Shrubs as ecosystem engineers across an environmental gradient: effects on species richness and exotic plant invasion

Ecosystem-engineering plants modify the physical environment and can increase species diversity and exotic species invasion. At the individual level, the effects of ecosystem engineers on other plants often become more positive in stressful environments. In this study, we investigated whether the community-level effects of ecosystem engineers also become stronger in more stressful environments. Using comparative and experimental approaches, we assessed the ability of a native shrub (Ericameria ericoides) to act as an ecosystem engineer across a stress gradient in a coastal dune in northern California, USA. We found increased coarse organic matter and lower wind speeds within shrub patches. Growth of a dominant invasive grass (Bromus diandrus) was facilitated both by aboveground shrub biomass and by growing in soil taken from shrub patches. Experimental removal of shrubs negatively affected species most associated with shrubs and positively affected species most often found outside of shrubs. Counter to the stress-gradient hypothesis, the effects of shrubs on the physical environment and individual plant growth did not increase across the established stress gradient at this site. At the community level, shrub patches increased beta diversity, and contained greater rarified richness and exotic plant cover than shrub-free patches. Shrub effects on rarified richness increased with environmental stress, but effects on exotic cover and beta diversity did not. Our study provides evidence for the community-level effects of shrubs as ecosystem engineers in this system, but shows that these effects do not necessarily become stronger in more stressful environments.     

Ecotypic diversity of a dominant grassland species resists exotic invasion

Many species are characterized by high levels of intraspecific or ecotypic diversity, yet we know little about how diversity within species influences ecosystem processes. Using a common garden experiment, we studied how intraspecific diversity within the widespread and often dominant North American native Pseudoroegneria spicata (Pursh) Á. Löve. affected invasion by Centaurea stoebe L. We experimentally manipulated Pseudoroegneria intraspecific diversity by changing the number of Pseudoroegneria ecotypes in common garden plots, using ecotypes collected throughout western North America. Invader biomass was 46% lower in mono-ecotype Pseudoroegneria plots than in control plots without any plants prior to invasion, and plots with 3–12 Pseudoroegneria ecotypes were 44% less invaded by Centaurea than the mono-ecotype plots. Across all plots, the total biomass of invading Centaurea plants was negatively correlated with total Pseudoroegneria biomass, but biotic resistance provided by high ecotypic diversity of Pseudoroegneria was not explained only by the increase in productivity that occurred with ecotypic diversity. Relative to Pseudoroegneria yield, Centaurea yield was lowest when Pseudoroegneria overyielded due to size-independent “complementarity” effects. This was not observed when overyielding was due to size-dependent effects. Our results suggest that the intraspecific diversity of a widespread and dominant species has the potential to impact invasion outcomes beyond its effects on native plant productivity and that mechanisms of biotic resistance to invaders may be to some degree independent of plant size.     

狙击生物入侵的对策探讨

我国加入 WTO后 ,随着世界经济贸易自由化进程和改革开放步伐的加快 ,国际生物水平的科技成果或产品将渗透进入我国生物界中 ,外来生物在丰富我们生物物种的同时 ,也会带来负面影响。近期生物界出现一个新的名词——生物入侵 ,是指生物由原生地经自然或人为的途径侵入到另一个新环境 ,对入侵地的生物多样性 ,农林牧渔业生产以及人类健康造成经济损失或生态灾难。通俗地说 ,就是外来物种“入侵并打败”当地物种 ,“反客为主”,导致当地生态失衡 ,进而引发一系列的问题。近年来 ,随着全球经济一体化步伐的加快 ,国内外贸易往来越来越频繁 ,…     

Effect of a native tree on seedling establishment of two exotic invasive species in a semiarid ecosystem

