Dynamical and statistical models of vertebrate population dynamics

Population dynamics methodology now powerfully combines discrete time models (with constant parameters, density dependence, random environment, and/or demographic stochasticity) and capture-recapture models for estimating demographic parameters. Vertebrate population dynamics has strongly benefited from this progress: survival estimates have been revised upwards, trade-offs between life history traits have been demonstrated, analyses of population viability and management are more and more realistic. Promising developments concern random effects, multistate and integrated models. Some biological questions (density dependence, links between individual and population levels, and diversification of life histories) can now be efficiently attacked.     

Sociality, individual fitness and population dynamics of yellow-bellied marmots

Social behaviour was proposed as a density-dependent intrinsic mechanism that could regulate an animal population by affecting reproduction and dispersal. Populations of the polygynous yellow-bellied marmot (Marmota flaviventris) fluctuate widely from year to year primarily driven by the number of weaned young. The temporal variation in projected population growth rate was driven mainly by changes in the age of first reproduction and fertility, which are affected by reproductive suppression. Dispersal is unrelated to population density, or the presence of the father; hence, neither of these limits population growth or acts as an intrinsic mechanism of population regulation; overall, intrinsic regulation seems unlikely. Sociality affects the likelihood of reproduction in that the annual probability of reproducing and the lifetime number of offspring are decreased by the number of older females and by the number of same-aged females present, but are increased by the number of younger adult females present. Recruitment of a yearling female is most likely when her mother is present; recruitment of philopatric females is much more important than immigration for increasing the number of adult female residents. Predation and overwinter mortality are the major factors limiting the number of resident adults. Social behaviour is not directed towards population regulation, but is best interpreted as functioning to maximize direct fitness.     

Genetic variability and viral seroconversion in an outcrossing vertebrate population

Inverse correlations between genetic variability and parasitism are important concerns for conservation biologists. We examined correlations between neutral genetic variability and the presence of antibodies to canine distemper virus (CDV) and feline parvovirus (FPV) in a free-ranging population of raccoons. Over 3 years there was a strong relationship between age and seroprevalence rates. Most young animals were seronegative to CDV and FPV, but the oldest age class was greater than 80 per cent seropositive to both viruses. CDV-seropositive animals had greater heterozygosity and lower measures of inbreeding compared with CDV-seronegative animals. This relationship was strongest among the youngest animals and did not occur during a 1 year CDV epidemic. In contrast, FPV-seropositive animals only had significantly lower measures of inbreeding in 1 year, perhaps because FPV-associated mortality is relatively low or primarily occurs among very young individuals that were under-represented in our sampling. These results suggest that even in large outcrossing populations, animals with lower heterozygosity and higher measures of inbreeding are less likely to successfully mount an immune response when challenged by highly pathogenic parasites.     

Relating individual behaviour to population dynamics

How do the behavioural interactions between individuals in an ecological system produce the global population dynamics of that system? We present a stochastic individual-based model of the reproductive cycle of the mite Varroa jacobsoni, a parasite of honeybees. The model has the interesting property in that its population level behaviour is approximated extremely accurately by the exponential logistic equation or Ricker map. We demonstrated how this approximation is obtained mathematically and how the parameters of the exponential logistic equation can be written in terms of the parameters of the individual-based model. Our procedure demonstrates, in at least one case, how study of animal ecology at an individual level can be used to derive global models which predict population change over time.     

Evolution of exploitation ecosystems I. Predation,foraging ecology and population dynamics in herbivores

Summary The hypothesis of exploitation ecosystems was reanalysed using the model of Armstrong (1979) which simultaneously deals with population dynamics and evolution. The results indicate that the prediction of Oksanenet al. (1981) of strict predation limitation of herbivores in productive ecosystems does not hold for coevolved systems. Depending on the nature of herbivore-carnivore coevolution, herbivore biomass may level off at a threshold productivity value or increase monotonously with increasing primary productivity, though at a strongly reduced rate in productive ecosystems. Under both circumstances, increasing primary productivity is predicted to be accompanied by gradual replacement of genuine folivores by semi-granivores and true granivores. The dominating guild members are predicted to show some degree of resource-limitation, although only granivores are predicted to be chiefly resource-limited even in the most productive ecosystems. Data on arctic-to-temperate patterns in the community structure of herbivorous vertebrates conform to the implications of the analysis.     

