Shifting dispersal modes at an expanding species’ range margin

While it is generally recognized that noncontiguous (long‐distance) dispersal of small numbers of individuals is important for range expansion over large geographic areas, it is often assumed that colonization on more local scales proceeds by population expansion and diffusion dispersal (larger numbers of individuals colonizing adjacent sites). There are few empirical studies of dispersal modes at the front of expanding ranges, and very little information is available on dispersal dynamics at smaller geographic scales where we expect contiguous (diffusion) dispersal to be prevalent. We used highly polymorphic genetic markers to characterize dispersal modes at a local geographic scale for populations at the edge of the range of a newly invasive grass species (Brachypodium sylvaticum) that is undergoing rapid range expansion in the Pacific Northwest of North America. Comparisons of Bayesian clustering of populations, patterns of genetic diversity, and gametic disequilibrium indicate that new populations are colonized ahead of the invasion front by noncontiguous dispersal from source populations, with admixture occurring as populations age. This pattern of noncontiguous colonization was maintained even at a local scale. Absence of evidence for dispersal among adjacent pioneer sites at the edge of the expanding range of this species suggests that pioneer populations undergo an establishment phase during which they do not contribute emigrants for colonization of neighbouring sites. Our data indicate that dispersal modes change as the invasion matures: initial colonization processes appear to be dominated by noncontiguous dispersal from only a few sources, while contiguous dispersal may play a greater role once populations become established.     

Micro-colinearity between rice, <Emphasis Type="Italic">Brachypodium</Emphasis>, and <Emphasis Type="Italic">Triticum monococcum</Emphasis> at the wheat domestication locus <Emphasis Type="Italic">Q</Emphasis>

Brachypodium, a wild temperate grass with a small genome, was recently proposed as a new model organism for the large-genome grasses. In this study, we evaluated gene content and microcolinearity between diploid wheat (Triticum monococcum), Brachypodium sylvaticum, and rice at a local genomic region harboring the major wheat domestication gene Q. Gene density was much lower in T. monococcum (one per 41 kb) because of gene duplication and an abundance of transposable elements within intergenic regions as compared to B. sylvaticum (one per 14 kb) and rice (one per 10 kb). For the Q gene region, microcolinearity was more conserved between wheat and rice than between wheat and Brachypodium because B. sylvaticum contained two genes apparently not present within the orthologous regions of T. monococcum and rice. However, phylogenetic analysis of Q and leukotriene A-4 hydrolase-like gene orthologs, which were colinear among the three species, showed that Brachypodium is more closely related to wheat than rice, which agrees with previous studies. We conclude that Brachypodium will be a useful tool for gene discovery, comparative genomics, and the study of evolutionary relationships among the grasses but will not preclude the need to conduct large-scale genomics experiments in the Triticeae.     

Salt tolerance of two perennial grass <Emphasis Type="Italic">Brachypodium sylvaticum</Emphasis> accessions

Key message

We studied the salt stress tolerance of two accessions isolated from different areas of the world (Norway and Tunisia) and characterized the mechanism(s) regulating salt stress in Brachypodium sylvaticum Osl1 and Ain1.

Abstract

Perennial grasses are widely grown in different parts of the world as an important feedstock for renewable energy. Their perennial nature that reduces management practices and use of energy and agrochemicals give these biomass crops advantages when dealing with modern agriculture challenges such as soil erosion, increase in salinized marginal lands and the runoff of nutrients. Brachypodium sylvaticum is a perennial grass that was recently suggested as a suitable model for the study of biomass plant production and renewable energy. However, its plasticity to abiotic stress is not yet clear. We studied the salt stress tolerance of two accessions isolated from different areas of the world and characterized the mechanism(s) regulating salt stress in B. sylvaticum Osl1, originated from Oslo, Norway and Ain1, originated from Ain-Durham, Tunisia. Osl1 limited sodium transport from root to shoot, maintaining a better K/Na homeostasis and preventing toxicity damage in the shoot. This was accompanied by higher expression of HKT8 and SOS1 transporters in Osl1 as compared to Ain1. In addition, Osl1 salt tolerance was accompanied by higher abundance of the vacuolar proton pump pyrophosphatase and Na⁺/H⁺ antiporters (NHXs) leading to a better vacuolar pH homeostasis, efficient compartmentation of Na⁺ in the root vacuoles and salt tolerance. Although preliminary, our results further support previous results highlighting the role of Na⁺ transport systems in plant salt tolerance. The identification of salt tolerant and sensitive B. sylvaticum accessions can provide an experimental system for the study of the mechanisms and regulatory networks associated with stress tolerance in perennials grass.

