Ecological explanations for successful invasion of exotic plants

The intentional introduction of exotic species can increase the level of local biodiversity, enrich people’s material lives, and bring significant social and economic benefits that are also the symbols of human progress. However, along with the frequent intercourse among countries and regions, the frequency of uncontrolled crossregional migration of species is increased and there is a lack of scientific management strategy for the intentional introduction of exotic species. Exotic species invasion, which is behind habitat fragmentation, has become the second largest threatening factor to the maintenance of the global-scale level of biological diversity. Exotic species invasion can destroy the structure of an ecosystem, disturb the economic life of a society, and do harm to human health. In this paper, the authors review some of the ecological explanations for issues such as “what causes or mechanisms have led to the successful invasion of exotic species”, including the “ideal weeds characteristics”, “biodiversity resistance hypothesis”, “enemies release hypothesis”, “evolution of increased competitive ability hypothesis”, “niche opportunity hypothesis”, and “novel weapon hypothesis”. The authors also analyze and evaluate the background and theoretical basis of the hypotheses, providing explanations for some phenomena, as well as the deficiencies of these explanations.     

Problems with the current deinococcal hypothesis: an alternative theory

All theories related to the evolution of Deinococcus radiodurans have a common denominator: the strong positive correlation between ionizing-radiation resistance and desiccation tolerance. Currently, the widespread hypothesis is that D. radiodurans’ ionizing-radiation resistance is a consequence of this organism’s adaptation to desiccation (desiccation adaptation hypothesis). Here, we draw attention to major discrepancy that has emerged between the “desiccation adaptation hypothesis” and recent findings in computational biology, experimental research, and terrestrial subsurface surveys. We explain why the alternative hypothesis, suggesting that D. radiodurans’ desiccation tolerance could be a consequence of this organism’s adaptation to ionizing radiation (radiation adaptation hypothesis), should be considered on equal basis with the “desiccation adaptation hypothesis”.     

Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

Monitoring and manipulation of a sublittoral hard bottom biocoenosis in Balsfjord,northern Norway

Sublittoral hard bottom biocoenoses in Balsfjord, Norway (69°31′ N, 19°1′ E), were monitored using underwater stereophotogrammetry. The study includes manipulation of natural densities of organisms and testing the importance of biological interactions and “key species ” for the structure of biocoenoses. Underwater photography has the advantages of being a non-destructive method, but it is selective because small or hidden organisms cannot always be observed. Field experiments with exclusion of organisms from cages seem suitable for testing hypotheses concerning which animals are “key species ” in certain biocoenoses. Sea-urchins(Strongylocentrotus droebachiensis, S. pallidus) were suspected to be “key species ” in the present study, and their removal from cages caused an increase in abundance of barnacles(Balanus balanoides), the limpetAcmaea testudinalis and algal cover.     

The elephant in the room: the role of failed invasions in understanding invasion biology

Most species introductions are not expected to result in invasion, and species that are invasive in one area are frequently not invasive in others. However, cases of introduced organisms that failed to invade are reported in many instances as anecdotes or are simply ignored. In this analysis, we aimed to find common characteristics between non‐invasive populations of known invasive species and evaluated how the study of failed invasions can contribute to research on biological invasions. We found intraspecific variation in invasion success and several recurring explanations for why non‐native species fail to invade; these included low propagule pressure, abiotic resistance, biotic resistance, genetic constraints and mutualist release. Furthermore, we identified key research topics where ignoring failed invasions could produce misleading results; these include studies on historical factors associated with invasions, distribution models of invasive species, the effect of species traits on invasiveness, genetic effects, biotic resistance and habitat invasibility. In conclusion, we found failed invasions can provide fundamental information on the relative importance of factors determining invasions and might be a key component of several research topics. Therefore, our analysis suggests that more specific and detailed studies on invasion failures are necessary.     

Birds from the Far East in Central Europe: some side-notes to Pfeifer et al. (2007)

The “reverse migration hypothesis” as the explanation for the vagrancy of Far Eastern birds in Europe is commented on. It is concluded that, as the postulated switch of 180° along a great circle line would lead the eastern or even central Siberian birds to North America rather than to Europe, the application of the “reverse migration hypothesis” to them seems untenable (which, however, does not preclude its plausibility in case of transatlantic, southern European, or even southwestern Siberian vagrants). On the other hand, patterns of appearance of the Turdus [naumanni] superspecies (Machalska et al. 1967) suggest the relationship of the phenomenon of vagrancy to post-breeding nomadic movements and/or periodical invasions of non-migrating species.     

