Historical perspectives on human-assisted biological invasions

Of all the things we have done to the Earth so far, which one is likely to make the most indelible mark? I have been posing this question to students and colleagues for some time, and usually receive one of three reasonable replies: habitat destruction, human overpopulation, or global climate change. Yet as bad as all these maladies are, they are, to some extent, reversible on time scales to which we can relate. The remedies for these problems, despite mammoth cultural and economic obstacles, can be visualized generally in terms of technology and scientific theory. Only the will is lacking. On longer geological and evolutionary time scales, most such human impacts, if corrected soon, would be mere blips, perturbations of the ecosphere that would leave only a few detectable traces in the fossil record a million years hence.     

Impacts of disturbance on migratory waterfowl

It is well known that disturbance from human activities can cause temporary changes in behaviour and locally affect temporal and spatial distribution of migratory and wintering waterfowl. But it is also known that, to some extent, birds can compensate for disturbance by altering their behaviour or habituating to human activities. Comparatively little is known about how these reactions to disturbance may impact on the large-scale dispersion of waterfowl and, ultimately, on their population dynamics. To be able to answer these questions, a better theoretical framework, based on optimal foraging theory incorporating predation risk, and field experiments are required. Furthermore, we need to study the waterfowl throughout their winter ranges to interpret the overall impacts of disturbance. This paper examines two cases where the impacts of disturbance have been assessed from field experiments. In one study, disturbance effects of shooting were tested by setting up experimental reserves in two Danish coastal wetlands. Over a 5-year period, these became two of the most important staging areas for coastal waterfowl, and the national totals of key species were significantly increased. A national management plan which will establish more than 50 new shooting-free refuges on Danish coastal areas within the next 5 years is likely to boost waterfowl numbers even more. Such retention of birds at more northerly sites on the fiyway should result in a more efficient resource utilization and may positively affect the population dynamics where numbers are affected by winter resources. In a second study, the impacts of disturbance by farmers on spring fattening of Pink-footed Geese Anser brachyrhynchus were analysed. In undisturbed areas in northern Norway, abdominal profiles of the geese increased rapidly, whereas in disturbed sites they did not. Subsequently, geese that had used undisturbed sites reproduced better than geese from disturbed sites.     

Effects of biological invasions on forest carbon sequestration

There has been a rapidly developing literature on the effects of some of the major drivers of global change on carbon (C) sequestration, particularly carbon dioxide (CO₂) enrichment, land use change, nitrogen (N) deposition and climate change. However, remarkably little attention has been given to one major global change driver, namely biological invasions. This is despite growing evidence that invasive species can dramatically alter a range of aboveground and belowground ecosystem processes, including those that affect C sequestration. In this review, we assess the evidence for the impacts of biological invaders on forest C stocks and C sequestration by biological invaders. We first present case studies that highlight a range of invader impacts on C sequestration in forest ecosystems, and draw on examples that involve invasive primary producers, decomposers, herbivores, plant pathogens, mutualists and predators. We then develop a conceptual framework for assessing the effects of invasive species on C sequestration impacts more generally, by identifying the features of biological invaders and invaded ecosystems that are thought to most strongly regulate C in forests. Finally we assess the implications of managing invasive species on C sequestration. An important principle that emerges from this review is that the direct effects of invaders on forest C are often smaller and shorter‐term than their indirect effects caused by altered nutrient availability, primary productivity or species composition, all of which regulate long‐term C pools and fluxes. This review provides a conceptual basis for improving our general understanding of biological invaders on ecosystem C, but also points to a paucity of primary data that are needed to determine the quantitative effects of invaders on ecosystem processes that drive C sequestration.     

Impacts of extreme climatic events on competition during grassland invasions

Phytometers of five C3 and five C4 species were transplanted into three different grasslands to study the effects of extreme climatic events on community invasibility and competition. Single extreme heating (eight hours at 52.5 °C) and rainfall (the equivalent of 100 mm) events in factorial combinations were superimposed on the grassland communities. A novel technique involving portable computer‐controlled chambers was used to create the heating events. In order to generate predictions of response to the extreme climatic events, the 10 phytometer species were categorized on the basis of 12 key plant functional traits. Using principal component analysis, two functional types (FTs) were identified as most likely to be advantaged (FT1, fast‐growing C4 annuals) and disadvantaged (FT2, slower‐growing C3 perennials) by an extreme climatic event. Competition between the resident vegetation and FT1 plus other C4 phytometers was consistently more intense within the exclusively C3 community compared to the dry C3/C4 community or moist C3/C4 community. The single extreme heating event had the greatest impact on competition, lowering the intensity of competition between the phytometers and resident vegetation. Our results indicate that competition is highly important in limiting the invasion of C3 grasslands by C4 species. The FT1 and FT2 responses confirmed predictions based on plant functional traits, whether growing as phytometers or as part of the resident vegetation. Future increases in climatic variability and the incidence of extreme climatic events are expected to suppress C3 competitive dominance and promote invasion of C4 species, in particular, the FT1 species.     

