Seed banks of invasive Australian Acacia species in South Africa: Role in invasiveness and options for management

Despite impressive efforts at clearing stands of invasive Australian Acacia species in South Africa, insufficient attention has been given to understanding the role of seed banks in the invasiveness and long-term persistence of populations. We review information on seeds of these species, considering seed production, seed rain, and the dynamics of seeds in three layers: leaf litter, and upper and lower seed banks in the soil. Many factors affect the accumulation and susceptibility to destruction of seed banks and thus the opportunities for intervention to reduce seed numbers for each of these components. Reduction of seed banks is crucial for the overall success of the multi-million dollar management initiatives against these species. Classical biological control of buds, flower and young pods has reduced the seed production of many Australian acacias in South Africa. Fire can be applied to reduce seed numbers in the leaf litter and upper seed bank in some cases, although there are serious problems associated with high fire intensities in dense acacia stands. Other options, e.g. soil inversion and solarisation, exist to exercise limited reduction of seed numbers in some situations. There is little prospect of meaningful reduction of seed numbers in the lower seed bank. Preventing the accumulation of seed banks by limiting seed production through biological control is by far the most effective means, and in almost all cases the only practical means, of reducing seed numbers. This must be an integral part of management strategies. Several invasive Australian acacias are already under effective biological control, and further work to identify additional potential agents for all the currently invasive species and potentially invasive alien species is the top priority for improving the efficiency of management programmes.     

Mate‐finding failure as an important cause of Allee effects along the leading edge of an invading insect population

The movement of humans and goods has facilitated the arrival of non‐native insects, some of which successfully establish and cause negative consequences to the composition, services, and functioning of ecosystems. The gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), is currently invading North American forests at variable rates, spreading by local and long‐distance movement in a process known as stratified dispersal. Newly arriving colonizers often occur considerably ahead of the population front, and a key question is the degree to which they successfully establish. Prior research has highlighted mate‐finding failures in sparse populations as a cause of an Allee effect (positive density dependence). We explored this mechanism by measuring the relationship between female mating success and background male moth densities along the gypsy moth western front in Northern Wisconsin (USA) over 2 years. The mating results were then compared with analogous previous studies in southern Wisconsin, and the southern front in West Virginia and Virginia (USA). Mate‐finding failures in low‐density populations were consistently observed to be density‐dependent across all years and locations. Mate‐finding failures in low‐density populations have important ramifications to invasive species management, particularly in predicting species invasiveness, preventing successful establishment by small founder populations, and concentrating eradication efforts where they are most likely to succeed.     

Modelling the impact of an invasive insect via reaction-diffusion

An exotic, specialist seed chalcid, Megastigmus schimitscheki, has been introduced along with its cedar host seeds from Turkey to southeastern France during the early 1990s. It is now expanding in plantations of Atlas Cedar (Cedrus atlantica). We propose a model to predict the expansion and impact of this insect. This model couples a time-discrete equation for the ovo-larval stage with a two-dimensional reaction-diffusion equation for the adult stage, through a formula linking the solution of the reaction-diffusion equation to a seed attack rate. Two main diffusion operators, of Fokker-Planck and Fickian types, are tested. We show that taking account of the dependence of the insect mobility with respect to spatial heterogeneity, and choosing the appropriate diffusion operator, are critical factors for obtaining good predictions.     

Multiscale responses of soil stability and invasive plants to removal of non-native grazers from an arid conservation reserve

