No evidence for local adaptation in an invasive alien plant: field and greenhouse experiments tracing a colonization sequence

Background and Aims

Local adaptation enables plant species to persist under different environmental conditions. Evolutionary change can occur rapidly in invasive annual species and has been shown to lead to local adaptation. However, the patterns and mechanisms of local adaptation in invasive species along colonization sequences are not yet understood. Thus, in this study the alien annual Impatiens glandulifera was used to investigate local adaptation to distinct habitats that have been consecutively invaded in central Europe.

Methods

A reciprocal transplant experiment was performed using 15 populations from alluvial deciduous forests, fallow meadows and coniferous upland forests, and a greenhouse experiment was performed in which plants from these habitats were grown under treatments reflecting the main habitat differentiators (shade, soil acidity, competition).

Key Results

Biomass production, specific leaf area, plant height and relative growth rate differed between habitats in the field experiment and between treatments in the greenhouse, but not between seed origins. Overall, there was no indication of local adaptation in either experiment.

Conclusions

Since I. glandulifera is a successful invader in many habitats without showing local adaptation, it is suggested that the species is coping with environmental variation by means of high phenotypic plasticity. The species seems to follow a ‘jack-and-master’ strategy, i.e. it is able to maintain high fitness under a wide range of environmental conditions, but performs particularly well in favourable habitats. Therefore, the proposed colonization sequence is likely to be based primarily on changes in propagule pressure. It is concluded that invasive alien plants can become dominant in distinct habitats without local adaptation.     

No evidence for local adaptation and an epigenetic underpinning in native and non‐native ruderal plant species in Germany

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The importance of biotic interactions and local adaptation for plant response to environmental changes: field evidence along an elevational gradient

Predicting the response of species to environmental changes is a great and on‐going challenge for ecologists, and this requires a more in‐depth understanding of the importance of biotic interactions and the population structuration in the landscape. Using a reciprocal transplantation experiment, we tested the response of five species to an elevational gradient. This was combined to a neighbour removal treatment to test the importance of local adaptation and biotic interactions. The trait studied was performance measured as survival and biomass. Species response varied along the elevational gradient, but with no consistent pattern. Performance of species was influenced by environmental conditions occurring locally at each site, as well as by positive or negative effects of the surrounding vegetation. Indeed, we observed a shift from competition for biomass to facilitation for survival as a response to the increase in environmental stress occurring in the different sites. Unlike previous studies pointing out an increase of stress along the elevation gradient, our results supported a stress gradient related to water availability, which was not strictly parallel to the elevational gradient. For three of our species, we observed a greater biomass production for the population coming from the site where the species was dominant (central population) compared to population sampled at the limit of the distribution (marginal population). Nevertheless, we did not observe any pattern of local adaptation that could indicate adaptation of populations to a particular habitat. Altogether, our results highlighted the great ability of plant species to cope with environmental changes, with no local adaptation and great variability in response to local conditions. Our study confirms the importance of taking into account biotic interactions and population structure occurring at local scale in the prediction of communities’ responses to global environmental changes.     

Geographical variation in germination traits of the salt-marsh cordgrass Spartina alterniflora in its invasive and native ranges

入侵地和原产地盐沼植物互花米草种子萌发性状的地理变异 种子萌发是植物早期生活史中最重要的阶段,决定了植物的生态位和地理分布范围,对外来植物的入侵潜力有重要影响。盐沼植物互花米草(Spartina alterniflora)在中国沿海滩涂的入侵范围最大,并已入侵到比原产地更低的纬度范围,这为我们研究互花米草在不同地理区域之间以及沿纬度梯度的萌发性状差异和适应提供了契机。在控温培养箱中淡水培养条件下,我们比较研究了来自入侵地(19°–40° N)10 个地点和原产地(27°–43° N)16个地点不同纬度互花米草种群的种子萌发性状差异,以及这种差异与各种 群来源地潮差和气候因素的相关性。原产地互花米草种群种子的萌发率和萌发指数比入侵地种群分别高10%和20%,但入侵地互花米草种群的萌发速度比原产地快3 d。入侵地互花米草种群的萌发率和萌发 指数随着纬度升高呈现线性递增的变化趋势,而原产地呈现线性递减的变化趋势。入侵地和原产地互花米草种群的平均萌发时间都与纬度呈现线性负相关。入侵地互花米草种群的萌发率和萌发指数与年日均温、年日最低均温、和年日最高均温呈现负相关,而在原产地呈现相反的相关关系。入侵地和原产地互花米草种群的平均萌发时间分别与年日均温、年日最低均温和年日最高均温呈现正相关关系。我们的研究结果表明,入侵地和原产地互花米草种群的萌发率和萌发指数已沿纬度进化出不同的渐变群格局,但平均萌发时间进化出与原产地一致的纬度渐变群格局,即在生物入侵过程中沿纬度梯度种子萌发策略会随着入侵时间和过程而发生变化。     

