and consequent effects on the abundance of rodents. 3. The finite rate of increase (lambda) of stoat populations in New Zealand beech forests varied substantially, from 1.98 during seedfall years to 0.58 during post-seedfall years. Predicted mean growth rates for stoat populations in continuous 3-, 4- or 5-year cycles are 0.85, 1.00 and 1.13. The variation in population growth was a consequence of high reproductive success of females during seedfall years combined with low survival and fertility of females of the post-seedfall cohort. 4. Variation in population growth was consistently more sensitive to changes in survival rates both when each matrix was evaluated in isolation and when matrices were linked into cycles. Relative contributions to variation in population growth from survival and fertility, especially in 0-1-year-old stoats, also depend on the year of the cycle and the number of transitional years before a new cycle is initiated. 5. Consequently, management strategies aimed at reducing stoat populations that may be best during one phase of the beech seedfall cycle may not be the most efficient during other phases of the cycle. We suggest that management strategies based on elasticities of vital rates need to consider how population growth rates vary so as to meet appropriate economic and conservation targets.     

Analysis of interspecific competition in perennial plants using Life Table Response Experiments

The impact of interspecific competition is usually measured by its effect upon plant growth, neglecting impacts upon other stages of the life cycle such as fecundity which have a direct influence upon individual fitness and the asymptotic population growth rate of a population (λ). We used parameterized matrix models for three perennial plant species grown with and without interspecific competition to illustrate how the methodology of Life Table Response Experiments (LTRE) can be used to link any change in population dynamics to changes in any part of the life cycle. Plants were herbaceous grassland species grown for two years in a field experiment at Rothamsted Experimental Station, England. Interspecific competition reduced λ by over 90% in all species. Survival and growth were slightly affected by competition whereas plant fecundity was greatly reduced. Nearly all of the observed difference in λ between the competition treatments was explained by the fecundity terms, and more precisely by a large difference in the number of seeds, and a high sensitivity of λ to the germination rate. Whereas most competition studies focus on the measurement of change in individual fitness, our study illustrates how informative it is to take account not only of the effect of competition upon vital rates but also of how different vital rates affect population growth rate.     

Inferring population trends for the world's largest fish from mark-recapture estimates of survival

1. Precise estimates of demographic rates are key components of population models used to predict the effects of stochastic environmental processes, harvest scenarios and extinction probability. 2. We used a 12-year photographic identification library of whale sharks from Ningaloo Reef, Western Australia to construct Cormack-Jolly-Seber (CJS) model estimates of survival within a capture-mark-recapture (CMR) framework. Estimated survival rates, population structure and assumptions regarding age at maturity, longevity and reproduction frequency were combined in a series of age-classified Leslie matrices to infer the potential trajectory of the population. 3. Using data from 111 individuals, there was evidence for time variation in apparent survival (phi) and recapture probability (p). The null model gave a phi of 0.825 (95% CI: 0.727-0.893) and p = 0.184 (95% CI: 0.121-0.271). The model-averaged annual phi ranged from 0.737 to 0.890. There was little evidence for a sex effect on survival. 4. Using standardized total length as a covariate in the CMR models indicated a size bias in phi. Ignoring the effects of time, a 5-m shark has a phi = 0.59 and a 9 m shark has phi = 0.81. 5. Of the 16 model combinations considered, 10 (63%) indicated a decreasing population (lambda < 1). For models based on age at first reproduction (alpha) of 13 years, the mean age of reproducing females at the stable age distribution (A) ranged from 15 to 23 years, which increased to 29-37 years when alpha was assumed to be 25. 6. All model scenarios had higher total elasticities for non-reproductive female survival [E(s(nr))] compared to those for reproductive female survival [E(s(r))]. 7. Assuming relatively slow, but biologically realistic, vital rates (alpha = 25 and biennial reproduction) and size-biased survival probabilities, our results suggest that the Ningaloo Reef population of whale sharks is declining, although more reproductive data are clearly needed to confirm this conclusion. Combining relatively precise survival estimates from CMR studies with realistic assumptions of other vital rates provides a useful heuristic framework for determining the vulnerability of large oceanic predators for which few direct data exist.     