The consequences of individual variability in moulting probability and the aggregation of stages for modelling copepod population dynamics

Individual variability in development rates of the calanoidcopepod Eurytemora affinis was studied using previously publisheddata that are here normalised for the effects of food and temperature.The late-developing individuals died before reaching the adultstage. The development of individuals was studied using twodifferent stage aggregations. In the one case, the stages wereaggregated into four groups: early nauplii (N1–N3), laternauplii (N4–N6), early copepodites (C1–C3) and latercopepodites (C4–C5). In another case, only total nauplii(N1–N6) and total copepodites (C1–C5) were used.The distributions of development time in the first groups obtainedfrom the normalised data show distinct asymmetry. The gammadensity function is asymmetrical to fit the distributions ofdevelopment in the different groups under different experimentalconditions. For one experimental condition (10³ cells ml^–1,15°C) the development of E. affinis was simulated usingan age-within-stage model together with the fitted distributionsof moulting probabilities. We showed that the asymmetrical gammadistribution was an adequate parameterization of the transferprocess. It was also useful to simplify the life cycle representationby lumping developmental stages. However, the comparison ofthe simulated abundance with two life cycle representationsshowed another source of error due to the heterogeneity of stage-specificmortality rates. The error of the estimation increases in proportionto the difference of mortality rates between two successivestages.     

Predation and bark beetle dynamics

Bark beetle populations may undergo dramatic fluctuations and are often important pests in coniferous forests. Their dynamics are thought to be primarily driven by factors affecting the resistance of the host tree to attack, i.e., bottom-up forces, while natural enemies are usually assigned a minor role in these systems. I present behavioral experiments that suggest that the clerid beetle Thanasimus dubius may be an important source of mortality for the bark beetle Dendroctonus frontalis during attack of the host tree, and determine the nature of the functional response of T. dubius under conditions close to natural. I also examine the numerical response of T. dubius to large-scale fluctuations in D. frontalis density, and the relationship between bark beetle population trends and predator density, and find that beetle populations tend to decline when predator densities are high. Combined with the effects of clerid larvae on bark beetle broods, these results suggest that top-down forces generated by natural enemies could also be an important component of bark beetle dynamics. The implications of these results for bark beetle dynamics are discussed in relation to the prolonged life-cycle of clerid beetles. Received: 23 January 1997 / Accepted: 5 April 1997     

Predation of freshwater jellyfish on Bosmina : the consequences for population dynamics,body size,and morphology

Invertebrate predators may cause strong changes in behaviour, life-history, and morphology of prey species. However, little is known about the influence of jellyfish on such characteristics of their prey. This study analyses the impacts of the freshwater jellyfish Craspedacusta sowerbii on life history and morphological defenses in a population of the cladoceran Bosmina longirostris . Length of mucro and antennule, size-dependent number of eggs, size at maturity, and size of juveniles, adults, and egg-carrying females were investigated during a 23 days experiment using medusae-enriched and control enclosures filled with natural plankton populations. Significant differences in parameters investigated were found not only between treatments, but also within treatments over time. Changes in Bosmina life-history parameters and morphology in controls were probably due to predation by cyclopoid copepods. The significant increase in the size of adults and egg-carrying females as well as the increase in mucro and antennule length in medusae-enriched enclosures are discussed as defense strategies against the freshwater jellyfish.     

Predation of experimental nests is linked to local population dynamics in a fragmented bird population

Artificial nest experiments (ANEs) are widely used to obtain proxies of natural nest predation for testing a variety of hypotheses, from those dealing with variation in life-history strategies to those assessing the effects of habitat fragmentation on the persistence of bird populations. However, their applicability to real-world scenarios has been criticized owing to the many potential biases in comparing predation rates of artificial and natural nests. Here, we aimed to test the validity of estimates of ANEs using a novel approach. We related predation rates on artificial nests to population viability analyses in a songbird metapopulation as a way of predicting the real impact of predation events on the local populations studied. Predation intensity on artificial nests was negatively related to the species' annual population growth rate in small local populations, whereas the viability of large local populations did not seem to be influenced, even by high nest predation rates. The potential of extrapolation from ANEs to real-world scenarios is discussed, as these results suggest that artificial nest predation estimates may predict demographic processes in small structured populations.     

