Using integral projection models to compare population dynamics of four closely related species

Demographic processes, such as survival, growth, and reproduction, can inform us about invasion risk, extinction risk, and trade-offs in life history strategies. The population dynamics of four Amaranthaceae species in southern Illinois, USA were examined using integral projection models (IPMs) to determine whether vital rates reflect life history among these closely related species. Two of the species, Amaranthus palmeri and Amaranthus tuberculatus, are summer annuals and considered to be some of the most problematic agricultural weeds in the US Midwest. Achyranthes japonica is a relatively new invasive exotic species that primarily inhabits forests. Iresine rhizomatosa, is an endangered species in the study area, which also inhabits forests. Two populations of each species were studied from 2012 to 2014 in which height of individuals were measured and used as the state variable in the IPMs. The Amaranthus species and Achyranthes japonica had an estimated population growth rate >1, projecting increases in population size. By contrast, λ was <1 for I. rhizomatosa, projecting a decline in population size demonstrating its endangered status. Germination rates and seed viability were dependent on species and varied over time. Elasticity analyses showed that survival and growth contributed most to λ for the perennial species; whereas, for the annual species population dynamics were driven primarily by fecundity. Overall, Achyranthes japonica and the Amaranthus species show similar trends in demographic processes that align with their invasive nature and not with their life histories. Furthermore, this study demonstrates that more research on the competitive nature of Achyranthes japonica is needed.     

Parental investment in temporally varying environments

Summary Using a model that allows the mean and variance of investment by parents in offspring to evolve in response to change in degree of temporal environmental variation, this paper shows that both parental investment parameters should increase with increases in temporal variation. If offspring receiving greater parental investment are viable over a broader range of environmental conditions, then increased temporal environmental variation can select for increases in parental investment. The variance in parental investment also may increase with increases in temporal variation, but there is a threshold level of temporal variation that must be exceeded before variance in parental investment is adaptive. Thus phenotypic variance in parental investment is not adaptive in all temporally varying environments. Further, increased overlap among generations reduces the expected effects of temporal variation on the mean and variance in parental investment. Thus a negative correlation between length of reproductive life and both measures of investment is expected. There is support for the predictions of this model in some animal groups, but not among plants. Possible reasons for the lack of support among plants are discussed and directions for future research aimed at distinguishing adaptive and maladaptive phenotypic variance in parental investment are suggested.     

Alternate plant life history strategies and coexistence in randomly varying environments

Environmental fluctuations can in theory allow the coexistence ofecologically similar species by time-sharing a niche, as envisioned by Hutchinson. The evolution of this situation is studied in a competition model, using as an example the evolution of seed germination strategies. Coexistence occurs via the evolution of low-risk and high-risk strategies for dealing with the variability by different species. Coexistence is promoted by intermediate levels of variability or disturbance, and by a trade-off between seed yield and seed survivorship. These results may be applicable also to other low vs. high risk life history options in unpredictably varying environments, such as: stress resistance vs. potentially rapid growth, high adult survivorship vs. high reproductive output. The model's predictions differ from those obtained without consideration of life history evolution in response to environmental variability, and are consistent with some recent studies of plant strategies in intermittently stressed communities.I thank A. Shmida for many discussions on this topic, D. Cohen and I. Noy-Meir for comments after a seminar presentation of this paper, and H. de Kroon, H. During, and E. Van der Maarel for decreasing my ignorance of the empirical literature.Research conducted while the author was recipient of a Sir Charles Clore Postdoctoral Fellowship in the Department of Applied Mathematics, Weizmann Institute of Science, Rehovot, Israel.     

Reproductive strategy of insects as adaptation to temporally varying environments

Diversity of oviposition curve observed in 125 insects was analysed using the ratio of the length of period for development to that for reproduction. On the basis of this empirical data, two extreme reproductive patterns were selected: prolonged reproduction with early maturity and concentrated reproduction with late maturity. Population growth of the species with each reproductive pattern was calculated usingLeslie Matrix under some simulated fluctuating environments where the length of time during which the environments change is short compared with the time required to stabilize age structure. These simulation studies show that there is an optimal ratio of the length of period for development to that for reproduction in achieving high population densities and this ratio varies depending on the favorableness of environment for reproductive success: as the environmental favorableness decreases, the optimal ratio becomes larger.     

