Priority effects vary with species identity and origin in an experiment varying the timing of seed arrival

Exotic species are sometimes phenologically distinct from native species in the invaded community, allowing them to be active when there may be reduced competition for resources. In southern California, annual species are particularly problematic invaders, and prior work has shown that these species germinate earlier in the growing season, giving them a competitive advantage over later‐germinating native species. This result begs the question, if being active earlier is advantageous, why have not native species adapted earlier cues for germination? We hypothesized native species would benefit less from earlier germination than exotic species (potentially due to slower growth following germination), thus negating potential selection for early germination. Here we manipulated planting time for common native and exotic species, growing them in all possible species pairs, to evaluate how competitive outcomes were altered by the time of arrival and the origin of competing species. In contrast to our hypotheses, the exotic species often had lower biomass when planted first, potentially due to disturbance when the second species was planted. In contrast, three out of our four native species benefited from earlier planting (a priority effect). Unlike the potential benefit of arriving early, we found no evidence that being planted one week later resulted in a competitive disadvantage, when compared to being planted simultaneously with a competitor. Further, we found that the magnitude and even direction of priority effects varied depending on the identity of the interacting species. Together these results suggest that a lack of directional selection may prevent adaptation towards earlier germination times of native species. Although this experiment was conducted with a limited suite of species, the results show that the role of seasonal priority effects varies among species, and that native species could benefit from seasonal priority effects in restoration efforts even when in competition with fast‐growing exotic annual species.     

Differentiation between native and exotic plant species from a dry grassland: fundamental responses to resource availability,and growth rates

Abstract Species from a mixed native/exotic dry grassland community in New Zealand were experimentally assessed for three ecophysiological parameters: nutrient response, water response and maximum relative growth rate (RGR_max). These are parameters that relate to factors proposed as important in structuring plant communities in dry environments. Native and exotic species did not differ consistently in water response. Exotic species tended to have a greater response to nutrients, but there was considerable overlap between native and exotic guilds. However, exotic species did have a higher intrinsic growth rate, and this effect was not attributable to differences in life histories. The results suggest that the exotic species are more competitive and more generalist than the native species. These traits are compatible with the concept of the ‘ideal invader’, and suggest the C‐R strategy of Grime’s theory. The native species showed characteristics consistent with stress tolerance (sensu Grime). The paucity of evidence for ecophysiological differentiation between the native and exotic guilds, except in intrinsic growth rate, indicates that the exotic species were able to invade not because they had superior adaptation to the physical environment, but because they possessed, by pre‐adaptation, the same ways of coping with that environment as the existing species. However, their ability to invade can be related to their growth rates.     

Ingestion by an endemic frugivore enhances seed germination of endemic plant species but decreases seedling survival of exotics

Aim To test whether ingestion by endemic frugivores differentially affects the seed germination time, germination percentage and seedling survival of endemic, native and exotic fleshy fruited plant species, and to identify the principal processes and attributes driving such effects. Location Round Island, Mauritius. Methods We conducted a germination and seedling survival experiment for 3 months to test whether ingestion (gut passage and deposition in faeces) by the endemic Telfair’s skink (Leiolopisma telfairii) had a differential effect on the germination time, germination percentage and seedling survival of two endemic, four native and two exotic fleshy fruited plant species. To assess the importance of factors involved in the ingestion process, we used a factorial design with gut passage (gut‐passed vs. not gut‐passed), depulping (whole fruit vs. manually depulped seed) and the presence of faecal material (faeces vs. without faeces). In addition, the roles of species‐specific traits, seed size and deposition density (average number of seeds per faeces) were examined. Results Exotic species had a higher germination percentage than indigenous (native and endemic) species when not ingested. Following skink ingestion, there was no longer a difference, as ingestion enhanced germination percentage most in endemic species. The exotic species still germinated faster overall than the indigenous species, despite ingestion accelerating the germination time of endemics. However, ingestion strongly reduced seedling survival of the exotic species, while having no negative effect on the survival of indigenous seedlings. Overall, ingested indigenous seeds were more likely to germinate and the seedlings more likely to survive than ingested exotic seeds and seedlings. Seed size, deposition density and the removal of fruit pulp by either manual depulping or gut passage were important predictors of germination time, germination percentage and seedling survival. Main conclusions These endemic frugivores can enhance the competitiveness of endemic compared with exotic fleshy fruited plants at the critical germination and seedling establishment stage. Consequently, conservation and restoration of mutualistic endemic plant–animal interactions may be vital to mitigating the degradation of habitats invaded by exotic plants, which is of particular relevance for island ecosystems in which large numbers of endemics are threatened by exotic invaders.     