Assessing the speed and predictability of local adaptation in invasive California poppies (Eschscholzia californica)

Insight into the speed and predictability of local adaptation can be gained by studying organisms, such as invasive species, that have recently expanded their geographical ranges. Common garden studies were designed to address these issues with the California poppy, Eschscholzia californica, collected from a wide range of environments in both its native (California) and invasive (Chile) ranges. We found similar patterns of plant trait variation along similar abiotic gradients in plants collected from both areas. Multivariate analysis demonstrated that coastal plants from both areas tended to be shorter, smaller plants with smaller seeds and flowers that germinate and flower later than plants collected from inland locations. In addition, size and fecundity traits in both native and invasive poppies were correlated with average rainfall totals; the plants that grew the largest and were the most fecund during the first year of growth originated from the driest areas. This parallel variation suggests that these traits are adaptive and that these patterns have evolved in Chile during the 110-150 years since introduction.     

土著昆虫素毒蛾在本地植物芦苇与入侵植物互花米草上的生活史

互花米草(Spartina alterniflora)入侵盐沼生态系统导致了土著广食性昆虫素毒蛾(Laelia coenosa)发生宿主转移。但是, 目前对素毒蛾在互花米草和芦苇(Phragmites australis)上的生活史规律的比较尚未见报道, 而此信息对评价素毒蛾能否持续利用互花米草具有重要的参考价值。为了明确素毒蛾在其原始宿主芦苇和外来新宿主互花米草上的生活史动态, 我们在上海崇明长江口盐沼湿地中, 对素毒蛾在芦苇和互花米草生境中的发生情况进行了系统调查。结果显示, 虽然素毒蛾在芦苇和互花米草上均一年发生3代, 并都以幼虫越冬, 但在互花米草生境中, 越冬代幼虫开始越冬的时间要比芦苇中晚20 d左右, 其他世代中的多个虫态在互花米草上持续出现的时间也比芦苇上延迟了约10 d。这些结果表明, 互花米草入侵后素毒蛾在外来植物上出现的持续时间虽然发生了延长, 但其年世代数未变化。我们推测素毒蛾在互花米草和芦苇上生长发育规律的差异可能受生境的资源可利用性及两种植物的营养与防御水平调控。     

Changes in the realized niche of the invasive succulent CAM plant Furcraea foetida

Furcraea foetida (Asparagaceae) is a native plant of Central America and northern South America but there is no information about its country of origin. The species was introduced into Brazil and is now considered invasive, particularly in coastal ecosystems. To date, nothing is known about the environmental factors that constrain its distribution and there is only inconclusive information about its location of origin. We used reciprocal distribution models (RDM) to assess invasion risk of F. foetida across Brazil and to identify source regions in its native range. We also tested the niche conservatism hypothesis using Principal Components Analyses and statistical tests of niche equivalency and similarity between its native and invaded ranges. For RDM analysis, we built two models using maximum entropy, one using records in the native range to predict the invaded distribution (forward‐Ecological Niche Model or forward‐ENM) and one using records in the invaded range to predict the native distribution (reverse‐ENM). Forward‐ENM indicated invasion risk in the Cerrado region and the innermost region of the Atlantic Forest, however, failed to predict the current occurrence in southern Brazil. Reverse‐ENM supported an existing hypothesis that F. foetida originated in the Orinoco river basin, Amazon basin and Caribbean islands. Prediction errors in the RDM and multivariate analysis indicated that the species expanded its realized niche in Brazil. The niche similarity test further suggested that the niche differences are because of differences in habitat availability between the two ranges, not because of evolutionary changes. We hypothesize that physiological pre‐adaptation (especially, the crassulacean acid metabolism), human‐driven propagule pressure and high competitive ability are the main factors determining the current spatial distribution of the species in Brazil. Our study highlights the need to include F. foetida in plant invasion monitoring programs, especially in priority conservation areas where the species has still not been introduced.     

