Naturalization of plant populations: the role of cultivation and population size and density

Field experimentation is required to assess the effects of environmental stochasticity on small immigrant plant populations—a widely understood but largely unexplored aspect of predicting any species’ likelihood of naturalization and potential invasion. Cultivation can mitigate this stochasticity, although the outcome for a population under cultivation nevertheless varies enormously from extinction to persistence. Using factorial experiments, we investigated the effects of population size, density, and cultivation (irrigation) on the fate of founder populations for four alien species with different life history characteristics (Echinochloa frumentacea, Fagopyrum esculentum, Helianthus annuus, and Trifolium incarnatum) in eastern Washington, USA. The fate of founder populations was highly variable within and among the 3 years of experimentation and illustrates the often precarious environment encountered by plant immigrants. Larger founder populations produced more seeds (P < 0.001); the role of founder population size, however, differed among years. Irrigation resulted in higher percent survival (P < 0.001) and correspondingly larger net reproductive rate (R ₀; P < 0.001). But the minimum level of irrigation for establishment, R ₀ > 1, differed among years and species. Sowing density did not affect the likelihood of establishment for any species. Our results underscore the importance of environmental stochasticity in determining the fate of founder populations and the potential of cultivation and large population size in countering the long odds against naturalization. Any implementation of often proposed post-immigration field trials to assess the risk of an alien species becoming naturalized, a requisite step toward invasion, will need to assess different sizes of founder populations and the extent and character of cultivation (intentional or unintentional) that the immigrants might receive.     

种群统计随机性和环境随机性对种群绝灭的影响

影响种群绝灭的随机干扰可分为种群统计随机性、环境随机性和随机灾害三大类。在相对稳定的环境条件下和相对较短的时间内,以前两类随机干扰对种群绝灭的影响为生态学家关注的焦点。但是,由于自然种群动态及其影响因子的复杂特征,进一步深入研究随机干扰对种群绝灭的作用在理论上和实践上都必须发展新的技术手段。本文回顾了种群统计随机性与环境随机性的概念起源与发展,系统阐述了其分析方法。归纳了两类随机性在种群绝灭研究中的应用范围、作用方式和特点的异同和区别方法。各类随机作用与种群动态之间关系的理论研究与对种群绝灭机理的实践研究紧密相关。根据理论模型模拟和自然种群实际分析两方面的研究现状,作者提出了进一步深入研究随机作用与种群非线性动态方法的策略。指出了随机干扰影响种群绝灭过程的研究的方向：更多的研究将从单纯的定性分析随机干扰对种群动力学简单性质的作用,转向结合特定的种群非线性动态特征和各类随机力作用特点具体分析绝灭极端动态的成因,以期做出精确的预测。     

Naturalization of exotic plant species in north‐eastern North America: trends and detection capacity

Aim To reconstruct historical naturalization trends of exotic vascular plants in Quebec and Massachusetts, two regions that experienced fluctuations in their herbarium specimen collection efforts. We hypothesized that the observed trends are linked to the collection effort and that they differ according to the motive for plant introduction. Methods A checklist of naturalized exotic plant taxa for Quebec, including oldest proof of naturalization and motive for introduction, was built from herbaria, journals, dissertations, theses, bulletins, websites and unpublished records. For Massachusetts, we used the checklist of Sorrie (2005; Rhodora, 107 , 284–329). The collection effort was estimated from a pool of species with specimens from the five largest Quebec herbaria and from the Harvard University Herbarium. Results Naturalization trends of exotic plants in Quebec are similar to those observed in Massachusetts. A large part of the plants naturalized in the 17th, 18th or 19th centuries, with very little naturalization during the last 30–90 years. In general, there is a significant statistical link between herbarium specimen collection effort and the discovery of new naturalized plant taxa. Very few utilitarian plants naturalized in either region during the 20th century. In Quebec, a high number of ornamental plants naturalized during the last 100 years. Main conclusions The link between herbarium specimen collection effort and the discovery of new naturalized plants is real, although not as straightforward as it seems. Our analysis suggests that at least part of the decline in the number of new naturalized exotic plants observed in Quebec and Massachusetts during the last 30–90 years is a direct consequence of the low interest for traditional floristic studies. However, the possibility of a real decline cannot be ruled out. We nevertheless provide here one of the first pieces of evidence of the potential consequences of the decline of local plant collection for environmental management and especially for early detection systems of new invaders.     