Stress relief may promote the evolution of greater phenotypic plasticity in exotic invasive species: a hypothesis

Invasion ecologists have often found that exotic invaders evolve to be more plastic than conspecific populations from their native range. However, an open question is why some exotic invaders can even evolve to be more plastic given that there may be costs to being plastic. Investigation into the benefits and costs of plasticity suggests that stress may constrain the expression of plasticity (thereby reducing the benefits of plasticity) and exacerbate the costs of plasticity (although this possibility might not be generally applicable). Therefore, evolution of adaptive plasticity is more likely to be constrained in stressful environments. Upon introduction to a new range, exotic species may experience more favorable growth conditions (e.g., because of release from natural enemies). Therefore, we hypothesize that any factors mitigating stress in the introduced range may promote exotic invaders to evolve increased adaptive plasticity by reducing the costs and increasing the benefits of plasticity. Empirical evidence is largely consistent with this hypothesis. This hypothesis contributes to our understanding of why invasive species are often found to be more competitive in a subset of environments. Tests of this hypothesis may not only help us understand what caused increased plasticity in some exotic invaders, but could also tell us if costs (unless very small) are more likely to inhibit the evolution of adaptive plasticity in stressful environments in general.     

Phenotypic plasticity of reproductive traits in response to food availability in invasive and native species of nematode

Invasive species cause severe ecological and economic damage; however, the mechanisms underlying their successful invasion often remain elusive. In the case of Bursaphelenchus xylophilus, a global quarantine pest which invaded Asia and Europe, it has been suggested that this species possesses highly competitive abilities, which promotes its establishment and rapid spread. To explore biological traits that may explain its highly competitive abilities, we focused on expression of phenotypic plasticity in response to the food conditions experienced by the females during their development as juveniles in the invasive species B. xylophilus and native species Bursaphelenchus mucronatus. We report an unexpected significant difference of phenotypic trade-off between egg number and egg size in the invasive species B. xylophilus and native species B. mucronatus. This leads to superior propagation ability of invasive species, under high and low food conditions in culture. These effects reflect adaptive optimal resource allocation where more eggs are produced in favorable environments to enhance population viability. Furthermore, we show that B. xylophilus eggs hatched earlier than B. mucronatus when their parents experienced high food availability. Thus, this study revealed, for the first time, phenotypic plasticity of reproductive traits in B. xylophilus which empowers the species a competitive advantage relative to their native counterpart B. mucronatus when they are under different range of food availability. These results are a step towards answering the vital question of how an exotic invasive species exclude a native species from its original niche.     

A Comparison of Plastic Responses to Competition by Invasive and Non-invasive Congeners in the Commelinaceae

Evidence supporting an association between phenotypic plasticity and invasiveness across a range of plant taxa is based primarily on comparisons between invasive species and native species whose potential invasiveness is typically unknown. Comparison of invasive and non-invasive exotic species would provide a better test of whether plasticity promotes invasion. Such comparisons should distinguish between adaptive and non-adaptive plasticity because they have different consequences for invasiveness. Adaptive plasticity is expected to promote the invasion of multiple habitats, but non-adaptive plasticity may reflect specialization for invading more favorable habitats only. We grew four invasive and four non-invasive species of the Commelinaceae with and without competitors and compared their putatively adaptive plasticity of three traits related to competitive ability and non-adaptive plasticity in performance. The invasive species grew significantly more than the non-invasive species only in the non-competitive environment. The invasive species had greater plasticity of performance (total biomass) in response to competition than non-invasives, but there was no consistent difference in the plasticities of the traits related to competitive ability. These results are consistent with specialization of these invasive taxa for invading the more productive non-competitive environment rather than a superior ability to invade both competitive and non-competitive environments. A comprehensive understanding of the relationship between plasticity and invasiveness will require many more comparisons of the plasticity of invasive and non-invasive taxa in a range of traits in response to a variety of environments.     

