The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

Photosynthetic responses of native and introduced C4 grasses from Venezuelan savannas

Summary Introduced African grasses are invading the grasslands of the Venezuelan savannas and displacing the native grasses. This work, which is part of a program to understand the reasons for the success of the African grasses, specifically investigates whether introduced and native grasses differ in some photosynthetic characteristics.The responses to photon flux density, leaf temperature, leaf-air vapour pressure difference and leaf water potential of leaf photosynthetic rate of two introduced African C₄ grasses (Hyparrhenia rufa and Melinis minutiflora) and of a lowland and a highland population of a native Venezuelan grass (Trachypogon plumosus) grown under controlled conditions were compared. These responses in all three species were typical of tropical C₄ pasture grasses. The introduced grasses had higher maximum leaf conductance, net photosynthetic rates, and optimum temperature (H. rufa only) for photosynthesis than T. plumosus. However, T. plumosus was able to continue photosynthesis to lower leaf water potentials than the two introduced grasses, and the efficiency which it utilized water, light and mineral nutrients to fix carbon were similar to those of the introduced grasses.The higher rates of leaf photosynthesis of the introduced grasses contributed to, but only partially explained, the higher growth rates compared to T. plumosus. The higher growth rates and nutrient concentration of the introduced grasses are consistent with their ability to establish rapidly, compete successfully for resources, and displace T. plumosus from moist, fertile sites. Conversely, the slower growth rate, lower nutrient concentrations, and superior water relations characteristics are consistent with the capacity of T. plumosus to resist invasion by introduced grasses in poorer sites.     

Bud banks of perennial savanna grasses in Botswana

Three semi‐arid savanna grasses in Botswana (Stipagrostis uniplumis, Eragrostis lehmanniana, and Aristida stipitata) were sampled to quantify their belowground bud banks during the dormant season and to estimate their relative allocation to vegetative and sexual reproduction. Bud banks of these African perennial caespitose grasses were also compared with four perennial caespitose grasses of semi‐arid North American grasslands. The three African grasses each maintained approximately two buds per tiller and showed a high percentage (88–99%) of tillers producing seed. Only E. lehmanniana produced new aerial tillers from axillary buds at elevated nodes on the stem as well as from the belowground bud bank. Compared with species of North American grasslands, these African grasses produced fewer belowground buds but showed a much higher percentage of tillers producing seed. These patterns indicate relatively greater belowground meristem limitation, lower allocation to vegetative reproduction (tillering) and higher allocation to seed reproduction in these African grasses, although studies of more species are needed to assess the generality of this pattern. The management of savannas in ways that favour the maintenance of a reserve population of belowground buds may increase the ability of grasses to respond to pulses of resource availability, increase their compensatory growth capacity following grazing or drought, and decrease the invasibility of these plant communities by exotic species, whereas maintaining allocation to sexual reproduction may be important for conserving genetic variation and enhancing their capacity to adapt to environmental change.     

Responses of tropical native and invader C<Subscript>4</Subscript> grasses to water stress,clipping and increased atmospheric CO<Subscript>2</Subscript> concentration

The invasion of African grasses into Neotropical savannas has altered savanna composition, structure and function. The projected increase in atmospheric CO₂ concentration has the potential to further alter the competitive relationship between native and invader grasses. The objective of this study was to quantify the responses of two populations of a widespread native C₄ grass (Trachypogon plumosus) and two African C₄ grass invaders (Hyparrhenia rufa and Melinis minutiflora) to high CO₂ concentration interacting with two primary savanna stressors: drought and herbivory. Elevated CO₂ increased the competitive potential of invader grasses in several ways. Germination and seedling size was promoted in introduced grasses. Under high CO₂, the relative growth rate of young introduced grasses was twice that of native grass (0.58 g g⁻¹ week⁻¹ vs 0.25 g g⁻¹ week⁻¹). This initial growth advantage was maintained throughout the course of the study. Well-watered and unstressed African grasses also responded more to high CO₂ than did the native grass (biomass increases of 21–47% compared with decreases of 13–51%). Observed higher water and nitrogen use efficiency of invader grasses may aid their establishment and competitive strength in unfertile sites, specially if the climate becomes drier. In addition, high CO₂ promoted lower leaf N content more in the invader grasses. The more intensive land use, predicted to occur in this region, may interact with high CO₂ to fincreasesavor the African grasses, as they generally recovered faster after simulated herbivory. The superiority of invader grasses under high CO₂ suggests further in their competitive strength and a potential increased rate of displacement of the native savannas in the future by grasslands dominated by introduced African species.     