Disturbances and ecosystem recovery from disturbance both involve numerous processes that operate on multiple spatial and temporal scales. Few studies have investigated how gradients of disturbance intensity and ecosystem responses are distributed across multiple spatial resolutions and also how this relationship changes through time during recovery. We investigated how cover of non-native species and soil-aggregate stability (a measure of vulnerability to erosion by water) in surface and subsurface soils varied spatially during grazing by burros and cattle and whether patterns in these variables changed after grazer removal from Mojave National Preserve, California, USA. We compared distance from water and number of ungulate defecations — metrics of longer-term and recent grazing intensity, respectively, — as predictors of our response variables. We used information-theoretic analyses to compare hierarchical linear models that accounted for important covariates and allowed for interannual variation in the disturbance–response relationship at local and landscape scales. Soil stability was greater under perennial vegetation than in bare interspaces, and surface soil stability decreased with increasing numbers of ungulate defecations. Stability of surface samples was more affected by time since removal of grazers than was stability of subsurface samples, and subsurface soil stability in bare spaces was not related to grazing intensity, time since removal, or any of our other predictors. In the high rainfall year (2003) after cattle had been removed for 1–2 years, cover of all non-native plants averaged nine times higher than in the low-rainfall year (2002). Given the heterogeneity in distribution of large-herbivore impacts that we observed at several resolutions, hierarchical analyses provided a more complete understanding of the spatial and temporal complexities of disturbance and recovery processes in arid ecosystems.     

Impact assessment of an invasive flatworm, Convoluta convoluta, in the Southern Gulf of Maine

Convoluta convoluta (Abildgaard 1806) is a small (2-3 mm long) acoellous turbellarian flatworm from Europe that has invaded the Gulf of Maine within the last 5 years. Although it has been reported in densities of up to 19 individuals/cm², its ecological impact remains unknown. In its native habitat, it consumes harpacticoid copepods and primary settling mussels <0.5-mm shell length. This study estimated the impact of C. convoluta on juvenile blue mussel populations (Mytilus edulis Linnaeus 1758) around the Isles of Shoals in the southern Gulf of Maine, USA as well as looking at their distribution in their new habitat. We surveyed worm densities at sites of differing wave exposure over three substrates (hard substrates, bladed algae, filamentous algae) to quantify patterns of worm abundance. We found worms on all substrate types with their highest abundances occurring in areas of maximal sunlight exposure and minimal physical disturbance. We showed a definite pattern of consumption of mussels in the lab and found C. convoluta to consume up to 35% of primary settling mussels in the field, but only under certain conditions. Per capita impact on juvenile mussels was found to vary greatly in correlation with mussel recruitment rate and water temperature, but not with the consumption of harpacticoids. Our results also suggested that per capita interaction strength was reduced by intraspecific density-dependent competition and water temperature. The overall impact of C. convoluta on mussel populations in the southern Gulf of Maine is therefore estimated to be minimal.     

Space–time patterns during the establishment of a nonindigenous species

Increasing rates of global trade and travel have the invariable consequence of an increase in the likelihood of nonindigenous species arrival, and some new arrivals are successful in establishing themselves. Quantifying the pattern of establishment of nonindigenous species across both spatial and temporal scales is paramount in early detection efforts, yet very difficult to accomplish. Previous work in epidemiology has proposed methods for assessing the space–time properties of emerging infectious diseases by quantifying the degree of space–time clustering between individual cases. I tested the applicability of one such method commonly used in epidemiology, the Knox test for space–time interaction, to analyze rare abundance data from an isolated, newly-establishing gypsy moth, Lymantria dispar, population in Minnesota, USA, and incorporated a bootstrap approach to quantify the space–time pattern in a random process that can be used to compare with results from empirical data. The use of the Knox test in assessing the establishment phase of biological invasions could potentially serve as an early warning system against new invaders, particularly for those with a known history of a repeated number of arrivals.     

Invasion dynamics and potential spread of the invasive alien plant species Ageratina adenophora (Asteraceae) in China