Theory predicts that in more stressful environments, positive plant-plant interactions should be more important than negative ones. For instance, in arid and semiarid regions, amelioration of soil drought produced by the shade of established plants could facilitate establishment of other species, in spite of light reduction. However, this theory has not been tested widely in the context of plant invasion. In this paper we evaluated the hypothesis that in a semiarid ecosystem of central Chile, the native tree, Lithrea caustica, should facilitate through positive shading effects, the seedling establishment of two widely planted and invasive forestry species, Pinus radiata and Eucalyptus globulus. We assessed the seedling establishment examining two processes: seedling recruitment (including germination) and subsequent seedling survival. We sowed seeds (to assess recruitment) and planted 8 months old seedlings (to assess seedling survival) of each exotic species under Lithrea patches, open sites and under an artificial shade mimicking Lithrea shading. The study was repeated in a north-facing and a south-facing slope in the study area located in a xeric zone within the distribution range of plantations of these species in central Chile. Our results show that in a north-facing slope Lithrea had positive effects on recruitment of both species, which was produced by shading. These effects were counteracted by negative effects on seedling survival but through a different mechanism, which suggests that Lithrea would have no significant effect on the whole seedling establishment process of Pinus radiata nor Eucalyptus globulus in this habitat. In turn, in a south-facing slope Lithrea had no significant effect on recruitment but had a negative effect on seedling survival, which was not produced by shading. This suggests that in this habitat Lithrea has a negative effect on the seedling establishment of these exotic species. Our results suggest that the effect of the native Lithrea caustica on the seedling establishment of these exotic species is dependent upon the life-cycle phase (recruitment or seedling survival) and habitat even within the same semiarid ecosystem. In contrast to the expected positive effects Lithrea is unlikely to facilitate seedling establishment of these exotic species in this area, and in fact in some habitats this effect could be negative. However, our results also suggest that a common mechanism proposed to resist invasion in forest ecosystems such as shading, probably is not sufficient to inhibit invasion in a semiarid region.     

Allelopathy and exotic plant invasion

The primary hypothesis for the astonishing success of many exotics as community invaders relative to their importance in their native communities is that they have escaped the natural enemies that control their population growth – the `natural enemies hypothesis'. However, the frequent failure of introduced biocontrols, weak consumer effects on the growth and reproduction of some invaders, and the lack of consistent strong top-down regulation in many natural ecological systems indicate that other mechanisms must be involved in the success of some exotic plants. One mechanism may be the release by the invader of chemical compounds that have harmful effects on the members of the recipient plant community (i.e., allelopathy). Here, we provide an abbreviated compilation of evidence for allelopathy in general, present a detailed case study for Centaurea diffusa, an invasive Eurasian forb in western North America, and review general evidence for allelopathic effects of invasive plants in native communities. The primary rationale for considering allelopathy as a mechanism for the success of invaders is based on two premises. First, invaders often establish virtual monocultures where diverse communities once flourished, a phenomenon unusual in natural communities. Second, allelopathy may be more important in recipient than in origin communities because the former are more likely to be naïve to the chemicals possessed by newly arrived species. Indeed, results from experiments on C. diffusa suggest that this invader produces chemicals that long-term and familiar Eurasian neighbors have adapted to, but that C. diffusa's new North American neighbors have not. A large number of early studies demonstrated strong potential allelopathic effects of exotic invasive plants; however, most of this work rests on controversial methodology. Nevertheless, during the last 15 years, methodological approaches have improved. Allelopathic effects have been tested on native species, allelochemicals have been tested in varying resource conditions, models have been used to estimate comparisons of resource and allelopathic effects, and experimental techniques have been used to ameliorate chemical effects. We do not recommend allelopathy as a `unifying theory' for plant interactions, nor do we espouse the view that allelopathy is the dominant way that plants interact, but we argue that non-resource mechanisms should be returned to the discussion table as a potential mechanism for explaining the remarkable success of some invasive species. Ecologists should consider the possibility that resource and non-resource mechanisms may work simultaneously, but vary in their relative importance depending on the ecological context in which they are studied. One such context might be exotic plant invasion.     

Competition and species packing in patchy environments

In models of competition in which space is treated as a continuum, and population size as continuous, there are no limits to the number of species that can coexist. For a finite number of sites, N, the results are different. The answer will, of course, depend on the model used to ask the question. In the Tilman-May-Nowak ordinary differential equation model, the number of species is asymptotically C log N with most species packed in at the upper end of the competitive hierarchy. In contrast, for metapopulation models with discrete individuals and stochastic spatial systems with various competition neighborhoods, we find a traditional species area relationship CN(a), with no species clumping along the phenotypic gradient. The exponent a is larger by a factor of 2 for spatially explicit models. In words, a spatial distribution of competitors allows for greater diversity than a metapopulation model due to the effects of recruitment limitation in their competition.     

