Controlling species invasions is a leading problem for applied ecology. While controlling populations expanding linearly or exponentially is straightforward, intervention in systems with complex dynamics can have complicated, and sometimes counterintuitive, consequences. Most invasive plant populations are stage-structured and density-dependent--a recipe for complex dynamics--and yet few population models have been created to explore the effects of control efforts on such species. We examined the demography of the invasive biennial plant Alliaria petiolata (garlic mustard) on the front of its spread into a natural area and found evidence of strong density dependence in vital rates of first-year rosette and second-year adult stage classes. We parameterized a density-dependent, stage-structured projection model using field-collected data. This model produces two-point cycles with alternating years in which adults vs. rosettes are more prevalent. Such population dynamics match observations in natural populations, suggesting that these complicated population dynamics may result from deterministic rules. We used this model to evaluate simulated management strategies, including herbicide treatment of rosettes and clipping or pulling of adult plants. Management of A. petiolata by inducing mortality of either rosettes or adults will not be effective at reducing population density unless the induced mortality is very high (>95% for rosettes and >85% for adults) and repeated every year. Indeed, induced mortality of rosettes can be counterproductive, causing increases in the stationary distribution of A. petiolata density. This species is typical of many invasive plants (stage-structured, short-lived, high fertility) and exhibits common forms of density dependence. Thus, the management implications of our study should apply broadly to other species with similar life histories. We suggest that management should focus on managing adults rather than rosettes, and on creating efficient control in targeted areas of the population, rather than spreading less efficient efforts widely.
Invasive plants may compete with native plants by increasing the pressure of native consumers, a mechanism known as "apparent competition." Apparent competition can be as strong as or stronger than direct competition, but the role of apparent competition has rarely been examined in biological invasions. We used four years of demographic data and seed-removal experiments to determine if introduced grasses caused elevated levels of seed consumption on native plant species in a coastal dune system in California, USA. We show that the endangered, coastal dune plant Lupinus tidestromii experiences high levels of pre-dispersal seed consumption by the native rodent Peromyscus maniculatus due to its proximity to the invasive grass, Ammophila arenaria. We use stage-structured, stochastic population models to project that two of three study populations will decline toward extinction under ambient levels of consumption. For one of these declining populations, a relatively small decrease in consumption pressure should allow for persistence. We show that apparent competition with an invasive species significantly decreases the population growth rate and persistence of a native species. We expect that apparent competition is an important mechanism in other ecosystems because invasive plants often change habitat structure and plant-consumer interactions. Possible implications of the apparent-competition mechanism include selective extinction of species preferred by seed consumers in the presence of an invasive species and biological homogenization of communities toward non-preferred native plant species.
An understanding of the demographic processes contributing to invasions would improve our mechanistic understanding of the invasion process and improve the efficiency of prevention and control efforts. However, field comparisons of the demography of invasive and noninvasive species have not previously been conducted. We compared the in situ demography of 17 introduced plant species in St. Louis, Missouri, USA, to contrast the demographic patterns of invasive species with their less invasive relatives across a broad sample of angiosperms. Using herbarium records to estimate spread rates, we found higher maximum spread rates in the landscape for species classified a priori as invasive than for noninvasive introduced species, suggesting that expert classifications are an accurate reflection of invasion rate. Across 17 species, projected population growth was not significantly greater in invasive than in noninvasive introduced species. Among five taxonomic pairs of close relatives, however, four of the invasive species had higher projected population growth rates compared with their noninvasive relative. A Life Table Response Experiment suggested that the greater projected population growth rate of some invasive species relative to their noninvasive relatives was primarily a result of sexual reproduction. The greater sexual reproduction of invasive species is consistent with invaders having a life history strategy more reliant on fecundity than survival and is consistent with a large role of propagule pressure in invasion. Sexual reproduction is a key demographic correlate of invasiveness, suggesting that local processes influencing sexual reproduction, such as enemy escape, might be of general importance. However, the weak correlation of projected population growth with spread rates in the landscape suggests that regional processes, such as dispersal, may be equally important in determining invasion rate.
