Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.     

Restoration of Wet Dune Slacks on the Dutch Wadden Sea Islands: Recolonization After Large-Scale Sod Cutting

The effects of sod cutting were studied in a dune area on the Dutch Wadden Sea Island of Texel. Sod cutting was carried out in a range of different dune slacks in order to restore dune slack vegetation with many endangered Red List species. Sod cutting removed approximately 96% of the soil seed bank. Species abundant in the seed bank, notably Juncacea, also had a high frequency in the vegetation that established during the first year after the restoration measures. Many other species not registered in the seed bank or in the former vegetation also appeared. Species richness in the monitored plots exceeded that of uncut reference plots after a few years. Colonization rates were higher than extinction rates in most plots, indicating that a stable state has not been reached after 5 years. Differences in species richness between slacks appeared to be related to the occurrence of source areas nearby and availability of dispersal agents, such as flooding and animals.     

Effects on soil,microclimate and vegetation of the native-invasive <Emphasis Type="Italic">Retama monosperma</Emphasis> (L.) in coastal dunes

The aim of this study was to analyse and quantify the effects of the canopy of the native-invasive N-fixer woody shrub Retama monosperma in the dune ecosystem, affecting the structure and function of the dune environment as well as plant community, in the context of the facilitation mechanism. Air temperature and relative humidity; soil pH, electric conductivity, organic matter (OM) and nutrient content; above and below-ground vegetation biomass, litter mass, species richness and Shannon diversity were determined and compared from sampling plots below the R. monosperma canopy and in canopy gaps within a coastal dune system in SW Spain. The relationships between soil OM and nutrient contents and above and below-ground vegetation biomass, litter mass, species richness and Shannon diversity were also assessed. A predominance of positive interactions was confirmed. The canopy of R. monosperma ameliorated temperature extremes beneath, and soil OM and nutrient concentrations were increased by 188–466%, compared to those found in gaps. Plant biomass increased by 442% beneath the canopy and was composed almost exclusively of herbaceous annuals. Plant diversity was not affected. Plant communities were clearly structured as fertility islands, distributed in an environmentally stressful dune matrix characterized by scarce vegetation cover and low biomass.     

Environmental factors influencing herb layer productivity in Central European oak forests: insights from soil and biomass analyses and a phytometer experiment

Habitat productivity and vegetation biomass are important factors affecting species diversity and ecosystem function, but factors determining productivity are still insufficiently known, especially in the forest herb layer. These factors are difficult to identify because different methods often yield different results. We sampled the herb layer biomass and assessed soil nutrients, moisture and light availability in 100 m² plots in Czech oak forests. Habitat productivity was estimated independently from nutrient content in the soil, herb layer biomass and using a bioassay experiment (growing phytometer plants of Raphanus sativus under standardised conditions in soil samples taken from forest plots). The generalised linear model for herb layer biomass showed it to increase with light, soil phosphorus and moisture availability, but only 10.7% of its variation was explained by these factors. The phytometer biomass increased mainly with soil pH and phosphorus availability; together with soil C/N ratio these factors explained 56.1% of the phytometer biomass variability. Combined evidence based on different approaches indicates that canopy shading and soil phosphorus tend to be the most important factors influencing the herb layer productivity of the studied oak forests.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Sawdust Addition Reduces the Productivity of Nitrogen‐Enriched Mountain Grasslands

Anthropogenic nutrient enrichment of mountain grasslands has boosted grasses and fast‐growing unpalatable plants at the expense of slow‐growing species, resulting in a significant loss in biodiversity. A potential tool to reduce nutrient availability and aboveground productivity without destroying the perennial vegetation is carbon (C) addition. However, little is known about its suitability under severe climatic conditions. Here, we report the results of a 3‐year field study assessing the effects of sawdust addition on soil nutrients, aboveground productivity, and vegetational composition of 10 grazed and ungrazed mountain grasslands. Of particular interest was the effect of C addition on grasses and on the tall unpalatable weed Veratrum album. After 3 years, soil pH, ammonium, and plant‐available phosphorus were not altered by sawdust application, and nitrate concentrations were marginally higher in treatment plots. However, the biomass of grasses and forbs (without V. album) was 20–25% lower in sawdust‐amended plots, whereas the biomass of V. album was marginally higher. Sawdust addition reduced the cover of grasses but did not affect evenness, vegetation diversity, or plant species richness, although species richness generally increased with decreasing biomass at our sites. Our results suggest that sawdust addition is a potent tool to reduce within a relatively short time the aboveground productivity and grass cover in both grazed and ungrazed mountain grasslands as long as they are not dominated by tall unpalatable weeds. The technique has the advantage that it preserves the topsoil and the perennial soil seed bank.     