Abundance and Distribution of Native and Alien Grasses in a “Cerrado” (Brazilian Savanna) Biological Reserve1

Several species of African grasses brought to Brazil as cattle forage have spread widely, outcompeting native herb species. The open forms of Brazilian savanna (“campo cerrado” and “campo sujo”) are the most affected by such invasions, because their structure is open, permitting enough sunlight into the lower strata. The invasion of alien forage grasses occurs in almost every cerrado nature reserve. This study was carried out in the “Cerrado de Hmas Biological Reserve”, Pirassununga, São Paulo State, Brazil, with the following objectives: (a) to compare the abundance of native and alien grass species; (b) to verify the importance of such alien grasses in the community; (c) to identity distribution patterns for the alien grass species in a gradient from the edge (highly disturbed) to the center (less disturbed) of the reserve; and (d) to explore the distribution of native and alien grasses in the search for possible competitive interactions. Using the “point method,” a total of 260 points was sampled and 52 species were recorded. The four most frequent species (FA = absolute frequency) were two native (Echinolaena inflexa [Poir.J Chase [FA = 38.85%] and Diandrostachya chrysotrix [Nees] Jacues Felix [FA = 15.38%]) and two alien African species (Melinis minutiflora Beauv. [FA = 33.08%] and Brachiaria decumbens Stapf [FA = 13.85%]). M. minutiflora and E. inflexa had higher values of absolute vigor (67.69 and 59.62%, respectively), relative vigor (28.16 and 24.80%, respectively), and cover (100.77 and 98.47, respectively), indicating higher biomasses and densities and their dominance in the community. B. decumbens presented the highest number of contacts per point, showing the highest stratification. To detect possible edge–center distribution gradients, correspondence analysis was done, initially using all the recorded species and subsequently only the four more frequent grasses, with similar results: (a) the alien grasses, especially M. minutiflora, did not show a distinct distribution gradient from edge to center, but occurred over the whole reserve; (b) no distinct ecotonal band around the reserve (edge–belt) was detected, the whole reserve seeming to be “ecotonal”; and (c) E. inflexa and M. minutiflora showed similar phytosociological patterns, and spatial distribution; association between these two species was statistically significant.     

Water relations of native and introduced C4 grasses in a neotropical savanna

Introduced African grasses are invading Neotropical savannas and displacing the native herbaceous community. This work, which is part of a program to understand the success of the African grasses, specifically investigates whether introduced and native grasses differ in their water relations. The water relations of the native Trachypogon plumosus and the successful invader Hyparrhenia rufa were studied in the field during two consecutive years in the seasonal savannas of Venezuela. The two C₄ grasses differed clearly in their responses to water stress. H. rufa consistently had higher stomatal conductance, transpiration rate, leaf water and osmotic potential and osmotic adjustment than the native T. plumosus. Also, leaf senescence occurred much earlier during the dry season in H. rufa. Both grasses showed a combination of water stress evasion and tolerance mechanisms such as stomatal sensitivity to atmospheric or soil water stress, decreased transpiring area and osmotic adjustment. Evasion mechanisms are more conspicuous in H. rufa whereas T. plumosus is more drought tolerant and uses water more conservatively. The evasion mechanisms and oportunistic use of water by H. rufa, characteristic of invading species, contribute to, but only partially explain, the success of this grass in the Neotropical savannas where it displaces native plants from sites with better water and nutrient status. Conversely, the higher water stress tolerance of t. plumosus is consistent with its capacity to resist invasion by alien grasses on shallow soils and sites with poorer nutrient and water status.     