Comparative analyses of spatial genetic structure (SGS) among species, populations, or cohorts give insight into the genetic consequences of seed dispersal in plants. We analysed SGS of a weedy tree in populations with known and unknown recruitment histories to first establish patterns in populations with single vs. multiple founders, and then to infer possible recruitment scenarios in populations with unknown histories. We analysed SGS in six populations of the colonizing tree Albizia julibrissin Durazz. (Fabaceae) in Athens, Georgia. Study sites included two large populations with multiple, known founders, two small populations with a single, known founder, and two large populations with unknown recruitment histories. Eleven allozyme loci were used to genotype 1385 individuals. Insights about the effects of colonization history from the SGS analyses were obtained from correlograms and Sp statistics. Distinct differences in patterns of SGS were identified between populations with multiple founders vs. a single founder. We observed significant, positive SGS, which decayed with increasing distance in the populations with multiple colonists, but little to no SGS in populations founded by one colonist. Because relatedness among individuals is estimated relative to a local reference population, which usually consists of those individuals sampled in the study population, SGS in populations with high background relatedness, such as those with a single founder, may be obscured. We performed additional analyses using a regional reference population and, in populations with a single founder, detected significant, positive SGS at all distances, indicating that these populations consist of highly related descendants and receive little seed immigration. Subsequent analyses of SGS in size cohorts in the four large study populations showed significant SGS in both juveniles and adults, probably because of a relative lack of intraspecific demographic thinning. SGS in populations of this colonizing tree is pronounced and persistent and is determined by the number and relatedness of founding individuals and adjacent seed sources. Patterns of SGS in populations with known histories may be used to indirectly infer possible colonization scenarios for populations where it is unknown.
Many species are adapted to disturbance and occur within dynamic, mosaic landscapes that contain early and late successional microhabitats. Human modification of disturbance regimes alters the availability of microhabitats and may affect the viability of species in these ecosystems. Because restoring historical disturbance regimes is typically expensive and requires action at large spatial scales, such restoration projects must be justified by linking the persistence of species with successional microhabitats. Coastal sand dune ecosystems worldwide are characterized by their endemic biodiversity and frequent disturbance. Dune-stabilizing invasive plants alter successional dynamics and may threaten species in these ecosystems. We examined the distribution and population dynamics of two federally endangered plant species, the annual Layia carnosa and the perennial Lupinus tidestromii, within a dune ecosystem in northern California, USA. We parameterized a matrix population model for L. tidestromii and examined the magnitude by which the successional stage of the habitat (early or late) influenced population dynamics. Both species had higher frequencies and L. tidestromii had higher frequency of seedlings in early successional habitats. Lupinus tidestromii plants in early successional microhabitats had higher projected rates of population growth than those associated with stabilized, late successional habitats, due primarily to higher rates of recruitment in early successional microhabitats. These results support the idea that restoration of disturbance is critical in historically dynamic landscapes. Our results suggest that large-scale restorations are necessary to allow persistence of the endemic plant species that characterize these ecosystems.
Abstract. The population effects of harvest depend on complex interactions between density dependence, seasonality, stage structure, and management timing. Here we present a periodic nonlinear matrix population model that incorporates seasonal density dependence with stage-selective and seasonally selective harvest. To this model, we apply newly developed perturbation analyses to determine how population densities respond to changes in harvest and demographic parameters. We use the model to examine the effects of popular control strategies and demographic perturbations on the invasive weed garlic mustard (Alliaria petiolata). We find that seasonality is a major factor in harvest outcomes, because population dynamics may depend significantly on both the season of management and the season of observation. Strategies that reduce densities in one season can drive increases in another, with strategies giving positive sensitivities of density in the target seasons leading to compensatory effects that invasive species managers should avoid. Conversely, demographic parameters to which density is very elastic (e.g., seeding survival, second-year rosette spring survival, and the flowering to fruiting adult transition for maximum summer densities) may indicate promising management targets.
Studies of invasive species often demonstrate that exotic or invasive plants have reduced levels of predation when first introduced into novel geographic areas. In some systems, native predators may subsequently be introduced and the plant-predator association is re-established. In this study, we analyzed a recent reassociation between the introduced tree, Albizia julibrissin, and its native seed predator, Bruchidius terrenus, which has also been introduced, in the vicinity of Athens, GA, USA. We documented a rapid increase in the levels of bruchid beetle predation on seeds of A. julibrissin from 2001 to 2002 and substantial predation in the following 5 years. From a six-year survey, we found that relative susceptibility to beetle predation among individual trees remained consistent over time. Furthermore, we found that phenology had significant effects on beetle predation on fruits and seeds, with fruits developed from early flowers having higher levels of beetle attacks than ones developed from later flowers in the season. Finally, seeds and fruits attacked by beetles were more likely to be attacked by a Fusarium fungus, probably due to coinfestation of the two predators. Host-predator reassociation that causes substantial loss of fertility or shows a relationship with reproductive phenology may have population level demographic effects or evolutionary consequences and deserve further research attention.
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