     

The effects of genetic and environmental factors on disease expression (stroma formation) and plant growth in Brachypodium sylvaticum infected by Epichloë sylvatica

Fungi in the genus Epichloë (Clavicipitaceae, Ascomycota) are endophytic and often mutualistic symbionts of many grasses in temperate areas. Species with a sexual cycle suppress host flowering and seed formation, whereas asexual fungi remain asymptomatic and transmit vertically by seed. Thus, the mode of reproduction may determine whether the symbiosis is mutualistic or parasitic. The level of sexual reproduction (disease expression) varies among different endophytes and on different grass hosts, but factors responsible for this variation, and evolutionary mechanisms leading to one or the other life strategy are not understood. As experimental system, we chose Brachypodium sylvaticum in which the endophyte E. sylvatica can express both reproduction modes. A field experiment was done in plots of a free air carbon dioxide enrichment (FACE) facility. We investigated the effects of three environmental factors (elevated CO2 concentration, shading and fertilisation) and one genetic factor (plant and fungal genotype combination), on plant growth and disease expression. Variation in plant growth was mainly dependant on the genotype and was increased by fertilisation. Elevated CO2 and shading slightly stimulated plant growth, but only in fertilised plants. Disease expression was overwhelmingly dependent on the genotype, hence genetic factors. Fertilisation slightly stimulated disease expression in some genotypes, while the effect of elevated CO2 was negligible, and by interaction with fertilisation inconsistent in the two years. Horizontal transmissions during the experiment, presumably mediated by ascospores, confounded the original infection status of the plants. Contagious infections occurred more frequently in the shade, and in endophyte‐free host plants. The latter suggests that pre‐existing infections render host plants less susceptible to superinfection by choke forming strains. Although our results clearly indicate that disease expression of E. sylvatica has a genetic basis, it is still unclear whether selection on the plants or the fungi is driving the evolution of this association.     

Phylogenetic reconstruction of the genusBrachypodium P. Beauv. (Poaceae) from combined sequences of chloroplastndhF gene and nuclear ITS

A phylogenetic reconstruction of eight species of the genusBrachypodium P. Beauv. (Poaceae) has been obtained combining sequence data from the chloroplastndhF gene and the nuclear ITS, and using five representatives of tribes Triticeae (Secale), Poeae (Lolium), Meliceae (Melica, Glyceria) and Oryzeae (Oryza) as out-groups. Similar numbers of informative substitutions for the ingroup species were provided by both the 3 region of the chloroplastndhF gene and the nuclear ITS region. The Mediterranean annualBrachypodium distachyon appears to be the basal lineage, followed by the divergence of the New World non-rhizomatousB. mexicanum, which antedates the separation of a core of six European and Eurosiberian rhizomatous perennials (Brachypodium arbuscula, B. retusum, B. rupestre, B. phoenicoides, B. pinnatum, andB. sylvaticum). The evolutionary reconstruction based on sequences of the chloroplast and the nuclear genomes is congruent with topologies obtained from analysis of RAPD data.     