Effects of eutrophication and snails on Eurasian watermilfoil (Myriophyllum spicatum) invasion

Exotic species can invade and establish new habitats both as a result of their own traits, and as a result of the characteristics of the environment they invade. Here, we show that the abundance of the invasive submerged aquatic plant, Myriophyllum spicatum (Eurasian watermilfoil) is highly dependent on the conditions of the environment in a mesocosm experiment. M. spicatum is allelopathic towards epiphytic algae, and in the absence of algivorous snails, we found that the abundance of both algae and M. spicatum significantly increased with experimentally increased nutrient loading, while the abundance of native submerged macrophytes declined. However, when snails were present, snail biomass increased with increasing nutrient loading, and M. spicatum biomass was consistently low while native submerged macrophyte biomass was consistently high. Our results stress the importance of the interaction between species traits and environmental conditions when considering the invasiveness of certain exotic species and the invasibility of certain environments.     

Alterations in flood frequency increase exotic and native species richness of understorey vegetation in a temperate floodplain eucalypt forest

The delivery of environmental flows for biodiversity benefits within regulated river systems can potentially contribute to exotic weed spread. This study explores whether exotic plants of a floodplain forest in Victoria, Australia, are characterised by specific functional groups and associated plant traits linked to altering hydrological conditions over time. Permanently marked 20 × 20 m² plots from five wetland sites in Eucalyptus camaldulensis floodplain forest were sampled twice, first in the early 1990s (1993–1994) and then 15 years later (2007–2008). Species cover abundance data for understorey vegetation communities were segregated by season and analysed using ordination analysis. Exotic species richness was modelled as a function of site flooding history and native species richness using general linear models. Site ordinations by detrended correspondence analysis showed differential community compositions between survey dates, but native and exotic species were not clearly differentiated in terms of DCA1 scores. Most exotics belonged to functional groups containing annual species that germinate and reproduce under drier conditions. Exotics reproducing under wetter conditions were in the minority, predominantly perennial and capable of both sexual and asexual reproduction. Site flooding history and native species richness significantly predicted exotic species richness. Vegetation changes are partially structured by reduced flood frequency favouring increased abundance of exotic, sexually reproducing annuals at drier sites. Sites of low flood frequency are more sensitive to future exotic weed invasion and will require targeted management effort. Flow restoration is predicted to benefit propagule dispersal of species adopting dual regeneration strategies, which are predominantly natives in this system.     

Life history of peracarid species in South‐western Atlantic: Comparison of population traits between native and exotic species

Life‐history traits of exotic species are important to understand the process involved in their settlement and their potential impact on native biodiversity. In this context, the seasonal density, the population structure and the reproductive patterns of exotic and native peracarid species of two natural marine environments of Southwestern Atlantic were studied in order to determine the traits that favour the invasion success of exotic species. Five samples, consisted of algal patches (0.20 × 0.20 m quadrants), were collected seasonally from 2016 to 2017 in two intertidal environments (La Estafeta and Cerro Avanzado). Both environments presented high richness of cryptogenic and exotic species (Tanais dulongii, Monocorophium insidiosum, Ampithoe valida, Melita palmata and Jassa marmorata), and only two native species were recorded (Apohyale grandicornis and Exosphaeroma lanceolatum). The comparison of life‐history traits suggested that the distribution, dominance and the highest densities of some exotic species are closely related to their continuous reproductive and recruitment periods, their capability to adapt their life‐history strategies to different environmental conditions and to a more efficient distribution of resources during reproduction; however, in native species, only A. grandicornis registered similar life‐history traits than exotic populations, suggesting that their distribution could be limited by a latitudinal gradient. We expect that these results provide essential information to understand the invasion pattern of exotic species and their potential impact on native biodiversity in Southwestern Atlantic.     