Invasion of natural and agricultural cranberry bogs by introduced and native plants

Plant species invasions, i.e., the entry of additional plant species into a habitat with negative effects on species already there, are a major ecological problem in natural habitats and a major economic problem in agricultural habitats. Nutrient availability, disturbance, and proximity to other habitats are likely factors that may interact to control invasion in both types of habitat. We hypothesized (1) that elevated nutrient availability can promote the abundance of introduced species even when high cover of the existing plant community is maintained, and (2) that higher levels of invasion on the edges than in the interiors of habitats are due to differences in resource availability between edges and interiors. To test these hypotheses, we measured soil characteristics and the abundances of plant species in natural and agricultural cranberry (Vaccinium macrocarpon Ait.) bogs in southeastern Massachusetts. Contrary to the first hypothesis, agricultural bogs did not have higher cover or richness of introduced species than natural bogs, despite having higher levels of soil nutrients. Contrary to the second hypothesis, the edges of both agricultural and natural bogs had a higher cover and richness of introduced species than the interiors, even though only natural bogs showed differences in resource availabilities between edges and interiors. Results suggest that having a high cover of existing species can counter positive effects of elevated nutrients on the spread of introduced and non-crop species. However, maintaining similar resource availabilities on the edges and interiors of habitats may not prevent greater invasion of edges. Avoiding disturbances to natural communities, maintaining high crop cover, and focusing active control of introduced or non-crop species on the edges of habitats could help limit plant invasions into natural and agricultural habitats alike.     

Non-host volatiles mediate associational resistance to the pine processionary moth

An increasing body of evidence indicates that the association between different plant species may lead to a reduction in insect herbivory, i.e. associational resistance. This might be due to a top–down regulation of herbivores by increased numbers of natural enemies or to a disruptive bottom–up influence of lower host plant accessibility. In particular, the richer plant communities release more diverse plant odours that may disturb olfactory-guided host choice and mating behaviour of insect herbivores, i.e. the “semiochemical diversity hypothesis”. However, this hypothesis has been rarely tested experimentally in natural habitats, notably forest ecosystems. We tested the effects of non-host volatiles (NHV) on mate and host location by the pine processionary moth (PPM) at the scale of individual pine trees with branches of non-host tree (birch) at their base. Pheromone trap catches and the numbers of larval nests were both reduced by non-host presence under treated pine trees, confirming an associational resistance mediated by NHV. In both males and females, the antenna could detect several birch volatiles, including methyl salicylate (MeSa). MeSa inhibited the attraction of the PPM male to pheromone traps, as did bark and leaf chips from birch trees. Our test of three doses of MeSa at the habitat scale (50 m forest edges) showed that the reduction in the numbers of male PPM captured in traps and in larval nests was MeSa dose-dependent. These results show that odours released by deciduous non-host trees can reduce herbivory by a forest defoliator in conifers, providing support to the “semiochemical diversity hypothesis” as a mechanism of associational resistance.     

Modelling invasibility in endogenously oscillating tree populations: timing of invasion matters

The timing of introduction of a new species into an ecosystem can be critical in determining the invasibility (i.e. the sensitivity to invasion) of a resident population. Here, we use an individual-based model to test how (1) the type of competition (symmetric versus asymmetric) and (2) seed masting influence the success of invasion by producing oscillatory dynamics in resident tree populations. We focus on a case where two species (one resident, one invader introduced at low density) do not differ in terms of competitive abilities. By varying the time of introduction of the invader, we show that oscillations in the resident population favour invasion, by creating “invasibility windows” during which resource is available for the invader due to transiently depressed resident population density. We discuss this result in the context of current knowledge on forest dynamics and invasions, emphasizing the importance of variability in population dynamics.     