Ageratina adenophora (Sprengel) R. King & H. Robinson (=Eupatorium adenophorum Sprengel) is one of the worst invasive alien species in China. Since A. adenophora was first noticed in Yunnan Province of China in the 1940s, its rapid spread has caused an ecological problem in south‐western China. Understanding its historical invasion pattern and its potential for further spread is needed to plan the management of the species. We reconstructed the historical process of its invasion and analysed its ecological preferences in the invaded region. After a lag phase of 20 years (1940–60), A. adenophora spread rapidly throughout the south and middle subtropical zones in Yunnan, Guizhou, Sichuan, and Guangxi, China, with an average expansion rate of 20 km per year. It spread relatively slowly in north subtropical areas, with an average expansion rate of 6.8 km per year. It has not established in warm temperate areas within the invaded regions. Although range expansion in Yunnan stopped after 1990, the expansion of its range into neighbouring provinces indicates that A. adenophora has not reached the full potential of its distribution and its range is still rapidly expanding within China. We applied ecological niche modelling (GARP — Genetic Algorithm for Rule‐set Prediction) to predict potential invasion areas in mainland China on the basis of occurrence points within colonized areas where A. adenophora has reached equilibrium. The predictions, confirmed by the range of values of four key environmental parameters, generally match the parameters of the geography and ecology in the invaded region. Southern and south‐central China have climatic conditions suggestive of a high potential for invasion by A. adenophora. Climatic conditions in northern and western China appear unsuitable for A. adenophora. Urgent measures should be taken to prevent this species from further spreading into the vast areas of potential habitat in southern and south‐central China.     

Potential changes to the biology and challenges to the management of invasive sea lamprey Petromyzon marinus in the Laurentian Great Lakes due to climate change

Control programs are implemented to mitigate the damage caused by invasive species worldwide. In the highly invaded Great Lakes, the climate is expected to become warmer with more extreme weather and variable precipitation, resulting in shorter iced‐over periods and variable tributary flows as well as changes to pH and river hydrology and hydrogeomorphology. We review how climate change influences physiology, behavior, and demography of a damaging invasive species, sea lamprey (Petromyzon marinus), in the Great Lakes, and the consequences for sea lamprey control efforts. Sea lamprey control relies on surveys to monitor abundance of larval sea lamprey in Great Lakes tributaries. The abundance of parasitic, juvenile sea lampreys in the lakes is calculated by surveying wounding rates on lake trout (Salvelinus namaycush), and trap surveys are used to enumerate adult spawning runs. Chemical control using lampricides (i.e., lamprey pesticides) to target larval sea lamprey and barriers to prevent adult lamprey from reaching spawning grounds are the most important tools used for sea lamprey population control. We describe how climate change could affect larval survival in rivers, growth and maturation in lakes, phenology and the spawning migration as adults return to rivers, and the overall abundance and distribution of sea lamprey in the Great Lakes. Our review suggests that Great Lakes sea lamprey may benefit from climate change with longer growing seasons, more rapid growth, and greater access to spawning habitat, but uncertainties remain about the future availability and suitability of larval habitats. Consideration of the biology of invasive species and adaptation of the timing, intensity, and frequency of control efforts is critical to the management of biological invasions in a changing world, such as sea lamprey in the Great Lakes.     

Increased resistance to generalist herbivores in invasive populations of the California poppy (Eschscholzia californica)

Escape from enemies in the native range is often assumed to contribute to the successful invasion of exotic species. Following optimal defence theory, which assumes a trade‐off between herbivore resistance and plant growth, some have predicted that the success of invasive species could be the result of the evolution of lower resistance to herbivores and increased allocation of resources to growth and reproduction. Lack of evidence for ubiquitous costs of producing plant toxins, and the recognition that invasive species may escape specialist, but not generalist enemies, has led to a new prediction: invasive species may escape ecological trade‐offs associated with specialist herbivores, and evolve increased, rather than decreased, production of defensive compounds that are effective at deterring generalist herbivores in the introduced range. We tested the performance of two generalist lepidopteran herbivores, Trichoplusia ni and Orgyia vetusta, when raised on diets of native and invasive populations of the California poppy, Eschscholzia californica. Pupae of T. ni were significantly larger when reared on native populations. Similarly, caterpillars of O. vetusta performed significantly better when raised on native populations, indicating that invasive populations of the California poppy are more resistant to herbivores than native populations. The chance of successful establishment of some non‐indigenous plant species may be increased by retaining resistance to generalist herbivores, and in some cases, invasive species may be able to escape ecological trade‐offs in their new range and evolve, as we observed, even greater resistance to generalist herbivores than native plants.     