Demography_Lab,an educational application to evaluate population growth: Unstructured and matrix models

Training in Population Ecology asks for scalable applications capable of embarking students on a trip from basic concepts to the projection of populations under the various effects of density dependence and stochasticity. Demography_Lab is an educational tool for teaching Population Ecology aspiring to cover such a wide range of objectives. The application uses stochastic models to evaluate the future of populations. Demography_Lab may accommodate a wide range of life cycles and can construct models for populations with and without an age or stage structure. Difference equations are used for unstructured populations and matrix models for structured populations. Both types of models operate in discrete time. Models can be very simple, constructed with very limited demographic information or parameter‐rich, with a complex density‐dependence structure and detailed effects of the different sources of stochasticity. Demography_Lab allows for deterministic projections, asymptotic analysis, the extraction of confidence intervals for demographic parameters, and stochastic projections. Stochastic population growth is evaluated using up to three sources of stochasticity: environmental and demographic stochasticity and sampling error in obtaining the projection matrix. The user has full control on the effect of stochasticity on vital rates. The effect of the three sources of stochasticity may be evaluated independently for each vital rate. The user has also full control on density dependence. It may be included as a ceiling population size controlling the number of individuals in the population or it may be evaluated independently for each vital rate. Sensitivity analysis can be done for the asymptotic population growth rate or for the probability of extinction. Elasticity of the probability of extinction may be evaluated in response to changes in vital rates, and in response to changes in the intensity of density dependence and environmental stochasticity.     

Evolutionary rescue under demographic and environmental stochasticity

Populations suffer two types of stochasticity: demographic stochasticity, from sampling error in offspring number, and environmental stochasticity, from temporal variation in the growth rate. By modelling evolution through phenotypic selection following an abrupt environmental change, we investigate how genetic and demographic dynamics, as well as effects on population survival of the genetic variance and of the strength of stabilizing selection, differ under the two types of stochasticity. We show that population survival probability declines sharply with stronger stabilizing selection under demographic stochasticity, but declines more continuously when environmental stochasticity is strengthened. However, the genetic variance that confers the highest population survival probability differs little under demographic and environmental stochasticity. Since the influence of demographic stochasticity is stronger when population size is smaller, a slow initial decline of genetic variance, which allows quicker evolution, is important for population persistence. In contrast, the influence of environmental stochasticity is population-size-independent, so higher initial fitness becomes important for survival under strong environmental stochasticity. The two types of stochasticity interact in a more than multiplicative way in reducing the population survival probability. Our work suggests the importance of explicitly distinguishing and measuring the forms of stochasticity during evolutionary rescue.     

Effects of habitat size and quality on equilibrium density and extinction time of Sorex araneus populations

1. The effects of changes in habitat size and quality on the expected population density and the expected time to extinction of Sorex araneus are studied by means of mathematical models that incorporate demographic stochasticity.
2. Habitat size is characterized by the number of territories, while habitat quality is represented by the expected number of offspring produced during the lifetime of an individual.
3. The expected population density of S. araneus is shown to be mainly influenced by the habitat size. The expected time to extinction of S. araneus populations due to demographic stochasticity, on the other hand, is much more affected by the habitat quality.
4. In a more general setting we demonstrate that, irrespective of the actual species under consideration, the likelihood of extinction as a consequence of demographic stochasticity is more effectively countered by increasing the reproductive success and survival of individuals then by increasing total population size.     