Variation in the impact of exotic grasses on native plant composition in relation to fire across an elevation gradient in Hawaii

The impact that an exotic species can have on the composition of the community it enters is a function of its abundance, its particular species traits and characteristics of the recipient community. In this study we examined species composition in 14 sites burned in fires fuelled by non‐indigenous C4 grasses in Hawaii Volcanoes National Park, Hawaii. We considered fire intensity, time since fire, climatic zone of site, unburned grass cover, unburned native cover and identity of the most abundant exotic grass in the adjacent unburned site as potential predictor variables of the impact of fire upon native species. We found that climatic zone was the single best variable for explaining variation in native cover among burned sites and between burned and unburned pairs. Fire in the eastern coastal lowlands had a very small effect on native plant cover and often stimulated native species regeneration, whereas fire in the seasonal submontane zone consistently caused a decline in native species cover and almost no species were fire tolerant. The dominant shrub, Styphelia tameiameia, in particular was fire intolerant. The number of years since fire, fire intensity and native cover in reference sites were not significantly correlated with native species cover in burned sites. The particular species of grass that carried the fire did however, have a significant effect on native species recovery. Where the African grass Melinis minutiflora was a dominant or codominant species, fire impacts were more severe than where it was absent regardless of climate zone. Overall, the impacts of exotic grass‐fuelled fires on native species composition and cover in seasonally dry Hawaiian ecosystems was context specific. This specificity is best explained by differences between the climatic zones in which fire occurred. Elevation was the main physical variable that differed among the climatic zones and it alone could explain a large percentage of the variation in native cover among sites. Rainfall, by contrast, did not vary systematically with elevation. Elevation is associated with differences in composition of the native species assemblages. In the coastal lowlands, the native grass Heteropogon contortus, was largely responsible for positive changes in native cover after fire although other native species also increased. Like the exotic grasses, this species is a perennial C4 grass. It is lacking in the submontane zone and there are no comparable native species there and almost all native species in the submontane zone were reduced by fire. The lack of fire tolerant species in the submontane zone thus clearly contributes to the devastating impact of fire upon native cover there.     

Dynamics of energy and nutrient concentration and construction cost in a native and two alien C4 grasses from two neotropical savannas

In Venezuela, the alien grasses Melinis minutiflora Beauv. and Hyparrhenia rufa (Nees.) Stapf tend to displace the native savanna plant community dominated by Trachypogon plumosus (Humb. and Bonpl.) Nees. This occurs in either relatively wetter and fertile highland savannas or in drier and less fertile lowland savannas. Although the native and aliens are perennial C₄ grasses, higher net assimilation leaf biomass per plant and germination rate of the latter are some causes for their higher growth rates and for their competitive success. The objective of this study is to compare seasonal tissue energy, N, P and K concentrations and the calculated construction costs (CC) between the native grass and either one of the alien grasses from lowland and highland savannas. We predict that, in order to out-compete native plants, alien grasses should be more efficient in resource use as evidenced by lower tissue energy and nutrient concentrations and CC.Tissue energy and nutrient concentration were measured throughout the year and compared between M. minutiflora and the co-occurring local population of T. plumosus in a highland savanna and between H. rufa and its neighbor local population of T. plumosus in a lowland savanna. CC was calculated from energy, N and ash concentrations considering ammonium as the sole N source. Differences between co-occurring species, T. plumosus populations, seasons, and organs were analyzed with ANOVA.Highland and lowland grasses differed in concentration and allocation of energy and nutrients whereas the differences between alien and native grasses were specific for each pair considered. Highland grasses had higher energy, N, P and CC than lowland grasses. These variables were always lowest in the culms. In the more stressed lowland site, tissue energy and nutrient concentrations decreased significantly during the dry season except in the roots of both grasses which had the highest energy and nutrients concentrations during the drought. This seasonal response was more marked in the local lowland population of T. plumosus in which maximum CC alternated seasonally between leaves and roots. Energy and nutrient concentrations and CC were the lowest in H. rufa. In the lowland savannas, the higher efficiency of resource use in the invader grass contributes to its higher competitive success through increased growth rate. In the highlands, overall tissue energy concentration and CC, but not N nor P concentration, were lower in the fast growing M. minutiflora but seasonal differences were lacking. The higher leaf CC in T. plumosus can be attributed to the higher proportion of sclerenchyma tissue which is more expensive to construct. Considering CC, both fast growing alien grasses are more efficient in resource use than the co-occurring native grass. However, the role of CC explaining the competitive success of the former, through higher growth rates, is more evident in the more stressful environment of the lowland savanna.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

