Consumers and establishment limitations contribute more than competitive interactions in sustaining dominance of the exotic herb garlic mustard in a Wisconsin, USA forest

The understory is a diverse component of temperate forest ecosystems, contributing significantly to forest ecosystem services. Despite their importance, many native understories face stresses from current and past land use, habitat fragmentation, invasive species, and overabundant herbivores. We established a four block, three factor experiment to evaluate the relative contribution of native plant establishment, competitive effects from the invasive herb garlic mustard (Alliaria petiolata), and herbivory from white-tailed deer (Odocoileus virginianus) to better understand the mechanisms promoting low native plant richness and cover and understory dominance by the biennial exotic herb garlic mustard in a NE Wisconsin, USA forest. Four years of garlic mustard removal failed to increase native plant richness or cover in non-restored plots. However, deer access and the introduction of native plants (restoration treatment) both significantly enhanced native plant cover and richness, with restored species cover in fenced plots approximately 216 % that of open-access plots, and the majority of these species flowered at significantly higher proportions inside of fenced areas. In contrast, deer access did not significantly alter the cover, or seed production of garlic mustard. We also found no significant effect of garlic mustard presence on the cover or flowering of restored native species. We conclude that multiple factors, including limited natural establishment by native species and selective herbivory drove low native, high exotic dominance at our site, suggesting that a shift in focus from invasive plant removal to combined native plant restoration and herbivore control is needed to maximize the recovery of this degraded forest understory.     

The role of plant–mycorrhizal mutualisms in deterring plant invasions: Insights from an individual‐based model

Understanding the factors that determine invasion success for non‐native plants is crucial for maintaining global biodiversity and ecosystem functioning. One hypothesized mechanism by which many exotic plants can become invasive is through the disruption of key plant–mycorrhizal mutualisms, yet few studies have investigated how these disruptions can lead to invader success. We present an individual‐based model to examine how mutualism strengths between a native plant (Impatiens capensis) and mycorrhizal fungus can influence invasion success for a widespread plant invader, Alliaria petiolata (garlic mustard). Two questions were investigated as follows: (a) How does the strength of the mutualism between the native I. capensis and a mycorrhizal fungus affect resistance (i.e., native plant maintaining >60% of final equilibrium plant density) to garlic mustard invasion? (b) Is there a non‐linear relationship between initial garlic mustard density and invasiveness (i.e., garlic mustard representing >60% of final equilibrium plant density)? Our findings indicate that either low (i.e., facultative) or high (i.e., obligate) mutualism strengths between the native plant and mycorrhizal fungus were more likely to lead to garlic mustard invasiveness than intermediate levels, which resulted in higher resistance to garlic mustard invasion. Intermediate mutualism strengths allowed I. capensis to take advantage of increased fitness when the fungus was present but remained competitive enough to sustain high numbers without the fungus. Though strong mutualisms had the highest fitness without the invader, they proved most susceptible to invasion because the loss of the mycorrhizal fungus resulted in a reproductive output too low to compete with garlic mustard. Weak mutualisms were more competitive than strong mutualisms but still led to garlic mustard invasion. Furthermore, we found that under intermediate mutualism strengths, the initial density of garlic mustard (as a proxy for different levels of plant invasion) did not influence its invasion success, as high initial densities of garlic mustard did not lead to it becoming dominant. Our results indicate that plants that form weak or strong mutualisms with mycorrhizal fungi are most vulnerable to invasion, whereas intermediate mutualisms provide the highest resistance to an allelopathic invader.     

