Do plant functional types based on leaf dry matter content allow characterizing native grass species and grasslands for herbage growth pattern?

Few studies have focused on vegetation characteristics of importance to feeding domestic herbivores, mainly the seasonal pattern of herbage growth at spring. Our objective is to establish and to evaluate a simple method of ranking grassland communities for these characteristics. We combined approaches at plant species level (comparison of grass species growing in a pure stand) and plant community level (comparison of grasslands differing mainly in their nutrient availability). Firstly, we ask if the ranking of species by leaf dry matter content (LDMC), a functional parameter used to assess the plant strategy for resource acquisition and use, is consistent with a classification of the species using three plant features that determine plant growth pattern at spring (beginning and ending of stem elongation, leaf lifespan). Secondly, for three networks of natural grasslands, we test whether there is consistency when ranking them by their dominant plant functional type (PFT A, B or C) established previously at species level, and by the three agronomic characteristics. For species growing in pure stands, there was a significant effect of PFT for the three plant features. For species having a low LDMC (A and B PFT), there were earlier stem elongation in the season, earlier flowering and shorter leaf lifespan. The opposite was observed for species having a high LDMC (C and D PFT). For grassland communities dominated by A-PFT, the ceiling yield for leaves and stems occurred earlier in spring than for those dominated by C-PFT. Results were consistent at plant and community levels. Scaling up from plant to community was well mediated by PFT. Plant features which characterize species for resource acquisition and use are consistent with herbage growth patterns at plant community level. These results show that herbage growth pattern and composition depend on PFTs and that knowing the PFT dominance is of great importance to plan the use of grasslands. We can expect to use the PFT approach to perform vegetation diagnosis at field level when the objective is to rank grassland communities for their agronomic characteristics. Fernando L. F. de Quadros was supported by CAPES and CNPq (BP-2).     

Plant Functional Types of Woody Species Related to Fire Disturbance in Forest–Grassland Ecotones

The study of plant functional types (PFTs) has been widely emphasized when analysing plant community changes in relation to variations in climate and disturbance regime. In this study, we search for PFTs of woody species near forest–grassland boundaries in South Brazil where, due to climate, forests tend to expand over grassland but are being restricted by frequent fires. We aimed at answering the questions: (i) which plant functional types of forest woody species can establish in adjacent grassland subject to fire disturbance and (ii) which plant functional types of forest and grassland woody species are related to short-term community dynamics in frequently burned grassland. Traits were assessed in woody plants in 156 plots (6.75 m²) arranged in 12 transects across forest–grassland boundaries with different fire history in their grassland part. The analysis used a recursive algorithm to search for traits and PFTs maximally associated to spatial distance from forest limit in one analysis, and elapsed time since last fire in another. As a result, nine PFTs of forest woody species were identified that best described community patterns associated to distance from forest. Resprouting ability characterized forest plants able to colonize grasslands. PFT diversity was higher in border plots than inside forest or grassland. Four PFTs of forest and grassland woody species best described woody species community patterns in the grassland associated to elapsed time since fire. Taller individuals of single-stemmed shrubs predominated in late post-fire recovery (3–4 years), while shorter multi-stemmed shrubs in recently burned areas (3 months to 1 year). PFTs of forest trees occurred in border plots or, as established adults, in grassland, remaining unaffected by fire. We conclude that easily measurable structural plant traits, such as those used in our study, are sufficient to evaluate post-fire community dynamics. Forest PFTs in burned grassland are restricted to those with resprouting ability to survive recurrent fire events. Establishment success is highest on protected sites with lesser or low-intensity fire.     

