Post-pastoral changes in composition and guilds in a semi-arid conservation area, Central Otago, New Zealand

Changes in the vegetation of Flat Top Hill, a highly modified conservation area in semi;arid Central Otago, New Zealand, are described four years after the cessation of sheep and rabbit grazing. Unusually moist weather conditions coincide with the four-year period of change in response to the cessation of grazing. Between 1993 and 1997, the average richness and diversity (H') of species increased, and the average proportion of native species decreased significantly. The vegetation was significantly richer in exotic annual and perennial grass species, exotic perennial forbs, exotic woody species and native tussock grasses in 1997 than in 1993. Eight response guilds of species are identified. Most "remnant" native shrubs and forbs were stable, in that they remained restricted to local refugia and showed little change in local frequency. However, taller native grass species increased, some locally, and others over wide environmental ranges. Rare native annual forbs and several native perennial species from "induced" xeric communities decreased, and this may be a consequence of competition from exotic perennial grasses in the absence of grazing. The invasive exotic herb Sedum acre decreased in abundance between 1993 and 1997, but several other prominent exotic species increased substantially in range and local frequency over a wide range of sites. Exotic woody species, and dense, sward-forming grasses are identified as potential threats to native vegetation recovery.     

The vegetation of roadside verges with respect to environmental gradients in southern New Zealand

Abstract. A survey of the vegetation of roadside verges was made across the southern part of the South Island of New Zealand. Samples were taken at 10-km intervals along selected roads providing a climatic range from the suboceanic conditions of the east coast into semi-arid Central Otago, and from Central Otago through the Southern Alps to the hyper-oceanic areas of high rainfall on the west Coast. The variation in the floristic composition is associated mainly with variation in rainfall, continentality, altitude, soil acidity, soil organic matter, and presence of forest. Sites in the arable and pastoral regions on the eastern side of the Southern Alps support a herbaceous vegetation consisting mostly of exotic species of European origin, with a few native grasses scattered through the drier and less fertile sites. A greater proportion of native species is found at higher altitudes. Roadside vegetation in the area of high rainfall to the west is characterized by indigenous ferns and woody species, although vegetation adjacent to cleared areas is more similar to that on roadsides adjacent to farmland on the east coast. The pattern of distribution of both native and exotic species is strongly related to altitudinal and climatic gradients, and the environmental responses of the exotic species are similar to those recorded in Europe. This suggests a colonization of all available sites by the exotic species, despite the relatively short time since their introduction to New Zealand, rather than an incomplete invasion.     

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.     

Review of ecological restoration programme on subantarctic Macquarie Island: Pest management progress and future directions

Summary The establishment of exotic species of vascular flora and vertebrate fauna on subantarctic Macquarie Island since its discovery in 1810 has resulted in major changes in the biota. A management programme aims to reduce the numbers of exotic plant and animal species and assist with the recovery of pre-existing communities and processes. This paper reviews the integrated vertebrate pests management programme on Macquarie Island since 1974 and outlines future management considerations. As part of this programme, the responses of some native and exotic species of vascular flora and vertebrate fauna were monitored following control of European Rabbit ( Oryctolagus cuniculus ) numbers. Changes in the vegetation recorded over 10 years showed that approximately half of all the vascular species had benefited from rabbit grazing, including several which formed a major part of the rabbit's diet. After rabbit control, some adversely affected plants responded rapidly to a reduction in grazing pressure while others will require an almost total cessation of grazing in order to re-establish their former distributions. With the decrease in rabbit numbers it was also necessary to control Feral Cats ( Felis catus ) due to their increased predation on native burrow-nesting birds. Feral Cat predation on introduced fauna also increased, one result of which was the eradication from the island of the introduced Weka ( Gallirallus australis scotti ). Reduced rabbit grazing is leading to re-establishment of the native Tall Tussock ( Poa foliosa ) grassland and with it the spread of the introduced Ship Rat ( Rattus rattus ). This review indicates that an integrated approach to pest management, with monitoring of the responses of both target and non-target species, is the most effective way to restore pre-existing communities and processes.
Key words grazing pressure, introduced species, predation, recovery, vertebrate pest management.     