Light,allelopathy, and post-mortem invasive impact on native forest understory species

Extended leaf phenology (early budbreak and/or delayed leaf drop) and allelopathy are potentially key invasion mechanisms in North American deciduous forests. Because extended phenology confers increased access to light energy and allelochemical production is energetically costly, these traits may interact synergistically to determine invader impact. Garlic mustard (Alliaria petiolata) exhibits both traits, and may also exploit high light in open habitats. We manipulated seasonal light availability to examine effects of light on garlic mustard’s growth, allelochemical production, and impact on native species. Invaded and not-invaded woodland microcosms were exposed to three light treatments: shading year-round (‘extended shade’), shading when the local tree canopy was closed (‘natural shade’), and ambient light year-round (‘no-shade’). Regardless of native presence, garlic mustard biomass was highest under natural shade and, due to apparent irradiation damage, lowest under no-shade. Similarly, growth and fruit production of garlic mustard monocultures were reduced in unshaded conditions. Consistent with these results, garlic mustard reduced the growth of native woodland forbs Blephilia hirsuta and Ageratina altissima most under natural shade, suggesting that extended leaf phenology mediates impact on these herbaceous species. However, garlic mustard growth did not predict reduction of whole-community biomass: invasion reduced native community growth most under no-shade, where invader biomass was lowest but allelochemical production was highest. This result may be driven by a ‘post-mortem’ pulse of allelochemicals from decaying garlic mustard tissue. We conclude that extended leaf phenology may mediate garlic mustard’s impact on some native species, and that light and allelopathy may interact to drive invasion.     

Competitive Interactions among First‐Year and Second‐Year Plants of the Invasive,Biennial Garlic Mustard (Alliaria petiolata) and Native Ground Layer Vegetation

We studied the effects of hand weeding of second‐year plants of the biennial garlic mustard (Alliaria petiolata) on first‐year plants (seedlings) and native ground layer vegetation. Garlic mustard is a Eurasian species that has invaded deciduous forest ground layers in eastern North America. Treatments consisted of a control and an early or late weeding of second‐year garlic mustard. The early treatment (early March) was applied before garlic mustard seeds had germinated and when most native species were dormant. The late treatment (mid‐May) occurred after plants had bolted, flowering was occurring, and most native species and new garlic mustard seedlings were actively growing. Pre‐treatment data were obtained in 2004 and treated and control plots were sampled in 2005, 2006, and 2007. No significant treatment effects were observed in 2004 or 2005. In 2006, mean cover of first‐year plants was higher in the early weeding treatment than in the late weeding treatment and control. In 2007, mean cover of first‐year garlic mustard was higher in the control than in either of the two weeding treatments. There were no significant treatment effects in any year on native vegetation cover, bare ground, or the five most abundant native species. Our data indicate that (1) late weeding of garlic mustard provided more effective control than early weeding because late weeding allows second‐year plants to compete with garlic mustard seedlings for a longer period of time and (2) competition between first‐ and second‐year plants is responsible for alternating dominance of first‐year and second‐year garlic mustard plants.     

Competitive and allelopathic effects of garlic mustard (<Emphasis Type="Italic">Alliaria petiolata</Emphasis>) on American ginseng (<Emphasis Type="Italic">Panax quinquefolius</Emphasis>)

The purpose of this study was to test competitive and allelopathic effects of invasive garlic mustard on American ginseng (Panax quinquefolius) seedlings under natural conditions. For comparative purposes, we also examined the potential effects of the native striped violet (Viola striata). In order to partition effects of resource competition and chemical suppression via allelopathy, field soils were amended with activated carbon or left unamended. Activated carbon positively affected ginseng growth as well as biomass of competitors. Ginseng mortality tended to increase with garlic mustard presence, though activated carbon alleviated this response. Garlic mustard had no significant effect on ginseng seedling growth, while striped violet suppressed shoot length in the absence of activated carbon. Our results showed a surprising effect of activated carbon on plant growth, a potential allelopathic effect of the native striped violet and suggest that newly invaded ginseng populations with low densities of garlic mustard may be able to withstand its effects. However, recruitment within invaded populations may decline.     