<Emphasis Type="Italic">PRUNUS</Emphasis>: a spatially explicit demographic model to study plant invasions in stochastic,heterogeneous environments

To model the invasion of Prunus serotina invasion within a real forest landscape we built a spatially explicit, non-linear Markov chain which incorporated a stage-structured population matrix and dispersal functions. Sensitivity analyses were subsequently conducted to identify key processes controlling the spatial spread of the invader, testing the hypothesis that the landscape invasion patterns are driven in the most part by disturbance patterns, local demographical processes controlling propagule pressure, habitat suitability, and long-distance dispersal. When offspring emigration was considered as a density-dependent phenomenon, local demographic factors generated invasion patterns at larger spatial scales through three factors: adult longevity; adult fecundity; and the intensity of self-thinning during stand development. Three other factors acted at the landscape scale: habitat quality, which determined the proportion of the landscape mosaic which was potentially invasible; disturbances, which determined when suitable habitats became temporarily invasible; and the existence of long distance dispersal events, which determined how far from the existing source populations new founder populations could be created. As a flexible “all-in-one” model, PRUNUS offers perspectives for generalization to other plant invasions, and the study of interactions between key processes at multiple spatial scales.     

Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NW Greece)

We investigate the persistent soil seed bank composition and its relation to the above-ground flora of grazed and non-grazed sub-Mediterranean deciduous oak forests of NW Greece. Twenty-eight taxa were recorded in the soil seed bank and 83 taxa (70 taxa in plots of seed bank sampling) in the above-ground vegetation. The dominant tree species and many woodland species found in the above-ground vegetation were absent from the soil seed bank. Similarity between the soil seed bank and the above-ground vegetation decreased with grazing, and grazing led to a decrease of species richness in above-ground vegetation and soil seed bank. Beta diversity of vegetation among grazed and among non-grazed plots did not differ, but was significantly higher between grazed and non-grazed areas. Beta diversity of the soil seed bank declined with grazing. When applying classification tree and logistic regression analyses, non-grazed forest sites are clearly differentiated by the presence of Phillyrea latifolia, Euphorbia amygdaloides and Brachypodium sylvaticum. PCA ordination of above-ground species composition reflected a gradient from sites grazed by ruminants to non-grazed sites, but no clear structure was detected in the seed bank.     

Spatial autocorrelation and the analysis of invasion processes from distribution data: a study with the crayfish <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Complex spatial dynamics are frequent in invasive species; analyzing distribution patterns can help to understand the mechanisms driving invasions. We used different spatial regression techniques to evaluate processes determining the invasion of the red swamp crayfish Procambarus clarkii. We evaluated four a priori hypotheses on processes that may determine crayfish invasion: landscape alteration, connectivity, wetland suitability for abiotic and biotic features. We assessed the distribution of P. clarkii in 119 waterbodies in a recently invaded area. We used spatially explicit statistical techniques (spatial eigenvector mapping, generalized additive models, Bayesian intrinsic conditional autoregressive models) within an information-theoretic framework to assess the support of hypotheses; we also analyzed the pattern of spatial autocorrelation of data, model residuals, and eigenvectors. We found strong agreement between the results of spatial eigenvector mapping and Bayesian autoregressive models. Procambarus clarkii was significantly associated with the largest, permanent wetlands. Additive models suggested also association with human-dominated landscapes, but tended to overfit data. The results indicate that abiotic wetlands features and landscape alteration are major drivers of the species’ distribution. Species distribution data, residuals of ordinary least squares regression, and spatial eigenvectors all showed positive and significant spatial autocorrelation at distances up to 2,500 m; this may be caused by the dispersal ability of the species. Our analyses help to understand the processes determining the invasion and to identify the areas most at risk where screening and early management efforts can be focused. The comparison of multiple spatial techniques allows a robust assessment of factors determining complex distribution patterns.     

Unravelling the effects of temperature, latitude and local environment on the reproduction of forest herbs

Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number × seed mass, while germinable seed output (GSO) was expressed as seed number × germination percentage. We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.     