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     

Impact of the invasive alien grass Melinis minutiflora at the savanna-forest ecotone in the Brazilian Cerrado

Exotic grasses are a serious threat to biodiversity in the cerrado savannas of central Brazil. Of particular concern is the possible role they may have in impeding tree regeneration at gallery (riverine) forest edges and increasing fire intensity, thereby driving gallery forest retreat. Here we quantify the effect of roads and distance from gallery forests on the abundance of the African grass Melinis minutiflora Beauv. and test for an effect of this species on woody plant regeneration and leaf area index. Melinis was present at approximately 70% of the sites near gallery forest edges, with its frequency declining sharply at greater distances from the edge. Melinis frequency was 2.8 times greater where roads were present nearby. Leaf area index (LAI) of the ground layer was 38% higher where Melinis was present than where it was absent. LAI was strongly correlated to fine fuel mass (r² = 0.80), indicating higher fuel loads where Melinis was present. The abundance of tree and shrub species in the ground layer was negatively related to LAI and to the presence of Melinis. The greater fuel accumulation and reduced tree regeneration caused by Melinis may cause a net reduction in forest area by increasing fire intensity at the gallery forest edge and slowing the rate of forest expansion.     

Can Invasive Species Enhance Competitive Ability and Restoration Potential in Native Grass Populations?

Native plant individuals often persist within communities dominated by exotics but the influence of this exposure on native populations is poorly understood. Selection for traits contributing to competitive ability may lead to native plant populations that are more tolerant of the presence of exotic invaders. In this way, long‐term coexistence with an exotic may confer competitive advantages to remnant (experienced) native populations and be potentially beneficial to restoration. In past studies we have documented genetic differentiation within native grass populations exposed to the exotic invader Russian knapweed (Acroptilon repens). Here, we examine populations of a cool‐season grass, needle‐and‐thread (Hesperostipa comata [Trin. & Rupr.]) and a warm season, alkali sacaton (Sporobolus airoides [Torr.]) collected from Russian knapweed‐invaded sites and adjacent noninvaded sites to assess their relative competitive ability against a novel exotic neighbor, Canada thistle (Cirsium arvense). Experienced S. airoides (from within A. repens invasions) appear to better tolerate (accumulate biomass, leaf nitrogen content, and to initiate new tillers) the presence of a novel competitor (C. arvense). Experienced and inexperienced H. comata genets differ in their response to the presence of C. arvense. Relative neighbor effects of native grasses on C. arvense were generally greater from experienced grasses. The ability to compete with novel neighbors may be driven by general competitive traits rather than species‐specific coevolutionary trajectories. Irrespective of competitive mechanisms, the conservation of native species populations within weed invasions may provide an important restoration tool by retaining unique components of native gene pools selected by competitive interactions with exotics.     

Experimental Restoration of Native Vegetation in Indiana Dunes National Lakeshore

We investigated the effects of prescribed fire, herbicide treatment, and sod removal on the eradication of exotic grasses and the establishment of native plant species in 24 experimental restoration plots in three razed residential sites within the boundary of Indiana Dunes National Lakeshore. During 1992–1995, herbicide treatment and sod removal decreased the combined cover of Poa pratensis (Kentucky blue grass) and Agropyron repens (quackgrass) significantly (from 82% to 13%, and 85% to 8%, respectively), whereas fire did not suppress such exotic lawn grasses. In 1993, several opportunistic species, represented by Cyperus spp. (umbrella sedges), Digitaria sanguinalis (crab grass), and Ambrosia artemisiifolia (common ragweed), filled the gaps left by the removal of lawn grasses. For the same period, Detrended Correspondence Analysis revealed a clear vegetation divergence between the control-fire plots and the herbicide-sod removal plots. While Poa pratensis and Agropyron repens continued to dominate the control and fire plots, the planted native species, represented by Schizachyrium scoparium (little blue-stem), Sorghastrum nutans (Indian grass), Rudbeckia hirta (black-eyed Susan), and Monarda punctata (horsemint), began to dominate in the herbicide and sod removal plots from 1994. In both herbicide and sod removal plots, the ground cover of grasses (68%) was much higher than the forbs (10%). The herbicide plots, where exotic species were removed but nitrogen-rich top soils were not removed, showed a higher diversity of planted native species than the sod removal plots (where both exotic species and top soils were removed) and the control-fire plots (where neither was removed). This finding suggests that an optimum but not excessive concentration of soil nitrogen is needed to support a maximum species diversity in such infertile substrate as sandy soil. In addition, the decrease in potassium in all plots, regardless of treatment, suggests that potassium may become a limiting factor for our restored native vegetation.     