Fire-driven alien invasion in a fire-adapted ecosystem

Disturbance plays a key role in many alien plant invasions. However, often the main driver of invasion is not disturbance per se but alterations in the disturbance regime. In some fire-adapted shrublands, the community is highly resilient to infrequent, high-intensity fires, but changes in the fire regime that result in shorter fire intervals may make these communities more susceptible to alien plant invasions. This study examines several wildfire events that resulted in short fire intervals in California chaparral shrublands. In one study, we compared postfire recovery patterns in sites with different prefire stand ages (3 and 24 years), and in another study we compared sites that had burned once in four years with sites that had burned twice in this period. The population size of the dominant native shrub Adenostoma fasciculatum was drastically reduced following fire in the 3-year sites relative to the 24-year sites. The 3-year sites had much greater alien plant cover and significantly lower plant diversity than the 24-year sites. In a separate study, repeat fires four years apart on the same sites showed that annual species increased significantly after the second fire, and alien annuals far outnumbered native annuals. Aliens included both annual grasses and annual forbs and were negatively correlated with woody plant cover. Native woody species regenerated well after the first fire but declined after the second fire, and one obligate seeding shrub was extirpated from two sites by the repeat fires. It is concluded that some fire-adapted shrublands are vulnerable to changes in fire regime, and this can lead to a loss of native diversity and put the community on a trajectory towards type conversion from a woody to an herbaceous system. Such changes result in alterations in the proportion of natives to non-natives, changes in functional types from deeply rooted shrubs to shallow rooted grasses and forbs, increased fire frequency due to the increase in fine fuels, and changes in carbon storage.     

Dispersal characteristics of three odonate species in a patchy habitat

Dispersal has a potentially profound effect on the dynamics of populations especially when a population occupies a patchy habitat. Ponds surrounded by terrestrial landscape are an example of patchy distribution of physical conditions and constitute "islands" for odonates. Few studies have focussed on dispersal in odonates. We have used the direct method of dispersal observing (capture-mark-recapture technique) in order to estimate the degree of linkage in three patchy populations of odonate localised on three ponds. We also examined the differences in dispersal ability within and among three species ( Coenagrion puella , Coenagrion scitulum and Libellula depressa ). The ponds were situated in southwest France on a limestone plateau. In this arid area, these ponds constitute the only surface water available and are relatively sparsely distributed. The size of the ponds ranged from 48 to 79 m² and they were 200 and 775 m apart. We demonstrated that three factors influence the dispersal ability of these odonates. The first is represented by the abiotic factors and especially weather conditions. This determines the number of days that dispersal is possible. The second is interspecific differences. We showed that sensitivity to weather conditions, species size and species behaviour influence dispersal ability. The third factor is the intraspecific characteristics. We demonstrated that there are differences in dispersal ability according to sex and age. To conclude, we discuss the importance of pond management to maintain the existing odonate populations and to facilitate introduction of new populations in this region where little exchange occurs between ponds.     

Intraguild responses of aphid predators on apple to the invasion of an exotic species, Harmonia axyridis

The effects of the invasion ofan exotic predator, Harmonia axyridis(Pallas) (Coleoptera: Coccinellidae), wereinvestigated using three experiments on theecology of aphid predators on apple. Oneexperiment, 1992, was collected prior to theH. axyridis invasion, and two others,1996 to 1997 and 1999 to 2000, were collectedafter the invasion. Except for one year, 1999,H. axyridis was the dominant coccinellid,replacing the formerly dominant Coccinellaseptempunctata L. (Coleoptera: Coccinellidae),another exotic species. The dominance of H. axyridis was greater among larvae thanamong adults. There was no apparent effect ofthe H. axyridis invasion on abundance ofthe predator, Aphidoletes aphidimyza(Rondani) (Diptera: Cecidomyiidae) and apossible positive effect on the abundance ofchrysopids (Neuroptera: Chrysopidae). Principal component analysis indicated thatalthough individual species were affected, theoverall effect of H. axyridis invasion onthe predator guild as a whole was negligible. The data indicate that the interaction betweenthe two exotic species, H. axyridis andC. septempunctata, may be allowing nativecoccinellids to become more abundant on applethan when C. septempunctata was thedominant coccinellid.     

Thornbush invasion in a savanna ecosystem in eastern Botswana

The paper describes the thornbush invasion (bush encroachment) found in the savanna at Olifants Drift in eastern Botswana. Overgrazing by cattle is responsible for the opening up of the grass sward and has enabled woody species to establish. A quantification of this process between 1950 and 1975 in an area of ca. 100 km² is given in terms of plant density as well as aboveground biomass. The loss of herbaceous biomass which generally accompanies bush encroachment and the possibilities for pasture restoration are mentioned.Species nomenclature follows the Flora of Southern Africa and the system in use at the herbarium of the Botanical Research Institute at Pretoria, South Africa.     