Fish Invasions in the World's River Systems: When Natural Processes Are Blurred by Human Activities

Because species invasions are a principal driver of the human-induced biodiversity crisis, the identification of the major determinants of global invasions is a prerequisite for adopting sound conservation policies. Three major hypotheses, which are not necessarily mutually exclusive, have been proposed to explain the establishment of non-native species: the “human activity” hypothesis, which argues that human activities facilitate the establishment of non-native species by disturbing natural landscapes and by increasing propagule pressure; the “biotic resistance” hypothesis, predicting that species-rich communities will readily impede the establishment of non-native species; and the “biotic acceptance” hypothesis, predicting that environmentally suitable habitats for native species are also suitable for non-native species. We tested these hypotheses and report here a global map of fish invasions (i.e., the number of non-native fish species established per river basin) using an original worldwide dataset of freshwater fish occurrences, environmental variables, and human activity indicators for 1,055 river basins covering more than 80% of Earth's surface. First, we identified six major invasion hotspots where non-native species represent more than a quarter of the total number of species. According to the World Conservation Union, these areas are also characterised by the highest proportion of threatened fish species. Second, we show that the human activity indicators account for most of the global variation in non-native species richness, which is highly consistent with the “human activity” hypothesis. In contrast, our results do not provide support for either the “biotic acceptance” or the “biotic resistance” hypothesis. We show that the biogeography of fish invasions matches the geography of human impact at the global scale, which means that natural processes are blurred by human activities in driving fish invasions in the world's river systems. In view of our findings, we fear massive invasions in developing countries with a growing economy as already experienced in developed countries. Anticipating such potential biodiversity threats should therefore be a priority.     

Can model species be used to advance the field of invasion ecology?

Hypotheses for explaining plant invasions have focused on a variety of factors that may influence invasion success, including propagule pressure, interactions of the introduced species with the biotic, abiotic, or disturbance properties of the new ecosystem, or the genetic characteristics of the invader itself. Evaluating the relative importance of these factors has been difficult because for most invaders key information about the introduced population or the introduction event is not available. We propose that natural experiments using model species is an important tool to test multiple invasion hypotheses at the same time, providing a complementary approach to meta-analysis and literature review. By focusing on a single candidate species, Pinus contorta, we explore several attributes that we propose constitute a good model, including: (a) intentional and relatively well documented introduction into a wide range of environments and countries across the world during the past century, where invasion success or failure has already occurred, (b) conspicuous growth form that simplifies assessment of growth rates, and comparisons across native and introduced ecosystems around the world, and, (c) documented and replicated variability of introduction intensity, genetic characteristics of the introduced populations, contrasting biotic communities present at sites of introduction, and abiotic conditions within and across introduced ecosystems. We propose that identifying model species with these characteristics will provide opportunities to disentangle the relative importance of different mechanisms hypothesized to influence invasion success, and thereby advance the field of invasion ecology.     

Demographic Stochasticity, Environmental Variability, and Windows of Invasion Risk for Bythotrephes Longimanus in North America

Biological invasions are a leading threat to freshwater biodiversity worldwide. A central unanswered question of invasion ecology is why some introduced populations establish while most fail. Answering this question will allow resource managers to increase the specificity and effectiveness of control efforts and policy. We studied the establishment of spiny water flea (Bythotrephes longimanus) in the United States and Canada by modeling introduction failure caused by demographic stochasticity, environmental variation, and seasonal environmental forcing. We compared predicted establishment rates with observed invasions of inland lakes in Ontario, Canada. Our findings suggest that environmental forcing can cause “windows” of invasion opportunity so that timing of introductions might be a greater determinant of population establishment than demographic stochasticity and random environmental variation. We expect this phenomenon to be exhibited by species representing a wide range of life histories. For spiny water flea in North America, a large window of invasion opportunity opens around the fourth week of May, persists through the summer, and closes with decreasing water temperatures in autumn. These results show how timing of introductions with respect to seasonally forced environmental drivers can be a key determinant of establishment success. By focusing on introductions during windows of invasion opportunity, resource managers can more effectively control invasion rates.     