Response of endemic and exotic earthworm communities to ecological restoration

Land conversion and environmental changes associated with agronomic practices are believed to have led to the disappearance of New Zealand endemic earthworms from agricultural land. Introduced European earthworms have since largely replaced endemic species in farming systems. We investigated the impact of vegetation restoration on earthworm communities. Recolonization by endemic earthworms increased with time after restoration at two studied sites in the South Island of New Zealand. However, exotic species did not disappear with restoration of native vegetation, even after 30 years. The persistence of exotic species leads to the cohabitation of the two communities and potential for interspecific competition.     

Reintroduction of Nassella pulchra to California coastal grasslands: Effects of topsoil removal,plant neighbour removal and grazing

Question: What is the most appropriate combination of treatments to reintroduce Nassella pulchra, a perennial bunchgrass, into degraded mediterranean coastal grasslands? Location: Central coast of California, USA. Methods: N. pulchra was sown from seeds and transplanted into a degraded grassland in a multi‐factorial experiment testing the effects of (1) two grazing intensities (lightly grazed by native mammal species or ungrazed); (2) topsoil removal and (3) reduction of plant neighbours. The experiment was carried out on two types of surrounding vegetation (exotic annual grasses and exotic forbs). Results: Topsoil removal greatly enhanced establishment from seeds and transplant survival, mainly because it reduced the exotic vegetation and thus reduced competition. While removing neighbours was essential when topsoil was left intact, it had a negative effect on N. pulchra when surrounding species included exotic forbs (Brassica spec, and Asteraceae) at low density (after topsoil removal). Moderate grazing by native mammals (deer, rabbits and gophers) did not affect N. pulchra. Conclusion: Our results suggest that seeding after topsoil has been removed is a promising method to reintroduce N. pulchra to highly degraded sites where there is little to no native seed bank.     

Removal of livestock alters native plant and invasive mammal communities in a dry grassland–shrubland ecosystem

The impacts of domesticated herbivores on ecosystems that did not evolve with mammalian grazing can profoundly influence community composition and trophic interactions. Also, such impacts can occur over long time frames by altering successional vegetation trajectories. Removal of domesticated herbivores to protect native biota can therefore lead to unexpected consequences at multiple trophic levels for native and non-native species. In the eastern South Island of New Zealand large areas of seral grassland–shrubland have had livestock (sheep and cattle) removed following changes in land tenure. The long-term (>10 years) outcomes for these communities are complex and difficult to predict: land may return to a native-dominated woody plant community or be invaded by exotic plants and mammals. We quantified direct and indirect effects of livestock removal on this ecosystem by comparing plant and invasive mammal communities at sites where grazing by livestock ceased c.10–35 years ago (conservation sites) with paired sites where pastoralism has continued to the present (pastoral sites). There was higher total native plant richness and reduced richness of exotic plants on conservation sites compared with pastoral sites. Further, there were differences in the use of conservation and pastoral sites by invasive mammals: rabbits and hedgehogs favoured sites grazed by livestock whereas house mice, brushtail possums and hares favoured conservation sites. Changes in the relative abundance of invasive mammal species after removal of domesticated livestock may compromise positive outcomes for conservation in successional plant communities with no evolutionary history of mammalian grazing.     

Manipulating disturbance regimes and seeding to restore mesic Mediterranean grasslands

Question: Can managing disturbance regimes alone or in combination with seeding native species serve to shift the balance from exotic towards native species? Location: Central coast of California, USA. Methods: We measured vegetation composition for 10 yr in a manipulative experiment replicated at three sites. Treatments included no disturbance, grazing and clipping at three frequencies with and without litter removal. We seeded eight native species into clipped plots and compared cover in comparable plots with no seeding. Results: Regardless of frequency, clipping generally shifted community dominance from exotic annual grasses to exotic annual forbs, rather than consistently favoring native species. At one site, perennial grass cover decreased in no‐disturbance plots, but only after 4 yr. Litter removal had minimal impact on litter depth and plant community composition. Grazing had a highly variable effect on the abundance of different plant guilds across sites and years. Seeding increased abundance of only two of eight native species. Conclusions: Managing disturbance regimes alone is insufficient to restore native species guilds in highly‐invaded grasslands and seeding native species has highly variable success.     

