Stéphane Legendre

Demographic Stochasticity and Social Mating System in the Process of Extinction of Small Populations: The Case of Passerines Introduced to New Zealand

Underlying the many causes of extinction of small populations is the random fate of each constituent individual or, in other words, demographic stochasticity. Demographic stochasticity is inherent to any demographic process, regardless of the environment, and its strength increases as population size gets smaller. In particular, random fluctuations in the proportion of males and females and the way they pair for reproduction (i.e., the social mating system) are usually neglected. To assess the potential importance of demographic stochasticity to the extinction process, a two‐sex model with an explicit mating system was built. Extinction probabilities computed via Monte Carlo simulation were compared to real data, the case of passerines introduced to New Zealand a century ago. This minimal model of extinction allowed assessment of the importance of the mating system in the colonization process. Monogamous mating led to a higher extinction risk than did polygynous mating. Demographic uncertainty imposes high extinction probabilities on short‐lived bird species as compared to long‐lived bird species. Theoretical results for two‐sex models are provided.

ULM, a software for conservation and evolutionary biologists

Matrix models for population dynamics have recently been studied intensively and have many applications to theoretical and applied problems (conservation, management). The computer program ULM (Unified Life Models) collects a good part of the actual knowledge on the subject. It is a powerful tool to study the life cycle of species and meta-populations. In the general framework of discrete dynamical systems and symbolic computation, simple commands and convenient graphics are provided to assist the biologist. The main features of the program are shown through detailed examples: a simple model of a starling population life cycle is first presented leading to basic concepts (growth rates, stable age distribution, sensitivities); the same model is used to study competing strategies in a varying environment (extinction probabilities, stochastic sensitivities); a meta-population model with migrations is then presented; some results on migration strategies and evolutionary stable strategies are eventually proposed.

Eco-evolutionary feedbacks, adaptive dynamics and evolutionary rescue theory.

Adaptive dynamics theory has been devised to account for feedbacks between ecological and evolutionary processes. Doing so opens new dimensions to and raises new challenges about evolutionary rescue. Adaptive dynamics theory predicts that successive trait substitutions driven by eco-evolutionary feedbacks can gradually erode population size or growth rate, thus potentially raising the extinction risk. Even a single trait substitution can suffice to degrade population viability drastically at once and cause ‘evolutionary suicide’. In a changing environment, a population may track a viable evolutionary attractor that leads to evolutionary suicide, a phenomenon called ‘evolutionary trapping’. Evolutionary trapping and suicide are commonly observed in adaptive dynamics models in which the smooth variation of traits causes catastrophic changes in ecological state. In the face of trapping and suicide, evolutionary rescue requires that the population overcome evolutionary threats generated by the adaptive process itself. Evolutionary repellors play an important role in determining how variation in environmental conditions correlates with the occurrence of evolutionary trapping and suicide, and what evolutionary pathways rescue may follow. In contrast with standard predictions of evolutionary rescue theory, low genetic variation may attenuate the threat of evolutionary suicide and small population sizes may facilitate escape from evolutionary traps.

Conservation and control strategies for the wolf (Canis lupus) in western Europe based on demographic models

Securing the long-term acceptance of large carnivores such as the wolf (Canis lupus) in Europe and North America raises a difficult challenge to conservation biologists: planning removals to reduce depredations on livestock while ensuring population viability. We use stochastic-stage-structured population models to investigate wolf population dynamics and to assess alternative management strategies. Among the various management strategies advocated by agencies, zoning that involves eliminating wolves outside a restricted area should be designed with caution, because probabilities of extinction are extremely sensitive to the maximum number of packs that a zone can support and to slight changes in stage specific survival probabilities. In a zoned population, viability is enhanced more by decreasing mortality rates in all classes than by increasing wolf zone size. An alternative to zoning is adaptive management, where there is no limit on pack number but population control can be operated whenever some predefined demographic conditions are met. It turns out that an adaptive management strategy that removes a moderate percentage (10%) of the population following each year of more than 5% of total population growth would provide visible actions addressing public concerns while keeping extinction probability low.

Evolutionary Entropy: A Predictor of Body Size, Metabolic Rate and Maximal Life Span

Body size of organisms spans 24 orders of magnitude, and metabolic rate and life span present comparable differences across species. This article shows that this variation can be explained in terms of evolutionary entropy, a statistical parameter which characterizes the robustness of a population, and describes the uncertainty in the age of the mother of a randomly chosen newborn. We show that entropy also has a macroscopic description: It is linearly related to the logarithm of the variables body size, metabolic rate, and life span. Furthermore, entropy characterizes Darwinian fitness, the efficiency with which a population acquires and converts resources into viable offspring. Accordingly, entropy predicts the outcome of natural selection in populations subject to different classes of ecological constraints. This predictive property, when integrated with the macroscopic representation of entropy, is the basis for enormous differences in morphometric and life-history parameters across species.