Native jewelweed,but not other native species,displays post‐invasion trait divergence

Invasive exotic plants reduce the diversity of native communities by displacing native species. According to the coexistence theory, native plants are able to coexist with invaders only when their fitness is not significantly smaller than that of the exotics or when they occupy a different niche. It has therefore been hypothesized that the survival of some native species at invaded sites is due to post‐invasion evolutionary changes in fitness and/or niche traits. In common garden experiments, we tested whether plants from invaded sites of two native species, Impatiens noli‐tangere and Galeopsis speciosa, outperform conspecifics from non‐invaded sites when grown in competition with the invader (Impatiens parviflora). We further examined whether the expected superior performance of the plants from the invaded sites is due to changes in the plant size (fitness proxy) and/or changes in the germination phenology and phenotypic plasticity (niche proxies). Invasion history did not influence the performance of any native species when grown with the exotic competitor. In I. noli‐tangere, however, we found significant trait divergence with regard to plant size, germination phenology and phenotypic plasticity. In the absence of a competitor, plants of I. noli‐tangere from invaded sites were larger than plants from non‐invaded sites. The former plants germinated earlier than inexperienced conspecifics or an exotic congener. Invasion experience was also associated with increased phenotypic plasticity and an improved shade‐avoidance syndrome. Although these changes indicate fitness and niche differentiation of I. noli‐tangere at invaded sites, future research should examine more closely the adaptive value of these changes and their genetic basis.     

Early bird catches the worm: germination as a critical step in plant invasion

The germination behavior of a plant influences its fitness, persistence, and evolutionary potential, as well as its biotic environment. This can have major effects on the invasive potential of a species. We review the findings of four types of experimental studies comparing basic germination characteristics of invasive versus non-invasive congeners, in their non-native or native distribution range; invasive alien versus native species; and invasive species in their native versus non-native distribution range. Early and/or rapid germination is typical of invasive species rather than their non-invasive congeners, and represents a pre-adaptation from which many invasive and naturalized species benefit. It also occurs more often in invasive than native species, suggesting that competition mitigation or avoidance in the early stages of a plant’s life, via the exploitation of vacant germination niches, might be more useful than a superior competitive ability in novel environments. This is further supported by a tendency of invasive species to germinate earlier and/or faster and have broader germination cues in their non-native than in their native range. It is also supported by broader germination requirements being reported for invasive species than their non-invasive or native congeners. In contrast, high percentage germination is not a consistent predictor of invasiveness, suggesting that the incorporation of a larger fraction of seed production into the soil seed bank rather than high germination is a better (or safer) strategy in novel environments. These patterns indicate that differences in the germination behavior of alien and native species contribute to the invasiveness of many species, although evidence under natural conditions is needed. The role of such differences in the establishment and spread of invasive species in novel environments and their long-term impact on community dynamics requires further study.     

Niche construction through germination cueing: life-history responses to timing of germination in Arabidopsis thaliana

Germination responses to seasonal conditions determine the environment experienced by postgermination life stages, and this ability has potential consequences for the evolution of plant life histories. Using recombinant inbred lines of Arabidopsis thaliana, we tested whether life-history characters exhibited plasticity to germination timing, whether germination timing influenced the strength and mode of natural selection on life-history traits, and whether germination timing influenced the expression of genetic variation for life-history traits. Adult life-history traits exhibited strong plasticity to season of germination, and season of germination significantly altered the strength, mode, and even direction of selection on life-history traits under some conditions. None of the average plastic responses to season of germination or season of dispersal were adaptive, although some genotypes within our sample did exhibit adaptive responses. Thus, recombination between inbred lineages created some novel adaptive genotypes with improved responses to the seasonal timing of germination under some, but not all, conditions. Genetically based variation in germination time tended to augment genetic variances of adult life-history traits, but it did not increase the heritabilities because it also increased environmentally induced variance. Under some conditions, plasticity of life-history traits in response to genetically variable germination timing actually obscured genetic variation for those traits. Therefore, the evolution of germination responses can influence the evolution of life histories in a general manner by altering natural selection on life-history traits and the genetic variation of these traits.     

Plant invasions differentially affected by diversity and dominant species in native‐ and exotic‐dominated grasslands

Plant invasions are an increasingly serious global concern, especially as the climate changes. Here, we explored how plant invasions differed between native‐ and novel exotic‐dominated grasslands with experimental addition of summer precipitation in Texas in 2009. Exotic species greened up earlier than natives by an average of 18 days. This was associated with a lower invasion rate early in the growing season compared to native communities. However, invasion rate did not differ significantly between native and exotic communities across all sampling times. The predictors of invasion rate differed between native and exotic communities, with invasion being negatively influenced by species richness in natives and by dominant species in exotics. Interestingly, plant invasions matched the bimodal pattern of precipitation in Temple, Texas, and did not respond to the pulse of precipitation during the summer. Our results suggest that we will need to take different approaches in understanding of invasion between native and exotic grasslands. Moreover, with anticipated increasing variability in precipitation under global climate change, plant invasions may be constrained in their response if the precipitation pulses fall outside the normal growing period of invaders.     