Low allelochemical concentrations detected in garlic mustard-invaded forest soils inhibit fungal growth and AMF spore germination

Garlic mustard’s (Alliaria petiolata, Brassicaceae) invasive success is attributed in part to its release of allyl isothiocyanate (AITC) into the soil. AITC can disrupt beneficial arbuscular mycorrhizal fungi (AMF) associated with native plant roots, which limits their soil resource uptake. However, AITC and its precursor, sinigrin, have never been detected in garlic mustard-invaded forest soils. Here, we use high performance liquid chromatography (HPLC) and gas chromatography-mass spectrometry (GC–MS) to assess the concentration and bioactivity of these putative allelochemicals in paired forest plots uninvaded or invaded by garlic mustard. Our methods detected AITC and sinigrin only where garlic mustard was present and our recovery of AITC/sinigrin coincided with adult senescence. A bioassay of in situ fungal hyphae abundance revealed significantly reduced hyphal abundance in the presence of garlic mustard relative to uninvaded soils. Finally, the lowest concentration of AITC measured in the field (~0.001 mM) is highly inhibitory to the spore germination of a forest AMF species, Glomus clarum. Together, our data provide the first direct evidence of garlic mustard-produced sinigrin and AITC in forest soils and demonstrate that even low levels of these chemicals have the potential to significantly suppress AMF growth and spore germination, strengthening their status as allelopathic novel weapons.     

Declines in a ground-dwelling arthropod community during an invasion by Sahara mustard (Brassica tournefortii) in aeolian sand habitats

Sahara Mustard (Brassica tournefortii; hereafter mustard), an exotic plant species, has invaded habitats throughout the arid southwestern United States. Mustard has reached high densities across aeolian sand habitats of southwestern deserts, including five distinct sand habitats in the eastern Coachella Valley, California. We examined trends in ground-dwelling arthropod community structure concurrent with mustard invasion in 90 plots within those habitats from 2003 to 2011 (n = 773 plot·years). We expected arthropod communities to respond negatively to mustard invasion because previous work documented significant negative impacts of mustard on diversity and biomass of native plants, the primary resource base for many of the arthropods. Arthropod abundance and species richness declined during the study period while mustard cover increased, and arthropod metrics were negatively related to mustard cover across all plots. When controlling for non-target environmental correlates (e.g. perennial frequency and precipitation) and for potential factors that we suspected of mediating mustard effects (e.g. native cover and sand compaction), negative relationships with mustard remained statistically supported. Nevertheless, arthropod richness’s relationship decreased slightly in strength and significance suggesting that mechanistic pathways may be both direct (via habitat structure) and indirect (via native cover suppression and sand compaction). However, mechanistic pathways for mustard effects, particularly on arthropod abundance, remain unclear. Most arthropod taxa, including most detritivores, decreased through time and were negatively related to mustard cover. In contrast, many predators were positively related to mustard. In total, our study provides substantial evidence for a negative effect of Sahara mustard on the structure of a ground-dwelling arthropod community.     

Seed rain and environmental controls on invasion of <Emphasis Type="Italic">Picea abies</Emphasis> into grassland

Although changes in land-use, climate, and the spread of introduced tree species have increased the global importance of tree invasions into grasslands, our ability to predict any particular invasion is limited. To elucidate mechanisms driving tree invasions of grasslands, we studied in detail how seed dispersal and fine-scale environment control the expansion of an introduced Picea abies Karst. (Norway spruce) population into Western Carpathian grassland. We mapped invading trees and measured tree size, fecundity, seed rain, seedling density, plant community composition, and light and soil environment within a 200 × 60 m belt across the invasion front. Maximum likelihood estimates of dispersal kernels suggested peak seed deposition directly underneath tree crowns where germination was poor, but mean dispersal distances were sufficiently large to generate overlapping seed shadows from multiple trees that saturated the invasion front with seeds further away from seed-dispersing trees. Partial Mantel tests indicated that germinant density was affected considerably less by seed rain than by moss cover (r = 0.54), overstory tree influence (r = −0.32), soil moisture (r = 0.21), grass cover (r = −0.15), and diffuse radiation (r = 0.13). However, these variables were not independent but formed complex multivariate gradients within the invasion front. Moss cover and soil moisture were negatively correlated with overstory tree influence and the resulting gradient clearly affected germinant density (partial Mantel r = 0.45). In contrast, positively correlated light and grass cover defined a gradient related weakly to germinant density (partial Mantel r = 0.05) as it integrated opposing effects of these variables on germinants. Seedlings had similar environmental associations, but except for the lasting positive effects of moss these tended to weaken with seedling size. Although a few seedlings may establish and survive in the more adverse environment of the outer edges of the invasion front, a significant population expansion may require a gradual build-up of the critical density of invading trees to reduce grass cover and facilitate germination on moist mossy seedbeds within uncolonized areas. Thus, Picea abies appears more likely to spread within temperate grasslands by gradual expansion of its population frontier rather than by advanced groups.     