A new hybrid horsetail Equisetum×lofotense (E. arvense×E. sylvaticum,Equisetaceae) from Norway

On the Island of Austvågøya (Lofoten, Norway), a new hybrid horsetail (E. arvense×E. sylvaticum) was found, which is described as E.×lofotense. Its macro‐ and micro‐morphological characters are discussed and compared with the similar hybrid E.×mildeanum (E. pratense×E. sylvaticum) with which it shares the E. sylvaticum parentage. The hybrid nature is verified by its completely aborted spores, and the micro‐morphology of the stem ridges and stomata, illustrated by SEM images, confirm the proposed origin. An overview of all known hybrids in the genus Equisetum is given and critically commented. It becomes evident that in subgenus Equisetum E. arvense is the species most frequently involved in hybridization events.     

EVIDENCE FOR GENETIC DIFFERENTIATION BETWEEN CHOKE-INDUCING AND ASYMPTOMATIC STRAINS OF THE EPICHLOË GRASS ENDOPHYTE FROM BRACHYPODIUM SYLVATICUM

Life cycle and breeding system variation in Epichloë grass endophytes (choke disease) is tightly linked to the degree of stroma formation. It is not known whether this variation results from differences in host resistance, fungal virulence, or environmental conditions. We found genetic differentiation between 173 asymptomatic (NS) and 93 stromata-forming (S) Epichloë strains isolated from one grass species, Brachypodium sylvaticum, based on 13 presumed allozyme loci, of which six were variable. The fungal strains originated from 10 sites in Switzerland, three sites of which were represented by both NS and S subpopulations. In total, 19 allozyme genotypes, that were nonrandomly distributed among S and NS were detected. Genetic variation measured as G_ST between S and NS strains isolated from the same site ranged from 0.73 to 0.98. Clonality, measured as linkage disequilibrium at one site, was significant in the NS subpopulation (P ? 0.001), but not in the S subpopulation (P = 0.21), implying asexual reproduction by NS strains as well as successful horizontal transmission of S strains. Since all seeds are usually infected vegetatively, horizontal transmission implies the occurrence of multiple host infections. Altogether, these results provide indirect evidence that NS and S strains do not belong to one panmictic population and that differentiation patterns of stroma formation found in nature are due to genetic differences among fungi in associations with their host plants. We discuss the direction of evolution of disease expression in this system. The distribution of genetic variability suggests that the asymptomatic strains were derived from stromata-forming populations.     

<Emphasis Type="Italic">Prunus serotina</Emphasis> unleashed: invader dominance after 70 years of forest development

Propagule pressure and disturbance have both been found to facilitate invasion. Therefore, knowledge on the history of introduction and disturbance is vital for understanding an invasion process, and research should focus on areas in which the invasive species has not been deliberately introduced or managed to study unconfounded colonization patterns. Comparing the outcome of such spontaneous colonization processes for different ecosystems might provide a useful framework for setting management priorities for invasive species that enter new, uninvaded areas. We focused on the 70-year spontaneous spread of the invasive tree species Prunus serotina in a pine forest in the Netherlands. To reconstruct the invasion pattern, we combined historical maps, tree ring analysis, spatially explicit tree inventory data, seed density data, and regeneration data for both native and non-native species. Prunus serotina was the only species that showed successful regeneration: the species was present throughout the forest in the tree, shrub, and herb layer. Native species were not able to outgrow the seedling stage. Our data demonstrate that P. serotina is a gap-dependent species with high seed production that builds up a seedling bank. We also compared the results of this study with a similar study on P. serotina colonization in a deciduous forest in Belgium, where P. serotina invasion was not successful. The sharp contrast between the outcomes of the two invasion processes shows the importance of studying an invasive species and the recipient ecosystem jointly and made us raise the hypothesis that herbivore pressure may facilitate P. serotina invasion.     