Patterns of plant naturalization show that facultative mycorrhizal plants are more likely to succeed outside their native Eurasian ranges

The naturalization of an introduced species is a key stage during the invasion process. Therefore, identifying the traits that favor the naturalization of non-native species can help understand why some species are more successful when introduced to new regions. The ability and the requirement of a plant species to form a mutualism with mycorrhizal fungi, together with the types of associations formed may play a central role in the naturalization success of different plant species. To test the relationship between plant naturalization success and their mycorrhizal associations we analysed a database composed of mycorrhizal status and type for 1981 species, covering 155 families and 822 genera of plants from Europe and Asia, and matched it with the most comprehensive database of naturalized alien species across the world (GloNAF). In mainland regions, we found that the number of naturalized regions was highest for facultative mycorrhizal, followed by obligate mycorrhizal and lowest for non-mycorrhizal plants, suggesting that the ability of forming mycorrhizas is an advantage for introduced plants. We considered the following mycorrhizal types: arbuscular, ectomycorrhizal, ericoid and orchid mycorrhizal plants. Further, dual mycorrhizal species were those that included observations of arbuscular mycorrhizas as well as observations of ectomycorrhizas. Naturalization success (based on the number of naturalized regions) was highest for arbuscular mycorrhizal and dual mycorrhizal plants, which may be related to the low host specificity of arbuscular mycorrhizal fungi and the consequent high availability of arbuscular mycorrhizal fungal partners. However, these patterns of naturalization success were erased in islands, suggesting that the ability to form mycorrhizas may not be an advantage for establishing self-sustaining populations in isolated regions. Taken together our results show that mycorrhizal status and type play a central role in the naturalization process of introduced plants in many regions, but that their effect is modulated by other factors.     

Extinction of populations by inbreeding depression under stochastic environments

Inbreeding depression may induce rapid extinction due to positive feedbacks between inbreeding depression and reduction of population size, which is often referred to as extinction vortex by inbreeding depression. The present analysis has demonstrated that the extinction vortex is likely to happen with realistic parameter values of genomic mutation rate of lethals or semilethals, equilibrium population size, intrinsic rate of natural increase, and rate of population decline caused by nongenetic extrinsic factors. Simulation models incorporating stochastic fluctuations of population size further indicated that extinction by inbreeding depression is facilitated by environmental fluctuations in population size. The results suggest that there is a positive interaction between genetic stochasticity and environmental stochasticity for extinction of populations by inbreeding depression. Received: May 10, 1999 / Accepted: November 5, 1999     

Naturalization of alien plants in China

Naturalization (the establishment of a self-sustaining population for at least a decade) is a fundamental precondition for plant invasion and so compiling a complete inventory of naturalized alien species is necessary for predicting and hence preventing such invasion. However, nationwide information on naturalized plants in China is still lacking. We compiled a nationwide list of the naturalized plant species of China, based on various literature reports. The list comprised a total of 861 naturalized plant species belonging to 110 families and 465 genera. The three most dominant families were Compositae, Poaceae, and Leguminosae, accounting for 16, 13 and 12% of naturalized plants, respectively. Among genera, Euphorbia and Solanum had the most naturalized species, followed by Ipomoea, Amaranthus, Oenothera, and Trifolium. Over half of all aliens were of American origin (52%), followed by those with European (14%) and Asian (13%) origins. Annuals and perennial herbs were prevalent among naturalized species; comparison to other studies suggests however that the invasive potential is higher among plants with longer life cycles than those of annuals. The taxonomic pattern of plant naturalization in China is similar to patterns worldwide. However, the low proportion of naturalized plants within the Chinese flora overall suggests that the potential for plant invasions in China may be high. Therefore, greater attention should be focused on naturalization of alien plants in China, especially concerning species of dominant families or genera, and those with a perennial life cycle.     

Naturalization of ornamental plant species in public green spaces and private gardens

Ornamental horticulture is the most important pathway for alien plant introductions worldwide, and consequently, invasive spread of introduced plants often begins in urban areas. Although most introduced ornamental garden-plant species are locally not naturalized yet, many of them have shown invasion potential elsewhere in the world, and might naturalize when climate changes. We inventoried the planted flora of 50 public and 61 private gardens in Radolfzell, a small city in southern Germany, to investigate whether local naturalization success of garden plants is associated with their current planting frequency, climatic suitability (as assessed with climatic niche modelling) and known naturalization status somewhere in the world. We identified 954 introduced garden-plant species, of which 48 are already naturalized in Radolfzell and 120 in other parts of Germany. All currently naturalized garden plants in Radolfzell have a climatic suitability probability of ≥ 0.75 and are naturalized in ≥ 13 out of 843 regions globally. These values are significantly higher than those of garden plants that have not become locally naturalized yet. Current planting frequencies, however, were not related to current naturalization success. Using the identified local naturalization thresholds of climatic suitability and global naturalization frequency, and climate projections for the years 2050 and 2070, we identified 45 garden-plant species that are currently not naturalized in Radolfzell but are likely to become so in the future. Although our approach cannot replace a full risk assessment, it is well-suited and applicable as one element of a screening or horizon scanning-type approach.     