Trophic network structure emerges through antagonistic coevolution in temporally varying environments

Understanding the mechanisms underlying ecological specialization is central to our understanding of community ecology and evolution. Although theoretical work has investigated how variable environments may affect specialization in single species, little is known about how such variation impacts bipartite network structure in antagonistically coevolving systems. Here, we develop and analyse a general model of victim-enemy coevolution that explicitly includes resource and population dynamics. We investigate how temporal environmental heterogeneity affects the evolution of specialization and associated community structure. Environmental productivity influences victim investment in resistance, which will shape patterns of specialization through its regulating effect on enemy investment in infectivity. We also investigate the epidemiological consequences of environmental variability and show that enemy population density is maximized for intermediate lengths of productive seasons, which corresponds to situations where enemies can evolve higher infectivity than victims can evolve defence. We discuss our results in the light of empirical studies, and further highlight ways in which our model applies to a range of natural systems.     

Fruit type,life form and origin determine the success of woody plant invaders in an urban landscape

The spread of alien plant species is a critical ecological event worldwide, but the forces that control this spread are not well documented. Alien plant species are well known to disrupt ecological services of native ecosystems, change the composition of native habitats, and often lead to the extirpation of native flora and fauna. Here, we report on life history patterns of plant species with rapidly spreading and declining ranges in North America’s major urban region. We tested for differences in life history traits between the 466 native and alien woody flora of the New York metropolitan area. We also examined the relationship between life history traits and change in distribution in the New York metropolitan area between 1900 and 2000. Native and alien species of the New York metropolitan area differ with respect to pollination vector and breeding system. However, pollination vector and breeding system are not associated with success, defined here as increasing range spread in the urban environment; instead, fruit type (dispersal), life form and origin are important determinants of success. Alien species that are deciduous trees, shrubs or vines with fleshy fruit are the most successful in increasing their distribution in this urban landscape. Newly introduced species with these characteristics are expected to have a better chance at establishing in similar urban landscapes and should be targets for intensive management. The ability to predict which alien species will become invasive is also a valuable tool for the prevention of invasions by newly introduced plant species.     

Demographic consequences of age-structure in extreme environments: population models for arctic and alpine ptarmigan

Organisms living in arctic and alpine environments are increasingly impacted by human activities. To evaluate the potential impacts of global change, a better understanding of the demography of organisms in extreme environments is needed. In this study, we compare the age-specific demography of willow ptarmigan (Lagopus lagopus) breeding at arctic and subalpine sites, and white-tailed ptarmigan (L. leucurus) breeding at an alpine site. Rates of egg production improved with age at the alpine and subalpine sites, but the stochastic effects of nest and brood predation led to similar rates of annual fecundity among 1-, 2-, and 3+-year-old females. All populations had short generation times (T<2.7 years) and low net reproductive rates (R ₀<1.2). Stable age distributions were weighted towards 1-year-old females in willow ptarmigan (>59%), and to 3+-year-old females in white-tailed ptarmigan (>47%). High damping ratios (ρ>3.2) indicated that asymptotic estimates were likely to match natural age distributions. Sensitivity and elasticity values indicated that changes in juvenile survival would have the greatest impact on the finite rate of population change (λ) in willow ptarmigan, whereas changes to the survival of 3+-year-old females would have a greater effect in white-tailed ptarmigan. High survivorship buffers white-tailed ptarmigan in alpine environments against the potential effects of climate change on annual fecundity, but may make the species more sensitive to the effects of pollutants or harvesting on adult survival. Conversely, processes that reduce annual fecundity would have a greater impact on the population viability of willow ptarmigan in arctic and subalpine environments. If these same demographic patterns prove to be widespread among organisms in extreme environments, it may be possible to develop general recommendations for conservation of the biological resources of arctic and alpine ecosystems.     