The effects of grazing and nutrient inputs on grass-heather competition

Summary

A combination of field and pot experiments was used to investigate the relative importance of grazing pressure and resource availability in grass-heather competition. In pots where nutrients were added, Nardus acquired the additional nutrients more rapidly than Calluna and increased its above-ground biomass to a greater extent. However, in the field, when fencing and fertilizer treatments were applied to a fine-scale mosaic of Calluna and grasses, including Nardus, nutrient addition only benefited Nardus in unfenced areas. In areas where sheep and deer were excluded, nitrogen addition increased heather cover; Nardus cover declined despite the added nutrients, possibly because the absence of grazing allowed heather to close its canopy above the Nardus, a shade-intolerant species. Effects were site dependent, with fencing increasing heather cover more rapidly on some plots than others; sites where grass cover was declining were the ones where grasses showed the smallest uptake of fertilizer. The results demonstrate that nitrogen additions well above the suggested critical load for heather moorland do not lead to a significant loss of heather cover if herbivory is absent, at least not within three years. It is suggested that the loss of heather moorlands to grass-dominated vegetation in Scotland may principally be driven by grazing, rather than nutrients.     

Impact of grazing and atmospheric nitrogen deposition on the vegetation of dry coastal dune grasslands

Abstract. A five-year experimental study was carried out to examine the combined effects of grazing and atmospheric nitrogen deposition on the vegetation of three dry dune grasslands: one short species-rich, one short species-poor, and one predominated by tall graminoids. Additional fertilization with nitrogen had no significant effect, neither in grazed nor in non-grazed plots. Exclusion of grazing by rabbits resulted in an increase in the frequency of perennial graminoids and a decrease in the frequency of annual graminoids and herbs. Nevertheless, species diversity remained the same in the species-rich grassland. During the experiment, the above-ground biomass increased in all nongrazed plots and the amount of bare soil and mosses decreased. The vegetation changes occurred mainly within one year after the exclusion of grazing. An exception is the grass-dominated site where the amount of Calamagrostis epigejos increased gradually from ca. 20 % in the first two years to about 50 % in the fourth and fifth year. Grazing by rabbits seems essential to prevent graminoids to become predominant in the dry dunes. If graminoids are dominant, grazing by horses can be an appropriate method to restore the original grassland vegetation. After six months of grazing by horses the grass-dominated site showed a decrease of the frequency of perennial graminoids, from 95 % to 80 %, and an increase of the frequency of perennial herbs, from 2.5 % to between 13 and 20 %.     

Propagule pressure and resource availability determine plant community invasibility in a temperate forest understorey

Few field experiments have examined the effects of both resource availability and propagule pressure on plant community invasibility. Two non-native forest species, a herb and a shrub ( Hesperis matronalis and Rhamnus cathartica , respectively), were sown into 60 1-m² sub-plots distributed across three plots. These contained reconstructed native plant communities in a replaced surface soil layer in a North American forest interior. Resource availability and propagule pressure were manipulated as follows: understorey light level (shaded/unshaded), nutrient availability (control/fertilized), and seed pressures of the two non-native species (control/low/high). Hesperis and Rhamnus cover and the above-ground biomass of Hesperis were significantly higher in shaded sub-plots and at greater propagule pressures. Similarly, the above-ground biomass of Rhamnus was significantly increased with propagule pressure, although this was a function of density. In contrast, of species that seeded into plots from the surrounding forest during the growing season, the non-native species had significantly greater cover in unshaded sub-plots. Plants in these unshaded sub-plots were significantly taller than plants in shaded sub-plots, suggesting a greater fitness. Total and non-native species richness varied significantly among plots indicating the importance of fine-scale dispersal patterns. None of the experimental treatments influenced native species. Since the forest seed bank in our study was colonized primarily by non-native ruderal species that dominated understorey vegetation, the management of invasions by non-native species in forest understoreys will have to address factors that influence light levels and dispersal pathways.     

