The role of Allee effects in gypsy moth, <Emphasis Type="Italic">Lymantria dispar</Emphasis> (L.), invasions

Allee effects have been applied historically in efforts to understand the low-density population dynamics of rare and endangered species. Many biological invasions likewise experience the phenomenon of decreasing population growth rates at low population densities because most founding populations of introduced nonnative species occur at low densities. In range expansion of established species, the initial colonizers of habitat beyond the organism’s current range are usually at low density, and thus could be subject to Allee dynamics. There has been consistent empirical and theoretical evidence demonstrating, and in some cases quantifying, the role of Allee dynamics in the gypsy moth, Lymantria dispar (L.), invasion of North America. In this review, we examine the potential causes of the Allee effect in the gypsy moth and highlight the importance of mate-finding failure as a primary mechanism behind an Allee effect, while the degree to which generalist predators induce an Allee effect remains unclear. We then explore the role of Allee effects in the establishment and spread dynamics of the gypsy moth system, which conceptually could serve as a model system for understanding how Allee effects manifest themselves in the dynamics of biological invasions.     

Early warning signals of demographic regime shifts in invading populations

The Allee effect can cause alternative stable states in population abundance of invasive species. Sudden eruption of invading populations from low to high abundance may be viewed as a regime shift from one alternative state to another. Previous research proposed several types of early warning signals to predict regime shifts in ecological systems such as polluted lakes and semiarid grasslands. This paper explores theoretically the potential of such indicators in predicting demographic regime shifts of invading populations. I analyzed a stochastic differential equation model for the population dynamics of an invasive species subject to Allee effects and propagule pressure. Diffusion approximation to the stochastic model suggests that persistent propagule pressure makes demographic regime shifts inevitable, but Allee effects can lengthen the mean time until regime shifts virtually indefinitely. To compare the potential of indicators, I examined standard deviation, skewness, and estimated return rates of longitudinal population abundance. I found that standard deviation showed a distinct increase as regime shifts became more likely, but skewness and return rates showed no clear trends. This result suggests that standard deviation might be a useful warning signal for forecasting an impending demographic regime shift of invading populations during the period when their abundance is still low.     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

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.     

Climate suitability and management of the gypsy moth invasion into Canada

The gypsy moth has become established throughout southern Canada east of Lake Superior where the climate is suitable for the completion of its univoltine life cycle. The spread of the gypsy moth to the north and west in Canada has so far been prevented by climatic barriers and host plant availability as well as by aggressive eradication of incipient populations. Climate change is expected to increase the area of climatic suitability and result in greater overlap with susceptible forest types throughout Canada, especially in the west. At the same time, the gypsy moth is spreading west in the USA into states bordering western Canadian provinces. These circumstances all lead to a greatly increased risk of further invasion into Canadian forests by the gypsy moth. Management actions need to be intensified in different ways in different parts of the country to reduce the impacts of spread in eastern Canada and to prevent the gypsy moth from invading western regions.     

The renowned cactus moth, Cactoblastis cactorum: its natural history and threat to native Opuntia floras in Mexico and the United States of America

Abstract. The cactus moth, Cactoblastis cactorum (Berg) (Phycitidae) is native to South America. It was released as a biological control agent against alien Opuntia- cacti in Australia in the 1920s, then in southern Africa, and latterly on several islands, including those in the Caribbean. In 1989, the cactus moth was discovered in Florida, in the United States of America, where it is now threatening the survival of indigenous Opuntia species. In this paper we identify some of the attributes that have contributed to the success of C. cactorum as a weed biological control agent. Many of these same qualities account for the problems that C. cactorum has caused in Florida and predispose it as a major threat to the speciose, native Opuntia- floras of Central and North America. An estimated 79 platyopuntia (prickly pear) species are at risk: 51 species endemic to Mexico; nine species endemic to the United States; and 19 species common to both countries. Many cultivated and wild Opuntia species, that are used in various ways, are also vulnerable to attack by C. cactorum , including at least 25 species in Mexico and three species in the United States, particularly the widely exploited and culturally important cultivars of O. ficus-indica . Some control strategies are suggested that may minimize the risk and consequences of invasion by the cactus moth. The wider implications of this threat to the practice of weed biological control and to conservation are discussed.     

Invasion speed is affected by geographical variation in the strength of Allee effects

Allee effects can play a critical role in slowing or preventing the establishment of low density founder populations of non-indigenous species. Similarly, the spread of established invaders into new habitats can be influenced by the degree to which small founder populations ahead of the invasion front are suppressed through Allee effects. We develop an approach to use empirical data on the gypsy moth, a non-indigenous invader in North America, to quantify the Allee threshold across geographical regions, and we report that the strength of the Allee effect is subject to spatial and temporal variability. Moreover, we present what is to our knowledge the first empirical evidence that geographical regions with higher Allee thresholds are associated with slower speeds of invasion.     