Empirical and virtual investigation of the population dynamics of an alien plant under the constraints of local carrying capacity: Heracleum mantegazzianum in the Czech Republic

During the last decades, invasive alien species have become a global concern because of their ecological and economic impact. Heracleum mantegazzianum (Apiaceae), is a tall monocarpic perennial native to Caucasus and invasive in Europe since the 1950s. Within an interdisciplinary EU project aimed at assessing suitable management strategy, we analysed the demography and ecology of this species in its invasive range. The monitoring of population dynamics in the Czech Republic led to the result that in the observed sites the species showed decreasing populations. To find an explanation for this unexpected result, two types of models were parameterized, based on the empirical data: (1) a stage-based transition matrix model, which projected a continuous negative development, and (2) a spatially explicit individual-based model (IBM), including individual variation. This second model was able to create a population with steady individual numbers. Analyses of the simulation showed that in more than 54% of the simulated years (n=5000) the growth rate was smaller than one. Still, population increase in the remaining years was sufficient to sustain a population. Nevertheless long-term observations document an invasive behaviour of the observed populations. Hence, we could assume temporal changes in the course of an invasion and thus wanted to evaluate the probability of sampling negative growth in dependence of time since first invasion. By using a method from ‘Virtual Ecology’, we approached the question: first we create an invasive population, based on the empirical data of H. mantegazzianum and second empirical sampling techniques were mimicked using the Virtual Ecologist approach. The results demonstrate how the probability of sampling negative growth increases with time since first invasion. Hence, we assume that the studied populations have already reached a maximum of their local invasive potential and thus stagnate in their size.     

Demographic analysis of Hubbell's neutral theory of biodiversity

Hubbell's neutral model is increasingly applied in both theoretical and empirical studies but so far little attention has been paid to the ecological mechanisms that determine species diversity in neutral communities. In this contribution we use a stochastic individual-based Markovian model to provide an explicit derivation of Hubbell's local community model from the fundamental processes of reproduction, mortality, and immigration, and show that such derivation provides important insights on the mechanisms regulating species diversity that cannot be obtained from the original model and its previous extensions. One important insight is that the basic parameters of Hubbell's model, community size (J) and the probability that a dying individual will be replaced by an immigrant (m), cannot be considered independent and that their interdependency leads to a counterintuitive trade-off between community size and species diversity. We further demonstrate that Hubbell's treatment of community size as a free parameter hides fundamental mechanisms that influence species diversity through their effect on the size of the community. For example, while in Hubbell's model immigration can only increase species diversity by promoting colonization rates, the demographic derivation shows that immigration can also promote species diversity by reducing extinction rates. Our demographic derivation also unifies previous contrasting predictions about the effect of reproduction on species diversity by showing that both positive and negative effects are possible, and that the balance between the two effects depends on the size of the community. The demographic derivation also reconciles an apparent contradiction between Hubbell's theory and patch occupancy theory, and integrates three previously proposed mechanisms of species diversity, the More Individuals Hypothesis, the rescue effect, and the dilution effect, within a single, unified framework.     

Dispersal evolution in temporally variable environments: implications for plant range dynamics

Dispersal—the movement of an individual from the site of birth to a different site for reproduction—is an ecological and evolutionary driver of species ranges that shapes patterns of colonization, connectivity, gene flow, and adaptation. In plants, the traits that influence dispersal often vary within and among species, are heritable, and evolve in response to the fitness consequences of moving through heterogeneous landscapes. Spatial and temporal variation in the quality and quantity of habitat are important sources of selection on dispersal strategies across species ranges. While recent reviews have evaluated the interactions between spatial variation in habitat and dispersal dynamics, the extent to which geographic variation in temporal variability can also shape range-wide patterns in dispersal traits has not been synthesized. In this paper, we summarize key predictions from metapopulation models that evaluate how dispersal evolves in response to spatial and temporal habitat variability. Next, we compile empirical data that quantify temporal variability in plant demography and patterns of dispersal trait variation across species ranges to evaluate the hypothesis that higher temporal variability favors increased dispersal at plant range limits. We found some suggestive evidence supporting this hypothesis while more generally identifying a major gap in empirical work evaluating plant metapopulation dynamics across species ranges and geographic variation in dispersal traits. To address this gap, we propose several future research directions that would advance our understanding of the interplay between spatiotemporal variability and dispersal trait variation in shaping the dynamics of current and future species ranges.     