Differential predation drives overyielding of prey species in a patchy environment

In environments characterized by regional heterogeneity among patches, competitor diversity can enhance ecosystem functions such as biomass production. Studies that have addressed the strength of diversity effects in heterogeneous environments have primarily considered a patchy distribution of resources. However, in many systems, top–down effects influence competitor productivity and composition. We use a three‐trophic level consumer–resource model to ask how differential responses to predation influence consumer diversity effects at two scales; 1) in patches with and without predator populations, and 2) at a ‘regional’ scale, consisting of one patch with‐ and one patch without a predator population. At the local scale, the strength and direction of consumer diversity effects depended on the strength of the differential response to predation. Positive or negative influences of consumer richness on equilibrium consumer biomass were the result of a selection effect of diversity. At the regional scale, we observed transgressive overyielding driven by a positive complementarity effect for parameters that define a strong differential response to predation. Given the prevalence of spatially and temporally heterogeneous top–down effects on competitor composition in many ecosystems and trophic levels, we advocate consideration of differential predation as an important step towards incorporating realistic trophic complexity into diversity–function studies.     

Spatiotemporal complexity of patchy invasion in a predator-prey system with the Allee effect

Invasion of an exotic species initiated by its local introduction is considered subject to predator-prey interactions and the Allee effect when the prey growth becomes negative for small values of the prey density. Mathematically, the system dynamics is described by two nonlinear diffusion-reaction equations in two spatial dimensions. Regimes of invasion are studied by means of extensive numerical simulations. We show that, in this system, along with well-known scenarios of species spread via propagation of continuous population fronts, there exists an essentially different invasion regime which we call a patchy invasion. In this regime, the species spreads over space via irregular motion and interaction of separate population patches without formation of any continuous front, the population density between the patches being nearly zero. We show that this type of the system dynamics corresponds to spatiotemporal chaos and calculate the dominant Lyapunov exponent. We then show that, surprisingly, in the regime of patchy invasion the spatially average prey density appears to be below the survival threshold. We also show that a variation of parameters can destroy this regime and either restore the usual invasion scenario via propagation of continuous fronts or brings the species to extinction; thus, the patchy spread can be qualified as the invasion at the edge of extinction. Finally, we discuss the implications of this phenomenon for invasive species management and control.     

Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem

Variation in the stable carbon and oxygen isotope composition (δ(13) C, Δ(18) O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ(13) C and Δ(18) O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ(18) O was tightly negatively correlated with stomatal conductance (g(s) ), whereas δ(13) C was positively correlated with intrinsic water use efficiency (WUE(i) ). Broad interspecific variation in Δ(18) O, δ(13) C and WUE(i) was largely determined by differences in g(s) , as indicated by a strong positive correlation between leaf δ(13) C and Δ(18) O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ(13) C and Δ(18) O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from 'profligate/opportunistic' (high g(s) , low WUE(i) ) to 'conservative' (low g(s) , high WUE(i) ). Coexisting species maintained their relative isotopic rankings in 2?yr with contrasting rainfall, suggesting the existence of species-specific 'isotopic niches' that reflect ecophysiological niche segregation in dryland plant communities.     

Gene flow among established Puerto Rican populations of the exotic tree species, Albizia lebbeck

We estimate gene flow and patterns of genetic diversity in Albizia lebbeck, an invasive leguminous tree in the dry forest of southwestern Puerto Rico. Genetic diversity estimates calculated for 10 populations of 24 trees each indicated that these populations may have been formed from multiple introductions. The presence of unique genotypes in the northernmost populations suggests that novel genotypes are still immigrating into the area. This combination of individuals from disparate locations led to high estimates of genetic diversity (He = 0.266, P = 0.67). Indirect estimates of gene flow indicate that only 0.69 migrants per generation move between populations, suggesting that genetic diversity within populations should decrease due to genetic drift. Since migration-drift equilibrium was not found, however, this estimate needs to be viewed with caution. The regular production of pods in this outcrossing species (tm = 0.979) indicates that sufficient outcross pollen is received to insure successful reproduction. Direct estimates of gene flow indicate that between 44 and 100% of pollen received by trees in four small stands of trees (n < 11) was foreign. The role of gene flow in facilitating the spread of this invasive plant species is discussed.     