Predator foraging behaviour drives food‐web topological structure

1. The structure and dynamics of prey populations are shaped by the foraging behaviours of their predators. Yet, there is still little documentation on how distinct predator foraging types control biodiversity, food-web architecture and ecosystem functioning. 2. We experimentally compared the effects of model fish species of two major foraging types of lake planktivores: a size-selective visual feeder (bluegill), and a filter feeder (gizzard shad). The visual feeder forages on individually captured consumer prey, whereas the filter feeder forages on various prey simultaneously, not only consumers but also primary producers. We ran a 1-month mesocosm experiment cross-classifying a biomass gradient of each predator type. We analysed the effect of each fish on food-web architecture by computing major topological descriptors over time (connectance, link density, omnivory index, etc.). These descriptors were computed from 80 predator-prey binary matrices, using taxa mostly identified at the species level. 3. We found that the visual feeder induced more trophic cul-de-sac (inedible) primary-producer species, lower link density and connectance, and lower levels of food-web omnivory and generalism than the filter feeder. Yet, predator biomass did not affect food-web topology. 4. Our results highlight that top-predator foraging behaviour is a key functional trait that can drive food-web topology and ultimately ecosystem functioning.

Ecological emergence of thermal clines in body size

The unprecedented rate of global warming requires a better understanding of how ecosystems will respond. Organisms often have smaller body sizes under warmer climates (Bergmanns rule and the temperature-size rule), and body size is a major determinant of life histories, demography, population size, nutrient turnover rate, and food-web structure. Therefore, by altering body sizes in whole communities, current warming can potentially disrupt ecosystem function and services. However, the underlying drivers of warming-induced body downsizing remain far from clear. Here, we show that thermal clines in body size are predicted from universal laws of ecology and metabolism, so that size-dependent selection from competition (both intra and interspecific) and predation favors smaller individuals under warmer conditions. We validate this prediction using 4.1 × 10(6) individual body size measurements from French river fish spanning 29 years and 52 species. Our results suggest that warming-induced body downsizing is an emergent property of size-structured food webs, and highlight the need to consider trophic interactions when predicting biosphere reorganizations under global warming.

Demography of a savanna palm tree in Ivory Coast (Lamto): population persistence and life-history

AFKIXACT. Burassus aetlziokurn is a dioecious palm tree. of African S~V~~IXLS. A stage-classified matrix population model has been parametrizcd with field data (Lamto reserve, Ivory Coast). It enabled the study of the persistence of the population, and analysis ol’ its sensitivity to diff’crcnt vital rates. Age of’ palms in each stage were estimated to complete the description of the palm life-history. The main results arc: (1) The studied populations arc very close to the equilibrium but the stable stage distribution (predicted by the model) and the observed distribution arc significantly different indicating a former change in the vital rates. (2) Reproduction seems to be highly delayed (first reproduction on average at 114 y), while the estimated duration of the reproductive part of the life-cycle is relatively short (22 y).

Which future for the French Pyrenean brown bear (Ursus arctos) population? An approach using stage-structured deterministic and stochastic models

The Pyrenean brown bear (Ursus arctos) population is considered as one of the most seriously threatened with extinction in Western Europe. To assess its viability and possible needs of augmentation, we develop deterministic and stochastic stage-structured demographic models. The deterministic model reveals that a bear population cannot have a high annual growth rate and is particularly sensitive to breeder survival. High demographic parameters appear to be crucial to population persistence, especially for a small population that remains vulnerable to demographic and environmental stochasticities. The Pyrenean population cannot therefore be considered as viable. Successful conservation strategies for this population would require releasing more bears in both sub-populations in the near future.

Density‐dependent life history and the dynamics of small populations

1. Small population dynamics depend importantly on the strength and shape of density dependence. Unfortunately, the lack of reliable life-history data often prevents to make accurate demographic predictions for populations regulated by density dependence. 2. We created a gradient from low to high densities in small experimental populations of common lizards (Zootoca vivipara) and investigated the shape and strength of the density dependence of life-history traits during a yearly cycle. We then analysed stochastic population dynamics using one-sex and two-sex age-structured matrix models. 3. Body growth and reproductive performances decreased with density, yearling and adult survival and body size at birth were density-independent, and juvenile survival increased with density. The density dependence of reproduction was partly explained by positive effects of body size on age at first reproduction and clutch size. 4. Parturition date decreased with density in sparse populations and then increased, providing one of the first empirical evidence of a component Allee effect in the phenology of reproduction. 5. Population growth rate (λ) was most affected by variations in juvenile and yearling survival. However, density at equilibrium was most affected by juvenile access to reproduction and yearling clutch size. 6. Stochastic simulations revealed that negative density dependence buffers the effects of initial density on extinction probability, has positive effects on the persistence of sparse populations and interacts with sex ratio fluctuations to shape extinction dynamics. 7. This study demonstrates that negative density dependence modifies the dynamics of small populations and should be investigated together with Allee effects to predict extinction risks.