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     

Long-term patterns of shrub expansion in a C4-dominated grassland: fire frequency and the dynamics of shrub cover and abundance

Worldwide, grassland ecosystems have experienced a major shift in growth-form dominance as woody plant species have expanded and replaced native grasses. In the C(4)-dominated grasslands of central North America, a reduction in fire frequency is the most cited cause of this shift in growth forms as fire both enhances grass productivity and constrains the establishment and expansion of native woody vegetation. Using an 18-yr plant species composition data set, we quantified patterns of change in shrub cover, frequency, and species richness associated with three distinct fire regimes. During the study period (1983-2000), shrub cover increased most dramatically in sites in which the frequency of fire was once every 4 yr (intermediate frequency; 28.6%) followed by sites in which fire occurred only once during the 18-yr period (low frequency; 23.7%). Annual fire effectively prevented the recruitment of new woody species, but even with this high fire frequency, shrub cover increased slightly (3.7%). Comparatively, shrub species richness increased by three and six, respectively, in the intermediate- and low-frequency fire sites. These data indicate that within this grassland, periods without fire are necessary for recruitment of both new individuals and additional shrub species; however, once established, shrub cover will increase regardless of fire frequency and even annual fire will not reduce shrub abundance.     

Dominance not richness determines invasibility of tallgrass prairie

Many recent studies suggest that more diverse communities are more resistant to invasion. Community characteristics that most strongly influence invasion are uncertain, however, due to covariation of diversity with competition and crowding. We examined separately the effects of species richness and dominance on invasion by an exotic legume, Melilotus officinalis , in intact, native Kansas grassland. We manipulated dominance of C₄ grasses by reducing their abundance (i.e. ramet densities) by ∼25 and 50%. In addition, richness was reduced by removing species that were mainly rare and uncommon as might be expected with environmental changes such as drought and fragmentation. In both years of the study (2001–2002), invasibility, measured as peak establishment of Melilotus , was not affected by a 3-fold reduction in species richness, nor was there an interaction between loss of species and reduced dominance on invasion. In contrast, reductions in abundance of the dominants significantly reduced invasibility of the grassland plots in both years. Because the abundance of dominants was highly correlated with measures of competition (i.e. ratio of dominant biomass to total biomass) and crowding (total stem densities), this pattern was opposite to that expected if competition were indeed limiting invasion. Rather, invasion appeared to be facilitated by the dominant species, most likely because reduced dominance increased environmental stress. Our results suggest that dominance is the key community characteristic determining invasibility, because highly competitive and space-filling species can either enhance or reduce susceptibility to invasion depending on whether dominants create a more competitive environment or alleviate stressful conditions.     

Shrub succession and invasibility in a New Zealand montane grassland

Abstract Two successive shrub invasions of a short tussock grassland induced by grazing and burning were examined in montane South Island, New Zealand. The first invasion was by a native shrub, matagouri (Discaria toumatou Raoul). The second invasion was by an exotic shrub, Scotch broom (Cytisus scoparius (L.) Link), which invaded the matagouri shrubland that had developed over the grassland. The invasions were investigated using analysis of spatial patterns of both shrubs and tussocks, and age, growth rates and size structure of the shrubs. Competition between the two shrub species was examined using spatial patterns and comparing allometric relationships. After initial invasion by matagouri of the grasslands, stand density increased by consolidation about its initial colonization points. Current matagouri distribution is often negatively associated with tussocks. Scotch broom occurs most frequently in a dense sward of introduced grasses and occasionally in tussocks in interstices among matagouri shrubs. Despite the palatability of Scotch broom to sheep that graze the site, there was no evidence that the spiny matagouri facilitates invasion by protecting Scotch broom seedlings; rather there was negative association between the shrub species. The two species probably compete for above-ground space. However, diameter and height growth rates of Scotch broom far exceed those of matagouri so Scotch broom is likely to increase in biomass rapidly at the site. The autogenic organization and disturbance history of the resident plant communities have rendered each vulnerable to successive invasions.     

A phylogenetic approach to community assembly from a local species pool

Ecological theory provides two contrasting predictions about the characteristics of the species combining to form communities. Classical competition theory states that they will be less similar than expected by chance, whilst the environmental structuring hypothesis states that they will be more similar. We investigated these predictions by applying phylogenetic methods of analysis (PICs) to a grassland community, examining species on the basis of their traits. At the scale of investigation most useful in making predictions about the presence and abundance of species (the community level, the species forming the community were more similar than would be expected by chance. The use of PICs resulted in a more sensitive test than if phylogeny had been ignored, allowing the detection of effects that would otherwise have been overlooked or underestimated. Selected traits from the PICs analysis were used to develop a predictive model of community membership using discriminant analysis. This correctly identified species in the pool which were present in the community but failed to predict absences accurately, implying that dispersal limitation may operate in the community.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Overyielding in experimental grassland communities – irrespective of species pool or spatial scale

