The effects of demographic stochasticity and parameter uncertainty on predicting the establishment of introduced species

Invasive species are a serious threat to biodiversity worldwide and predicting whether an introduced species will first establish and then become invasive can be useful to preserve ecosystem services. Establishment is influenced by multiple factors, such as the interactions between the introduced individuals and the resident community, and demographic and environmental stochasticity. Field observations are often incomplete or biased. This, together with an imperfect knowledge of the ecological traits of the introduced species, makes the prediction of establishment challenging. Methods that consider the combined effects of these factors on our ability to predict the establishment of an introduced species are currently lacking. We develop an inference framework to assess the combined effects of demographic stochasticity and parameter uncertainty on our ability to predict the probability of establishment following the introduction of a small number of individuals. We find that even moderate levels of demographic stochasticity influence both the probability of establishment, and, crucially, our ability to correctly predict that probability. We also find that estimation of the demographic parameters of an introduced species is fundamental to obtain precise estimates of the interaction parameters. For typical values of demographic stochasticity, the drop in our ability to predict an establishment can be 30% when having priors on the demographic parameters compared to having their accurate values. The results from our study illustrate how demographic stochasticity may bias the prediction of the probability of establishment. Our method can be applied to estimate probability of establishment of introduced species in field scenarios, where time series data and prior information on the demographic traits of the introduced species are available.     

Reciprocally beneficial interactions between introduced plants and ants are induced by the presence of a third introduced species

Interspecific interactions play an important role in the success of introduced species. For example, the ‘enemy release’ hypothesis posits that introduced species become invasive because they escape top–down regulation by natural enemies while the ‘invasional meltdown’ hypothesis posits that invasions may be facilitated by synergistic interactions between introduced species. Here, we explore how facilitation and enemy release interact to moderate the potential effect of a large category of positive interactions – protection mutualisms. We use the interactions between an introduced plant (Japanese knotweed Fallopia japonica), an introduced herbivore (Japanese beetle Popillia japonica), an introduced ant (European red ant Myrmica rubra), and native ants and herbivores in riparian zones of the northeastern United States as a model system. Japanese knotweed produces sugary extrafloral nectar that is attractive to ants, and we show that both sugar reward production and ant attendance increase when plants experience a level of leaf damage that is typical in the plants’ native range. Using manipulative experiments at six sites, we demonstrate low levels of ant patrolling, little effect of ants on herbivory rates, and low herbivore pressure during midsummer. Herbivory rates and the capacity of ants to protect plants (as evidenced by effects of ant exclusion) increased significantly when plants were exposed to introduced Japanese beetles that attack plants in the late summer. Beetles were also associated with greater on‐plant foraging by ants, and among‐plant differences in ant‐foraging were correlated with the magnitude of damage inflicted on plants by the beetles. Last, we found that sites occupied by introduced M. rubra ants almost invariably included Japanese knotweed. Thus, underlying variation in the spatiotemporal distribution of the introduced herbivore influences the provision of benefits to the introduced plant and to the introduced ant. More specifically, the presence of the introduced herbivore converts an otherwise weak interaction between two introduced species into a reciprocally beneficial mutualism. Because the prospects for facilitation are linked to the prospects for enemy release in protection mutualisms, species introductions can have complex effects on existing species interactions, between both native and introduced species.     