Conservation and restoration of Nardus grasslands in the Swiss northern Alps

Aim: Species‐rich Nardus grasslands are high nature‐value habitats. In Switzerland, many of these grasslands are degraded even though they have been under protection since the 1980s. Degradation shows two divergent trends: Nardus grasslands are either dominated by Nardus stricta or by eutrophic plants, both trends leading to the disappearance of typical Nardus grassland species. With this study, we aim to identify the factors that could be adjusted to conserve the integrity of this habitat. Location: Bernese Alps, Switzerland. Methods: In 2016, we investigated the underlying causes of this degradation process by assessing vegetation composition in 48 Nardus grasslands located in the Swiss northern Alps of canton Bern and linking it to soil, management and environmental variables. To explore the effect of the degradation on higher trophic levels, orthopteran species richness and densities were assessed. Results: Results show that Nardus meadows (mown) are rarely degraded compared to Nardus pastures (grazed). Within pastures, eutrophic plants are most abundant on small pastures with low soil carbon/nitrogen ratio, indicating high nutrient availability. Nardus stricta dominance is most problematic on north‐exposed slopes and in summer pastures. A plausible driver of both degradation trends is the grazing management regime: within small pastures at low elevation where the grazing periods are short but intense, soil carbon/nitrogen ratio is low because of high dung deposition, thus the eutrophic species become dominant. Contrastingly, on large summer pastures with low‐intensity and long‐term grazing, N. stricta becomes dominant due to selective grazing. Both degradation trends show a negative impact on the orthopteran density. Conclusion: Species‐rich Nardus grasslands are a precious alpine habitat for specialised plant species and orthopterans. With an extensive mowing regime or a more controlled grazing regime that homogenises intensity in time and space, species‐rich Nardus grasslands can be conserved in Switzerland.     

Woody Plant Encroachment into Grasslands: Spatial Patterns of Functional Group Distribution and Community Development

Woody plant encroachment into grasslands has been globally widespread. The woody species invading grasslands represent a variety of contrasting plant functional groups and growth forms. Are some woody plant functional types (PFTs) better suited to invade grasslands than others? To what extent do local patterns of distribution and abundance of woody PFTs invading grasslands reflect intrinsic topoedaphic properties versus plant-induced changes in soil properties? We addressed these questions in the Southern Great Plains, United States at a subtropical grassland known to have been encroached upon by woody species over the past 50-100 years. A total of 20 woody species (9 tree-statured; 11 shrub-statured) were encountered along a transect extending from an upland into a playa basin. About half of the encroaching woody plants were potential N₂-fixers (55% of species), but they contributed only 7% to 16 % of the total basal area. Most species and the PFTs they represent were ubiquitously distributed along the topoedaphic gradient, but with varying abundances. Overstory-understory comparisons suggest that while future species composition of these woody communities is likely to change, PFT composition is not. Canonical correspondence analysis (CCA) ordination and variance partitioning (Partial CCA) indicated that woody species and PFT composition in developing woody communities was primarily influenced by intrinsic landscape location variables (e.g., soil texture) and secondarily by plant-induced changes in soil organic carbon and total nitrogen content. The ubiquitous distribution of species and PFTs suggests that woody plants are generally well-suited to a broad range of grassland topoedaphic settings. However, here we only examined categorical and non-quantitative functional traits. Although intrinsic soil properties exerted more control over the floristics of grassland-to-woodland succession did plant modifications of soil carbon and nitrogen concentrations, the latter are likely to influence productivity and nutrient cycling and may, over longer time-frames, feed back to influence PFT distributions.     

Reciprocal interactions between plants and soil in an upland grassland

Through the production of litter, plants with different life history strategies are predicted to both affect and be affected by the properties of soil. Competitive species are expected to increase the fertility of, and have a positive growth feedback with, soil, whereas stress-tolerant species should decrease fertility but show no growth feedback. We maintained monocultures of competitive (Lolium perenne and Agrostis capillaris) and stress-tolerant (Festuca ovina and Nardus stricta) grasses on an unproductive grassland for six years. The Nardus soil developed significantly greater inorganic nitrogen than the Agrostis and Festuca soil, and significantly greater soil moisture content than the Festuca soil. However, there were no differences in organic matter content, phosphate or bulk density between the soil types. In a greenhouse assay, each species was grown in soil cores from the different monocultures as well as natural turf. There were significant differences in growth between plant species and soil types. As expected, L. perenne produced the greatest amount of biomass. However, plants grown on Nardus soil were twice as large and had a 21% lower root allocation than plants grown on any of the other soil types. Lolium perenne, A. capillaris and F. ovina had significant negative growth feedbacks with their own soil (−0.460, −0.821 and −0.792, respectively) and N. stricta had a significant positive feedback (0.560). This study highlights the difficulties of predicting how plant traits will affect soil properties.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Resprouting after disturbance: an experimental study with short-lived monocarpic herbs