Evolutionary indirect effects of biological invasions

Just as ecological indirect effects can have a wide range of consequences for community structure and ecosystem function, theory suggests that evolutionary indirect effects can also influence community dynamics and the outcome of species interactions. There is little empirical evidence documenting such effects, however. Here, I use a multi-generation selection experiment in the field to investigate: (1) how the exotic plant Medicago polymorpha and the exotic insect herbivore Hypera brunneipennis affect the evolution of anti-herbivore resistance traits in the native plant Lotus wrangelianus and (2) how observed Lotus evolutionary responses to Hypera alter interactions between Lotus and other members of the herbivore community. In one of two study populations, I document rapid evolutionary changes in Lotus resistance to Hypera in response to insecticide treatments that experimentally reduced Hypera abundance, and in response to Medicago-removal treatments that also reduced Hypera abundance. These evolutionary changes in response to Hypera result in reduced attack by aphids. Thus, an evolutionary change caused by one herbivore species alters interactions with other herbivore taxa, an example of an eco-evolutionary feedback. Given that many traits mediate interactions with multiple species, the effects of evolutionary changes in response to one key biotic selective agent may often cascade through interaction webs to influence additional community members.     

The fungal dimension of biological invasions

Fungi represent an essential component of biodiversity, not only because of the large number of species, but also for their ecological, evolutionary and socio-economic significance. Yet, until recently, fungi received scant consideration in ecology, especially invasion ecology. Their under-representation is largely the result of a lack of scientific knowledge of fungal biodiversity and ecology. With the exception of pathogenic fungi, which cause emergent infectious diseases, the impact of fungal invasions is often difficult to quantify owing to limited baseline data on fungal communities. Here, we aim to raise awareness among mycologists and ecologists of the fungal dimension of invasions and of the need to intensify research in fungal ecology to address issues of future introductions.     

An evolutionary perspective of biological invasions

The workshop on the Evolutionary Perspective of Biological Invasions in Terrestrial Ecosystems was held in Halle, Germany from 30 September to 3 October 2002.     

The evolutionary consequences of biological invasions

A major challenge of invasion biology is the development of a predictive framework that prevents new invasions. This is inherently difficult because different biological characteristics are important at the different stages of invasion: opportunity/transport, establishment and spread. Here, we draw from recent research on a variety of taxa to examine the evolutionary causes and consequences of biological invasions. The process of introduction may favour species with characteristics that promote success in highly disturbed, human-dominated landscapes, thus exerting novel forms of selection on introduced populations. Moreover, evidence is accumulating that multiple introductions can often be critical to the successful establishment and spread of introduced species, as they may be important sources of genetic variation necessary for adaptation in new environments or may permit the introduction of novel traits. Thus, not only should the introduction of new species be prevented, but substantial effort should also be directed to preventing the secondary introduction of previously established species (and even movement of individuals among introduced populations). Modern molecular techniques can take advantage of genetic changes postintroduction to determine the source of introduced populations and their vectors of spread, and to elucidate the mechanisms of success of some invasive species. Moreover, the growing availability of genomic tools will permit the identification of underlying genetic causes of invasive success.     

对生物入侵的几点思考

生物入侵涉及异地迁徙及建群等复杂问题,涉及个体、种群、群落、生态系统等生态学的各个层次,又必不可少地要深入地去探索生物入侵过程中的快速进化和分子生态学基础。生物系统的超级复杂性加上环境和人为因素的极度随机性,充分显示了在这一领域从事研究的艰巨性和挑战性。入侵种具有两面性,因此,预防比治理更为关键。为了应对生物入侵,有待建立完善的国家和地区的防控系统、提高技术水平并加强有关诸多经济因素对生物入侵的影响的研究。     

Speciation genetics of biological invasions with hybridization

The negative effects of human‐induced habitat disturbance and modification on multiple dimensions of biological diversity are well chronicled ( Turner 1996 ; Harding et al. 1998 ; Lawton et al. 1998 ; Sakai et al. 2001 ). Among the more insidious consequences is secondary contact between formerly allopatric taxa ( Anderson & Hubricht 1938 ; Perry et al. 2002 ; Seehausen 2006 ). How the secondary contact will play out is unpredictable ( Ellstrand et al. 2010 ), but if the taxa are not fully reproductively isolated, hybridization is likely, and if the resulting progeny are fertile, the eventual outcome is often devastating from a conservation perspective ( Rhymer & Simberloff 1996 ; Wolf et al. 2001 ; McDonald et al. 2008 ). In this issue of Molecular Ecology, Steeves et al. (2010) present an analysis of hybridization between two avian species, one of which is critically endangered and the other of which is invasive. Their discovery that the endangered species has not yet been hybridized to extinction is promising and not what one would necessarily expect from theory.     

How to manage biological invasions under globalization

Protecting national borders against biological invasions is becoming increasingly difficult because those whose actions result in invasions seldom bear legal responsibility for those actions. Invasion costs are often an externality (an unintended side effect) of international trade. Externalities are best dealt with by internalizing them; that is, by getting those who harm society to meet the cost. This is the 'polluter pays principle', which, under current trade rules, is difficult to implement. Tariffs could, however, be used to confront exporters with the costs of their actions, and the right to do this should be embedded in trade agreements. At the same time, international aid could be used to protect donor societies against the inability of some other countries to take appropriate biosecurity measures. The impact of invasions can thus be reduced by tackling their economic externalities.     