Domestic exotics and the perception of invasibility

Susceptibility of an area to invasion by exotic species is often judged by the fraction of introduced species in the local biota. However, the degree of invasion, particularly in mainland areas, has often been underestimated because of the exclusion of ‘domestic exotics’ (those introduced to internal units from within the national border) in calculations. Because all introduced species on islands are considered as exotics, this contributes to the perception that islands are more susceptible to invasion than are continental regions. Here, we determine the contribution of domestic exotic species to the degree of invasion (exotic fraction) in mainland areas. We quantify the relationships of exotic fraction to the area, human population density and land use within each of the 48 conterminous US states to identify mechanisms that potentially influence the degree of invasion. For each of the 48 conterminous US states, we compiled the number of species introduced from outside the United States (‘foreign exotics’) and the number of exotics introduced from other conterminous US states (‘domestic exotics’). The status of each species as foreign or domestic was determined for each state by researching its precise origins through vouchered herbarium records, supplemented by literature ( Kartesz, 2010 ). We found that (1) the exotic fraction inevitably decreases with increasing area as the pool of potential exotic species decreases; (2) exotic richness of areas within large mainland regions is underestimated to the extent that species introduced among areas within a region are considered as natives; and (3) human activities contribute disproportionately more exotics to smaller than to larger administrative areas. How we define ‘exotic’ influences how we count non‐native species and perceive invasibility. Excluding domestic exotics in mainland regions leads to a biased perception of increased invasibility on islands, where all introduced species are considered exotic. Thus, future documentation and interpretation of invasion patterns and management of exotics should account for these biases in quantifying the exotic fraction.     

Rapid synthesis and introduction of a protected EDTA-like group during the solid-phase assembly of peptides.

A simple two-step procedure is reported for the synthesis of a tert-butyl ester protected form of an EDTA-like bifunctional chelating agent. This reagent can be easily introduced on any available amino group during the assembly of peptides on solid-phase supports. Using the model tetradecapeptide OVA(323-336), we have introduced an EDTA group at the N-terminus of this T-cell epitope and confirmed that the EDTA group is present on the molecule, can chelate metals, and does not affect the biological activity of the peptide.     

Island biogeography and the species richness of introduced mammals on New Zealand offshore islands

Aim To investigate and establish the significance of various island biogeographic relationships (geographical, ecological and anthropological) with the species richness of introduced mammals on offshore islands. Location The 297 offshore islands of the New Zealand archipelago (latitude: 34–47°S; longitude: 166–179°E). Methods Data on New Zealand offshore islands and the introduced mammals on them were collated from published surveys and maps. The species richness of small and large introduced mammals were calculated for islands with complete censuses and regressed on island characteristics using a Poisson distributed error generalized linear model. To estimate the ‘z‐value’ for introduced mammals on New Zealand islands, least‐squares regression was used [log₁₀ S vs. log₁₀ A]. Results High collinearity was found between the area, habitat diversity and elevation of islands. The island characteristics related to the species richness of introduced mammals differed predictably between large and small mammals. The species richness of introduced large mammals was mostly related to human activities on islands, whereas species richness of introduced small mammals was mostly related to island biogeographical parameters. The ‘z‐value’ for total species richness is found to be expectedly low for introduced mammals. Main conclusions Distance appears to have become ecologically trivial as a filter for introduced mammal presence on New Zealand offshore islands. There is strong evidence of a ‘small island’ effect on New Zealand offshore islands. The species richness of both small and large introduced mammals on these islands appears to be most predominantly related to human use, although there is some evidence of natural dispersal for smaller species. The ecological complexity of some islands appears to make them less invasible to introduced mammals. Some human activities have an interactive effect on species richness. A small number of islands have outlying species richness values above what the models predict, suggesting that the presence of some species may be related to events not accounted for in the models.     

Patterns of introduction of non-indigenous non-marine snails and slugs in the Hawaiian Islands

The native snails of the Hawaiian Islands are disappearing. One cause is predation by introduced carnivorous snails. Habitat destruction/modification is also important, facilitating the spread of other non-indigenous snails and slugs. Eighty-one species of snails and slugs are recorded as having been introduced. Thirty-three are established: 12 freshwater, 21 terrestrial. Two or three species arrived before western discovery of the islands (1778). During the nineteenth century about one species per decade, on average, was introduced. The rate rose to about four per decade during the twentieth century, with the exception of an especially large number introduced in the 1950s as putative biocontrol agents against the giant African snail, Achatina fulica. The geographical origins of these introductions reflect changing patterns of commerce and travel. Early arrivals were generally Pacific or Pacific rim species. Increasing trade and tourism with the USA, following its annexation of Hawaii, led to an increasing proportion of American species. More general facilitation of travel and commerce later in the twentieth century led to a significant number of European species being introduced. African species dominated the 1950s biological control introductions. The process continues and is just part of the homogenization of the unique faunas of tropical Pacific islands.     