Lengths and correlates of lag phases in upper-Midwest plant invasions

There are often lag phases in plant invasions, seemingly dormant periods between arrival in a new range and rapid population growth. Lags impede prioritization of invasive-species control efforts: when eradication is most feasible, it is often unclear whether a species is benign or a potentially harmful “sleeper weed.” I used herbarium records to estimate lag phases for invasive or potentially invasive plant species in three regions of the upper Midwest. I tested whether factors related to species’ invasion epidemiology, traits, or the habitats they invade were correlated with lag lengths. From an initial pool of 151 species, there were sufficient records to test for lags in 76 for northern Wisconsin, 90 for southern Wisconsin, and 91 for the southern Lake Michigan region. Lags were identified in 77% (197) of these 257 datasets and ranged from 3–140 years with a mean of 47.3 ± 34.6 (SD). Lags differed by native range, introduction pathway, growth form and habit, dispersal mode, flowering phenology, pollination mode for a subset of species, and breadth, light availability, and water availability of invaded habitats. However, estimated lags were highly variable and tested factors did not have strong explanatory power. Exotic species comprised an increasing proportion of total herbarium records. Of the species with known introduction pathways, 85% were intentionally introduced, mainly as ornamentals. The long durations, high variability, and low predictability of lags, along with human culpability for an increasingly non-native flora, support a cautious approach to species introductions.     

Predation contributes to invasion resistance of benthic communities against the non-indigenous tunicate <Emphasis Type="Italic">Ciona intestinalis</Emphasis>

Marine anthropogenic structures offer novel niches for introduced species but their role in the subsequent invasion to natural habitats remains unknown. Upon arrival in new environments, invaders must overcome biotic resistance from native competitors and predators if they are to establish successfully in natural habitats. We tested the hypotheses that (1) artificial structures (e.g., suspended aquaculture installations) present a niche opportunity for invasive species by providing a refuge from native benthic predators, and (2) native predators in natural benthic habitats suppress successful colonization by invaders. A recruitment experiment showed that the ascidians Pyura chilensis (native) and Ciona intestinalis (invasive) could recruit to both suspended artificial structures and natural benthic habitats. Ciona, however, was only able to establish adult populations on artificial structures. In natural benthic habitats Ciona only recruited and grew in predator-exclusion cages, because without this protection predation prevented its establishment. In predation experiments, native invertebrate and fish predators removed all invasive ascidians (recruits and adults) in benthic habitats, which contrasted with the high adult survival of the native ascidian P. chilensis. The refuge from a number of benthic predators facilitates the establishment of large populations of invasive species on suspended structures. We present a conceptual model of the invasion processes that includes the anthropogenic structures as a transitional stepping-stone that facilitates invasion by enhancing and prolonging propagule supply to surrounding natural communities. Those established invaders might then overcome biotic resistance during time periods when populations of consumers or competitors are weakened by natural or anthropogenic disturbances. Our results suggest that the conservation of natural habitats with a high diversity of native predators can be an effective means to prevent the spread of invasive species growing on suspended structures.     

Ecological Patterns and Biological Invasions: Using Regional Species Inventories in Macroecology

Macroecology depends heavily on a comparative methodology in order to identify large-scale patterns and to test alternative hypotheses that might generate such patterns. With the advent and accessibility of large electronic databases of species and their life history and ecological attributes, ecologists have begun seeking generalities, and examining large-scale ecological hypotheses involving core themes of range, abundance and diversity. For example, combinations of ecological, life history and phylogenetic data have been analysed using large species sets to test hypotheses in invasion biology. Analysis of regional species inventories can contribute cogently to our understanding of invasions. Here we examine several ways in which database analysis is effective. We review 19 studies of comparative invasions biology, each using >100 species of plants in their analyses, and show that invader success is linked to seven correlates: short life cycle, abiotic (mostly wind) dispersal, large native range size, non-random taxonomic patterns (emphasizing certain families or orders), presence of clonal organs, occupying disturbed habitats, and earlier time of introduction. These phylogenetically influenced, comparative analyses using regional species inventories are only just beginning and have much potential.     

The invasiveness of an introduced species does not predict its impact

Inconsistent use of terminology plagues the study and management of biological invasions. The term “invasive” has been used to describe inter alia (1) any introduced non-indigenous species; (2) introduced species that spread rapidly in a new region; and (3) introduced species that have harmful environmental impacts, particularly on native species. The second definition in various forms is more commonly used by ecologists, while the third definition is pervasive in policy papers and legislation. We tested the relationship between the invasiveness of an introduced species and its impact on native biodiversity. We quantified a species’ invasiveness by both its rate of establishment and its rate of spread, while its impact was assigned a categorical ranking based on the documented effects of the invader on native species populations. We found no correlations between these variables for introduced plants, mammals, fishes, invertebrates, amphibians and reptiles, suggesting that the mechanisms of invasion and impact are not strongly linked. Our results support the view that the term “invasive” should not be used to connote negative environmental impact.     