Effect of population size on the prospect of species survival

Many recent studies have demonstrated a negative effect of small population size on single plant traits. However, not much is known about the actual consequences of reduced plant performance on the long-term prospect of species survival. I studied the effect of population size on population growth rate and survival probability in the rare perennial herbScorzonera hispanica occurring in fragmented grasslands. Its performance was measured using several traits related to reproduction in 21 populations ranging in size from 3 to 2475 plants. These data were then connected with data on full demography of the species from three of the studied populations. Two different matrix models differing in the number of transitions based on measurements in the populations differing in size were used to explore the relationship between population size and population growth rate. Both matrix models showed that despite the decline in seed production in small populations, population growth rate is never significantly different from one, and the populations could thus be expected to survive in the long run. Calculations of extinction probabilities that take into account demographic and environmental stochasticity, however, showed that populations below 100 flowering individuals have a high probability to become extinct. This demonstrates that demographic and environmental stochasticity is an important driver of the fate of small populations in this system. This study demonstrates that estimation of population growth rate can provide new insights into the effect of population size on population growth and survival. It also shows how matrix models enable the combination various pieces of information about the single populations into one overall measure, and may provide a useful tool for the standardization of studies on the effects of population size on population performance.     

Demographic factors and genetic variation influence population persistence under environmental change

Population persistence has been studied in a conservation context to predict the fate of small or declining populations. Persistence models have explored effects on extinction of random demographic and environmental fluctuations, but in the face of directional environmental change they should also integrate factors affecting whether a population can adapt. Here, we examine the population‐size dependence of demographic and genetic factors and their likely contributions to extinction time under scenarios of environmental change. Parameter estimates were derived from experimental populations of the rainforest species, Drosophila birchii, held in the lab for 10 generations at census sizes of 20, 100 and 1000, and later exposed to five generations of heat‐knockdown selection. Under a model of directional change in the thermal environment, rapid extinction of populations of size 20 was caused by a combination of low growth rate (r) and high stochasticity in r. Populations of 100 had significantly higher reproductive output, lower stochasticity in r and more additive genetic variance (V_A) than populations of 20, but they were predicted to persist less well than the largest size class. Even populations of 1000 persisted only a few hundred generations under realistic estimates of environmental change because of low V_A for heat‐knockdown resistance. The experimental results document population‐size dependence of demographic and adaptability factors. The simulations illustrate a threshold influence of demographic factors on population persistence, while genetic variance has a more elastic impact on persistence under environmental change.     

The extended Moran effect and large-scale synchronous fluctuations in the size of great tit and blue tit populations

1. Synchronous fluctuations of geographically separated populations are in general explained by the Moran effect, i.e. a common influence on the local population dynamics of environmental variables that are correlated in space. Empirical support for such a Moran effect has been difficult to provide, mainly due to problems separating out effects of local population dynamics, demographic stochasticity and dispersal that also influence the spatial scaling of population processes. Here we generalize the Moran effect by decomposing the spatial autocorrelation function for fluctuations in the size of great tit Parus major and blue tit Cyanistes caeruleus populations into components due to spatial correlations in the environmental noise, local differences in the strength of density regulation and the effects of demographic stochasticity. 2. Differences between localities in the strength of density dependence and nonlinearity in the density regulation had a small effect on population synchrony, whereas demographic stochasticity reduced the effects of the spatial correlation in environmental noise on the spatial correlations in population size by 21.7% and 23.3% in the great tit and blue tit, respectively. 3. Different environmental variables, such as beech mast and climate, induce a common environmental forcing on the dynamics of central European great and blue tit populations. This generates synchronous fluctuations in the size of populations located several hundred kilometres apart. 4. Although these environmental variables were autocorrelated over large areas, their contribution to the spatial synchrony in the population fluctuations differed, dependent on the spatial scaling of their effects on the local population dynamics. We also demonstrate that this effect can lead to the paradoxical result that a common environmental variable can induce spatial desynchronization of the population fluctuations. 5. This demonstrates that a proper understanding of the ecological consequences of environmental changes, especially those that occur simultaneously over large areas, will require information about the spatial scaling of their effects on local population dynamics.     