Observations on patterns of granular fertiliser deposition beside hedges and its likely effects on the botanical composition of field margins

Patterns of granular fertiliser deposition differed markedly between a pneumatic boom applicator and a spinning disk machine, both of which were tractor-mounted. The pneumatic applicator gave relatively even distribution across the boom width. A spinning disk gave more variable deposition and significant amounts were spread further than the expected 12 m overlapping pattern. When operating with a deflector plate on the spinning disk, significant amounts of fertiliser were also spread further than the expected 6 m. However, when operating next to a hedge, there was evidence of fertiliser concentration at the base of the hedge and prevention of granules passing through to adjacent habitat. A peak concentration of up to 150 kg ha^-1 of fertiliser was deposited at the hedge/field edge interface. In a tray experiment on seedling competition and establishment, there was no evidence of nitrogen fertiliser effects, at rates found in the field, on early plant establishment or species diversity. However, in a competition experiment with established plants of four grass species grown in pots, the nitrophilous species Bromus sterilis was able to increase growth at increasing nitrogen level, at the expense of slow-growing Brachypodium sylvaticum and Anthoxanthum odoratum. Whilst hedges may buffer fertiliser contamination of habitats adjacent to agricultural fields, deposition of concentrated nitrogen fertiliser beside the hedge is likely to encourage the growth of nitrophilous plant species. In the short term, this may not affect botanical composition or diversity, but to reduce the likely long-term adverse effects of fertiliser misplacement, we recommend that farmers are encouraged to use pneumatic fertiliser applicators and to introduce vegetation buffer strips at field edges.     

Prediction and error in multi‐stage models for spread of aquatic non‐indigenous species

Aim Predictions of spread of non‐indigenous species allow for greater efficiency in managing invasions by targeting areas for preventative measures. The invasion sequence is a useful concept in predictions of spread, as it allows us to test hypotheses about the transport and establishment of propagules in novel habitats. Our aims are twofold: (1) to develop and validate multi‐stage invasion models for the introduced fishhook waterflea, Cercopagis pengoi, and (2) to assess how variability in the transport patterns of the propagules influences the accuracy and spatial extent for predictions of spread. Location New York State, USA. Methods We developed a two‐stage model for the spread of C. pengoi. First, we developed a stochastic gravity model for dispersal based on surveys of recreational boat traffic in New York State as a proxy for propagule pressure. We then modelled the probability of establishment based on predicted levels of propagule pressure and measures of lakes’ physicochemistry. In addition, we used Monte Carlo simulations based on the gravity model to propagate variability in boater traffic through the establishment model to assess how uncertainty in dispersal influenced predictions of spread. Results The amount recreationalists were willing to spend, lake area and population size of the city nearest to the destination lake were significant factors affecting boater traffic. In turn, boater traffic, lake area, specific conductance and turbidity were significant predictors of establishment. The inclusion of stochastic dispersal reduced the rate of false positives (i.e. incorrect prediction of an invasion) in detecting invasions at the upper 95% prediction interval for the probability of establishment. Main conclusions Combinations of measures of propagule pressure, habitat suitability and stochastic dispersal allow for the most accurate predictions of spread. Further, multi‐stage spread models may overestimate the extent of spread if stochasticity in early stages of the models is not considered.     

Seed predation of two alien Opuntia species invading Mediterranean communities

Invasibility depends on the interaction of the introduced species with the abiotic and biotic factors of the recipient community. In particular, the biotic resistance posed by native herbivores has been claimed to be of great importance in controlling plant invasion. We investigated fruit and seed predation of two exotic Opuntia species within and between Mediterranean communities in order to determine how patterns of predation matched patterns of invasion. Predators were small mammals, presumably mice, which could consume more than 50% of the seeds produced. Predators could be equally effective in consuming fruit and single seeds. O. maxima fruits were slightly preferred to O. stricta fruits, but predators did not distinguish between seeds. Seed predation was more intense in invaded than in non-invaded communities. However, there was a high spatial variation in seed predation that did not always match patterns of invasion, suggesting that seed predation alone is not a good predictor of community invasibility to Opuntia. According to these results invasibility to Opuntia is limited in some (but not all) communities by native mice. Seed losses by predation were high for both species. However, we estimated that more than 75% of seeds dispersed by birds to non-invaded areas are not predated.     