Cattle and Weedy Shrubs as Restoration Tools of Tropical Montane Rainforest

Over the last 150 years, a large proportion of forests in Latin America have been converted to pastures. When these pastures are abandoned, grasses may slow re‐establishment of woody species and limit forest regeneration. In this study, we explored the use of cattle in facilitating the establishment of woody vegetation in Colombian montane pastures, dominated by the African grasses Pennisetum clandestinum (Kikuyo) and Melinis minutiflora (Yaraguá). First, we described woody and herbaceous vegetation in grazed and non‐grazed pastures. Second, we tested the effect of grazing and seed addition on the establishment and growth of woody species. We also determined if the effect of grazing was different in P. clandestinum and M. minutiflora pastures. We found that low stocking density of cattle greatly increased density, number of branches per individual (a measure of “shrubiness”), and basal area of woody species, but also reduced woody plant species richness and diversity. In the grazed area, the shrubs Baccharis latifolia (Chilca) and Salvia sp. (Salvia) were the most abundant. The combined effect of grazing and shading from the shrubs reduced herbaceous vegetation by 52 to 92%. In the grazing/seed addition experiment, grazing increased establishment of woody seedlings, particularly of the shrub Verbesina arborea (camargo), but the largest effect was seed addition. Where grasses are an important barrier to regeneration, grazing can facilitate the establishment of shrubs that create a microhabitat more suitable for the establishment of montane forest tree species.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Effectiveness of repeated autumn and spring fires for understorey restoration in weed‐invaded temperate eucalypt woodlands

Question. Can strategic burning, targeting differing ecological characteristics of native and exotic species, facilitate restoration of native understorey in weed‐invaded temperate grassy eucalypt woodlands? Location. Gippsland Plains, eastern Victoria, Australia. Methods. In a replicated, 5‐year experimental trial, the effects of repeated spring or autumn burning were evaluated for native and exotic plants in a representative, degraded Eucalyptus tereticornis grassy woodland. Treatments aimed to reduce seed banks and modify establishment conditions of exotic annual grasses, and to exhaust vegetative reserves of exotic perennial grasses. Treatments were applied to three grassland patch types, dominated by the native grass Austrodanthonia caespitosa, ubiquitous exotic annuals, or the common exotic perennial grass Paspalum dilatatum. Results. The dominant native grass Austrodanthonia caespitosa and native forbs were resilient to repeated fires, and target exotic annuals and perennials were suppressed differentially by autumn and spring fires. Exotic annuals were also suppressed by drought, reducing the overall treatment effects but indicating important opportunities for restoration. The initially sparse exotic geophyte Romulea rosea increased in cover with fire and the impact of this species on native forbs requires further investigation. There was minimal increase in diversity of subsidiary natives with fire, probably owing to lack of propagules. Conclusions. While fire is often considered to increase ecosystem invasibility, our study showed that strategic use of fire, informed by the relative responses of available native and exotic taxa, is potentially an effective step towards restoration of weed‐invaded temperate eucalypt woodlands.     

Assessing Fire Potential in a Brazilian Savanna Nature Reserve1

Fire is a natural ecological force in the cerrado. However, the increasing use of fire by people means that conservation areas are subject to frequent burns. The aim of this study was to assess the potential of fire in an ecological reserve in the Brazilian savannas (cerrado) of central Brazil. Data about vegetation type, topography, climate, and fuel characteristics were input into the fire prediction models BEHAVE and FARSITE to simulate fire behavior during different weather conditions and from different entry points into the conservation area. The results indicated that there is a higher probability of fire entry from particular border regions as a result of the fuel characteristics. The presence of invasive grasses, such as Melinis minutiflora, within parts of the reserve also significantly affected the pattern of fire spread. Wind speed greatly increased the spread and extent of fire. The study showed that significant improvements in modeling fire behavior in savannas still need to be made. This study was the initial stage in the development of a decision support system for fire management in the cerrado.     