Naturalization of introduced plants: ecological drivers of biogeographical patterns

CONTENTS: Summary 383 I. Introduction 383 II. The introduction-naturalization-invasion continuum for conceptualizing biological invasions 384 III. The biogeographical background for studying naturalization: variation among populations and regions 385 IV. Factors determining naturalization in plants 388 Acknowledgements 392 References 392 SUMMARY: The literature on biological invasions is biased in favour of invasive species - those that spread and often reach high abundance following introduction by humans. It is, however, also important to understand previous stages in the introduction-naturalization-invasion continuum ('the continuum'), especially the factors that mediate naturalization. The emphasis on invasiveness is partly because most invasions are only recognized once species occupy large adventive ranges or start to spread. Also, many studies lump all alien species, and fail to separate introduced, naturalized and invasive populations and species. These biases impede our ability to elucidate the full suite of drivers of invasion and to predict invasion dynamics, because different factors mediate progression along different sections of the continuum. A better understanding of the determinants of naturalization is important because all naturalized species are potential invaders. Processes leading to naturalization act differently in different regions and global biogeographical patterns of plant invasions result from the interaction of population-biological, macroecological and human-induced factors. We explore what is known about how determinants of naturalization in plants interact at various scales, and how their importance varies along the continuum. Research that is explicitly linked to particular stages of the continuum can generate new information that is appropriate for improving the management of biological invasions if, for example, potentially invasive species are identified before they exert an impact.     

EVOLUTION AND EXTINCTION IN A CHANGING ENVIRONMENT: A QUANTITATIVE-GENETIC ANALYSIS

Because of the ubiquity of genetic variation for quantitative traits, virtually all populations have some capacity to respond evolutionarily to selective challenges. However, natural selection imposes demographic costs on a population, and if these costs are sufficiently large, the likelihood of extinction will be high. We consider how the mean time to extinction depends on selective pressures (rate and stochasticity of environmental change, and strength of selection), population parameters (carrying capacity, and reproductive capacity), and genetics (rate of polygenic mutation). We assume that in a randomly mating, finite population subject to density-dependent population growth, individual fitness is determined by a single quantitative-genetic character under Gaussian stabilizing selection with the optimum phenotype exhibiting directional change, or random fluctuations, or both. The quantitative trait is determined by a finite number of freely recombining, mutationally equivalent, additive loci. The dynamics of evolution and extinction are investigated, assuming that the population is initially under mutation-selection-drift balance. Under this model, in a directionally changing environment, the mean phenotype lags behind the optimum, but on the average evolves parallel to it. The magnitude of the lag determines the vulnerability to extinction. In finite populations, stochastic variation in the genetic variance can be quite pronounced, and bottlenecks in the genetic variance temporarily can impair the population's adaptive capacity enough to cause extinction when it would otherwise be unlikely in an effectively infinite population. We find that maximum sustainable rates of evolution or, equivalently, critical rates of environmental change, may be considerably less than 10% of a phenotypic standard deviation per generation.     

Estimating population trends using population viability analyses for the conservation ofCapra pyrenaica

Large herbivore populations can suffer important oscillations with considerable effects on ecosystem functions and services, yet our capacity to predict population fate is limited and conditional upon the availability of data. This study investigated the interannual variation in the growth rate of populations ofCapra pyrenaica Schinz, 1838, and its extinction risk by comparing the dynamics of populations that were stable for more than two decades (Gredos and Tortosa-Beceite), populations that had increased recently (Tejeda-Almijara), and populations that were in decline (Cazorla-Segura) or extinct (the Pyrenees population; hereafter, bucardo). To estimate quasi-extinction threshold assessments (50% of population extinct in this study), which have implications for the conservation of the species, we used empirical data and the predictions derived from several theoretical models. The results indicate that when variance of log population growth rate reaches a specific threshold, the probability of quasi-extinction increased drastically. ForC. pyrenaica, we recommend keeping population variance < 0.05, which will reduce the likelihood that the irruptive oscillations caused by environmental and demographic stochasticity will put the population at risk. Models to predict the dynamics ofC. pyrenaica populations should incorporate temporal stochasticity because, in this study, it strongly increased the likelihood that a population declined.     