In a large integrated biodiversity project (‘The Jena Experiment’ in Germany) we established two experiments, one with a pool of 60 plant species that ranged broadly from dominant to subordinate competitors on large 20 × 20 m and small 3.5 × 3.5 m plots (= main experiment), and one with a pool of nine potentially dominant species on small 3.5 × 3.5 m plots (= dominance experiment). We found identical positive species richness–aboveground productivity relationships in the main experiment at both scales. This result suggests that scaling up, at least over the short term, is appropriate in interpreting the implications of such experiments for larger‐scale patterns. The species richness–productivity relationship was more pronounced in the experiment with dominant species (46.7 and 82.6% yield increase compared to mean monoculture, respectively). Additionally, transgressive overyielding occurred more frequently in the dominance experiment (67.7% of cases) than in the main experiment (23.4% of cases). Additive partitioning and relative yield total analyses showed that both complementarity and selection effects contributed to the positive net biodiversity effect.     

The role of plant species size in invasibility: a field experiment

Large plant species self-thin to disproportionately lower densities than smaller plant species, and therefore may leave more patches of unused space suitable for invasion. Using experimental monocultures of 11 old-field perennial plant species differing in maximum size, as well as mixtures composed of all monoculture species, we tested our primary hypothesis that monocultures of larger species will be more susceptible to natural invasion. After 3 years, monocultures of larger species were invaded by a significantly greater number of species, and more ramets, from the surrounding vegetation. Invading plant species were significantly smaller than the monoculture species being invaded, suggesting that smaller plant species may be better invaders. Thus, we quantified a trade-off between species size, which is frequently associated with increased competitive ability for light, and invasibility, suggesting one reason why large and small species coexist in virtually all plant communities. Although we expected that invasion would enhance biomass production by more fully capturing available resources, we found that the most highly invaded plots of each species produced significantly less biomass. This suggests that increased diversity resulting from invasion did not result in complementary resource use. Mixture plots containing all experimental species did not admit a significantly different number of invading ramets or species than most monocultures, indicating no obvious role for diversity in resistance to invasion, or complementary resource use. Our results suggest that relatively large species may be limited in their capacity to competitively exclude other, smaller species from communities because pure stands of the former are more susceptible to invasion by the latter.     

Native and alien species diversity at the local and regional scales in a grazed California grassland

Serpentine meadows in Northern California supported higher species richness at the 1-m² scale than adjacent nonserpentine meadows, and had a considerably higher proportion of native species. Within each soil type, total species richness (natives plus aliens) was unrelated to biomass, cover, soil depth, or soil characteristics (N, P, Ca⁺⁺, Mg⁺⁺, water-holding capacity). However, the proportion of native species on serpentine was higher in meadows with lower levels of phosphorus and a lower calcium/magnesium ratio; the proportion of native species in nonserpentine meadows was higher on cool (north to northeast facing) slopes. At a regional scale, some of these effects were partly reversed; the rate at which new species accumulated with the addition of new sites, or beta diversity, was highest for native plant species in nonserpentine meadows. All of the above effects were independent of whether grazing by cattle was absent (removed 13 years ago) or present. The status of low-productivity serpentine soils as a refuge for native grassland species appears to be the result of their abiotic resistance to alien species, but not of a negative relationship between productivity and total species richness. Received: 10 April 1999 / Accepted: 4 June 1999     

Dominant species identity regulates invasibility of old-field plant communities

Dominant species are known to exert strong influence over community dynamics, although little work has addressed how they affect invasibility. In this study, we examined whether dominant species identity and abundance affected invasibility of old-field plant communities. To quantify invasibility, we added seeds of 19 plant species into plots dominated by one of four different herbaceous perennial species ( Andropogon virginicus , Bromus inermis , Centaurea maculosa , or Solidago canadensis ) . We found that, independent of species richness and abiotic variables, plots dominated by Andropogon were the least invasible, while Bromus and Centaurea plots had the highest invasibility. We examined several potential mechanisms by which these dominant species might influence invasibility, and found invasion to increase with decreasing litter biomass and increasing community species richness. The abundance of the dominant species was not a significant predictor of invasion. These results indicate that dominant species identity plays an important role in determining invasibility of plant communities, though exact mechanisms underlying these effects still need to be explored.     