Role of glucosinolates in plant invasiveness

Many plants have been intentionally or accidentally introduced to new habitats where some of them now cause major ecological and economic threats to natural and agricultural ecosystems. The potential to become invasive might depend on plant characteristics, as well as on specific interactions with other organisms acting as symbionts or antagonists, including other plants, microbes, herbivores, or pollinators. The invasion potential furthermore depends on abiotic conditions in the habitat. Several species of the Brassicaceae, well known for their glucosinolate–myrosinase defence system, are invasive species. Various factors are reviewed here that might explain why these species were so successful in colonising new areas. Specific emphasis is laid on the role of glucosinolates and their hydrolysis products in the invasion potential. This particular defence system is involved specifically in plant–plant, plant–microbe and plant–insect interactions. Most research has been done on the mechanisms underlying invasion success of Alliaria petiolata and Brassica spp., followed by Bunias orientalis and Lepidium draba. Some examples are also given for plants that are not necessarily considered as invasives, but which were well investigated with respect to their interference potential with their biotic environment. For each species, most likely a combination of different plant characteristics enhanced the competitive abilities and led to diverse invasive phenotypes.     

A Global History of Australian Trees

Scholars studying the globalization of Australian trees have previously emphasized the rapid natural propagation of Australian trees outside of their native habitats, believing their success to be a reversal of “ecological imperialism” from the “new world” to the “old world.” This article argues that the expansion of Australian trees should not be viewed as a biological phenomenon, but as the result of a long-term attempt by powerful states and state-sponsored scientists to select and breed Australian species that could grow in a variety of climates and ecological conditions. Five non-biological factors largely determined the success of these attempts to grow Australian trees: the abundance or paucity of natural forests, state power, the amount of scientific research directed to planting Australian trees, the cost of labor, and the ability to utilize hardwood timbers and bark. This paper compares the use of Australian trees in Australia, India, and South Africa to demonstrate that biology was not the determining factor in the long-term success of many Australian genera and species.     

Predicting invasion success in complex ecological networks

A central and perhaps insurmountable challenge of invasion ecology is to predict which combinations of species and habitats most effectively promote and prevent biological invasions. Here, we integrate models of network structure and nonlinear population dynamics to search for potential generalities among trophic factors that may drive invasion success and failure. We simulate invasions where 100 different species attempt to invade 150 different food webs with 15–26 species and a wide range (0.06–0.32) of connectance. These simulations yield 11 438 invasion attempts by non-basal species, 47 per cent of which are successful. At the time of introduction, whether or not the invader is a generalist best predicts final invasion success; however, once the invader establishes itself, it is best distinguished from unsuccessful invaders by occupying a lower trophic position and being relatively invulnerable to predation. In general, variables that reflect the interaction between an invading species and its new community, such as generality and trophic position, best predict invasion success; however, for some trophic categories of invaders, fundamental species traits, such as having the centre of the feeding range low on the theoretical niche axis (for non-omnivorous and omnivorous herbivores), or the topology of the food web (for tertiary carnivores), best predict invasion success. Across all invasion scenarios, a discriminant analysis model predicted successful and failed invasions with 76.5 per cent accuracy for properties at the time of introduction or 100 per cent accuracy for properties at the time of establishment. More generally, our results suggest that tackling the challenge of predicting the properties of species and habitats that promote or inhibit invasions from food web perspective may aid ecologists in identifying rules that govern invasions in natural ecosystems.     

What would T. H. Huxley have made of prion diseases?

T. H. Huxley was “Darwin’s bulldog,” and took the offensive in championing the cause of evolution against skeptical scientists and outraged theologians. As such, he took part in one of the great “paradigm shifts” of biology, at the end of the nineteenth century. Huxley was a rigorous scientist and wrote important articles on scientific method, as well as publishing extensively on a wide range of subjects in natural history. In the second half of the twentieth century, the “prion hypothesis” was put forward to explain the pathogenesis of a curious group of diseases known as the transmissible spongiform encephalopathies. This also involved a “paradigm shift” because the prion hypothesis postulated that biologically relevant information could be enciphered in protein conformation (rather than encoded in nucleic acid base sequences), and could be transmitted from one molecule to another, thereby causing infectious disease. This article examines a few of Huxley’s remarks to speculate on how he might have responded to the scientific debate about prion disease had he lived a century later.