Palm Community Structure and Land Cover Changes in the San Juan Biological Corridor, Costa Rica

Biological corridors connect forest fragments, minimize the isolation of remaining habitats, and reduce the negative impact of fragmentation. Important studies in a corridor include documentation of land cover and the relationships between the plant community and land cover changes. In this study, density, species richness, and species diversity of palms of different life stages and growth forms were documented in forty 10 × 10 m census plots in the San Juan Biological Corridor (SJBC), Costa Rica. Changes in the percent cover of agricultural, partially forested, and forested lands from 1979 to 2001 were quantified based on existing land classifications. Associations between these palm community characteristics and the reduction in forest cover in 200-m and 1000-m annuli surrounding the census plots were investigated. These palm community characteristics were spatially interpolated to examine their distributions in SJBC. The community characteristics of palms of all growth forms varied among census plots. Forested lands in SJBC decreased from 88 percent to 26 percent during 1979–2001, while the percentage cover of agricultural and partially forested lands increased. The annual deforestation rate was 0.8 percent from 1979 to 1992, 4.3 percent from 1992 to 1997, and 7.5 percent from 1997 to 2001. Overall, palm community characteristics of small-canopy palms, dwarf palms, and cotyledonous seedlings had lower values with a lower percentage forest cover. The spatially interpolated palm community characteristics were consistently greatest within or adjacent to protected areas. This study demonstrates the complexity of the landscape and the effect of loss of adjacent forests on the palm community.     

Escaping an evolutionary trap: preference and performance of a native insect on an exotic invasive host

Exotic plants may act as population sinks or evolutionary traps for native herbivores. The native butterfly Pieris oleracea lays eggs on garlic mustard, Alliaria petiolata, but larvae develop very poorly on this exotic invasive plant. We examined oviposition preference of individual females and larval performance of their offspring for individuals from one area where garlic mustard is well established and one where it is absent. These data were used to assess whether garlic mustard is being incorporated into or excluded from the diet. Females from the area without garlic mustard showed a wide range of preference, families had low larval survival on garlic mustard, and larval survivorship showed no correlation with mothers’ preferences. Females from the area with garlic mustard preferred it to the native host, and larval survivorship on garlic mustard was positively correlated with the mother’s preference. Individuals surviving on garlic mustard took longer to pupate and weighed >30% less compared to pupae reared on normal hosts. Our results suggest that where garlic mustard is well established P. oleracea may be adapting to this plant by both improved larval performance and increased adult female oviposition preference for it.     

Allelopathic inhibition of germination by Alliaria petiolata (Brassicaceae)

Garlic mustard (Alliaria petiolata, Brassicaceae) is an invasive, nonindigenous species currently invading the understory of North American woodlands where it is a serious threat to the native flora. Part of this success might be due to allelopathic interference by garlic mustard. Two congeneric species, the European Geum urbanum and the North American Geum laciniatum, were tested for allelopathic inhibition of germination by garlic mustard. Seeds were germinated either on substrate contaminated by garlic mustard or on substrate with contamination neutralized by activated carbon. Allelopathic effects of native European and invasive North American garlic mustard populations were also compared. Activated carbon increased germination by 14%, indicating that garlic mustard contaminated the substrate through root exudates. Activated carbon in turn counteracted this effect. The two test species differed in their sensitivity to allelopathic interference. North American G. laciniatum had a much stronger increase in germination when activated carbon was added to the substrate, independent of the origin of garlic mustard. In contrast, the European G. urbanum germinated better in substrate precultivated with North American garlic mustard, whereas activated carbon increased its germination only in substrate precultivated with European garlic mustard.     