Competition strength of two significant invasive species in coastal dunes

To investigate the effect of increased nutrient availability on competition amongst invasive and native plants, I measured changes in above and below ground biomass of Chrysanthemoides monilifera spp. rotundata (bitou bush) and Asparagus aethiopicus (asparagus fern) competing with two native species, Banksia integrifolia and Ficinia nodosa, under high- and low-nutrient regimes. Bitou bush, as a primary invader, was competitive under all conditions lowering the growth of native species in both high and low nutrients. Asparagus fern as a secondary invader, did not influence growth of native species but responded, like bitou bush, to high nutrients. Native species were generally negatively affected by increases in nutrients. With bitou bush soils often providing higher nutrients, the chance of secondary invasion by asparagus fern is more likely, although asparagus fern is unlikely to invade low nutrient soils quickly. The invasive species, therefore, differed in their competitive ability in these coastal dune communities.     

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.     

Effects of light and nutrient availability on dry matter and N allocation in six successional grassland species

Summary Recent discussions on determinants of competitive success during succession require the study of the combined effect of light and nutrient availability on growth and allocation. These effects can be used to predict the outcome of competition at changing resource availabilities. This work is part of a study on the successional sequence in permanent grassland starting after fertilizer application is stopped, but with continued mowing, in order to restore former species-rich communities. This yields a successional sequence which proceeds from grasslands with a high nutrient availability and a closed canopy, to grasslands with a low nutrient availability and an open canopy. If allocation is related to competitive ability, species from the productive stages would be expected to allocate more biomass and nitrogen to leaves, which could make them better competitors for light, while species from the unproductive stages would allocate more biomass to roots, which could make them better nutrient competitors. This study reports on growth, specific leaf area (SLA), vertical display of leaves, and allocation of biomass and nitrogen of six grassland species from this successional sequence at 16 combinations of light and nutrient supply. Species from the poorer successional stages reached a lower final dry weight than species from the richer stages, over all treatment combinations. The experimental design made it possible to test for unique effects of the resource ratio effect of light and nutrients on allocation characteristics. This resource-ratio effect was defined as the ratio light intensity/(light intensity + nutrient supply rate), using standardized levels for the treatments. The within-species variation (plasticity) in both allocation of dry matter and nitrogen was linearly related to this resource-ratio effect. Some interspecific differences in this relationship were found which could be related to the position of the species along the successional gradient. However, the range of plasticity in allocation pattern expressed within each species was much larger than the differences between species. It was concluded that allocation differences between these grassland species are relatively unimportant, given the large amount of plasticity in these traits. Interspecific differences in SLA and vertical stature seemed to be more important in explaining the position of species along the successional gradient.     

科尔沁沙地封育恢复过程中植物群落特征变化及影响因素

封育是退化沙地植被恢复与生态重建的重要措施, 理解长期处于封育状态下不同类型沙地植物群落特征变化及其影响因素有利于沙地植被恢复和生态重建。该文基于对科尔沁沙地长期封育的流动沙丘(2005年封育)、固定沙丘(1985年封育)和沙质草地(1997年封育)连续多年(2005-2017年)的植物群落调查, 结合土壤种子库、土壤养分以及气象数据, 分析了植物群落特征变化及其对环境变化的响应。研究结果表明流动沙丘植被盖度显著增加, 群落生物量和物种多样性年际间波动变化, 但无明显趋势; 固定沙丘植物群落存在逆行演替趋势, 具体表现为群落生物量、灌木和半灌木以及豆科优势度显著下降, 而一年生和多年生杂类草优势度显著增加; 沙质草地群落物种丰富度和多年生禾草优势度存在降低趋势, 并且一年生杂类草优势度明显高于其他功能群, 群落存在退化现象。3类沙地土壤种子密度变化不显著, 而种子丰富度在流动沙丘显著增加, 在固定沙丘和沙质草地有下降趋势, 土壤养分仅有有效氮和有效磷含量增加。回归分析结果表明气温和降水是影响年内生物量积累的主要因素, 但对年际间群落生物量和物种丰富度变化影响不大。除趋势对应分析结果显示土壤种子库与植物群落之间存在很高的相似性, 典型相关分析结果表明沙质草地植物群落与土壤养分紧密相关, 而固定沙丘群落主要与土壤水分紧密相关。综合以上结果可知, 封育33年的固定沙丘群落和封育21年的沙质草地群落都存在退化现象, 而封育11年的流动沙丘群落正在缓慢恢复, 因此封育年限的设定对退化沙地植被恢复至关重要, 封育时间过长不仅不利于植物群落恢复, 反而会使群落发生逆行演替, 建议封育年限的设定应综合考虑植被退化程度、土壤养分状况、土壤种子库基础以及气候条件等因素的影响。     