Impact of the microscale distribution of a Pseudomonas strain introduced into soil on potential contacts with indigenous bacteria

Soil bioaugmentation is a promising approach in soil bioremediation and agriculture. Nevertheless, our knowledge of the fate and activity of introduced bacteria in soil and thus of their impact on the soil environment is still limited. The microscale spatial distribution of introduced bacteria has rarely been studied, although it determines the encounter probability between introduced cells and any components of the soil ecosystem and thus plays a role in the ecology of introduced bacteria. For example, conjugal gene transfer from introduced bacteria to indigenous bacteria requires cell-to-cell contact, the probability of which depends on their spatial distribution. To quantitatively characterize the microscale distribution of an introduced bacterial population and its dynamics, a gfp-tagged derivative of Pseudomonas putida KT2440 was introduced by percolation in repacked soil columns. Initially, the introduced population was less widely spread at the microscale level than two model indigenous functional communities: the 2,4-dichlorophenoxyacetic acid degraders and the nitrifiers (each at 10(6) CFU g(-1) soil). When the soil was percolated with a substrate metabolizable by P. putida or incubated for 1 month, the microscale distribution of introduced bacteria was modified towards a more widely dispersed distribution. The quantitative data indicate that the microscale spatial distribution of an introduced strain may strongly limit its contacts with the members of an indigenous bacterial community. This could constitute an explanation to the low number of indigenous transconjugants found most of time when a plasmid-donor strain is introduced into soil.     

Human races and the problem of variability--a recommendation for the solution by Franz Weidenreich

In parts of anthropology race concepts were playing a central role for evolutionary reconstruction as a means for assessment and study of human variability. This method was criticised already quite early. In this paper the arguments of one of those early critics, Franz Weidenreich, are introduced and reconstructed on a methodological basis. While avoiding race concepts, Franz Weidenreich developed other methods for reconstructing lines of evolutionary development. He suggested to use an extended concept of physical constitution. This concept is also introduced and studied with reference to its methodological foundations.     

Impact of the Microscale Distribution of a Pseudomonas Strain Introduced into Soil on Potential Contacts with Indigenous Bacteria

Soil bioaugmentation is a promising approach in soil bioremediation and agriculture. Nevertheless, our knowledge of the fate and activity of introduced bacteria in soil and thus of their impact on the soil environment is still limited. The microscale spatial distribution of introduced bacteria has rarely been studied, although it determines the encounter probability between introduced cells and any components of the soil ecosystem and thus plays a role in the ecology of introduced bacteria. For example, conjugal gene transfer from introduced bacteria to indigenous bacteria requires cell-to-cell contact, the probability of which depends on their spatial distribution. To quantitatively characterize the microscale distribution of an introduced bacterial population and its dynamics, a gfp-tagged derivative of Pseudomonas putida KT2440 was introduced by percolation in repacked soil columns. Initially, the introduced population was less widely spread at the microscale level than two model indigenous functional communities: the 2,4-dichlorophenoxyacetic acid degraders and the nitrifiers (each at 10⁶ CFU g⁻¹ soil). When the soil was percolated with a substrate metabolizable by P. putida or incubated for 1 month, the microscale distribution of introduced bacteria was modified towards a more widely dispersed distribution. The quantitative data indicate that the microscale spatial distribution of an introduced strain may strongly limit its contacts with the members of an indigenous bacterial community. This could constitute an explanation to the low number of indigenous transconjugants found most of time when a plasmid-donor strain is introduced into soil.     

New Record and Biology of the Molly Poecilia sphenops (Poeciliidae), Discovered in the Northern Sahara of Algeria

Journal of Ichthyology - This study reports data on the first observation and biology of the Molly Poecilia sphenops, an exotic fish species recently introduced into two natural and artificial...     