Demographic models to simulate the stable ratio between ecologically similar gametophytes and tetrasporophytes in populations of the Gigartinaceae (Rhodophyta)

In populations of the Gigartinaceae (Rhodophyta), gametophytes often predominate numerically over tetrasporophytes. Several hypotheses have been proposed to explain this dominance, based on the usually implicit assumption that the stable ratio between gametophytes and tetrasporophytes (G:T ratio) should be 1 if both reproductive phases are ecologically similar. We developed demographic models to test this assumption, for which we considered that both phases are ecologically similar. Defining ecologic similarity for most demographic rates is relatively straightforward, except for rates of spore output. The first set of models considered the same spore output per thallus of both phases as representing ecologic similarity. Model iterations led to stable G:T ratios of 1 for triennial and for perennial thalli, regardless of the initial G:T ratio, but not for annual thalli with initial G:T ratios different from 1. However, equal spore output may not represent ecologic similarity, due to size differences between carpospores and tetraspores. The second set of models considered the lowest possible spore output for each phase, according to the life history of this family: only one carposporangium, with one carpospore, is produced from every two gametophytes and only one tetrasporangium, with four tetraspores, is produced by every tetrasporophyte. Model iterations led to stable G:T ratios of 2.8 for most cases, a ratio of 1 being obtained only every 2 years for annual thalli with an initial G:T ratio of 1. Increasing absolute spore output, without altering the relative output between phases and incorporating density-independent mortality through a matrix model, given the same mortality rate for both phases, did not modify results. We suggest that the combination of both modeling and field research may uncover more rapidly than otherwise the most relevant ecologic differences between phases, if any, that underlie the G:T ratio observed for a given population.     

Environmental extremes versus ecological extremes: impact of a massive iceberg on the population dynamics of a high-level Antarctic marine predator

Extreme events have been suggested to play a disproportionate role in shaping ecological processes, but our understanding of the types of environmental conditions that elicit extreme consequences in natural ecosystems is limited. Here, we investigated the impact of a massive iceberg on the dynamics of a population of Weddell seals. Reproductive rates of females were reduced, but survival appeared unaffected. We also found suggestive evidence for a prolonged shift towards higher variability in reproductive rates. The annual number of females attending colonies showed unusual swings during the iceberg period, a pattern that was apparently the consequence of changes in sea-ice conditions. In contrast to the dramatic effects that were recorded in nearby populations of emperor penguins, our results suggest that this unusual environmental event did not have an extreme impact on the population of seals in the short-term, as they managed to avoid survival costs and were able to rapidly re-achieve high levels of reproduction by the end of the perturbation. Nevertheless, population projections suggest that even this modest impact on reproductive rates could negatively affect the population in the long run if such events were to occur more frequently, as is predicted by models of climate change.     

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.     

Stochastic matrix models for conservation and management: a comparative review of methods

Stochastic matrix models are frequently used by conservation biologists to measure the viability of species and to explore various management actions. Models are typically parameterized using two or more sets of estimated transition rates between age/size/stage classes. While standard methods exist for analyzing a single set of transition rates, a variety of methods have been employed to analyze multiple sets of transition rates. We review applications of stochastic matrix models to problems in conservation and use simulation studies to compare the performance of different analytic methods currently in use. We find that model conclusions are likely to be robust to the choice of parametric distribution used to model vital rate fluctuations over time. However, conclusions can be highly sensitive to the within-year correlation structure among vital rates, and therefore we suggest using analytical methods that provide a means of conducting a sensitivity analysis with respect to correlation parameters. Our simulation results also suggest that the precision of population viability estimates can be improved by using matrix models that incorporate environmental covariates in conjunction with experiments to estimate transition rates under a range of environmental conditions.