Release from native herbivores facilitates the persistence of invasive marine algae: a biogeographical comparison of the relative contribution of nutrients and herbivory to invasion success

The effect of herbivory and nutrient enrichment on the growth of invasive and native macroalgal species was simultaneously studied in two biogeographic regions: the Caribbean and Hawaii. Herbivores suppressed growth of invasive algae in their native (Caribbean) and invaded range (Hawaii), but despite similar levels of herbivore biomass, the intensity of herbivory was lower in Hawaii. Algal species with a circumtropical distribution did not show a similar effect of herbivores on their growth. Nutrient enrichment did not enhance growth of any algal species in either region. The reduction in herbivore intensity experienced by invasive algae in Hawaii rather than an escape from (native) herbivores provided invasive macroalgae with “enemy release” sensu the Enemy Release Hypothesis (ERH). Since native, Hawaiian herbivores still feed and even prefer invasive algae over native species, invasion scenario’s that involve predation (e.g. the ERH) could be falsely dismissed when invasive species are only studied in their invasive range. We therefore argue that escape from herbivores (i.e. enemy release) can only effectively be determined with additional information on the intensity of predation experienced by an invasive species in its native range.     

A hitchhiker’s guide to the Maritimes: anthropogenic transport facilitates long‐distance dispersal of an invasive marine crab to Newfoundland

Aim To determine timing, source and vector for the recent introduction of the European green crab, Carcinus maenas (Linnaeus, 1758), to Newfoundland using multiple lines of evidence. Location Founding populations in Placentia Bay, Newfoundland, Canada and potential source populations in the north‐west Atlantic (NWA) and Europe. Methods We analysed mitochondrial and microsatellite genetic data from European and NWA populations sampled during 1999–2002 to determine probable source locations and vectors for the Placentia Bay introduction discovered in 2007. We also analysed Placentia Bay demographic data and shipping records to look for congruent patterns with genetic analyses. Results Demographic data and surveys suggested that C. maenas populations are established and were in Placentia Bay for several years (c. 2002) prior to discovery. Genetic data corroboratively suggested central/western Scotian Shelf populations (e.g., Halifax) as the likely source area for the anthropogenic introduction. These Scotian Shelf populations were within an admixture zone made up of genotypes from both the earlier (early 1800s) and later (late 1900s) introductions of the crab to the NWA from Europe. Placentia Bay also exhibited this mixed ancestry. Probable introduction vectors included vessel traffic and shipping, especially vessels carrying ballast water. Main conclusions Carcinus maenas overcame considerable natural barriers (i.e., coastal and ocean currents) via anthropogenic transport to become established and abundant in Newfoundland. Our study thus demonstrates how non‐native populations can be important secondary sources of introduction especially when aided by human transport. Inference of source populations was possible owing to the existence of an admixture zone in central/western Nova Scotia made up of southern and northern genotypes corresponding with the crab’s two historical introductions. Coastal vessel traffic was found to be a likely vector for the crab’s spread to Newfoundland. Our study demonstrates that there is considerable risk for continued introduction or reintroduction of C. maenas throughout the NWA.     

Boom‐bust dynamics in biological invasions: towards an improved application of the concept

Boom‐bust dynamics – the rise of a population to outbreak levels, followed by a dramatic decline – have been associated with biological invasions and offered as a reason not to manage troublesome invaders. However, boom‐bust dynamics rarely have been critically defined, analyzed, or interpreted. Here, we define boom‐bust dynamics and provide specific suggestions for improving the application of the boom‐bust concept. Boom‐bust dynamics can arise from many causes, some closely associated with invasions, but others occurring across a wide range of ecological settings, especially when environmental conditions are changing rapidly. As a result, it is difficult to infer cause or predict future trajectories merely by observing the dynamic. We use tests with simulated data to show that a common metric for detecting and describing boom‐bust dynamics, decline from an observed peak to a subsequent trough, tends to severely overestimate the frequency and severity of busts, and should be used cautiously if at all. We review and test other metrics that are better suited to describe boom‐bust dynamics. Understanding the frequency and importance of boom‐bust dynamics requires empirical studies of large, representative, long‐term data sets that use clear definitions of boom‐bust, appropriate analytical methods, and careful interpretations.     