Wet dune slacks: decline and new opportunities

For a number of infiltrated coastal dune areas it is discussed to what extent artificial infiltration for the public water supply affects the quality of soil, groundwater and vegetation around pools and ponds, and what its effect is on the vegetation. Further, the results of investigations into the quality of vegetation, soil and water of a number of non-infiltrated, less affected dune areas are presented. The emphasis is on changes in groundwater flow pattern and on changes in the chemical composition of groundwater on the vegetation of wet dune slacks. Finally, recommendations for the management of wet dune slacks are presented. It can be concluded that the introduction of nutrients through infiltration causes an abundance of nitrophilous herbaceous vegetation along the banks of all infiltration ponds and most dune pools. Of the three investigated macro-nutrients, nitrate, potassium and phosphate, the latter shows the most significant correlation with the composition, cover and biomass of the vegetation. The moist biotopes of non-infiltrated dunes have largely disappeared because of desiccation, mainly as a consequence of water withdrawal, afforestation and coastal erosion. Relatively unaffected dune slacks can be found in the dunes on the Dutch Wadden Sea islands and a small number of dune areas on the mainland. In most areas, however, a serious decline in many rare species has been observed during the past twenty years because of eutrophic and acid precipitation, often in combination with disturbances of the groundwater regime.     

Density Effects of a Dominant Understory Herb,Isoglossa woodii (Acanthaceae), on Tree Seedlings of a Subtropical Coastal Dune Forest

Suppression of tree seedlings by the understory is an important ecological filter with implications for tree diversity and dynamics. In a greenhouse competition experiment, we used seedlings of four canopy species from coastal dune forest (Diospyros natalensis, Euclea racemosa, Sideroxylon inerme and Apodytes dimidiata) to examine the relative competitive effects of the dominant understory herb Isoglossa woodii on seedling performance. We manipulated I. woodii density, light and nutrient levels and measured growth responses. Total seedling biomass decreased with density of I. woodii. The magnitude of biomass suppression with competitor density was similar among tree species. Consequently there was no discernable hierarchy of competitive ranking among tree species. The relative growth rate of seedlings decreased at higher densities of I. woodii and increased at higher nutrient levels but was unaffected by variation in light conditions. Aboveground biomass decreased at higher densities of I. woodii and at higher light levels but increased at higher nutrient levels. Size asymmetric competition for light and nutrients may be the major driver of aboveground interactions between tree seedling and I. woodii. While tree species showed no hierarchy of competitive ability their seedlings exhibited equivalent responses to competition from an understory dominant, permitting species coexistence and the maintenance of species diversity.     

Light, nutrients and the growth of herbaceous forest species

The herb layer of forests planted on former agricultural land often differs from that of old-growth forest. This study investigates if the expected increased nutrient availability in the shaded conditions of newly planted forests and the plasticity of the species to adjust their biomass allocation to different levels of light and nutrients help to explain these differences in the herb layers of the two forest types. In a greenhouse experiment biomass distribution and production of two species characteristic for the highly shaded forest floor, Circaea lutetiana and Mercurialis perennis, and two species more common in the forest-edge, Aegopodium podagraria and Impatiens parviflora were studied at different levels of light (2%, 8% and 66% of the full light level) and nutrients (30 and 300 kg N ha^–1 per year). The main factor affecting allocation and biomass production was light availability. Nutrient supply only had a significant effect at the higher light levels. Species were mainly plastic to changes in light and the two species from the forest floor showed to be more rigid in allocation pattern than the species from the forest-edge. So, although the species from the forest-edge were more plastic, they did not profit from the increased nutrient supply because the main factor affecting biomass distribution and production was light availability.     