Stress avoidance in a common annual: reproductive timing is important for local adaptation and geographic distribution

Adaptation to local environments may be an important determinant of species' geographic range. However, little is known about which traits contribute to adaptation or whether their further evolution would facilitate range expansion. In this study, we assessed the adaptive value of stress avoidance traits in the common annual Cocklebur (Xanthium strumarium) by performing a reciprocal transplant across a broad latitudinal gradient extending to the species' northern border. Populations were locally adapted and stress avoidance traits accounted for most fitness differences between populations. At the northern border where growing seasons are cooler and shorter, native populations had evolved to reproduce earlier than native populations in the lower latitude gardens. This clinal pattern in reproductive timing corresponded to a shift in selection from favouring later to earlier reproduction. Thus, earlier reproduction is an important adaptation to northern latitudes and constraint on the further evolution of this trait in marginal populations could potentially limit distribution.     

Limits to local adaptation in six populations of the annual plant Diodia teres

* Local adaptation is common, but tests for adaptive differentiation frequently compare populations from strongly divergent environments, making it unlikely that any influence of stochastic processes such as drift or mutation on local adaptation will be detected. Here, the hypothesis that local adaptation is more likely to develop when the native environments of populations are more distinct than when they are similar was tested. * A reciprocal transplant experiment including two populations from each of three habitats was conducted to determine the pattern of local adaptation. In addition to testing for local adaptation at the population level, the hypothesis was tested that local adaptation is more common between populations from different habitats than between populations from the same habitat. * Local adaptation was not common, but more evidence was found of local adaptation between populations from different habitats than between populations from the same habitat. Two instances of foreign genotype fitness advantage confirm that stochastic processes such as drift can limit local adaptation. * These results are consistent with the hypothesis that stochastic processes can inhibit local adaptation but are more likely to be overwhelmed by natural selection when populations occur in divergent environments.     

Establishment,spread, and impact of an invasive planthopper on its invasive host plant: Prokelisia marginata (Homoptera: Delphacidae) exploiting Spartina anglica (Poales: Poaceae) in Britain

1. Since its recent arrival in Britain, the planthopper Prokelisia marginata has spread widely around saltmarshes on the east and south coast of England and south Wales, feeding on Common Cordgrass, Spartina anglica, itself an invasive non-native species. 2. Results suggest that P. marginata populations in Britain benefit from a degree of natural enemy release. No evidence of parasitism was found in over 71 000 eggs, nymphs, and adults inspected. The only potential natural enemy control was suggested by a positive correlation between the densities of planthoppers and generalist spiders. 3. Experimental exposure under both glasshouse and field conditions to typical field densities of planthoppers resulted in significant negative effects on a number of host plant performance metrics. 4. Spartina anglica is important for stabilising estuarine sediments and has been deliberately planted for this purpose in the past. Its weakening as a result of heavy planthopper herbivory could have serious consequences for the long-term stability of Britain's vulnerable saltmarsh habitats.     

Methods to test the interactive effects of drought and plant invasion on ecosystem structure and function using complementary common garden and field experiments

Abiotic global change drivers affect ecosystem structure and function, but how they interact with biotic factors such as invasive plants is understudied. Such interactions may be additive, synergistic, or offsetting, and difficult to predict. We present methods to test the individual and interactive effects of drought and plant invasion on native ecosystems. We coupled a factorial common garden experiment containing resident communities exposed to drought (imposed with rainout shelters) and invasion with a field experiment where the invader was removed from sites spanning a natural soil moisture gradient. We detail treatments and their effects on abiotic conditions, including soil moisture, light, temperature, and humidity, which shape community and ecosystem responses. Ambient precipitation during the garden experiment exceeded historic norms despite severe drought in prior years. Soil moisture was 48% lower in drought than ambient plots, but the invader largely offset drought effects. Additionally, temperature and light were lower and humidity higher in invaded plots. Field sites spanned up to a 10‐fold range in soil moisture and up to a 2.5‐fold range in light availability. Invaded and resident vegetation did not differentially mediate soil moisture, unlike in the garden experiment. Herbicide effectively removed invaded and resident vegetation, with removal having site‐specific effects on soil moisture and light availability. However, light was generally higher in invader‐removal than control plots, whereas resident removal had less effect on light, similar to the garden experiment. Invasion mitigated a constellation of abiotic conditions associated with drought stress in the garden experiment. In the field, where other factors co‐varied, these patterns did not emerge. Still, neither experiment suggested that drought and invasion will have synergistic negative effects on ecosystems, although invasion can limit light availability. Coupling factorial garden experiments with field experiments across environmental gradients will be effective for predicting how multiple stressors interact in natural systems.     