Dispersion in time and space affect mating success and Allee effects in invading gypsy moth populations

1. Understanding why invading populations sometimes fail to establish is of considerable relevance to the development of strategies for managing biological invasions. 2. Newly arriving populations tend to be sparse and are often influenced by Allee effects. Mating failure is a typical cause of Allee effects in low-density insect populations, and dispersion of individuals in space and time can exacerbate mate-location failure in invading populations. 3. Here we evaluate the relative importance of dispersal and sexual asynchrony as contributors to Allee effects in invading populations by adopting as a case study the gypsy moth (Lymantria dispar L.), an important insect defoliator for which considerable demographic information is available. 4. We used release-recapture experiments to parameterize a model that describes probabilities that males locate females along various spatial and temporal offsets between male and female adult emergence. 5. Based on these experimental results, we developed a generalized model of mating success that demonstrates the existence of an Allee threshold, below which introduced gypsy moth populations are likely to go extinct without any management intervention.     

Is the exotic bumblebee Bombus terrestris really invading Tasmanian native vegetation?

There has been a great deal of disagreement surrounding the capacity of Bombus terrestris to invade Tasmanian native vegetation. This paper reviews the conflicting findings of previous surveys of the invasion of Tasmania by B. terrestris, and presents new data from the 2004–2005 austral summer. From this, it is clear that B. terrestris has extensively invaded Tasmanian native vegetation. The new data provide strong evidence that B. terrestris is breeding in native vegetation in every region of Tasmania. More than 10 bumblebees were seen in one day at 153 locations in native vegetation, including 42 locations within 10 National Parks and 38 locations within the Tasmanian Wilderness World Heritage Area. Nests of B. terrestris were also found within two National Parks. These findings suggest that B. terrestris would also invade native vegetation in non-arid temperate regions of the Australian mainland, if it is introduced there.     

Spatial analysis of gypsy moth populations in Sardinia using geostatistical and climate models

• 1 Spatial fluctuations of the Sardinian population of the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae) were characterized using geostatistical and climate models. Data on gypsy moth egg mass abundance recorded at 282 permanent monitoring sites from 1980 to 2004 were incorporated in a geographic information system with the vegetational, geomorphological and pedological features of the sites.
• 2 Statistical analyses revealed that the relative outbreak frequency was related to the predominant host tree, slope and elevation of the monitoring sites, whereas there was no correlation between outbreak frequency and exposure and soil type.
• 3 By using bioclimatic modelling, probability maps of gypsy moth outbreaks were generated. The model identified a probability surface with climatic conditions favourable to gypsy moth outbreaks and thus potentially subject to defoliation. The maps included 92 sites where outbreaks never occurred, suggesting that the Sardinian climate may not be a determinant factor for gypsy moth outbreaks.
• 4 The geostatistical method cokriging with outbreak frequency as a covariate was found to be the most suitable technique to estimate gypsy moth egg mass abundance. Semivariograms showed spatial correlation of egg mass abundance within the range 18.5–53 km. The results obtained were used to create regional gypsy moth distribution maps by cokriging, which demonstrated the outbreak foci and different infestation levels at each monitoring area. These results can help to delimit the treatment areas and develop rational gypsy moth management programmes.

     

The invasion by the Yellow-legged hornet: A systematic review

The Yellow-legged hornet (Vespa velutina nigrithorax), native to regions of Southeast Asia, was accidentally introduced in Europe, South Korea, and Japan, where is has often become invasive. Due to its potential negative impacts at ecologic, economic and social levels, this hornet was included in the “Union list” of the EU legislation for invasive alien species. This means that measures are urgently needed to prevent further introductions, as well as to early-detect and control spread to avoid new populations. In this study we aim to identify the main reported drivers of distribution, ecological preferences, impacts, and methods for preventing introduction, controlling, and managing this invasive species. The supporting information was obtained from a comprehensive literature search. Then, a literature review was performed to classify the records gathered and to extract the relevant information following an adapted Drivers-Pressures-State-Impacts-Responses framework. The achieved results show a growing interest of researchers on V. velutina nigrithorax through time due to its quick spread and impacts on new ecosystems. They also revealed that there is much information on the State of invasions, whereas more knowledge is needed regarding the Drivers of those invasions. Biological traits such as life history traits, morphology, and the sting venom properties are some of the most studied topics regarding V. velutina nigrithorax. In the future, research should focus on the topics that lack information, analyse other Response solutions that meet the intended measures by the EU legislation, and use new methodology to study the impacts caused by this invader under new perspectives.     