Impacts of introduced species on the biota of an oceanic archipelago: the relative importance of competitive and trophic interactions

Introduced species negatively impact native species through competitive and trophic interactions, particularly on oceanic islands that have never been connected to any continental landmass. However, there are few studies on the relative importance of competitive interactions (resource competition with introduced species) and trophic interactions (predation or herbivory by introduced species) with respect to the negative impacts on native organisms on oceanic islands. A literature review on introduced and native species of the oceanic Ogasawara (Bonin) Islands in the western Pacific Ocean indicated that many native species (e.g., bees, beetles, damselflies, butterflies, land snails, birds, and plants) have been negatively impacted by introduced predators and herbivores (e.g., lizards, rats, flatworms, and goats). Several native plants and bees have been negatively affected by introduced competitors. However, the native species that have competed with introduced species have also suffered from either intense herbivory or predation by other introduced species. Thus, introduced predators and herbivores have had greater impacts on native species than introduced competitors in the Ogasawara Islands.     

Invasion trajectory of alien trees: the role of introduction pathway and planting history

Global change is driving a massive rearrangement of the world's biota. Trajectories of distributional shifts are shaped by species traits, the recipient environment and driving forces with many of the driving forces directly due to human activities. The relative importance of each in determining the distributions of introduced species is poorly understood. We consider 11 Australian Acacia species introduced to South Africa for different reasons (commercial forestry, dune stabilization and ornamentation) to determine how features of the introduction pathway have shaped their invasion history. Projections from species distribution models (SDMs) were developed to assess how the reason for introduction influences the similarity between climatic envelopes in native and alien ranges. A lattice model for an idealized invasion was developed to assess the relative contribution of intrinsic traits and introduction dynamics on the abundance and extent over the course of simulated invasions. SDMs show that alien populations of ornamental species in South Africa occupy substantially different climate space from their native ranges, whereas species introduced for forestry occupy a similar climate space in native and introduced ranges. This may partly explain the slow spread rates observed for some alien ornamental plants. Such mismatches are likely to become less pronounced with the current drive towards ‘eco gardens’ resulting in more introductions of ornamental species with a close climate match between native and newly introduced regions. The results from the lattice model showed that the conditions associated with the introduction pathway (especially introduction pressure) dominate early invasion dynamics. The placement of introduction foci in urban areas limited the extent and abundance of invasive populations. Features of introduction events appear to initially mask the influence of intrinsic species traits on invasions and help to explain the relative success of species introduced for different purposes. Introduction dynamics therefore can have long‐lasting influences on the outcomes of species redistributions, and must be explicitly considered in management plans.     

The paradox of the parasites: implications for biological invasion

The enemy-release hypothesis for biological invasions supposes that invasive species may be more successful in their introduced ranges than in their native ranges owing to the absence of coevolved natural enemies. Recent studies supporting this hypothesis have found that introduced plants and animals are less parasitized in their introduced ranges than in their native ranges. Expanding on this theory, I hypothesize that the role of enemy release may differ among the introduction, establishment and spread phases of an invasion. I present a simple model indicating that parasite release is unlikely to greatly affect the chance of establishment in populations with and without an immune subpopulation. The specific numerical relationship between the number of individuals introduced and the chance of establishment depends on a relationship between virulence, here conceptualized as the chance for the extinction of a lineage, and the fraction of the population infected at introduction. These results support the idea of a 'filter effect' in which different biological processes regulate the different phases of an invasion.     