We experimentally demonstrated the ability of three short-lived monocarpic species to vegetatively regenerate (resprout) from roots after severe disturbance. We assessed the relationship between resprouting ability and (1) timing of injury with respect to life-cycle stage (reproductive vs. vegetative plant), life-history mode (annual vs. winter annual) and phenological stage (flowering vs. fruiting plant), (2) nutrient availability, and (3) disturbance severity (removal of all axillary buds Yes/No). In a chamber experiment with the annual or potentially winter annualRorippa palustris, all injured plants resprouted in all nutrient levels and day-length regimes (day-length regimes simulated conditions of an annual and a winter annual cohort). The number of adventitious buds on roots was positively affected only by injury. The extent of regeneration and amount of regenerated biomass were higher at high nutrient level and long-day regime.     

Effect of two AMF life strategies on the tripartite symbiosis with Bradyrhizobium japonicum and soybean

This study is the first in assessing the effect of soil disturbance on the contribution of arbuscular mycorrhizal fungi (AMF) with different life-history strategies to the tripartite symbiosis with soybeans and Bradyrhizobium japonicum (Kirchner) Jordan. We hypothesized that Gigaspora margarita Becker and Hall would be more affected by soil disturbance than Glomus clarum Nicol. and Schenck, and consequently, the tripartite symbiosis would develop more rapidly and lead to greater N₂ fixation in the presence of the latter. Soil pasteurization allowed the establishment of treatments with individual AMF species and soil disturbance enabled the development of contrasting root colonization potentials. In contrast, the colonization potential of B. japonicum was kept the same in all treatments. Soil disturbance significantly reduced root colonization by both AMF, with Gi. margarita being considerably more affected than G. clarum. Furthermore, the tripartite symbiosis progressed faster with G. clarum, and at 10 days after plant emergence, there was 30% more nodules when G. clarum was present compared to that when the bacterial symbiont alone was present. At flowering, the absence of soil disturbance stimulated N₂ fixation by 17% in mycorrhizal plants. However, this response was similar for both AMF.     

Effect of Manipulated Rainfall on Root Production and Plant Belowground Dry Mass of Different Grassland Ecosystems

A field experiment was established to quantify the effects of different amounts of rainfall on root growth and dry mass of belowground plant parts in three types of grassland ecosystems. Mountain (Nardus grassland), highland (wet Cirsium grassland), and lowland grassland (dry Festuca grassland) ecosystems were studied in 2006 and 2007. Roofs constructed above the canopy of grass stands and gravity irrigation systems simulated three climate scenarios: (1) rainfall reduced by 50%, (2) rainfall enhanced by 50%, and (3) the full natural rainfall of the current growing season. Experimentally reduced amounts of precipitation significantly affected both yearly root increments and total root dry mass in the highland grassland. Dry conditions in 2007 resulted in considerable reduction of total belowground dry mass in highland and mountain grasslands. Although not all differences in root biomass of studied grasslands were statistically significantly, some also showed a decrease in root increment and in the amount of belowground dry mass in dry conditions.     

Regeneration of different plant functional types in a Masson pine forest following pine wilt disease

Pine wilt disease is a severe threat to the native pine forests in East Asia. Understanding the natural regeneration of the forests disturbed by pine wilt disease is thus critical for the conservation of biodiversity in this realm. We studied the dynamics of composition and structure within different plant functional types (PFTs) in Masson pine forests affected by pine wilt disease (PWD). Based on plant traits, all species were assigned to four PFTs: evergreen woody species (PFT1), deciduous woody species (PFT2), herbs (PFT3), and ferns (PFT4). We analyzed the changes in these PFTs during the initial disturbance period and during post-disturbance regeneration. The species richness, abundance and basal area, as well as life-stage structure of the PFTs changed differently after pine wilt disease. The direction of plant community regeneration depended on the differential response of the PFTs. PFT1, which has a higher tolerance to disturbances, became dominant during the post-disturbance regeneration, and a young evergreen-broad-leaved forest developed quickly after PWD. Results also indicated that the impacts of PWD were dampened by the feedbacks between PFTs and the microclimate, in which PFT4 played an important ecological role. In conclusion, we propose management at the functional type level instead of at the population level as a promising approach in ecological restoration and biodiversity conservation.     