Persistence of structured populations in random environments

Environmental fluctuations often have different impacts on individuals that differ in size, age, or spatial location. To understand how population structure, environmental fluctuations, and density-dependent interactions influence population dynamics, we provide a general theory for persistence for density-dependent matrix models in random environments. For populations with compensating density dependence, exhibiting “bounded” dynamics, and living in a stationary environment, we show that persistence is determined by the stochastic growth rate (alternatively, dominant Lyapunov exponent) when the population is rare. If this stochastic growth rate is negative, then the total population abundance goes to zero with probability one. If this stochastic growth rate is positive, there is a unique positive stationary distribution. Provided there are initially some individuals in the population, the population converges in distribution to this stationary distribution and the empirical measures almost surely converge to the distribution of the stationary distribution. For models with overcompensating density-dependence, weaker results are proven. Methods to estimate stochastic growth rates are presented. To illustrate the utility of these results, applications to unstructured, spatially structured, and stage-structured population models are given. For instance, we show that diffusively coupled sink populations can persist provided that within patch fitness is sufficiently variable in time but not strongly correlated across space.     

The global invasion success of Central European plants is related to distribution characteristics in their native range and species traits

Aim Determining which traits predispose a species to become invasive is a fundamental question of invasion ecology, but traits affect invasiveness in concert with other factors that need to be controlled for. Here, we explore the relative effects of biological traits of plant species and their distributional characteristics in the native range on invasion success at two stages of invasion. Location Czech Republic (for native species); and the world (for alien species). Methods The source pool of 1218 species of seed plants native to Central Europe was derived from the flora of the Czech Republic, and their occurrence in 706 alien floras all over the world was recorded, distinguishing whether they were listed as an ‘alien’ or a ‘weed’ in the latest version of Randall’s ‘Global compendium of weeds’ database. The latter type of occurrence was considered to indicate species ability to invade and cause economic impact, i.e. a more advanced stage of invasion. Using the statistical technique of regression trees, we tested whether 19 biological traits and five distributional characteristics of the species in their native range can be used to predict species success in two stages of invasion. Results The probability of a species becoming alien outside its native distribution range is determined by the size of its native range, and its tolerance of a wide range of climates acquired in the region of origin. Biological traits play only an indirect role at this stage of invasion via determining the size of the native range. However, the ability of species to become a weed is determined not only by the above characteristics of native distribution, but also directly by biological traits (life form and strategy, early flowering, tall stature, generative reproduction, number of ploidy levels and opportunistic dispersal by a number of vectors). Species phylogenetic relatedness plays only a minor role; it is more important at the lowest taxonomic levels and at the later stage of invasion. Main conclusion The global success of Central European species as ‘weeds’ is determined by their distributional characteristics in the native ranges and by biological traits, but the relative importance of these determinants depends on the stage of invasion. Species which have large native ranges and are common within these ranges should be paid increased attention upon introductions, and the above biological traits should be taken into account in screening systems applied to evaluate deliberate introductions of alien plants to new regions.     

Allee effects in biological invasions

Understanding the dynamics of small populations is obviously important for declining or rare species but is also particularly important for invading species. The Allee effect, where fitness is reduced when conspecific density is low, can dramatically affect the dynamics of biological invasions. Here, we summarize the literature of Allee effects in biological invasions, revealing an extensive theory of the consequences of the Allee effect in invading species and some empirical support for the theory. Allee effects cause longer lag times, slower spread and decreased establishment likelihood of invasive species. Expected spatial ranges, distributions and patterns of species may be altered when an Allee effect is present. We examine how the theory can and has been used to detect Allee effects in invasive species and we discuss how the presence of an Allee effect and its successful or unsuccessful detection may affect management of invasives. The Allee effect has been shown to change optimal control decisions, costs of control and the estimation of the risk posed by potentially invasive species. Numerous ways in which the Allee effect can influence the efficacy of biological control are discussed.     

Crayfish invading Europe: the case study of Procambarus clarkii

The red swamp crayfish, Procambarus clarkii, native to northeastern Mexico and southcentral USA, is today the dominant macroinvertebrate in several European countries. While the first introduction of this species into Spain is well-documented, little is known about its pathways of invasion and the reason for its rapid spread in several European countries. Study of the biology of the species has revealed a number of properties that makes this crayfish a successful invader. Procambarus clarkii exhibits properties characteristic of an r-selected species, including early maturity at small body size, rapid growth rates, large numbers of offspring at a given parental size, and relatively short life spans. It is also plastic in its life cycle, able to disperse widely in the habitat and to tolerate environmental extremes. It displays generalist and opportunistic feeding habits, consuming macrophytes and preying on amphibians. Procambarus clarkii can also replace indigenous crayfish by a combination of mechanisms, including competitive exclusion and transmission of the fungus-like Aphanomyces astaci, responsible for the crayfish plague. Finally, this species shows a wide behavioral flexibility when coping with new types of predators. The results of these studies, combined with the increasing information available in the scientific literature on this and other crayfish species, will help us understand invasions in this taxon and make predictions about the identity of future crayfish invaders.     