Infection by powdery mildew Erysiphe cruciferarum (Erysiphaceae) strongly affects growth and fitness of Alliaria petiolata (Brassicaceae)

Garlic mustard (Alliaria petiolata) is an invasive biennial that negatively impacts plant and animal communities throughout North America and lacks significant herbivory in its invasive range. Throughout Ohio, many garlic mustard populations support the powdery mildew fungus Erysiphe cruciferarum, although disease incidence varies among populations and environments. Effects of infection on plant growth, as well as both plant and fungal responses to drought and light conditions, were examined on greenhouse-grown, first-year garlic mustard plants. Also, the effects of the fungus on plant growth and fitness were studied in a naturally growing population of second-year plants in the field. Powdery mildew significantly reduced growth of first-year plants in the greenhouse, eventually causing complete mortality. Simulated drought slowed both plant growth and disease development, independent of light conditions. In the field, plants with little incidence of disease after their first year grew taller during their second year, producing significantly more siliques and twice as many seeds as heavily diseased plants did. Seed germination rates did not differ between plants with different levels of disease severity. Consistent reductions in survival, growth, and fitness caused by fungal infection may reduce populations of garlic mustard. These effects may be more evident in moist sites that favor fungal development.     

Influence of litter removal and mineral soil disturbance on the spread of an invasive grass in a Central Hardwood forest

Soil and litter disturbances within Central Hardwood forests may facilitate exotic plant species invasion of interior forest areas. Microstegium vimineum is an annual exotic grass that has become common throughout the Southeastern United States. Three replicates of three different mineral soil and litter disturbance treatments, plus a control with no disturbance, were established on the leading edge of M. vimineum patches prior to seed fall. All patches were located in areas with similar forest canopy structure and slope in three Central Hardwood forest stands prior to seed fall. At the beginning of the following growing season, each individual M. vimineum seedling was mapped within the treatment plots. The mean number of M. vimineum individuals that established within each treatment did not differ significantly from the control. The distance at which 90% of the individuals had spread, and the overall mean distance spread were significantly farther for the litter removal treatment than the control. The farthest individual seedling from the boundary of existing patches in both the litter removal and the mineral soil disturbance and litter removal treatments were significantly farther than the control. The individuals that spread the farthest are of most concern due to the large number of viable seed that a single M. vimineum plant can produce. These results suggest that disturbance of the litter layer may increase the spread rate of M. vimineum and facilitate its invasion of new habitats, and that leaving litter layers intact may slow the spread of M. vimineum.     

Disturbance,drought and dynamics of desert dune grassland,South Africa

A seven-year study of marked plants and plots in Stipagrostis ciliata (Desf.) de Winter dune grassland, in the arid (<100 mm yr^–1) Bushmanland area of the Northern Cape province of South Africa, was designed to test the hypothesis that establishment of ephemeral plants, and recruitment of perennial grasses was dependent upon disturbances that reduced the density of living perennial grass tussocks. In 1989, eight 4 m² plots were cleared of perennial vegetation by uprooting and removing all plants so as to resemble small-scale disturbances made by burrowing mammals or territorial antelope. The vegetation on the cleared plots and surroundings was monitored until 1996. Initial results supported our hypothesis. In wet years, when ephemeral plants were abundant, their average fresh mass was 2–3 times greater per unit area on the cleared plots than in control plots in adjacent, undisturbed grassland. Many Stipagrostis seedlings established in the cleared plots over the two years following clearing but were rare in adjacent areas among established conspecifics. However, a drought in 1992 (11 mm of rain over 12 months) lead to widespread mortality of the perennial grass, killing 56% (range 22–79%) of established tufts. High densities of Stipagrostis seedlings appeared following the drought-breaking rains in January 1993, both in the disturbed plots and in the surrounding `undisturbed' dune grassland. Ephemeral plants established in large numbers throughout the area during the high rainfall year of 1996 and were generally more numerous in the old disturbances than in control plots. Seven years after clearing the biomass of grass on the cleared plots was approximately 34% of the mass removed from the plots in 1989 whereas in the undisturbed grassland biomass was 66% of 1989 levels. Drought had little long-term effect on community composition, and Stipagrostis ciliata constituted 94–98% of plant community before and after drought. Cleared plots were recolonised by S. ciliata, but the contribution of other grass species increased by 6–9%. Synchronous recruitment following occasional drought-induced mortality can generate even-aged populations of the dominant desert dune grasses.     