Oviposition strategy as a means of local adaptation to plant defence in native and invasive populations of the viburnum leaf beetle

Herbivores have been hypothesized to adapt locally to variation in plant defences and such adaptation could facilitate novel associations in the context of biological invasions. Here, we show that in the native range of the viburnum leaf beetle (VLB, Pyrrhalta viburni), two populations of geographically isolated hosts-Viburnum opulus and Viburnum tinus-have divergent defences against VLB oviposition: negative versus positive density-dependent egg-crushing wound responses, respectively. Populations of beetles coexisting with each host show an adaptive behavioural response: aggregative versus non-aggregative oviposition on V. opulus and V. tinus, respectively. In parallel, we show that in North America, where VLB is invasive, defences of three novel hosts are negatively density-dependent, and beetles' oviposition behaviour is aggregative. Thus, local adaptation to plant defences has the potential to facilitate the invasion of herbivores onto novel hosts.     

Cuticular melanisation and immune response in a butterfly: local adaptation and lack of correlation

1. Despite growing knowledge on the relationship between ecological variables and individual immune function, data on the spatial variability of immune defence in invertebrate natural populations are scarce. 2. Here, we use replicated populations of the butterfly Lycaena tityrus from different altitudes to investigate genetic variation in the melanin‐based encapsulation response. As high‐ and low‐altitude populations differ in cuticular pupal melanisation, we further tested for any associations between pupal melanisation and parasite resistance. 3. Although pupal melanisation was higher at higher compared with lower altitudes (and at a higher compared with a lower rearing temperature), any obvious relations to the encapsulation response were absent. Further phenotypic correlations within groups were significant in one out of four cases only, suggesting that in L. tityrus encapsulation operates largely independent of cuticular melanisation. 4. A significant interaction between altitude and temperature indicated that high‐altitude animals show a stronger melanisation response than low‐altitude ones at the lower temperature and vice versa, indicating local adaptation to different climates.     

Enhanced fitness due to higher fecundity,increased defence against a specialist and tolerance towards a generalist herbivore in an invasive annual plant

Aims The superior performance of many non-indigenous species in a new range can be attributed to different factors such as pre-adaptation to environmental conditions in new areas or to factors inherent to displacement mechanisms such as loss of co-evolved pathogens and herbivores that increase the speed of evolutionary change towards a shift in allocation from defence to growth and reproduction. To assess the importance of the different mechanisms governing the success of Conyza canadensis, a globally successful invader, we simultaneously tested several recent hypotheses potentially explaining the factors leading to biological invasion.Methods We tested (i) whether plants from the non-native range showed a higher fitness than plants from the native North American range, (ii) whether they differed in resistance against an invasive generalist herbivore, the slug Arion lusitanicus and against a recently established specialist aphid herbivore, Uroleucon erigeronense and (iii) experimentally assessed whether C. canadensis releases allelopathic chemicals that have harmful effects on competing species in the non-native range. We compared populations along a similar latitudinal gradient both in the native North American and invasive European range and analysed patterns of adaptive clinal variation in biomass production.Important findings The invasion success of C. canadensis in Europe cannot be attributed to a single trait, but to a combination of factors. Invasive plants benefited from increased growth and above all, increased reproduction (a key trait in an annual plant) and were less attacked by a co-migrated specialist enemy. The observed loss of defence against generalist slugs did not translate into a decreased fitness as invasive C. canadensis plants showed a high re-growth potential. In contrast to earlier in vitro studies, we detected no allelopathic effects on the competing flora in the non-native range. The latitudinal cline in vegetative biomass production in the non-native range observed in our common garden study indicates a high adaptive potential. However, only further genetic studies will provide conclusive evidence whether the differentiation in the non-native range is caused by pre-adaptation and sorting-out processes of putatively repeatedly introduced populations of this composite, long-distance disperser with highly volatile seeds or evolved de novo as a rapid response to new selection pressures in the non-native range.     