The Rengen Grassland Experiment: Effect of Soil Chemical Properties on Biomass Production, Plant Species Composition and Species Richness

The Rengen Grassland Experiment (RGE), set up on a Nardus grassland in 1941, consists of a control and five fertilizer treatments (Ca, CaN, CaNP, CaNP-KCl and CaNP-K₂SO₄). In 2005, soil chemical properties were analyzed to investigate the effect of soil variables on biomass production, plant species composition and species richness of vascular plants. Further, the effect of sampling scale (from 0.02 to 5.76 m²) on species richness was investigated. Soil properties (plant-available contents of K, P, C:N ratio, and pH) and biomass production were found to be strictly dependent on the fertilizers applied. Diversification of soil P content between treatments with and without P application is still in progress. Biomass production was most positively affected by P and K soil contents under N application. Furthermore, pH had a small positive effect on biomass production, and C:N ratio a moderately negative one. Two types of nutrient limitation were recognized: (1) limitation of total biomass production and (2) limitation of individual plant species. Long-term addition of a limiting nutrient affected the grassland ecosystem in three ways: (1) causing a change in plant species composition without significant increase in total biomass production, (2) causing no change in species composition but with significant increase in total biomass production, and (3) causing substantial change in plant species composition accompanied by significant increase in total biomass production. The explanatory power of all measured soil properties on plant species composition was almost the same as the power of the treatment effect (61.7% versus 62% of explained variability in RDA). The most powerful predictors of plant species composition were soil P, K and Mg contents, pH, and biomass production. The soil P content and biomass production were the only variables leading to a significant negative effect on species richness. An almost parallel increase in species richness with increasing sampling area was detected in all treatments. Constant differences among treatments were independent of sampling area.     

A STUDY OF TRIPLOID AND 3 X – 1 ANEUPLOID PLANTS IN THE GENUS GOSSYPIUM

Cytological comparisons were made of triploid and 3x – 1 plants of Gossypium hirsutum (haplo 17 and 18) X G. aridum, G. armourianum, G. harknessii, and G. raimondii. Tests and observations led to these conclusions: (1) Chromosome conjugation varied significantly from plant to plant and date to date within plants. (2) The D genome chromosomes of G. hirsutum are closer in homology to G. raimondii than to the other species tested. (3) The chromosomes of G. aridum, have closer homology to the A genome of G. hirsutum than do the chromosomes of the other D species tested. (4) The D genome of G. hirsutum has a small translocation as compared to the genomes of the four D species studied.     

Is analysing the nitrogen use at the plant canopy level a matter of choosing the right optimization criterion?

Herbivores can have strong deleterious effects on plant growth, reproduction, and even survival. Because these effects might be strongly interrelated, the direct consumptive effects of herbivores and a variety of indirect effects are difficult to untangle. Reductions in growth, for example, may strongly impact the flowering behaviour of plant species in the current season, but at the same time incur costs to survival, growth and reproduction in the next growing season(s). To get better insights in the effects of herbivory on the flowering behaviour of the long-lived polycarpic grassland herb Primula veris L., flowering patterns were monitored over ten consecutive years under two treatments (grazing and control mowing regimes). We tested the hypothesis that the size at flowering was affected by the presence of herbivores, and whether this translated into costs to future reproduction and survival. Overall, grazed plants were significantly smaller than control plants, and the size at which plants flowered was also significantly smaller when herbivores were present. The transition probability of flowering and of surviving into the next year was significantly smaller for all plants in the current year if they had been grazed than if they had been mown, indicating that herbivory incurred costs to both flowering and survival. Grazed plants also needed longer to start flowering, had fewer flowers and flowered less frequently, causing a significantly lower proportion of flowering adults in the population. These results suggest that the observed regression in plant size due to herbivory does not allow plants to capture enough resources to guarantee regular flowering in the longer run.     