Population Genetic Consequences of the Allee Effect and the Role of Offspring-Number Variation

A strong demographic Allee effect in which the expected population growth rate is negative below a certain critical population size can cause high extinction probabilities in small introduced populations. But many species are repeatedly introduced to the same location and eventually one population may overcome the Allee effect by chance. With the help of stochastic models, we investigate how much genetic diversity such successful populations harbor on average and how this depends on offspring-number variation, an important source of stochastic variability in population size. We find that with increasing variability, the Allee effect increasingly promotes genetic diversity in successful populations. Successful Allee-effect populations with highly variable population dynamics escape rapidly from the region of small population sizes and do not linger around the critical population size. Therefore, they are exposed to relatively little genetic drift. It is also conceivable, however, that an Allee effect itself leads to an increase in offspring-number variation. In this case, successful populations with an Allee effect can exhibit less genetic diversity despite growing faster at small population sizes. Unlike in many classical population genetics models, the role of offspring-number variation for the population genetic consequences of the Allee effect cannot be accounted for by an effective-population-size correction. Thus, our results highlight the importance of detailed biological knowledge, in this case on the probability distribution of family sizes, when predicting the evolutionary potential of newly founded populations or when using genetic data to reconstruct their demographic history.     

Metabolic activity of Flavobacterium strain P25 during starvation and after introduction into bulk soil and the rhizosphere of wheat

Abstract: The physiological state of introduced Flavobacterium strain P25 cells was determined in starvation cultures, in bulk soil, and in the rhizosphere of wheat using direct viable counts (DVC; based on cell elongation after use of nalidixic acid and substrate addition, resulting in a potential activity measurement) and the redox dye 5-cyano-2, 3-ditolyl tetrazolium chloride (CTC; based on respiration without substrate additions, resulting in an in situ activity measurement). Both methods clearly demonstrated that the metabolic activity of Flavobacterium P25 cells decreased during starvation, followed by increased activity after amendment with substrate. This confirmed the applicability of DVC and CTC methods to Flavobacterium P25. Both DVC and CTC methods showed that the percentage of active cells in an introduced Flavobacterium P25 population in rhizosphere soil was lower than that in bulk soil in the first 1–2 weeks after planting wheat seedlings. After two weeks, the percentage of metabolically active cells in the P25 population in rhizosphere soil was higher than in bulk soil. Since different aspects of cellular physiology are measured when applying DVC and CTC, the impact of variations in environmental factors on the metabolic state of introduced strains may be monitored closely by these methods.     

Contrasting patterns of genome‐wide polymorphism in the native and invasive range of the marine mollusc Crepidula fornicata

Selection processes are believed to be an important evolutionary driver behind the successful establishment of nonindigenous species, for instance through adaptation for invasiveness (e.g. dispersal mechanisms and reproductive allocation). However, evidence supporting this assumption is still scarce. Genome scans have often identified loci with atypical patterns of genetic differentiation (i.e. outliers) indicative of selection processes. Using microsatellite‐ and AFLP‐based genome scans, we looked for evidence of selection following the introduction of the mollusc Crepidula fornicata. Native to the northwestern Atlantic, this gastropod has become an emblematic invader since its introduction during the 19th and 20th centuries in the northeastern Atlantic and northeastern Pacific. We examined 683 individuals from seven native and 15 introduced populations spanning the latitudinal introduction and native ranges of the species. Our results confirmed the previously documented high genetic diversity in native and introduced populations with little genetic structure between the two ranges, a pattern typical of marine invaders. Analysing 344 loci, no outliers were detected between the introduced and native populations or in the introduced range. The genomic sampling may have been insufficient to reveal selection especially if it acts on traits determined by a few genes. Eight outliers were, however, identified within the native range, underlining a genetic singularity congruent with a well‐known biogeographical break along the Florida. Our results call into question the relevance of AFLP genome scans in detecting adaptation on the timescale of biological invasions: genome scans often reveal long‐term adaptation involving numerous genes throughout the genome but seem less effective in detecting recent adaptation from pre‐existing variation on polygenic traits. This study advocates other methods to detect selection effects during biological invasions—for example on phenotypic traits, although genome scans may remain useful for elucidating introduction histories.     