Aboveground nutrient cycling and forest development on poor sandy soil

The aboveground nutrient turnover of three ecosystems representing the main stages of heathland forest succession in NW Germany was investigated in a comparative study with regard to nutrient availability of the soil and light availability below the canopy. It was expected that nutrient availability would play a decisive role in forest development on nutrient poor acidic soil. The results show that the input of nutrients into the Calluna heathland is higher than the annual aboveground turnover of N, P, Ca, Mg, and K via litterfall. Compared to the pioneering birch-pine forest, the annual aboveground turnover of nutrients within the Calluna heathland, and therefore the nutrient availability is very low, while the light availability below the canopy is high. The increasing productivity of the growing successional forest is combined with an increasing nutrient turnover, mainly via litter fall. As a result, the increasing nutrient availability favours shade tolerant species with a higher nutrient demand at the late stages of succession. Consequently, the presumed terminal stage of succession, the oak-beech forest, is characterized by low light availability below the canopy and higher nutrient supply according to the resource-ratio hypothesis of Tilman (1985, 1986, 1988).     

Nutrient and Carbon Additions to the Microbial Soil Community and its Impact on Tree Seedlings in a Boreal Spruce Forest

We have added glucose and nutrients to manipulate soil microbial activity and nutrient availability in a boreal spruce forest to study the performance of birch and spruce seedlings in relation to the soil microbial community. The proportion of aboveground biomass in the seedlings was largest in plots amended with extra nutrients, while ectomycorrhizal (ECM) colonisation was low in these plots. ECM appeared beneficial for growth of both species, but only at low levels of colonisation (<25% ECM colonised root-tips). The soil microbial biomass, as determined by total PLFA, was largest in plots treated with glucose and there was a significant negative relationship between birch seedling size and levels of total PLFA in soil. This could be taken to suggest that poor seedling growth was due to nutrient limitation caused by microbial assimilation. However, the treatment response of the birch seedlings was generally weak, and spruce often showed no response at all to the addition of nutrients and glucose. The most consistent parameter for the variation in plant performance, as well as for the microbial soil community, was the block-effect. This suggests a strong spatial structure in the soil microbial community, and that this structure was robust with respect to our treatments even though they continued over a 3-year period.     

Nutrient status,disturbance and competition: an experimental test of relationships in a wet meadow

Abstract. In many species-rich hay meadows in Central Europe, the traditional extensive (low input, low output) management is no longer economical and meadows are either fertilized or abandoned. Both these practices lead to changes in species composition and usually to a loss in species diversity. The response of a species-rich meadow plant community to fertilization, mowing and removal of dominant species was studied in a manipulative factorial experiment over four years. Both species richness and seedling recruitment were positively influenced by mowing and to a lesser extent by removal of the dominant species, Molinia caerulea, and were negatively influenced by fertilization. Fertilization caused an immediate increase in community biomass. Response to removal of the dominant species was delayed by one season, continued over the whole period, and by the fourth year the biomass reached a similar value as in the plots with Molinia present. Changes in species composition were evaluated by RDA for repeated measurements. The best and only significant predictor of species response to fertilization was plant height. This shows that with increased nutrient availability, nutrient limitation weakens and competition for light becomes the decisive factor. Competition for light appears to be more asymmetric than competition for nutrients, and consequently, it is more likely to drive inferior species to extinction. This is, together with seedling recruitment limitations, the most important cause of a decrease in species richness under high nutrient levels.     

How Does Restoration of Native Canopy Affect Understory Vegetation Composition? Evidence from Riparian Communities of the Hunter Valley Australia

Restoration of native vegetation often focuses on the canopy layer species, with the assumption that regeneration of the understory elements will occur as a consequence. The goal of this study was to assess the influence of canopy restoration on the composition and abundance of understory plant species assemblages along riparian margins in the Hunter Valley, NSW, Australia. We compared the floristic composition (richness, abundance, and diversity) of understory species between nonrevegetated (open) and canopy revegetated plots across five sites. A number of other factors that may also influence understory vegetation, including soil nutrients, proximity to main channel, and light availability, were also measured. We found that sites where the canopy had been restored had lower exotic species richness and abundance, as well as higher native species cover, but not native species richness, compared with open sites. Multivariate analysis of plots based on plant community composition showed that revegetated sites were associated with lower total species diversity, light availability, and exotic cover. This study has found that the restoration of the canopy layer does result in lower exotic species richness and cover, and higher native species cover and diversity in the understory, a desirable restoration outcome. Our results provide evidence that restoration of native canopy species may facilitate restoration of native understory species; however, other interventions to increase native species richness of the understory should also be considered as part of management practice.