Montane outcrop vegetation of Banks Peninsula, South Island, New Zealand

Species composition patterns and vegetation?environment relationships were quantified for montane volcanic outcrops on Banks Peninsula. The flora of these habitat islands comprises 346 vascular plant species including 82 exotic species and 52 species that are nationally rare, regionally rare, or regional endemics. Both MDS ordination analysis and TWINSPAN results illustrated the high compositional and environmental heterogeneity across the outcrops. Multi-dimensional scaling revealed that primary environmental factors related to community composition comprise both regional-scale gradients of altitude and outcrop-scale gradients of slope steepness, soil pH, area available to plants, maximum vegetation height, and the percentage of the surrounding vegetation that is forest. Accordingly, TWINSPAN separated four outcrop communities that occur on steeper slopes, have less fertile soils and tend not to face north from three outcrop communities that have shallower slopes, more fertile soils and tend to face north. Types in the first group are more likely to be bordered by forest or taller shrublands, whereas those in the second group occur on outcrops primarily bordered by grasslands and support more exotic species. Within these broader groups, communities differ in their altitude and the size, soil depth and shading of the outcrops on which they occur. We describe the vegetation of the seven communities; this ranges from predominance of stunted trees and taller statured species such as Podocarpus hallii and Phormium cookianum to vegetation of shrubby species such as Heliohebe lavaudiana and Hebe strictissima, to short vegetation of native woodland and grassland species such as Polystichum vestitum and Rytidosperma corinum, to exotic pasture of clovers and exotic grasses. The percentage of species on an outcrop face that are exotic is well modelled by site factors, with exotics increasing as the surrounding matrix becomes more disturbed, slopes become more gentle, the percentage of shade on the outcrop decreases, and soil fertility increases. In contrast, nearby disturbance has little influence on the percentage or number of species that are rare on an outcrop face; rather rare species richness is more strongly related to outcrop area and lack of shade, echoing patterns observed for rare outcrop species elsewhere in the world. These results highlight the importance of considering the high compositional heterogeneity among outcrops and the influence of disturbance to surrounding ecosystems in guiding conservation planning.     

Afforestation causes changes in post‐fire regeneration in native shrubland communities of northwestern Patagonia,Argentina

Question: What are the effects of fire in native shrubland communities and in pine plantations established in these shrublands? Location: Northern Patagonia, Argentina. Methods: We surveyed four sites in Chall‐Huaco valley, located in northwest Patagonia. Each site was a vegetation mosaic composed of an unburned Pinus ponderosa plantation, a plantation burned in 1996, and an unburned matorral and a matorral burned by the same fire. We recorded the cover of all vascular plant species. We also analysed species richness, total cover, proportion of exotic species, abundance of woody species and herb species, cover of exotic species, abundance of woody and herb species and differences in composition of species. For both shrubs and tree species we recorded the main strategy of regeneration (by resprouting or by seed). Results: We found that fire had different effects on native matorral and pine plantations. Five years after fire, plantations came to be dominated by herbs and exotic species, showing differences in floristic composition. In contrast, matorral communities remained very similar to unburned matorral in terms of species richness, proportion of woody species, and herb species and proportion of exotics. Also, pine plantations were primarily colonized by seedlings, while matorrals were primarily colonized by resprouting. Conclusions: Matorrals are highly fire resilient communities, and the practice of establishing plantations on matorrals produces a strong reduction in the capacity of matorral to return to its original state. The elimination of shrubs owing to the effect of plantations can hinder regeneration of native ecosystems. Burned plantations may slowly develop into ecosystems similar to the native ones, or they may produce a new ecosystem dominated by exotic herbs. This study shows that plantations of exotic conifers affect native vegetation even after they have been removed, as in this case by fire.     