Role of nurse plants in Araucaria Forest expansion over grassland in south Brazil

Abstract Araucaria Forest expansion over grassland takes place under wet climate conditions and low disturbance and it is hypothesized that isolated trees established on grassland facilitate the establishment of forest woody species beneath their canopies. Forest with Araucaria angustifolia is a particular type of Brazilian Atlantic Forest and the main forest type on the highland plateau in south Brazil, often forming mosaics with natural Campos grassland. The objectives of this paper were to evaluate the role of isolated shrubs and trees as colonization sites for seedlings of Araucaria Forest woody species on grassland, to determine which species function as preferential nurse plants in the process and the importance of vertebrate diaspore dispersal on the structure of seedling communities beneath nurse plants. The study was conducted in São Francisco de Paula, Rio Grande do Sul State, where we sampled isolated shrubs and trees in natural grassland near Araucaria Forest edges. Seedlings were counted and identified, and seedling diaspore dispersal syndromes, size and colour were registered. We detected 11 woody species with a potential role in nucleating grassland colonization by forest species. Beneath the canopies of nurse plants more forest species seedlings were found compared with open field grassland and the seedlings had diaspores mostly dispersed by vertebrates. Also, more seedlings were found under the canopy of A. angustifolia than beneath other nurse plant species. We conclude that A. angustifolia trees established on grassland act as nurse plants, by attracting disperser birds that promote colonization of the site by other forest species seedlings, and that under low level of grassland disturbance, conservation of frugivorous vertebrate assemblages may increase forest expansion over natural grassland and also facilitate the regeneration of degraded forest areas.     

Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries. Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related, perennial orchid species that differ in (a) floral traits important for pollination, including flowering phenology, floral display, and spur length, and (b) dominant pollinators. If plant–pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species. We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers, and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the expected direction, with selection for earlier flowering in two populations of the early‐flowering G. conopsea s.s. and for later flowering in one population of the late‐flowering G. densiflora. No divergent selection on floral morphology was detected, and there was no significant stabilizing selection on any trait in the two species. The results suggest ongoing adaptive differentiation of flowering phenology, strengthening this premating reproductive barrier between the two species. Synthesis: This study is among the first to test whether divergent selection on floral traits contribute to the maintenance of species differences between closely related plants. Phenological isolation confers a substantial potential for reproductive isolation, and divergent selection on flowering time can thus greatly influence reproductive isolation and adaptive differentiation.     

31种木本植物开花物候与系统发育的关系

植物物候通常被认为是由环境因素,如降水、温度和日照长度所决定,然而环境因素是否是物候唯一的决定因素仍然存在很大争议。谱系结构表征了植物在进化上的顺序,该发育时序是否对物候产生影响,当前仍然未知。在调查2016年春季新疆乌鲁木齐市最常见的31种木本植物的初始开花时间、败花时间和开花持续时间的基础上,通过分析植物开花物候的分布特征、开花物候在乔灌木间的差别、以及植物谱系距离与开花物候距离间的关系,试图揭示植物的开花物候和物种谱系(进化)顺序间的关系。结果表明:(1)新疆乌鲁木齐市31种木本植物的初始开花时间为4月18日±9d、败花时间为5月5日±12d、开花持续时间为(16±8)d;(2)乔木的初始开花时间和败花时间的标准差分别均低于灌木,乔木开花物候相对灌木更稳定;(3)乔木的初始开花和败花时间均显著早于灌木(P0.05),但开花持续时间在两者间未有显著性差异(P0.05);(3)31种木本植物间的初始开花时间距离、败花时间距离和开花持续时间距离均与物种谱系距离存在显著线性回归关系(P0.05)。综上可知:乔灌木在垂直空间上的分化使得木本植物的开花物候在植物生活型间存在不同。对植物的开花物候,除已被证明的降水、温度和日照长度等环境因素的影响外,物种进化顺序也可能造成了它在植物种间、时间和空间上的变异。     

Abundances of thrips on plants in vegetative and flowering stages are related to plant volatiles