Subtle Land‐Use Change and Tropical Biodiversity: Dung Beetle Communities in Cerrado Grasslands and Exotic Pastures

Although many tropical savannas are highly influenced by humans, the patterns of biodiversity loss in these systems remain poorly understood. In particular, the biodiversity consequences of replacing native grasslands with exotic pastures have not been studied. Here we examine how the conversion of the native savanna grasslands affects dung beetle communities. Our study was conducted in 14 native (grassland: campo limpo), and 21 exotic (Urochloa spp. monoculture) pastures in Carrancas, Minas Gerais, Brazil. We collected 4996 dung beetle individuals from 66 species: 3139 individuals from 50 species in native pastures and 1857 individuals from 55 species in the exotic pastures. Exotic pastures had lower dung beetle richness, abundance and biomass than native pastures. Species composition between the two pasture types was significantly different and exotic pastures were dominated by few abundant species. Indicator species analysis detected 16 species indicators of native pastures and three of exotic pastures, according to relative abundance and frequency in each pasture system. Our results show that the conversion of native pastures to exotic pastures leads to a predictable loss of local species richness, increasing dominance and changes in species composition. These results highlight the importance of maintaining native pastures in the Cerrado agro‐pastoral landscape. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Exotic herbivores directly facilitate the exotic grasses they graze: mechanisms for an unexpected positive feedback between invaders

The ability of an exotic species to establish in a system may depend not only on the invasibility of the native community, but also on its interactions with other exotic species. Though examples of mutually beneficial interactions between exotic species are known, few studies have quantified these effects or identified specific mechanisms. We used the co-invasion of an endangered island ecosystem by exotic Canada geese (Branta canadensis) and nine exotic annual grasses to study the effects of an invading herbivore on the success of invading grasses. On our study islands in southwestern Canada, we found that geese fed selectively on the exotic grasses and avoided native forbs. Counter to current theory suggesting that the grasses should be limited by a selective enemy, however, the grasses increased in proportional abundance under grazing whereas forbs showed declining abundance. Testing potential mechanisms for the effects of grazing on grasses, we found that the grasses produced more stems per area when grazing reduced vegetation height and prevented litter accumulation. Forming dense mats of short stems appeared to be an efficient reproductive and competitive strategy that the Eurasian grasses have evolved in the presence of grazers, conferring a competitive advantage in a system where the native species pool has very few annual grasses and no grazers. Germination trials further demonstrated that selective herbivory by geese enables their dispersal of exotic grass seed between heavily invaded feeding areas and the small islands used for nesting. In summary, the exotic geese facilitated both the local increase and the spatial spread of exotic grasses, which in turn provided the majority of their diet. This unexpected case of positive feedback between exotic species suggests that invasion success may depend on the overall differences between the evolutionary histories of the invaders and the evolutionary history of the native community they enter. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Invasion by Non-Native Annual Grasses: The Importance of Species Biomass, Composition, and Time Among California Native Grasses of the Central Valley

The Central Valley of California is noted for its dearth of remnant native grass populations and for low native grass seedling establishment within grasslands now dominated by non‐native annual species. In contrast, remnant populations are common along the coast, and studies have shown an ability for seedlings and adults to compete with non‐native annual grasses. The invasibility of well‐established populations of native grasses in the Central Valley remains unclear. The objectives of this study were to compare the invasibility of native grasses differing in density and species composition and, given the species in this study, to assess the ability of mixes with greater species richness to resist invasion relative to their abilities in monoculture. In the Sacramento Valley of California, six species of native grasses were planted at three densities in monospecific and mixed‐species plots. Percent cover of native perennial and non‐native annual grasses was measured in years 2 and 3, and biomass was sampled in year 5. Native grass biomass and, to a lesser extent, species composition were important in explaining variation in non‐native grass invasibility in the fifth year. Species‐rich treatments did not experience less invasion than would be expected by the proportional invasibility of each species in monoculture. However, invasibility of plots consisting of slower growing, shorter statured species decreased over time, suggesting a successional benefit to diverse communities. This study demonstrates that established stands of native grasses in the Sacramento Valley can resist invasion by non‐native annual grasses and that stand biomass is a particularly important factor in determining invasibility.