Areas of high conservation value at risk by plant invaders in Georgia under climate change

Invasive alien plants (IAP) are a threat to biodiversity worldwide. Understanding and anticipating invasions allow for more efficient management. In this regard, predicting potential invasion risks by IAPs is essential to support conservation planning into areas of high conservation value (AHCV) such as sites exhibiting exceptional botanical richness, assemblage of rare, and threatened and/or endemic plant species. Here, we identified AHCV in Georgia, a country showing high plant richness, and assessed the susceptibility of these areas to colonization by IAPs under present and future climatic conditions. We used actual protected areas and areas of high plant endemism (identified using occurrences of 114 Georgian endemic plant species) as proxies for AHCV. Then, we assessed present and future potential distribution of 27 IAPs using species distribution models under four climate change scenarios and stacked single‐species potential distribution into a consensus map representing IAPs richness. We evaluated present and future invasion risks in AHCV using IAPs richness as a metric of susceptibility. We show that the actual protected areas cover only 9.4% of the areas of high plant endemism in Georgia. IAPs are presently located at lower elevations around the large urban centers and in western Georgia. We predict a shift of IAPs toward eastern Georgia and higher altitudes and an increased susceptibility of AHCV to IAPs under future climate change. Our study provides a good baseline for decision makers and stakeholders on where and how resources should be invested in the most efficient way to protect Georgia's high plant richness from IAPs.     

Embryos of non-native anoles are robust to urban thermal environments

The transformation of natural habitats into urban landscapes dramatically alters thermal environments, which in turn, can impact local biota. Ectothermic organisms that are oviparous are particularly sensitive to these altered environments because their embryos cannot behaviorally thermoregulate and the surrounding environment determines the temperature experienced during development. We studied the effects of urban and forested thermal environments on embryo development and hatchling phenotypes in two non-native lizards (Anolis sagrei and A. cristatellus) in metropolitan Miami, Florida. To determine if embryos from urban and forested sites are adapted to their respective thermal environments, we incubated eggs from each site using temperatures that simulate likely nest conditions in both urban and forested environments. For both species, urban thermal environments accelerated embryonic development, but had no impact on egg survival or any of the phenotypic traits that were measured (e.g., body size, running performance, and locomotor behavior). Our results provide no evidence that embryos from urban and forested sites are adapted to their respective thermal environments. Instead, the lack of any major effects suggest that embryos of both species are physiologically robust with respect to novel environments, which could have facilitated their success in establishing in non-native ranges and in human-modified landscapes.     

Population structure and dynamics of the clonal dwarf-shrub Linnaea borealis

Abstract. The demography of the long-lived clonal dwarf-shrub Linnaea borealis was studied during four years in a coniferous forest in central Sweden. The main object was to infer patterns of temporal variation in population dynamics of this species. The shoot population is organized in fragments, i.e. physically connected systems of shoots partly covered by the moss carpet. The age and size structure of the fragment population is described, but shoots are more convenient units for a study of population dynamics. A stochastic model of shoot population dynamics was constructed, and simulations indicated a considerable temporal variation in population size and flowering. Hence, variability as such is an essential aspect of the dynamics of established populations of Linnaea. Simulations of extinction risks revealed that small-sized shoot populations (ca. 250 shoots) are likely to be long-lived when experiencing environmentally induced demographic variation of the range observed. Mortality agents for established genets, such as large-scale disturbances, were not incorporated in the models. Some implications of variable population growth rates in clonal plants in woodlands are discussed.     