Effects of Herbicide on the Invasive Biennial Alliaria petiolata (Garlic Mustard) and Initial Responses of Native Plants in a Southwestern Ohio Forest

Restoration often includes control of invasive plants, but little is known about how native plant communities respond to this control. The biennial Alliaria petiolata (M. Bieb.) Cavara and Grande (garlic mustard) is one of the most prevalent invasive plants in forests of eastern North America. We investigated the effects of the herbicide Round‐up (glyphosate) on Alliaria and the response of the forest floor plant community to the herbicide and the subsequent decline of Alliaria. In an old‐growth Acer–Fagus stand and a second‐growth Liriodendron‐dominated stand in Hueston Woods State Nature Preserve, Ohio, United States, we spot applied Round‐up in November 2000 and 2001 in 25 1 × 1–m plots and maintained 25 control plots. Herbicide decreased Alliaria density in both stands and reduced the density of other species in leaf during treatment (mostly exotic winter annuals) in the old‐growth stand. Treatment did not affect the initial density of the Alliaria cohort that germinated in the spring of 2001, but decreased the 2002 cohort. Community differences were found in the old‐growth stand after Alliaria reduction, specifically greater cover of spring ephemerals in the herbicide treatment. In the second‐growth stand, herbicide treatment increased reproduction of the late‐summer perennial, Phryma leptostachya. These results indicate that glyphosate reduces Alliaria without negatively impacting native species and that some native species respond positively to a single‐year reduction in this invasive biennial.     

Native species richness buffers invader impact in undisturbed but not disturbed grassland assemblages

Many systems are prone to both exotic plant invasion and frequent natural disturbances. Native species richness can buffer the effects of invasion or disturbance when imposed in isolation, but it is largely unknown whether richness provides substantial resistance against invader impact in the face of disturbance. We experimentally examined how disturbance (drought/burning) influenced the impact of three exotic invaders (Centaurea stoebe, Linaria dalmatica, or Potentilla recta) on native abundance across a gradient of species richness, using previously constructed grassland assemblages. We found that invaders had higher cover in experimentally disturbed plots than in undisturbed plots across all levels of native species richness. Although exotic species varied in cover, all three invaders had significant impacts on native cover in disturbed plots. Regardless of disturbance, however, invader cover diminished with increasing richness. Invader impacts on native cover also diminished at higher richness levels, but only in undisturbed plots. In disturbed plots, invaders strongly impacted native cover across all richness levels, as disturbance favoured invaders over native species. By examining these ecological processes concurrently, we found that disturbance exacerbated invader impacts on native abundance. Although diversity provided a buffering effect against invader impact without disturbance, the combination of invasion and disturbance markedly depressed native abundance, even in high richness assemblages.     

Impact of tree species on nutrient and light availability: evidence from a permanent plot study of old-field succession

This paper compares vegetation composition, light availability, carbon and nutrient pools and Ellenberg indicator values among four old-field successional permanent plots that have received an initial treatment (ploughing, herbicide or sterilisation) prior to being left undisturbed in 1969, a second set of six plots received additional treatments (continued ploughing or mulching until 1982). On all plots species rich pioneer forests developed. Vegetation still varies among plots with different initial treatments: Sterilised plots can be distinguished from the others by dominance of Betula pendula, ploughed plots by Fraxinus excelsior, whereas herbicide-treated plots are intermediate with proportions of both species. By affecting light availability at the ground, tree species in turn influences ground vegetation and soil properties. Light availability turned out to be the dominant factor determining the composition of the vegetation in old-field succession.     