The seasonal variability of Drosophila population according to the individual asymmetry

The natural Drosophila population is characterized by the presence of directional (DA) and fluctuating (FA) asymmetry of individuals. It was found that genotype has an effect on DA-level. FA evaluated in spring, summer and autumn periods had its maximum value in summer period. Genetically determined seasonal decrease in size of individuals was accompanied with increase in their FA. The structure of FA population variability is defined by genotypes of individuals. The phenotype and genotype structures of Drosophila population were investigated by FA of the individuals. There was regrouping of lineages number within of each frequency class in period from spring to autumn. Investigating central frequency class with least FA values in spring to summer period we observed the decrease in number of lineages for all traits with the exception of sternoupleral bristles (SB). At the same time the increase in number of lineages in the central and extreme frequency class with maximal FA values of sternoupleral bristles (SB). At the same time the increase in number of lineages in the central and extreme frequency class with maximal FA values of lines is observed. The number of lineages in the central frequency class of genotype structure is prevailing to all traits, without dependence on season. Individuals with rather high FA value acquire advantages in summer period whereas the individuals with low FA--in spring and autumn periods. Annual dynamics of FA is defined by this population parameters reorganization. The reasons of seasonal change of FA are discussed.     

Predation and the evolutionary dynamics of species ranges

Gene flow that hampers local adaptation can constrain species distributions and slow invasions. Predation as an ecological factor mainly limits prey species ranges, but a richer array of possibilities arises once one accounts for how predation alters the interplay of gene flow and selection. We extend previous single-species theory on the interplay of demography, gene flow, and selection by investigating how predation modifies the coupled demographic-evolutionary dynamics of the range and habitat use of prey. We consider a model for two discrete patches and a complementary model for species along continuous environmental gradients. We show that predation can strongly influence the evolutionary stability of prey habitat specialization and range limits. Predators can permit prey to expand in habitat or geographical range or, conversely, cause range collapses. Transient increases in predation can induce shifts in prey ranges that persist even if the predator itself later becomes extinct. Whether a predator tightens or loosens evolutionary constraints on the invasion speed and ultimate size of a prey range depends on the predator effectiveness, its mobility relative to its prey, and the prey's intraspecific density dependence, as well as the magnitude of environmental heterogeneity. Our results potentially provide a novel explanation for lags and reversals in invasions.     

Environmental variability uncovers disruptive effects of species' interactions on population dynamics

How species respond to changes in environmental variability has been shown for single species, but the question remains whether these results are transferable to species when incorporated in ecological communities. Here, we address this issue by analysing the same species exposed to a range of environmental variabilities when (i) isolated or (ii) embedded in a food web. We find that all species in food webs exposed to temporally uncorrelated environments (white noise) show the same type of dynamics as isolated species, whereas species in food webs exposed to positively autocorrelated environments (red noise) can respond completely differently compared with isolated species. This is owing to species following their equilibrium densities in a positively autocorrelated environment that in turn enables species–species interactions to come into play. Our results give new insights into species'' response to environmental variation. They especially highlight the importance of considering both species'' interactions and environmental autocorrelation when studying population dynamics in a fluctuating environment.     

What individual life histories can (and cannot) tell about population dynamics

We present an overview of a long-term research programme that is aimed at revealing the relations between individual feeding, growth, reproduction and mortality in Daphnia pulex and the state and dynamics of the population. We analyse a physiologically structured population model, in which individual performance is described using an energy budget model that incorporates a food dependence. The model predictions are shown to be at odds with experimental observations on populations of Daphnia. We argue that these discrepancies are primarily due to insufficient knowledge about the precise size-scaling of the food ingestion rate, which plays a central role in the competitive interaction among individuals. To a lesser extent, the discrepancies arise because details about the energy budget of individual Daphnia are not sufficiently known for the food conditions prevailing in population experiments.     