Using avatar species to model the potential distribution of emerging invaders

Aim Anticipating the potential distributions of emerging invasive species is complicated by the tendency for species distribution models to perform better when both native and invasive range data are available for model development. If invasive range data are lacking, species models are liable to under‐estimate distributions for emerging invaders, particularly for species that are not at equilibrium with their native range environment due to historical factors, dispersal limitation and/or ecological interactions. We demonstrate the potential to use well‐quantified niche shifts from established ‘avatar’ (i.e. the remote or virtual manifestation of an entity) invaders to develop plausible distributions for data‐poor emerging invaders contingent on niche shifts of similar magnitude or character. Location Global. Methods Using the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii as our avatar invaders, we quantify how niche position, size and structure differs between native and total ranges using Mahalanobis distance (a measure of multivariate similarity) and the climate predictors of annual minimum and maximum air temperature. We then generalize patterns of niche shift from these species to the emerging crayfish invader Cherax quadricarinatus. Results Some patterns of niche shifts were similar for Pacifastacus leniusculus and Procambarus clarkii, but niche shifts were of considerably greater magnitude for P. clarkii. When a native range model for C. quadricarinatus was modified with generalized niche shifts similar to Pacifastacus leniusculus and Procambarus clarkii, the potential global distribution for this species increased considerably, including many areas not identified by the native range model. Main conclusions We illustrate the potential to use avatar invaders to provide cautionary, niche shift‐assuming species distribution models for emerging invaders. Many theoretical and applied implications of the avatar species concept require additional investigation, including the development of frameworks to select appropriate avatar species and evaluate the performance of avatar‐derived models for emerging invaders. Despite these research needs, we believe this concept will have considerable utility for predicting vulnerability to invasion by data‐poor species; this is a critical management need because shifting pathways of introduction and climate change will produce many novel, emerging invasive species in the future.     

The role of migration in the genetic structure of populations in temporally and spatially varying environments II. Island Models

The Island Model introduced by Sewall Wright (1951) has proven to be a useful construction for studying the interaction of genetic drift, population subdivision, and mutation. Interest in the model has recently increased because of its relevance to certain questions involving the rate of differentiation of sub-populations under the neutral allele hypothesis (e.g., Smith, 1970; Latter, 1973). It is perhaps the only realistic population structure in which the test for neutrality proposed by Lewontin and Krakauer (1973) is valid (Lewontin and Krakauer, 1975). If data from natural populations is to be compared to the predictions of the Island Model, it is desirable to have an alternative model with the same migration pattern but with natural selection operating. In this paper one such model will be introduced where the stochastic element comes from random fluctuations in the environment rather than from genetic drift. The model is a direct extension of the one in the previous paper in this series (Gillespie, 1975) which dealt with a population which is subdivided into two patches with restricted migration between them.     

The importance of sampling scale in the assessment of intraspecific life-history differences in plains killifish,Fundulus zebrinus,populations (Pisces: Cyprinodontidae)

Synopsis The purpose of this study was to determine the effects of unpredictable environmental fluctuations on the demographic and genetic structure of Fundulus zebrinus populations. Collections of F. zebrinus were taken from three rivers in the Arkansas River basin: the Arkansas, Chikaskia, and Ninnescah. Fish were sampled from three sites on each river on nine collection dates throughout 1984 and 1985. Totals of 2100 fish and 6000 fish were included in electrophoretic and demographic analyses, respectively. The results of the study indicate that within a limited geographic region (i.e. within rivers) spatial differences and temporal changes in both demographic and genetic population characteristics occur frequently and are primarily stochastic. However, on a larger spatial scale (i.e. across rivers), general trends emerge for demographic and especially for genetic population characteristics. These results illustrate the importance of sampling scale for conclusions of life-history evolution in fluctuating environments. In addition, it was found that regulation of Fundulus zebrinus populations includes an important density-independent component. Stochastic demographic differences across space and changes through time and spatially and temporally heterogeneous allele frequencies, are both indicative of density-independent regulation. Variation in population parameters, both demographic and genetic, was observed between populations sampled from each river. These population differences were attributed to differences between the rivers themselves.     