Twelve years of repeated wild hog activity promotes population maintenance of an invasive clonal plant in a coastal dune ecosystem

Invasive animals can facilitate the success of invasive plant populations through disturbance. We examined the relationship between the repeated foraging disturbance of an invasive animal and the population maintenance of an invasive plant in a coastal dune ecosystem. We hypothesized that feral wild hog (Sus scrofa) populations repeatedly utilized tubers of the clonal perennial, yellow nutsedge (Cyperus esculentus) as a food source and evaluated whether hog activity promoted the long‐term maintenance of yellow nutsedge populations on St. Catherine's Island, Georgia, United States. Using generalized linear mixed models, we tested the effect of wild hog disturbance on permanent sites for yellow nutsedge culm density, tuber density, and percent cover of native plant species over a 12‐year period. We found that disturbance plots had a higher number of culms and tubers and a lower percentage of native live plant cover than undisturbed control plots. Wild hogs redisturbed the disturbed plots approximately every 5 years. Our research provides demographic evidence that repeated foraging disturbances by an invasive animal promote the long‐term population maintenance of an invasive clonal plant. Opportunistic facultative interactions such as we demonstrate in this study are likely to become more commonplace as greater numbers of introduced species are integrated into ecological communities around the world.     

Frequent burning promotes invasions of alien plants into a mesic African savanna

Fire is both inevitable and necessary for maintaining the structure and functioning of mesic savannas. Without disturbances such as fire and herbivory, tree cover can increase at the expense of grass cover and over time dominate mesic savannas. Consequently, repeated burning is widely used to suppress tree recruitment and control bush encroachment. However, the effect of regular burning on invasion by alien plant species is little understood. Here, vegetation data from a long-term fire experiment, which began in 1953 in a mesic Zimbabwean savanna, were used to test whether the frequency of burning promoted alien plant invasion. The fire treatments consisted of late season fires, lit at 1-, 2-, 3-, and 4-year intervals, and these regularly burnt plots were compared with unburnt plots. Results show that over half a century of frequent burning promoted the invasion by alien plants relative to areas where fire was excluded. More alien plant species became established in plots that had a higher frequency of burning. The proportion of alien species in the species assemblage was highest in the annually burnt plots followed by plots burnt biennially. Alien plant invasion was lowest in plots protected from fire but did not differ significantly between plots burnt triennially and quadrennially. Further, the abundance of five alien forbs increased significantly as the interval (in years) between fires became shorter. On average, the density of these alien forbs in annually burnt plots was at least ten times as high as the density of unburnt plots. Plant diversity was also altered by long-term burning. Total plant species richness was significantly lower in the unburnt plots compared to regularly burnt plots. These findings suggest that frequent burning of mesic savannas enhances invasion by alien plants, with short intervals between fires favouring alien forbs. Therefore, reducing the frequency of burning may be a key to minimising the risk of alien plant spread into mesic savannas, which is important because invasive plants pose a threat to native biodiversity and may alter savanna functioning.     

The impact of grazing pressure in clearfelled,burned and undisturbed eucalypt forest

An area of dry eucalypt forest on dolerite in southeastern Tasmania with comparable sites that were a) unlogged, b) clearfelled, c) clearfelled and slash-burned, was monitored over a 24 month period. Detailed records were made of plant species composition and cover on floristically similar fenced and unfenced permanent plots in two major vegetation types. Mammal faecal pellets were cleared from each of the unfenced plots and subsequently counted and attributed to species of origin. For the six combinations of treatment and vegetation type pellet deposition rate was highly correlated with differences in the rate of change of species cover between fenced and unfenced plots and the lifeform composition of the vegetation. The greatest changes were observed in the clearfelled, burned area, while there was relatively little change in the undisturbed forest. Grass and herb species were generally disadvantaged by mammal exclusion while shrubs and graminoids generally grew faster in the fenced plots.Nomenclature of plant species follows Curtis & Morris (1975).This project was supported financially by the Forest Ecology Research Fund, Tasmania.