Estimating the spatial distribution of wintering little brown bat populations in the eastern United States

Depicting the spatial distribution of wildlife species is an important first step in developing management and conservation programs for particular species. Accurate representation of a species distribution is important for predicting the effects of climate change, land‐use change, management activities, disease, and other landscape‐level processes on wildlife populations. We developed models to estimate the spatial distribution of little brown bat (Myotis lucifugus) wintering populations in the United States east of the 100th meridian, based on known hibernacula locations. From this data, we developed several scenarios of wintering population counts per county that incorporated uncertainty in the spatial distribution of the hibernacula as well as uncertainty in the size of the current little brown bat population. We assessed the variability in our results resulting from effects of uncertainty. Despite considerable uncertainty in the known locations of overwintering little brown bats in the eastern United States, we believe that models accurately depicting the effects of the uncertainty are useful for making management decisions as these models are a coherent organization of the best available information.     

Growth of newly established alien populations: comparison of North American gypsy moth colonies with invasion theory

A common characteristic observed in many biological invasions is the existence of a lag between the time of arrival by the alien population and the time when established populations are noticed. Considerable advances have been made in modeling the expansion of invading species, and there is often remarkable congruence between the behavior of these models with spread of actual populations. While these models have been used to characterize expansion of very newly founded colonies, there have been few attempts to compare the behavior predicted from theory with spread in actual newly founded populations, largely due to the difficulty of sampling sparse populations. Models predict that time lags in the radial expansion of newly invaded populations may be due to time requirements for the population to grow from founding to detectable levels. Models also indicate that these time lags can be predicted based upon population parameters such as the intrinsic rate of population growth and diffusion coefficient. In this paper, we compared the behavior of these models with historical data on gypsy moth, Lymantria dispar, establishment and spread to show similarities between model predictions and observed population spread, both of which exhibited temporal lags of expansion. However, actual populations exhibited certain behaviors that were not predicted, and this could be due, in part, to the existence of Allee effects and stochasticity. Further work that incorporates these effects is needed to more fully understand the growth of incipient colonies of invading species. Ultimately, this information can be of critical importance in the selection of effective strategies for their detection and eradication.     

Influence of foraging behavior and host spatial distribution on the localized spread of the emerald ash borer, <Emphasis Type="Italic">Agrilus planipennis</Emphasis>

Management programs for invasive species are often developed at a regional or national level, but physical intervention generally takes place over relatively small areas occupied by newly founded, isolated populations. The ability to predict how local habitat variation affects the expansion of such newly founded populations is essential for efficiently targeting resources to slow the spread of an invasive species. We assembled a coupled map lattice model that simulates the local spread of newly founded colonies of the emerald ash borer (Agrilus planipennis Fairmaire), a devastating forest insect pest of ash (Fraxinus spp.) trees. Using this model, we investigated the spread of A. planipennis in environments with different Fraxinus spp. distributions, and explored the consequences of ovipositional foraging behavior on the local spread of A. planipennis. Simulations indicate that increased larval density, resulting from lower host tree density or higher initial population sizes, can increase the spread rate during the first few years after colonization by increasing a density-dependent developmental rate and via host resource depletion. Both the radial spread rate and population size were greatly influenced by ovipositional foraging behavior. Two known behaviors of ovipositing A. planipennis females, attraction towards areas with high ash tree density and attraction to stressed trees, had opposing effects on spread. Results from this model illustrate the significant influence of resource distribution and foraging behavior on localized spread, and the importance of these factors when formulating strategies to monitor and manage invasive pests.     

Sexual reproduction of cordgrass hybrids (Spartina foliosa x alterniflora) invading tidal marshes in San Francisco Bay

We genetically analysed cordgrass plants and seedlings throughout the San Francisco, California, USA, estuary and found that hybrids between exotic Spartina alterniflora and native Spartina foliosa are the principal cordgrass invaders and colonizers. We hypothesized that this was due to higher seed set and siring ability by hybrids relative to the native species; too few alien parents remained in San Francisco Bay for our comparative studies. Hybrid seed comprised 91% to 98% of that set in the marsh study plants over the 2 years of the study. Total viable pollen production by hybrid plants was 400 times that of the native plants. Seed and pollen production were highly skewed towards a few hybrid genotypes. In addition to seed produced by hybrid plants, hybrid seed was produced by S. foliosa due to hybrid backcrossing. While the greatest advantage for hybrids was in pollen and seed production, hybrid seeds germinated, and seedlings survived and grew as well or better than the native species. As native S. foliosa becomes increasingly rare, hybrid seed floating on the tides will predominate, overwhelming recruitment sites and resulting in further colonization by hybrids. In an evolutionary context, hybrids with exceptional pollen and seed production will be initially favoured by natural selection, leading to the evolution of even more fertile hybrid genotypes.