Trade‐off between early emergence and herbivore susceptibility mediates exotic success in an experimental California plant community

Ecological trade‐offs are fundamental to theory in community ecology; critical for understanding species coexistence in diverse plant communities, as well as the evolution of diverse life‐history strategies. Invasions by exotic species can provide insights into the importance of trade‐offs in community assembly, because the ecological strategies of invading species often differ from those present in the native species pool. Exotic annual species have invaded many Mediterranean‐climate areas around the globe, and often germinate and emerge earlier in the growing season than native species. Early‐season growth can enable exotic annual species to preempt space and resources, competitively suppressing later‐emerging native species; however, early‐emerging individuals may also be more apparent to herbivores. This suggests a potential trade‐off between seasonal phenology and susceptibility to herbivory. To evaluate this hypothesis, we monitored the emergence and growth of 12 focal species (six each native and exotic) in monoculture and polyculture, while experimentally excluding generalist herbivores both early and later in the growing season. Consistent with past studies, the exotic species emerged earlier than native species. Regardless of species origin, earlier‐emerging species achieved greater biomass by the end of the experiment, but were more negatively impacted by herbivory, particularly in the early part of the growing season. This greater impact of early‐season herbivory on early‐active species led to a reduction in the competitive advantage of exotic species growing in polyculture, and improved the performance of later‐emerging natives. Such a trade‐off between early growth and susceptibility to herbivores could be an important force in community assembly in seasonal herbaceous‐dominated ecosystems. These results also show how herbivore exclusion favors early‐active exotic species in this system, with important implications for management in many areas invaded by early‐active exotic species.     

温带半干旱地区一年生植物种子的萌发特性

在实验室条件下研究了中国温带半干旱地区科尔沁沙地的23种1年生植物的种子萌发特性（新采集种子、冷藏和干藏种子）.大籽蒿、虎尾草、冠芒草、沙蓬和地锦的新种子萌发率达90%左右,11种植物新种子萌发率均低于70%,说明这些植物的新种子具有或多或少的休眠属性.经过150d的冷干藏后,大籽蒿、虎尾草萌发率保持在90%以上,说明这两种植物完全没有休眠机制;冠芒草、沙蓬和地锦的种子萌发率下降较多,可能是储藏的环境条件导致的2次休眠现象;冷藏和干藏处理均能使绿珠藜、毛马唐、细叶益母草、雾冰藜、金狗尾草、苋菜、马齿苋、碱地肤和水稗草的种子在生长季开始时完成生理后熟,萌发率达到80%以上;干藏有利于促进毛马唐、细叶益母草、马齿苋和鹤虱的种子成熟,冷藏有利于促进绿珠藜和金狗尾草的种子成熟;黄蒿、灰绿藜、画眉草和烛台虫实在不同处理下的萌发率都比较低,说明种子内在生理休眠作用较强,具有减少种子一次性萌发数量的风险分摊策略.大多数1年生植物均能在较短时间内达到最终萌发率的90%,表现出迅速萌发的特性;黄蒿、灰绿藜、碱地肤和沙蓬种子则在不同处理中表现出延长萌发时间的策略来适应半干旱地区不确定的环境条件.最后,探讨了几种主要1年生植物的种子萌发对策与其对环境适应机制之间的关系.     

Effects of exotic species on evolutionary diversification

Exotic species invasions create almost ideal conditions for promoting evolutionary diversification: establishment of allopatric populations in new environmental conditions; altered ecological opportunities for native species; and new opportunities for hybridization between previously allopatric taxa. Here, we review recent studies of the evolutionary consequences of species invasions, revealing abundant and widespread examples of exotic species promoting evolutionary diversification via increased genetic differentiation among populations of both exotic and native species and the creation of new hybrid lineages. Our review indicates that, although the well-documented reductions to biodiversity caused by exotic species might outweigh the increases resulting from diversification, a complete understanding of the net effects of exotic species on biodiversity in the long term will require consideration of both.     