Western flower thrips (WFTs), Frankliniella occidentalis Pergrande, and onion thrips (OTs), Thrips tabaci Lindeman, are two cosmopolitan insect pests of agricultural and horticultural plants. Understanding the occurrence and development of thrips on plants is crucial for identifying suitable plants that can be used for developing a “push-pull” strategy against thrips. In this study, the dynamics of WFTs and OTs on plants (Allium fistulosum L., Medicago sativa L., Luffa cylindrica (L.) Roem., Ocimum basilicum L., and Schizonepeta tenuifolia (Benth.) Briq.) were investigated for two consecutive years (2018–2019). Throughout the survey, the abundances of both thrips species were strongly associated with plant species and plant phenology; both thrips species were present at relatively high densities on M. sativa but very low densities on O. basilicum and S. tenuifolia. Populations of both thrips species greatly increased during plant flowering. A Y-tube olfactory test was used to study the effects of plant volatiles in mediating thrips behaviour and showed that volatiles of M. sativa were attractive to both thrips species whether emitted by the plant in the vegetative or flowering stage, while volatiles of O. basilicum and S. tenuifolia were repellent to thrips. Additionally, because of the presence of a high number of floral chemicals, both thrips species exhibited a greater preference for volatiles emitted by plants in the flowering period over those emitted by plants in the vegetative period. Our observations indicate that plant species and flowering status play an important role in the abundance dynamics of thrips and that the volatiles of flowering plants attract thrips more strongly than volatiles emitted by vegetative plants. These findings can facilitate the screening of attractive/unattractive plants for developing push-pull strategies to control thrips.     

Assessing the potential of candidate DNA barcodes for identifying non‐flowering seed plants

In plants, matK and rbcL have been selected as core barcodes by the Consortium for the Barcode of Life (CBOL) Plant Working Group (PWG), and ITS/ITS2 and psbA‐trnH were suggested as supplementary loci. Yet, research on DNA barcoding of non‐flowering seed plants has been less extensive, and the evaluation of DNA barcodes in this division has been limited thus far. Here, we evaluated seven markers (psbA‐trnH, matK, rbcL, rpoB, rpoC1, ITS and ITS2) from non‐flowering seed plants. The usefulness of each region was assessed using four criteria: the success rate of PCR amplification, the differential intra‐ and inter‐specific divergences, the DNA barcoding gap and the ability to discriminate species. Among the seven loci tested, ITS2 produced the best results in the barcoding of non‐flowering seed plants. In addition, we compared the abilities of the five most‐recommended markers (psbA‐trnH, matK, rbcL, ITS and ITS2) to identify additional species using a large database of gymnosperms from GenBank. ITS2 remained effective for species identification in a wide range of non‐flowering seed plants: for the 1531 samples from 608 species of 80 diverse genera, ITS2 correctly authenticated 66% of them at the species level. In conclusion, the ITS2 region can serve as a useful barcode to discriminate non‐flowering seed plants, and this study will contribute valuable information for the barcoding of plant species.     

Invasion of temperate grassland by a subtropical annual grass across an experimental matrix of water stress and disturbance

Abstract. An experimental matrix of water stress and disturbance was superimposed on a Lolium perenne-Trifolium repens grassland using a crossed-gradient design, and the annual subtropical grass Digitaria sanguinalis was introduced into the pasture as seeds and transplanted seedlings. Digitaria plants achieved maximum biomass at high water availability and high disturbance. Digitaria plants grown from transplanted seedlings attained greater biomass further from the conditions of high water availability and high disturbance, compared with those that had grown from seed. The biomass of the temperate species was maximized with high water availability and low to intermediate disturbance conditions. The reproductive effort of Digitaria was maximized under intermediate to high water availability and intermediate to low disturbance. Combinations of water stress and disturbance that gave rise to maximum growth of the temperate and subtropical species were consistent with those predicted by C-S-R theory. Results suggest that processes in the regenerative phase of the plant life cycle were constraining the success of Digitaria in New Zealand grassland. Addition of seed to the soil seed bank would probably be maximized in patches of grassland with high disturbance and water availability; these patches will subsequently act as foci for future invasions by Digitaria.