Micro-environmental factors and the endemism of bromeliad aquatic fauna

Tank bromeliads harbour aquatic microcosms with many endemic species among their leaves. We performed a set of experiments to determine which factors maintain the bromeliad aquatic fauna in isolation from neighbouring ponds. We cultivated three invertebrates species (an ostracod, an annelid and a cladoceran) from a pond surrounded by terrestrial bromeliads in Southeastern Brazil and introduced them inside cleaned bromeliads, using recipients with the same volume as controls. The pH, conductivity and organism densities were monitored in the bromeliad samples and controls for 41 days. The samples introduced inside the cleaned bromeliads showed a significant decrease in pH and conductivity compared to the controls. The pond organism populations introduced in the bromeliads presented a high extinction rate and a significant population decrease when compared to the ones introduced in the controls. We attributed the population decline experienced by the pond organisms to the oligotrophic conditions generated inside the tanks by the bromeliads due to the nutrient absorption. We suggest that the changes in water chemistry induced by the bromeliads could play an important role in isolating their microcosm communities from other freshwater systems. Other mechanisms that could produce the high rates of endemism in bromeliad fauna are discussed. Handling editor: K. Martens     

Introduced mammals in Australian rangelands: Future threats and the role of monitoring programmes in management strategies

Abstract In the present paper, we have provided an initial assessment of the current and future threats to biodiversity posed by introduced mammals (predators and herbivores) inhabiting the Australian rangelands, exploring trends in populations and options for management. Notably, rabbits have declined in recent years in the wake of rabbit haemorrhagic disease, populations of feral camels have increased dramatically and foxes appear to have moved northwards, thereby threatening native fauna within an expanded range. Following on, we developed a framework for monitoring the impacts of introduced mammals in the Australian rangelands. In doing so, we considered the key issues that needed to be considered in designing a monitoring programme for this purpose and critically evaluated the role of monitoring in pest animal management. Finally we have provided a brief inventory of current best‐practice methods of estimating the abundance of introduced mammal populations in the Australian rangelands with some comments on new approaches and their potential applications.     

Distribution of native and introduced fish in the Seven Creeks River system,Victoria

The distribution offish within the Seven Creeks River system, a tributary of the Goulburn River in the Murray–Darling basin, was determined primarily by a survey carried out during the summer of 1975–76. Information on the past occurrence of fish in the system was obtained from historical records. Seventeen species offish, eleven native and six exotic, have been recorded from the system. Habitat characteristics, cohabiting species and food habits are presented for each species recorded during the survey, and factors affecting present distributions are discussed. Siltation appears to have had adverse effects on native Murray cod, Maccullochella peeli, and Macquarie perch, Macquaria australasica, both of which were once common in the lower reaches of the system. The presence of Macquarie perch and the rare native trout cod, Maccullochella macquariensis, in the upper reaches of the system is traced to fish released in 1921 and 1922. A nother native species, the western carp gudgeon, Hypseleotris klunzingeri, was first observed in the system only after it had been introduced into farm dams in the area in the mid 1960s. Relationships between native and introduced fish are complex. Although the food requirements of some native and introduced species overlap and some native fish have been found in the stomachs of introduced species, the only evidence of a substantial effect of an introduced species on a native species is the apparent fragmentation of the range of common mountain galaxiids, Galaxias olidus, by brown trout, Salmo trutta, whose numbers in the system were, until recently, augmented by continual releases of hatchery-reared fish.     

Body size and the rate of spread of invasive ladybird beetles in North America

Invasive species are distinguished by their rate of spread and this is thought to be associated with the ability to produce many offspring. However, it is possible that many studies do not succeed in highlighting a positive correlation between invasiveness and reproductive rate because they lack an allometric perspective. Information on the ladybird beetles introduced into North America and data on life-history traits of 30 species of ladybird beetles were used to search for a relationship between ability to invade and traits related to reproduction and dispersal. We analyzed the mechanisms responsible for the rate of spread of invasive species of the aphidophagous species of ladybird introduced into North America that became established and spread. The two largest species extended their range an order of magnitude faster than the other species. The potential reproductive rate and the speed of movement are both positively correlated with body mass, which appears to be a good predictor of the ability to spread and colonize new territory. Further studies of invasive species should therefore include an allometric perspective in order to allow comparisons between species and an assessment of the influence of reproduction and dispersal potential on the rate with which they spread when exploiting highly suitable habitats.