Root Inputs Influence Soil Water Holding Capacity and Differentially Influence the Growth of Native versus Exotic Annual Species in an Arid Ecosystem

Invasion by exotic annual species is increasingly impacting Southern California arid lands, altering ecosystem processes and plant community composition. With climate change, the Southwestern United States is expected to experience increasingly variable rainfall. Larger rainfall events could facilitate invasion by exotic species that can capitalize on high resource conditions. Exotic annual species also have dense shallow root systems that could create positive feedbacks to further invasion by increasing soil organic matter and water holding capacity. Alternatively, fine root inputs could create negative feedbacks to exotic plant growth if they stimulate microbial nutrient immobilization. The dual influences of rainfall regime and fine root inputs on species performance were evaluated in an experiment where native and exotic species were grown individually and in combination under varying watering regimes (large infrequent or small frequent pulses, holding total rainfall constant) and root additions (with or without sterilized exotic roots). Mean soil moisture increased with larger infrequent watering events, and also with root addition. Plant growth (both native and exotic) increased with larger watering events, but declined with root addition. Exotic species growth declined more than native species growth with root additions. Mechanistically, root addition lowered inorganic nitrogen (N) availability, and microbial N immobilization increased with soil moisture content. Together these results show that increased fine root production promotes negative feedbacks to growth of exotic species via microbial N immobilization, especially under conditions of high soil moisture. Further, our results suggest that organic carbon additions are a potentially effective strategy for suppressing growth of problematic desert invaders.     

Ecophysiological basis of the Jack-and-Master strategy: Taraxacum officinale (dandelion) as an example of a successful invader

Aims Successful invasive plants are often assumed to display significant levels of phenotypic plasticity. Three possible strategies by which phenotypic plasticity may allow invasive plant species to thrive in changing environments have been suggested: (i) via plasticity in morphological or physiological traits, invasive plants are able to maintain a higher fitness than native plants in a range of environments, including stressful or low-resource habitats: a 'Jack-of-all-trades' strategy; (ii) phenotypic plasticity allows the invader to better exploit resources available in low stress or favorable habitats, showing higher fitness than native ones: a 'Master-of-some' strategy and (iii) a combination of these abilities, the 'Jack-and-Master' strategy.Methods We evaluated these strategies in the successful invader Taraxacum officinale in a controlled experiment mimicking natural environmental gradients. We set up three environmental gradients consisting of factorial arrays of two levels of temperature/light, temperature/water and light/water, respectively. We compared several ecophysiological traits, as well as the reaction norm in fitness-related traits, in both T. officinale and the closely related native Hypochaeris thrincioides subjected to these environmental scenarios.Important findings Overall, T. officinale showed significantly greater accumulation of biomass and higher survival than the native H. thrincioides, with this difference being more pronounced toward both ends of each gradient. T. officinale also showed significantly higher plasticity than its native counterpart in several ecophysiological traits. Therefore, T. officinale exhibits a Jack-and-Master strategy as it is able to maintain higher biomass and survival in unfavorable conditions, as well as to increase fitness when conditions are favorable. We suggest that this strategy is partly based on ecophysiological responses to the environment, and that it may contribute to explaining the successful invasion of T. officinale across different habitats.     

Playing smart vs. playing safe: the joint expression of phenotypic plasticity and potential bet hedging across and within thermal environments

Adaptive phenotypic plasticity evolves when cues reliably predict fitness consequences of life‐history decisions, whereas bet hedging evolves when environments are unpredictable. These modes of response should be jointly expressed, because environmental variance is composed of both predictable and unpredictable components. However, little attention has been paid to the joint expression of plasticity and bet hedging. Here, I examine the simultaneous expression of plasticity in germination rate and two potential bet‐hedging traits – germination fraction and within‐season diversification in timing of germination – in seeds from multiple seed families of five geographically distant populations of Lobelia inflata (L.) subjected to a thermal gradient. Populations differ in germination plasticity to temperature, in total germination fraction and in the expression of potential diversification in the timing of germination. The observation of a negative partial correlation between the expression of plasticity and germination variance (potential diversification), and a positive correlation between plasticity and germination fraction is suggestive of a trade‐off between modes of response to environmental variance. If the observed correlations are indicative of those between adaptive plasticity and bet hedging, we expect an optimal balance to exist and differ among populations. I discuss the challenges involved in testing whether the balance between plasticity and bet hedging depends on the relative predictability of environmental variance.