The impact of suburbanization on remnant coastal vegetation in Hobart,Tasmania

Abstract. Changes in vascular species composition and abundance were examined in coastal vegetation at Hobart, Tasmania, Australia over a period of 13 yr, during which suburbanization extended to cover a large proportion of its hinterland. There were significant increases in the richness and cover of exotic vascular plant species, most of which derived from the Mediterranean basin or southern Africa. While most of the introduced species were confined to vegetation with a high exotic component, and close to domestic gardens, some, including the South African coastal shrub, Chrysanthemoides monilifera, expanded in otherwise native vegetation relatively remote from gardens. The impact of proximity to gardens on the relative abundance of exotics and natives in coastal vegetation in 1997 best correlated with the pattern of distance from gardens in 1966, suggesting that there is, at least, a 30‐yr lag period in transformation from largely native to substantially exotic vegetation. This lag period provides an opportunity to reduce exotic invasion problems before they become severe.     

Invasion of exotic plants in nutrient‐enriched urban bushland

Abstract Invasion of exotic plants into vegetation communities on low‐nutrient soils in Sydney is often attributed to increased phosphorus in the soil. However, the composition of native vegetation in nearby national parks has been found to be more closely correlated to combinations of soil nutrients and other environmental variables than phosphorus alone. This study examined whether phosphorus or a range of variables better matched patterns of native and exotic plants in urban bushland. Sites in urban bushland and national parks were sampled, vegetation frequency recorded and soil samples collected. Soil samples were analysed for a range of chemical and physical attributes. A significantly greater number of exotic species were found at the urban sites, and significantly fewer native species than in the national parks. All measured soil nutrients were found to be of a significantly increased concentration in urban soil. Using regression analysis, an index of all measured soil nutrients was found to explain more of the variation in the percentage of exotic species at a site than phosphorus alone. Multivariate analysis showed a gradient of sites from minimal exotic invasion to heavy invasion. However, this gradient also corresponded to changes in native species. The gradient was found to match that of increasing soil nutrient levels. A combination of soil nutrients was found to correlate better with the multivariate species composition than was phosphorus alone. The results suggest that it is the increase of many soil nutrients, not phosphorus alone, that is contributing to the invasion of urban bushland by exotic plants and the alteration of the suite of native species.     

Colonization by alpine native plants of a stabilized road verge on the Bogong High Plains, Victoria

Summary Although many native species are now used in the revegetation of soil disturbances in Australian alpine areas, exotic species were, until recently, the only components of seed mixes. The use of exotic species and fertilizer was justified by their availability and low cost, and the prediction that native species would replace the exotic sward, presumably once soil nutrient levels dropped to those found in native vegetation. There was no apparent regard for the invasive capacities of the exotic species used. An investigation of a road verge on the Bogong High Plains, revegetated with a mixture of exotic species in the late 1950s, has shown that colonization by native plants can be very slow. Although many native species had colonized by 1993, exotic species still provided 50% of the cover. Agrostis capillaris accounted for most of this. The major native colonizers were Carex spp., Colobanthus affinis , Euchiton spp., Leptinella filicula , Poa hiemata , Ranunculus victoriensis and Scleranthus biflorus . Poa hiemata provided by far the most cover of the native colonizers. There have been considerable increases in the cover and number of native species on the verge since 1989. Cattle grazing and trampling are likely to have limited colonization of native plants prior to the removal of livestock in 1991. Recommendations are made on approaches to future revegetation in the alpine area.     

Long-term grazing impacts on vegetation diversity,composition, and exotic species presence across an aridity gradient in northern temperate grasslands