Effects of Rhinoncomimus latipes on the growth and reproduction of Persicaria perfoliata,an invasive plant in North America

Native to Asia, mile-a-minute Persicaria perfoliata, is an invasive weed in North America, and the weevil Rhinoncomimus latipes is a host-specific insect agent which occurs widely in China. We conducted a common garden experiment to compare P. perfoliata plant responses of native and invasive populations to herbivory by the weevils from different origins. We found weevils from Hunan, Hubei and Heilongjiang Provinces had strong, moderate and weak ability to suppress host plant, respectively. Weevils from Hunan and Hubei Provinces had stronger impact on the growth of both native and invasive plant populations than the weevil from Heilongjiang Province. The losses in seed output of invasive plants were also significantly greater than natives in the weevil treatments. Our results suggested that the weevil population from Hunan Province may be the most suitable for the control of mile-a-minute, while the population from Heilongjiang Province may be the least suitable due to climate matching.     

Rapid genetic adaptation precedes the spread of an exotic plant species

Human activities have increasingly introduced plant species far outside their native ranges under environmental conditions that can strongly differ from those originally met. Therefore, before spreading, and potentially causing ecological and economical damage, non‐native species may rapidly evolve. Evidence of genetically based adaptation during the process of becoming invasive is very scant, however, which is due to the lack of knowledge regarding the historical genetic makeup of the introduced populations and the lack of genomic resources. Capitalizing on the availability of old non‐native herbarium specimens, we examined frequency shifts in genic SNPs of the Pyrenean Rocket (Sisymbrium austriacum subsp. chrysanthum), comparing the (i) native, (ii) currently spreading non‐native and (iii) historically introduced gene pool. Results show strong divergence in flowering time genes during the establishment phase, indicating that rapid genetic adaptation preceded the spread of this species and possibly assisted in overcoming environmental constraints.     

The role of intraspecific competition in the dispersal of an invasive fish

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The chestnut blight fungus world tour: successive introduction events from diverse origins in an invasive plant fungal pathogen

Clonal expansion has been observed in several invasive fungal plant pathogens colonizing new areas, raising the question of the origin of clonal lineages. Using microsatellite markers, we retraced the evolutionary history of introduction of the chestnut blight fungus, Cryphonectria parasitica, in North America and western Europe. Combining discriminant analysis of principal components and approximate Bayesian computation analysis, we showed that several introduction events from genetically differentiated source populations have occurred in both invaded areas. In addition, a low signal of genetic recombination among different source populations was suggested in North America. Finally, two genetic lineages were present in both invaded areas as well as in the native areas, suggesting the existence of genetic lineages with a high capacity to establish in diverse environments and host species. This study confirmed the importance of multiple introductions, but questioned the role of genetic admixture in the success of introduction of a fungal plant pathogen.     

Lack of local adaptation of invasive crofton weed (Ageratina adenophora) in different climatic areas of Yunnan Province,China

Aims Crofton weed, with a subtropical origin, has successfully invaded in diverse habitats that belong to different climate zones in Southwest China. We tested whether local adaptation plays an important role in the successful invasion of crofton weed in heterogeneous environments.Methods Five populations from different habitats with an altitude ranging from 678 to 2356 m were selected. Plant height, biomass, seed yield and seed germination capability of these populations were investigated in the field. Greenhouse and reciprocal transplant experiments with the five populations were conducted, and all the above characters were measured and compared among these populations.Important findings Plant height, biomass, seed yield and seed germination rate were each significantly different among the five populations in field. However, there was no difference among these populations in the greenhouse experiment. In the reciprocal transplant experiment, plants from the five populations responded similarly to different habitats in the field, indicating lack of local adaptation. Instead, phenotypic plasticity likely plays a key role in the invasion success of crofton weed in different habitats.