The demography of native and non-native plant species in mountain systems: examples in the Greater Yellowstone Ecosystem

In mountainous areas, native and non-native plants will be exposed to climate change and increased disturbance in the future. Non-native plants may be more successful than natives in disturbed areas and thus be able to respond quicker to shifting climatic zones. In 2009, monitoring plots were established for populations of a non-native species (Linaria dalmatica) and a closely related native species (Castilleja miniata) on an elevation gradient in the Greater Yellowstone Ecosystem, USA. Population data were collected twice during the growing season for 3 years and used to calculate population vital rates for both species, and to construct population dynamics models for L. dalmatica. Linaria dalmatica vital rates were more associated with climatic/environmental factors than those of C. miniata. Population dynamics models for L. dalmatica showed no trend in population growth rate (λ) vs. elevation. The highest λ corresponded with the lowest vegetation and litter cover, and the highest bare ground cover. All populations with λ < 1 corresponded with the lowest measured winter minimum temperature. There was a negative association between λ and number of weeks of adequate soil moisture, and a weak positive association between λ and mean winter minimum temperature. Variance in vital rates and λ of L. dalmatica suggest broad adaptation within its current range, with the potential to spread further with or without future changes in climate. There is evidence that λ is negatively affected by persistent soil moisture which promotes the growth of other plant species, suggesting that it might expand further if other species were removed by disturbance.     

Optimal conservation strategy in fluctuating environments with species interactions: resource-enhancement of the native species versus extermination of the alien species

Alien species are often a major threat to native species. We consider optimal conservation strategies for a population whose viability is affected both by an alien species (such as a competitor, a predator, or a pathogen) and by random fluctuations of the environment (e.g. precipitation, temperature). We assume that the survivorship of the native population can be improved by providing resources such as food and shelter, and also by an extermination effort that decreases the abundance of the alien species. These efforts decrease the extinction probability of the native population, but they are accompanied by economic costs. We search for the optimal strategy that minimizes the weighted sum of the extinction probability and the economic costs over a single year. We derive conditions under which investment should be made in both resource-enhancement and extermination, and examine how the optimal effort levels change with parameters. When the optimal strategy includes both types of efforts, the optimal extermination effort level turns out to be independent of the density and economic value of the native species, or the variance of the environmental fluctuation. Furthermore, the optimal resource-enhancement effort is then independent of the density of the alien species. However, the parameter dependencies greatly change if one of the efforts becomes zero. We also examine the situation in which the impact of the alien species is uncertain. The optimal extermination effort increases with the uncertainty of this impact except when the cost of extermination is very high.     

Demographic consequences of drought in the herbaceous perennial Cryptantha flava: effects of density,associations with shrubs,and plant size

The demographic consequences of a severe drought year were examined for two experimental plantings of the herbaceous desert perennial Cryptantha flava(Boraginaceae) in northeastern Utah, United States. A total of 6680 nutlets were planted individually or in clusters of four both under shrubs and in open microhabitats within two natural populations. Survival, growth, and flowering as a function of density and microhabitat were followed for 7 years, including 1 year when precipitation just before and during the growing season was 74.5% below normal. The design permitted assessment of how intraspecific density and shrub cover affect demographic response to drought. Mortality increased and flowering decreased dramatically during drought but neither varied with density or between shrub and open microhabitats. For plants growing under shrubs, survival (at Site 1) and growth (at Site 2) varied with shrub species. Average aboveground plant size also decreased during drought. Population size hierarchies were rearranged because larger plants lost leaf rosettes while many smaller plants grew. Density and microhabitat affected plant performance in non-drought years but more often at Site 1 than at Site 2. Individuals growing alone often were more likely to flower and/or produced more inflorescences when they did flower than did individuals growing with at least one other plant. However, for 2 years, survival rates at Site 1 were higher for plants growing in clumps than for single individuals. Shrubs also had mixed effects on plant performance. In some years, survival was higher under shrubs, but at Site 1 plants in the open often were more likely to flower and/or produced more inflorescences. Thus despite severe demographic consequences of drought, the study provided no evidence that intraspecific competition, interference by shrubs, or facilitation by shrubs increases under limited soil water.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.