Eco-evolutionary litter feedback as a driver of exotic plant invasion

Many studies have examined positive feedbacks between invasive plant traits and nutrient cycling, but few have investigated whether feedbacks arise from introduction of pre-adapted species or from eco-evolutionary feedback that develops after introduction. Eco-evolutionary feedback could occur between an invader's leaf tissue C:N ratio and its response to litter accumulation. Previous modeling predicts that occurrence of this feedback would be reflected by: (1) field data showing higher litter:biomass ratios in the invasive range; (2) high C:N genotypes benefiting more from experimental litter additions than low C:N genotypes; (3) this beneficial effect on high C:N genotypes inducing a critical transition toward invader dominance when a critical amount of litter is added to a native species-dominated community experiencing low nutrient conditions. Here, we empirically tested these predictions for the invasive grass Phalaris arundinacea, which has undergone post-introduction evolutionary change toward attaining higher C:N ratios under high nutrient conditions. We performed a biogeographical comparison of litter:biomass ratios in the native (Europe) and invasive (USA) range, and an experiment with mesocosms from the invasive range under low nutrient conditions. Low and high C:N Phalaris genotypes were introduced into native-dominated and bare mesocosms, to which varying litter amounts were added. The biogeographical comparison revealed that litter:biomass ratios were higher in the invasive range. The mesocosm experiment showed that when grown in isolation, only high C:N genotypes responded positively to litter. This effect, however, was not strong enough to stimulate Phalaris when exposed to competition with native species. Our results suggest that eco-evolutionary feedback between Phalaris’ C:N ratio and litter accumulation could occur, but only under high nutrient conditions. Our experiments suggest that eco-evolutionary feedback may select for specialist rather than superior genotypes. Hence, genotypic variation induced by post-introduction admixture may be subject to context-dependent selection due to eco-evolutionary feedback, increasing trait variation within invasive populations.     

Genetic variability and phylogeographical patterns of a nonindigenous species invasion: a comparison of exotic vs. native zebra and quagga mussel populations

There have been few investigations of the number of founding sources and amount of genetic variability that lead to a successful nonindigenous species invasion, although genetic diversity is believed to play a central role. In the present study, population genetic structure, diversity and divergence patterns were analysed for the zebra mussel Dreissena polymorpha [n=280 samples and 63 putative randomly amplified polymorphic DNA (RAPDs) gene loci] and the quagga mussel D. bugensis (n=136 and 52 loci) from 10 nonindigenous North American and six Eurasian sampling sites, representing their present‐day ranges. Results showed that exotic populations of zebra and quagga mussels had surprisingly high genetic variability, similar to those in the Eurasian populations, suggesting large numbers of founding individuals and consistent with the hypothesis of multiple colonizations. Patterns of genetic relationships indicate that the North American populations of D. polymorpha likely were founded by multiple source populations from north‐western and northcentral Europe, but not from southcentral or eastern Europe. Sampling areas within North America also were significantly divergent, having levels of gene flow and migration about twice those separating long‐established Eurasian populations. Samples of D. bugensis in Lakes Erie and Ontario were significantly different, with the former being more closely related to a native population from the Dnieper River, Ukraine. No evidence for a founder effect was discerned for either species.     

Phenological complementarity, species diversity, and ecosystem function

Increasing species diversity frequently enhances ecosystem function. Phenological complementarity, the asynchrony of species resource use and growth, may explain how species diversity influences ecosystem function but remains largely untested. We used an early successional plant community containing species with a variety of phenologies to test whether increasing species diversity enhances ecosystem function by increasing phenological complementarity. Over a two-year period, we increased environmental heterogeneity within an abandoned field with variation in disturbance, soil nutrients, water, light availability, and disturbance in 160 permanent plots, and measured percent cover of each plant species three times in each growing season. We did not manipulate species composition directly, and thus diversity and complementarity in each plot were the result of pre-existing conditions and responses of individuals to experimental treatments. Species diversity was measured in two ways, as the total number of species per plot and as the evenness of species abundances. Phenological complementarity was measured as the negative logarithm of the variance ratio. We tested whether the number of plant species per plot, species evenness, and their phenological complementarity in the first year predicted total annual cover in the second year. Total annual cover increased only moderately with number of species and evenness, consistent with studies that randomize species composition among replicate plots. Any effect that species number or evenness had on total annual cover, however, was not due to phenological complementarity. Rather, diversity was unrelated to phenological complementarity. These results indicate that naturally occurring variation in species diversity had little effect on whether phenological complementarity can enhance ecosystem function.