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Abstract Background It has been suggested that climate change will lead to increased environmental fluctuations, which will undoubtedly have evolutionary consequences for all biota. For instance, fluctuations can directly increase the risk of invasions of alien species into new areas, as these species have repeatedly been proposed to benefit from disturbances. At the same time increased environmental fluctuations may also select for better invaders. However, selection by fluctuations may also influence the resistance of communities to invasions, which has rarely been tested.
We tested eco-evolutionary dynamics of invasion with bacterial clones, evolved either in constant or fluctuating temperatures, and conducted experimental invasions in both conditions. Results We found clear evidence that ecological fluctuations, as well as adaptation to fluctuations by both the invader and community, all affected invasions, but played different roles at different stages of invasion.
Ecological fluctuations clearly promoted invasions, especially into fluctuation mal-adapted communities. The evolutionary background of the invader played a smaller role.
Conclusions Our results indicate that climate change associated disturbances can directly increase the risk of invasions by altering ecological conditions during invasions, as well as via the evolution of both the invader and communities.
Our experiment provides novel information on the complex consequences of climate change on invasions in general, and also charts risk factors associated with the spread of environmentally growing opportunistic pathogens. Background Current climate change scenarios predict that in addition to the increase in temperature, fluctuations in temperature and other environmental conditions are also increasing [ 1 ] and creating selection pressures for biota.
Some species benefit from these changes, and invasions and range expansions have been documented for many taxa [ 2 — 4 ]. In particular, fluctuations in environmental conditions might lead to the evolution of invasive genotypes [ 5 ] aiding species invasions. Thus, global climate change, with increased environmental fluctuations, could bring evolutionary and ecological problems to native fauna and flora: they must cope both with the direct changes caused by environmental fluctuations and with freshly-evolved invasive genotypes.
This could be especially disastrous if native communities are mal-adapted to fluctuations. One possible evolutionary explanation for the emergence of invasive species and genotypes is that they have evolved in a disturbed and fluctuating environment.
It has been suggested that rapid fluctuations in particular, can select for traits that could promote invasion success, such as high population growth rate, plasticity and persistence [ 5 — 8 ]. Climate change has been suggested to lead to increased extreme events e.
If a community is adapted to fluctuations, environmental fluctuations should not cause repercussions in population size and hence benefit invaders. Although it can be argued that all environments are potentially prone to invasions, significant variation exists in the sensitivity of different environments to invasions.
Empirical evidence supports the theory that heterogeneous both in space and time and disturbed environments are more prone to invasions than stable environments [ 11 — 15 ]. Disturbances might facilitate invasions by altering community composition, species competitive interactions, free resources, ecosystem processes, and propagule supply [ 10 ]. Recent research has also recognized that invasions are interactions between the invader and the environment invaded, and as such the invasive traits might only make sense in certain environments [ 16 — 18 ].
Since disturbed environments can promote the evolution of invasive traits [ 5 ], studying the evolutionary background and pre-adaptations of the invader together with the effects of the current environment can also generate important information on the causes of invasions, especially in the context of current climate change [ 17 ].
For example; will the increased fluctuations predispose communities to an increased risk of invasions, and will the fluctuation-adapted invaders invade such environments with even greater likelihood, as is suggested in the anthropogenically induced adaptation to invade — hypothesis AIAI; [ 19 ]?
To test these key ideas about the ecological and evolutionary determinants of invasions [ 17 ], we used several species of microbes that had evolved for 2. These species and strains allowed us to make unique combinations of the invader Serratia marcescens and the community Pseudomonas chlororaphis, Enterobacter aerogenes and Leclercia adecarboxylata in which evolutionary adaptations either matched, or not, the environmental conditions during the invasions.
With microbes it is straightforward to test general ecological and evolutionary theories, which are not amenable to testing with higher organisms e. Consequently, microbial experiments have also become more popular in invasion biology [ 21 — 27 ]. However, the effects of environmental fluctuations, other than in resources, on invasion success [ 12 , 23 ] have rarely been tested [ 28 ].
The invader species S. Hence, our experiment provides also an important test on the determinants of spread of pathogenic bacteria and diseases facing climate change induced fluctuations.