Determinants of C3 forb growth and production in a C4 dominated grassland

Forbs are the most abundant species within the vascular flora of tallgrass prairie and they make the greatest contribution to biodiversity of all growth forms. However, little is known about the factors that determine their productivity and growth rates. The objective of this study was to assess the controls of forb growth (absolute and relative) and production in tallgrass prairie from a long-term burning experiment at the Konza Prairie in NE Kansas. Over the 14-year study, forbs comprised 9% of the total biomass production on sites with a high fire frequency vs. 29% on the low fire frequency site, with gramminoids accounting for the remainder. Although interannual variations in peak biomass of the grasses was strongly correlated with environmental variables related to water availability, there were no similar relationships for forbs, suggesting that production of forbs and grasses responded to interannual variations in climate in different ways. Multivariate analysis of climatic controls on growth rates of grasses and forbs yielded similar results. Although forbs had low biomass and absolute growth per unit ground area in frequently burned prairie, their relative growth rates were highest in such sites. Thus, it appears that reduced growth rates of individual forbs per se do not limit forb success in annually burned prairie. Instead, direct negative effects of fire on forbs (increased mortality) may be responsible. Determinants of forb growth and productivity in unburned prairie remain unresolved.     

Impacts of C4 grass introductions on soil carbon and nitrogen cycling in C3-dominated successional systems

While recent research has focused on the effects of exotic plant species on ecosystem properties, less is known about how restoring individual native plant species, differing in biomass and tissue chemistry, may impact ecosystems. We examined how three native C(4) prairie grasses affected soil C and N cycling 11 years after reintroduction into successional old-field communities dominated by non-native C(3) grasses. The species examined in this study differ in traits that are expected to influence soil C and N cycling (biomass and tissue chemistry). Thus, we hypothesized that cycling rates would decrease, thereby increasing pool sizes in soils under C(4) species compared under C(3) species. As predicted, the C(4) species had greater biomass and more recalcitrant tissue [higher C:N, acid detergent fiber (ADF):N] compared to the dominant C(3) species. The three C(4) species did not differ in tissue C:N, ADF:N, or root biomass, but Andropogon had more than twice the shoot biomass of Schizachyrium and Sorghastrum. Soils under the C(4) species did not differ in inorganic N levels, but levels were lower than in soils under the C(3) species, and soils under Andropogon had slightly lower in situ net N mineralization rates compared to those under C(3) species. We found little evidence of larger surface soil C pools under C(4) species versus C(3) species after 11 years and no differences in subsurface soil C or N among species. The C(4) species contributed a significant amount of C to both soil depths after 11 years. Our results demonstrate that C(4) species reintroduction into old-fields can alter C and N cycling on relatively short timescales, and that individual C(4) species differ in the magnitude of these effects. Improving our understanding of how species influence ecosystem properties is essential to predicting the ecosystem-level consequences of plant community alterations due to land use changes, global change, and species introductions.     

Associated growth of C3 and C4 desert plants helps the C3 species at the cost of the C4 species

C₃ desert plant Reaumuria soongorica (RS-C₃) and C₄ desert plant Salsola passerina (SP-C₄) may exist either in individual or in associated communities. Carbon isotope composition, leaf water potential, gas exchange and chlorophyll fluorescence characteristics of the individual and associated communities were compared with reveal, whether the associated growth represent an advantage under harsh habitat. The results showed that the ??¹³C values of leaves of RS-C₃ and SP-C₄ across different habitats fluctuated, respectively, from ?24 to ?27??? and from ?14 to ?16???. Leaf water potential of RS-C₃ was lower than SP-C₄ all day long, growing either individually or associated with the C₃ plant. When associated with the C₄ plant, the net photosynthetic rate of the RS-C₃ increased, and the photosynthetic rate of the partner SP-C₄ decreased. The transpiration rates of the associated RS-C₃ and SP-C₄ were both lower than in their individual colonies. In associated communities, in RS-C₃, the maximal photochemical efficiency, the effective photochemical efficiency, the relative electron transport rate, the photochemical quenching of PS II increased, and the non-photochemical quenching of PS II decreased; all these parameters changed oppositely in the SP-C₄ plant. This shows that, in the associated community, the C₄ plants might facilitate adaptation of the RS-C₃, while SP-C₄ plant can adapt to the harsh environment through their own specialties. The association favored the expression of natural photosynthetic characteristics and survival of RS-C₃, while retarded the growth of SP-C₄. Associated growth decreases the transpiration rate of the whole community; it is conducive to improve its water use efficiency.     

Introducing the biogeographic species pool

The species pool concept has played a central role in the development of ecological theory for at least 60 yr. Surprisingly, there is little consensus as to how one should define the species pool, and consequently, no systematic approach exists. Because the definition of the species pool is essential to infer the processes that shape ecological communities, there is a strong incentive to develop an ecologically realistic definition of the species pool based on repeatable and transparent analytical approaches. Recently, several methodological tools have become available to summarize repeated patterns in the geographic distribution of species, phylogenetic clades and taxonomically broad lineages. Here, we present three analytical approaches that can be used to define what we term ‘the biogeographic species pool’: distance‐based clustering analysis, network modularity analysis, and assemblage dispersion fields. The biogeographic species pool defines the pool of potential community members in a broad sense and represents a first step towards a standardized definition of the species pool for the purpose of comparative ecological, evolutionary and biogeographic studies.     

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.