Little is known about the specific role of exotic species on measures of grassland plant diversity, including how this may vary with climatic conditions or large mammal herbivory. This study examined vegetation responses to long-term livestock grazing, including plant richness and diversity, as well as the contribution of exotic species to these metrics, across a network of 107 northern temperate grasslands in Alberta, Canada, spanning a broad aridity gradient. Exposure to grazing modestly increased plant richness, but did not alter Shannon’s diversity, Simpson’s diversity, or evenness, suggesting stability in floral diversity relative to grazing. However, grazing did increase grass cover while reducing shrub cover, the latter of which was only apparent in mesic grasslands. Unlike total plant diversity, exotic species richness and cover, together with exotic plant contributions to diversity, varied jointly with grazing and aridity. While long-term grazing increased exotic species, this response was most apparent in wetter areas, and non-grazed grasslands remained more resistant to the presence of exotics. Several exotic species were positive indicators of grazing in wetter grasslands, and coincided with lower native species cover, indicating grazing may be facilitating a shift from native to exotic vegetation under these conditions. Overall, our results indicate that while long-term grazing has altered the composition and cover of certain functional groups, including favoring exotics and minimizing woody vegetation in mesic areas, overall changes to plant diversity were limited. Additionally, these findings suggest that semi-arid northern temperate grasslands remain relatively resistant to grazing effects, including their susceptibility to exotic plant encroachment. These results improve our understanding of how ongoing grazing exposure may impact grassland diversity, including efforts to conserve native vegetation, as well as the important role of climate in altering fundamental grassland responses to grazing.     

Site‐specific responses of native and exotic species to disturbances in a mesic grassland community

Abstract. Grassland communities are increasingly recognized as disturbance‐dependent ecosystems, yet there are few replicated, multi‐site studies documenting vegetation responses to varying frequencies and types of grassland disturbance. Even so, land managers frequently manipulate disturbance regimes in an attempt to favour native grassland plants over exotic species. We conducted a factorial experiment testing three frequencies of clipping combined with litter accumulation, litter removal, and soil disturbance within the highly threatened California coastal prairie plant community. We monitored the response of native/exotic, grass/forb plant guilds once a year for four years. More frequent clipping reduced cover of exotic grasses and favoured exotic forbs, whereas native species were largely unaffected by clipping frequency. Litter accumulation, litter removal, and soil disturbance did not affect vegetation composition. Effects of litter accumulation may take longer than our experiment allowed, and soil disturbance due to our treatments was not sufficiently strong to show consistent effects relative to mammalian soil disturbance. Treatment response of some plant guilds differed among sites, highlighting the importance of replicating experiments at several sites before recommending conservation management practices.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Long‐term changes in the vegetation after the cessation of livestock grazing in Eucalyptus marginata (jarrah) woodland remnants

This paper documents changes in the floristic composition of Eucalyptus marginata Donn (jarrah) woodlands over 7 years of recovery from continual, intensive livestock grazing. In remnants of native woodland left after agricultural clearing, which have been subjected to livestock grazing, comparisons were made between the floristics of fenced exclosure plots and open plots that continued to be grazed. The vegetation in nearby remnants, which had not been subjected to livestock grazing, was also surveyed. An initial increase in annual exotic pasture species after grazing relief was only temporary and highly influenced by fluctuations in annual climatic patterns, particularly rainfall distribution and abundance. Subsequent years saw a decrease in exotic annuals in exclosure plots and an increase in native perennials, in a trend towards becoming more floristically similar to the ungrazed sites. Germination of overstorey species was observed in the exclosure plots, however, development of seedlings and saplings was sparse. Results indicate that for jarrah woodland in southwestern Australia, natural regeneration is possible after the removal of livestock, with the return (within 6 years) of native species richness to levels similar to those found in ungrazed vegetation. Re‐establishment of cover, however, appears to take longer. The floristic dynamics are described in terms of a nonequilibrium model. Two vegetation states exist, degraded remnants with an understorey dominated by annual species, and ungrazed vegetation with an understorey dominated by perennial shrubs and herbs. The former state is maintained by continual heavy grazing by livestock. Upon relief from grazing, the vegetation undergoes a transition towards floristic similarity to ungrazed vegetation. After 6 years, vegetation change in the exclosure plots appears to be continuing and therefore it is still in transition.     