Ethnicity and naturalization

This article examines whether or not ethnicity has an independent effect on the likelihood of immigrant naturalization using the Public Use Microdata Sample [PUMS] data from the 1980 US Census. Ethnic differences in the propensity to become naturalized US citizens were analysed among four panethnic groups and across thirty‐three major ethnic groups. The results point to the continuing significance of ethnicity in the naturalization process. However, the effect of ethnicity is not as strong as the effects of other structural factors. Three hypotheses that attempt to explain ethnic differences in the propensity for naturalization were also tested. The evidence lends strong support to the forced self‐protection hypothesis, but it provides no support for the discrimination hypothesis and the cultural differences hypothesis.     

A method for validating stochastic models of population viability: a case study of the mountain pygmy-possum (Burramys parvus)

1.  A method of validating stochastic models of population viability is proposed, based on assessing the mean and variance of the predicted population size.
2.  The method is illustrated with a model of the population dynamics of the mountain pygmy-possum ( Burramys parvus Broom 1895), based on annual census data collected from a single population in the Snowy Mountains of New South Wales, Australia between 1986 and 1997. The model incorporates density-dependence in survivorship and recruitment, and demographic and environmental stochasticity.
3.  The model appeared to make reasonable predictions for the three populations that were used for validation, provided the equilibrium population size was estimated accurately. This may require that differences in habitat quality between populations be taken into account.
4.  Following validation, the model was given new parameters using the additional data from the three populations, and the risk of population decline within the next 100 years was assessed. Although populations as small as 15 females are predicted to be relatively safe from extinction caused by stochastic processes, B. parvus appears vulnerable to loss of habitat and reductions in the population growth rate.
5.  The approach used in this paper is one of few attempts to validate a model of population viability using field data, and demonstrates that some aspects of stochastic population models can be tested.     

Effective size of a fluctuating age-structured population

Previous theories on the effective size of age-structured populations assumed a constant environment and, usually, a constant population size and age structure. We derive formulas for the variance effective size of populations subject to fluctuations in age structure and total population size produced by a combination of demographic and environmental stochasticity. Haploid and monoecious or dioecious diploid populations are analyzed. Recent results from stochastic demography are employed to derive a two-dimensional diffusion approximation for the joint dynamics of the total population size, N, and the frequency of a selectively neutral allele, p. The infinitesimal variance for p, multiplied by the generation time, yields an expression for the effective population size per generation. This depends on the current value of N, the generation time, demographic stochasticity, and genetic stochasticity due to Mendelian segregation, but is independent of environmental stochasticity. A formula for the effective population size over longer time intervals incorporates deterministic growth and environmental stochasticity to account for changes in N.     

Phenotypic plasticity and population viability: the importance of environmental predictability

Phenotypic plasticity plays a key role in modulating how environmental variation influences population dynamics, but we have only rudimentary understanding of how plasticity interacts with the magnitude and predictability of environmental variation to affect population dynamics and persistence. We developed a stochastic individual-based model, in which phenotypes could respond to a temporally fluctuating environmental cue and fitness depended on the match between the phenotype and a randomly fluctuating trait optimum, to assess the absolute fitness and population dynamic consequences of plasticity under different levels of environmental stochasticity and cue reliability. When cue and optimum were tightly correlated, plasticity buffered absolute fitness from environmental variability, and population size remained high and relatively invariant. In contrast, when this correlation weakened and environmental variability was high, strong plasticity reduced population size, and populations with excessively strong plasticity had substantially greater extinction probability. Given that environments might become more variable and unpredictable in the future owing to anthropogenic influences, reaction norms that evolved under historic selective regimes could imperil populations in novel or changing environmental contexts. We suggest that demographic models (e.g. population viability analyses) would benefit from a more explicit consideration of how phenotypic plasticity influences population responses to environmental change.