Meristem activity and biomass production as response mechanisms in two forest herbs

Actaea spicata and Geranium sylvaticum are deciduous rosette herbs, frequently coexisting in nutrient-rich forests of boreal Sweden. A. spicata is restricted to this habitat whereas G. sylvaticum has a wider ecological amplitude. To explore the different distributions of these species we studied the growth mechanisms by which they responded to nutrient and light supply. Plants raised from seeds were grown under four combinations of light and nutrient supplies and growth was followed for 20 weeks. Growth responses were measured as biomass accumulation and meristem activity, i.e. leaf development by shoot meristems. Biomass accumulation and meristem activity were affected by nutrient and light treatments. However, A. spicata responded more to nutrients, whereas G. sylvaticum responded more to light, and low resource supply shortened the time during which shoot meristems of A. spicata developed leaves without a corresponding effect on biomass accumulation. In G. sylvaticum, the periods of leaf development and biomass accumulation were the same, but a relationship between high development rates and peak allocation of biomass to shoot meristems indicated a storage function of these during leaf development. We conclude that the inflexible growth of A. spicata, mainly affected by nutrient supply, makes it less competitive in open habitats and restricts it to shaded, nutrient-rich habitats. G. sylvaticum should be more successful in habitats subjected to unpredictable disturbances, due to a flexible growth morphology and a dynamic function of meristems that buffers against variation in nutrient supply and facilitates positive light responses. Meristematic data should be included in interspecific comparisons on growth responses to different resource supplies. Unless species have the same growth mechanism, it is not satisfactory to study biomass-related traits alone, as internal constraints concerning plant development might confuse the interpretation.     

Exploring valid reference genes for gene expression studies in <Emphasis Type="Italic">Brachypodium distachyon</Emphasis>by real-time PCR

Background  

The wild grass species Brachypodium distachyon (Brachypodium hereafter) is emerging as a new model system for grass crop genomics research and biofuel grass biology. A draft nuclear genome sequence is expected to be publicly available in the near future; an explosion of gene expression studies will undoubtedly follow. Therefore, stable reference genes are necessary to normalize the gene expression data.     

Contrasting community responses to fertilization and the role of the competitive ability of dominant species

Question: Do contrasting biotic contexts in nutrient‐poor grasslands affect the predictability of invasion by exploitative species following fertilization? Location: French Alps. Methods: We examined community responses after 2 years of nutrient addition for two nutrient‐poor European calcareous grasslands, a mesoxeric community dominated by the short bunchgrass Bromus erectus and a mesic community dominated by the tall rhizomatous grass Brachypodium rupestre. We also performed reciprocal transplantations of these two dominant slow‐growing species and Arrhenatherum elatius, a tall fast‐growing grass that dominates nutrient‐rich communities and is likely to invade nutrient‐poor communities after fertilization. Transplants were grown with or without neighbors, in order to measure their individual responses (without neighbors) and competition intensity (by comparing performances with and without neighbors using the Relative Neighbor Effect index – RNE) during one growing season in all three communities. Results: In the Bromus community, fertilization induced a strong increase in fast‐growing grasses (including A. elatius). Competition intensity was low for the three transplanted grasses, but strongly increased with resource addition, to reach values observed in the Arrhenatherum community. In the Brachypodium community, no change in competition intensity with fertilization was detected, because of the high mortality of the two “non‐resident” species, irrespective of the presence of neighbors. Conclusions: Community responses to nutrient improvement are context‐dependent and vary as a function of the biotic environment. Soil processes are proposed as the main drivers of community resistance to the invasion of fast‐growing species in the mesic, nutrient‐poor grassland dominated by the large conservative competitor B. rupestre.