Exploiting Allee effects for managing biological invasions

Biological invasions are a global and increasing threat to the function and diversity of ecosystems. Allee effects (positive density dependence) have been shown to play an important role in the establishment and spread of non-native species. Although Allee effects can be considered a bane in conservation efforts, they can be a benefit in attempts to manage non-native species. Many biological invaders are subject to some form of an Allee effect, whether due to a need to locate mates, cooperatively feed or reproduce or avoid becoming a meal, yet attempts to highlight the specific exploitation of Allee effects in biological invasions are surprisingly unprecedented. In this review, we highlight current strategies that effectively exploit an Allee effect, and propose novel means by which Allee effects can be manipulated to the detriment of biological invaders. We also illustrate how the concept of Allee effects can be integral in risk assessments and in the prioritization of resources allocated to manage non-native species, as some species beset by strong Allee effects could be less successful as invaders. We describe how tactics that strengthen an existing Allee effect or create new ones could be used to manage biological invasions more effectively.     

Modeling biological invasions into periodically fragmented environments

Range expansion of a single species in a regularly striped environment is studied by using an extended Fisher model, in which the rates of diffusion and reproduction periodically fluctuate between favorable and unfavorable habitats. The model is analyzed for two initial conditions: the initial population density is concentrated on a straight line or at the origin. For each case, we derive a mathematical formula which characterizes the spatio-temporal pattern of range expansion. When initial distribution starts from a straight line, it evolves to a traveling periodic wave (TPW), whose frontal speed is analytically determinable. When the range starts from the origin, it tends to expand radially at a constant average speed in each direction (ray speed) keeping its frontal envelope in a similar shape. By examining the relation between the ray speed and the TPW speed, we derive the ray speed in a parametric form, from which the envelope of the expanding range can be predicted. Thus we analyze how the pattern and speed of the range expansion are affected by the pattern and scale of fragmentation, and the qualities of favorable and unfavorable habitats. The major results include: (1). The envelope of the expanding range show a variety of patterns, nearly circular, oval-like, spindle-like, depending on parameter values; (2). All these patterns are elongated in the direction of stripes; (3). When the scale of fragmentation is enlarged without changing the relative spatial pattern, the ray speed in any direction increases, i.e., the rate of range expansion increases.     

A proposed unified framework for biological invasions

There has been a dramatic growth in research on biological invasions over the past 20 years, but a mature understanding of the field has been hampered because invasion biologists concerned with different taxa and different environments have largely adopted different model frameworks for the invasion process, resulting in a confusing range of concepts, terms and definitions. In this review, we propose a unified framework for biological invasions that reconciles and integrates the key features of the most commonly used invasion frameworks into a single conceptual model that can be applied to all human-mediated invasions. The unified framework combines previous stage-based and barrier models, and provides a terminology and categorisation for populations at different points in the invasion process.     

Dispersal polymorphism and the speed of biological invasions

The speed at which biological range expansions occur has important consequences for the conservation management of species experiencing climate change and for invasion by exotic organisms. Rates of dispersal and population growth are known to affect the speed of invasion, but little is known about the effect of having a community of dispersal phenotypes on the rate of range expansion. We use reaction-diffusion equations to model the invasion of a species with two dispersal phenotypes into a previously unoccupied landscape. These phenotypes differ in both their dispersal rate and population growth rate. We find that the presence of both phenotypes can result in faster range expansions than if only a single phenotype were present in the landscape. For biologically realistic parameters, the invasion can occur up to twice as fast as a result of this polymorphism. This has implications for predicting the speed of biological invasions, suggesting that speeds cannot just be predicted from looking at a single phenotype and that the full community of phenotypes needs to be taken into consideration.     

Resource-dependent dispersal and the speed of biological invasions

Many mobile organisms exhibit resource-dependent movement in which movement rates adjust to changes in local resource densities through changes in either the probability of moving or the distance moved. Such changes may have important consequences for invasions because reductions in resources behind an invasion front may cause higher dispersal while simultaneously reducing population growth behind the front and thus lowering the number of dispersers. Intuiting how the interplay between population growth and dispersal affects invasions is difficult without mathematical models, yet most models assume dispersal rates are constant. Here we present spatial-spread models that allow for consumer-resource interactions and resource-dependent dispersal. Our results show that when resources affect the probability of dispersal, then the invasion dynamics are no different than if resources did not affect dispersal. When resources instead affect the distance dispersed, however, the invasion dynamics are strongly affected by the strength of the consumer-resource interaction, and population cycles behind the wave front lead to fluctuating rates of spread. Our results suggest that for actively dispersing invaders, invasion dynamics can be determined by species interactions. More practically, our work suggests that reducing invader densities behind the front may be a useful method of slowing an invader's rate of spread.