A rapid technique for assessing the suitability of areas for invasive species applied to New Zealand's rivers

Early responses to incursions of non‐indigenous species (NIS) into new areas include modelling and surveillance to define the organisms’ potential and actual distributions. For well‐studied invasive species, predictive models can be developed based on quantitative data describing environmental tolerances. In late 2004, an invasive freshwater diatom Didymosphenia geminata, an NIS for which we had no such quantitative data, was detected in a New Zealand river. We describe a procedure used to rapidly develop a classification of suitability for all New Zealand's rivers, based on two sources of information. First, from a review of the limited available literature and unpublished data, we determined that temperature, hydrological and substrate stability, light availability, and water pH were the most important environmental gradients determining D. geminata's broad‐scale distribution and capacity for establishing and forming blooms in rivers. The second information source was a GIS‐based river network developed for a national classification of New Zealand's rivers, with associated data describing environmental characteristics of each section of the network. We used six variables that were available for every section of the network as surrogates for the environmental gradients that determine suitability. We then determined the environmental distance of all the river sections in the network from our assessment of the optimal conditions conducive to D. geminata blooms. The analysis suggested that > 70% of New Zealand's river sections (stream order > 3) fell into the two highest suitability categories (on a five‐point scale). At the time of writing, D. geminata had spread to 12 catchments, all of which were within these two categories. The technique is applicable in initial responses to incursions of NIS where quantitative information is limited, and makes optimal use of available qualitative information. Our assessment contributed to evaluations of the potential ecological, social, and economic impacts of D. geminata and is currently being used to stratify site selection for ongoing surveillance.     

Spatial dynamics of an invasive bird species assessed using robust design occupancy analysis: the case of the Eurasian collared dove (Streptopelia decaocto) in France

Aim The study of the spatial dynamics of invasive species is a key issue in invasion ecology. While mathematical models are useful for predicting the extent of population expansions, they are not suitable for measuring and characterizing spatial patterns of invasion unless the probability of detection is homogeneous across the distribution range. Here, we apply recently developed statistical approaches incorporating detection uncertainty to characterize the spatial dynamics of an invasive bird species, the Eurasian collared dove (Streptopelia decaocto). Location France. Methods Data on presence/absence of doves were recorded from 1996 to 2004 over 1045 grid cells (28 × 20 km) covering the entire country. Each grid cell included five point counts spaced along a route, which was visited twice a year, allowing for an estimation of detection probability. Each route was assigned to one of six geographical regions. We used robust design occupancy analysis to assess spatial and temporal variation in parameters related to the spatial dynamics of the species. These parameters included occupancy rate, colonization and local extinction probabilities. Our inference approach was based on the selection of the most parsimonious model among competitive models parametrized with conditional probabilities. Results The probability of detecting the presence of doves on a given route was high. However, we found evidence to incorporate detection uncertainty in inference processes about spatial dynamics, since detection probability was neither perfect (i.e. it was < 1), nor constant over space and time. Results showed a clear positive trend in occupancy rate over the study period, increasing from 55% in 1996 to 76% in 2004. In addition, occupancy rate differed among regions (range: 37–79%) and further analysis showed that colonization probability by region was positively related to occupancy rate. Finally, local extinction probability was lower than colonization probability and showed a tendency to decrease over the study period. Main conclusions Our results emphasize the importance of estimating detection probabilities in order to draw proper inferences about the spatial and temporal dynamics of the invasion pattern of the collared dove. In contrast to the perceived spatial dynamics from national atlas surveys, we provide evidence that the range of this species is currently increasing in France. Other results, such as regional specificity in colonization probabilities and time variation in local extinction are consistent with expectations from invasion and metapopulation theory.