Emergence of individual behaviour at the population level. Effects of density-dependent migration on population dynamics

The aim of this work is to study the effects of different individual behaviours on the overall growth of a spatially distributed population. The population can grow on two spatial patches, a source and a sink, that are connected by migrations. Two time scales are involved in the dynamics, a fast one corresponding to migrations and a slow one associated with the local growth on each patch. Different scenarios of density-dependent migration are proposed and their effects on the population growth are investigated. A general discussion on the use of aggregation methods for the study of integration of different ecological levels is proposed.     

High genetic variability in an ecologically variable vertebrate,Bufo viridis

Allozymic variation in proteins encoded by 26 loci was analyzed electrophoretically in 517 specimens of green toads from 11 populations from Israel and one population from Vis Island in the Adriatic Sea. Genetic variation in this toad is the highest yet reported in any vertebrate. All three genetic parameters, mean number of alleles per locus (A), mean proportion of loci polymorphic per population (P), and mean number of heterozygous loci per individual (H), are very high (A = 1.65, range 1.38-2.04; P = 0.423, range 0.346-0.615; H = 0.133, range 0.108-0.159). Central and marginal mainland populations are only slightly more variable than desert isolates, but much more variable than the Vis Island population. Genetic similarity is very high between mainland populations (S = 0.951, range 0.93-0.97). Frequencies of two alleles (Icd-lc and Tfa) are correlated with an ecological gradient of increasing aridity. Regulatory enzymes appeared to contribute more to overall polymorphism than non-regulatory enzymes. The genetic variation observed suggests that selection for heterozygosity as an adaptive strategy is operating in the ecologically variable environment in which green toads live.     

Photo-identification of individual weedy seadragons Phyllopteryx taeniolatus and its application in estimating population dynamics

Forty-three individual adult weedy seadragons Phyllopteryx taeniolatus were identified from underwater images using patterns of spots and blotches on the lateral surface of the abdomen. These patterns were unique and did not change over the 18 month course of the study and could therefore be used to identify individuals when estimating population variables using non-invasive capture-mark-recapture and accumulation curve methods. Two similar neighbouring sites in southern Tasmania showed considerable differences in their estimated populations of P. taeniolatus. Estimated annual survival was >80% at one site suggesting that P. taeniolatus may be considerably longer lived than other syngnathids with a maximum life span in excess of 10 years. Males incubating embryos were observed from October to March and at least two clutches could be borne during this period. This technique of photo-identification could provide a cheap and effective way to monitor populations of this iconic species across its range, particularly in conjunction with optimized pattern-recognition software.     

Predation impact of Leptodora kindtii on population dynamics and morphology of Bosmina fatalis and B. longirostris in mesocosms

1. We assessed the impact of predation by the invertebrate predator Leptodora kindtii on Bosmina longirostris and B. fatalis, which show seasonal and reciprocal succession patterns in Lake Suwa, in a mesocosm experiment using 20‐L tanks with or without the predator under different food conditions. We also analysed morphological responses of the two Bosmina species to the predator in the tanks. 2. Bosmina fatalis dominated B. longirostris regardless of predator presence under high food density. However, the presence of Leptodora induced the dominance of B. fatalis more rapidly than its absence. On the contrary, no dominance of B. fatalis was observed in tanks with low food density, irrespective of the presence of the predator. Only B. fatalis showed morphological changes in response to the presence of Leptodora. 3. Mucrone length and antennule shape (angle between body and antennule and angle between antennules) showed marked responses at both high and low food densities, but antennule length responded only at high food density. Mucrone length seems to be a more effective defence against Leptodora. 4. The results suggest that B. fatalis is a superior competitor against B. longirostris and is more resistant to Leptodora predation, especially in good food conditions. The repeatedly observed seasonal succession of the two Bosmina species in the eutrophic Lake Suwa – the replacement of B. longirostris by B. fatalis following the occurrence of abundant Leptodora– seems to be caused by the selective predation of Leptodora on B. longirostris as well as the competitive ability of B. fatalis.