Floristic composition and diversity changes over 60 years in eastern mulga communities of south central Queensland,Australia

Botanical surveys from prior to 1949 documented the plant species composition of the mulga (Acacia aneura) communities in the eastern Mulga Lands Bioregion of Queensland, Australia. These surveys recorded 282 vascular plant species including 268 natives and 14 exotics. Since 1949, mulga communities have experienced considerable modification through continuous grazing by domestic sheep and cattle even during drought periods and extensive vegetation clearance. To increase our knowledge on floristic composition and diversity change over this time, floristic data were recollected from mulga communities in the same region between 2007 and 2010 using a more systematic procedure. These surveys recorded 292 vascular plant species including 269 natives and 23 exotics. Species richness and composition within five life forms: tree, shrub, vine, forb and graminoid were compared between the recent survey data and the historical survey data. The majority of native plant species were either in the forb (45.4% of total native species) or graminoid (25.7% of total native species) life forms over the 60-year period. There was no significant difference in species richness between the historical and recent surveys within tree or shrub life forms, but there were differences in species richness in vine, forb and graminoid life forms over time. Similarities in native species between the historical and recent surveys were high. Sorensen Similarity Indices (ISS) were: 0.77 at species level, 0.84 at genus level and 0.87 at family level. In contrast, the similarity indices for exotic species was low (0.59) for species and genera (0.61), but high for families (0.86). More exotic species were recorded recently than historically. Among these, buffel grass (Pennisetum ciliare) and colocynth (Citrullus colocynthis) were more commonly encountered than other exotic species in the eastern mulga communities of south central Queensland, Australia.     

Promoting Native Vegetation and Diversity in Exotic Annual Grass Infestations

Exotic plant invasions are especially problematic because reestablishment of native perennial vegetation is rarely successful. It may be more appropriate to treat exotic plant infestations that still have some remaining native vegetation. We evaluated this restoration strategy by measuring the effects of spring burning, fall burning, fall applied imazapic, spring burning with fall applied imazapic, and fall burning with fall applied imazapic on the exotic annual grass, medusahead (Taeniatherum caput‐medusae (L.) Nevski), and native vegetation at six sites in Oregon for 2 years post‐treatment. Medusahead infestations included in this study had some residual native perennial bunchgrasses and forbs. Burning followed by imazapic application provided the best control of medusahead and resulted in the greatest increases in native perennial vegetation. However, imazapic application decreased native annual forb cover the first year post‐treatment and density the first and second year post‐treatment. The spring burn followed by imazapic application produced an almost 2‐fold increase in plant species diversity compared to the control. The fall burn followed by imazapic application also increased diversity compared to the control. Results of this study indicate that native plants can be promoted in medusahead invasions; however, responses vary by plant functional group and treatment. Our results compared to previous research suggest that restoration of plant communities invaded by exotic annual grass may be more successful if efforts focus on areas with some residual native perennial vegetation. Thus, invasive plant infestations with some native vegetation remaining should receive priority for restoration efforts over near monocultures of invasive plant species.     

How environmental and disturbance factors influence species composition in temperate Australian grasslands

Abstract. A distinctive feature of Australian vegetational history is the abruptness of change since European settlement, involving the influx of exotic species and the imposition of exogenous disturbances which are novel in both intensity and character. This can produce two sources of habitat variability: the natural patterns arising from environmental variation, as well as an overlying effect of disturbance. The relative importance of these two types of variables were compared in temperate herbaceous vegetation. Canonical Correspondence Analysis showed that environment and disturbance had similar contributions to floristic variability. Individually, lithology, altitude and soil disturbance were the strongest variables while slope position, grazing and water enrichment were slightly less important. Despite generally low levels of site specificity, groups of species associated with lithology, slope position, altitude and different disturbance regimes were identified. Exotic species were associated with higher levels of disturbance, but showed levels of environmental specialization similar to the native component. Through combination of this analysis with a previous analysis of species richness for the same data set, it became evident that environmental variation mostly resulted in species substitutions while disturbances led to losses of species, with partial replacement by exotics. Synthesizing these results, we identified three broad groups in relation to tolerance of levels of exogenous disturbance: (1) intolerant species - native taxa intolerant of severe disturbances and constituting the species - rich component of the vegetation; (2) tolerant species - exotic and native taxa occurring at both disturbed and undisturbed habitats and (3) disturbance specialists - predominantly exotic species, correlated with high levels of disturbance.