Ecosystem response to removal of exotic riparian shrubs and a transition to upland vegetation

Understanding plant community change over time is essential for managing important ecosystems such as riparian areas. This study analyzed historic vegetation using soil seed banks and the effects of riparian shrub removal treatments and channel incision on ecosystem and plant community dynamics in Canyon de Chelly National Monument, Arizona. We focused on how seeds, nutrients, and ground water influence the floristic composition of post-treatment vegetation and addressed three questions: (1) How does pre-treatment soil seed bank composition reflect post-treatment vegetation composition? (2) How does shrub removal affect post-treatment riparian vegetation composition, seed rain inputs, and ground water dynamics? and (3) Is available soil nitrogen increased near dead Russian olive plants following removal and does this influence post-treatment vegetation? We analyzed seed bank composition across the study area, analyzed differences in vegetation, ground water levels, and seed rain between control, cut-stump and whole-plant removal areas, and compared soil nitrogen and vegetation near removed Russian olive to areas lacking Russian olive. The soil seed bank contained more riparian plants, more native and fewer exotic plants than the extant vegetation. Both shrub removal methods decreased exotic plant cover, decreased tamarisk and Russian olive seed inputs, and increased native plant cover after 2 years. Neither method increased ground water levels. Soil near dead Russian olive trees indicated a short-term increase in soil nitrogen following plant removal but did not influence vegetation composition compared to areas without Russian olive. Following tamarisk and Russian olive removal, our study sites were colonized by upland plant species. Many western North American rivers have tamarisk and Russian olive on floodplains abandoned by channel incision, river regulation or both. Our results are widely applicable to sites where drying has occurred and vegetation establishment following shrub removal is likely to be by upland species.     

Plant–community responses to shrub cover in a páramo grassland released from grazing and burning

Shrub encroachment can follow grazing or burning release in páramo grasslands. While encroachment decreases herbaceous species richness in some grassland systems, the effects of this process on the herbaceous community in páramo grasslands are currently unknown. We collected data on shrub cover, herbaceous‐species cover and species composition in a páramo grassland 12 years after release from burning and cattle grazing near Zuleta, Ecuador. Topographic and soil measures were also included as predictor variables of differences in community composition. Contrary to studies in other systems, shrub cover did not have a significant effect on herbaceous‐species richness, whereas shrub‐species richness significantly increased with shrub cover. However, shrub cover was associated with significant shifts in herbaceous–community composition. Most notably, there was an increase in some shade‐tolerant forbs and tall‐statured wetland grasses with increasing shrub cover, and a corresponding decrease in some short‐statured grasses and early successional forbs. These results could indicate that the ameliorative effects of shrubs (e.g. frost and wind protection) in harsh alpine environments may partially compensate for the expected competitive effect of shrubs due to shading.     

Early environments drive diversity and floristic composition in Mediterranean old fields: Insights from a long-term experiment

While many studies have explored how previous and current environmental conditions affect the performance of individual organisms, their relative importance as drivers of current diversity and composition of communities is virtually unknown. We evaluated the response of herbaceous communities to previous (experienced during early establishment) and current environmental conditions by comparing their composition and diversity in an abandoned Mediterranean cropland planted with Quercus ilex subsp. ballota L. seedlings. These seedlings received different experimental treatments (summer irrigation and artificial shading) during the first three years after planting, and were interrupted from then on. We tested two complementary hypotheses: (i) the previous environments experienced by the herbaceous communities during their establishment have a long-term carry-over effect on diversity and composition of species assemblages and (ii) these communities are influenced by their current environment, particularly by the woody layer and the soil conditions. Overall, we observed an important contribution of initial environmental conditions in determining the current diversity and composition of herbaceous communities. Amelioration of environmental conditions, particularly water stress, during community establishment resulted in a decrease in alpha and beta diversity, possibly as a consequence of decreasing environmental heterogeneity. Previous environments accounted for 26.3% of the explained variance in current community composition. Annuals, legumes and forbs also responded significantly to previous environments, which explained 27.9%, 36.2% and 30.1%, respectively, of the variance in their composition. Our results suggest that those species present at a particular site early in succession pre-empt the site and influence vegetation dynamics on that site for a long time. This study provides important insights for understanding the mechanisms underlying the ecological effects of issues like cropland reforestation and woody vegetation encroachment.     

Vegetation dynamics and exotic plant invasion following high severity crown fire in a southern California conifer forest

Early post-fire vegetation dynamics following large, severe forest fires are largely unknown for the southern California mountains owing to historic fire suppression. Vegetation in 38 forest stands was surveyed (2004, 2005, and 2007) following the 2003 Cedar Fire in the Cuyamaca Mountains, Peninsular Ranges, San Diego County, California, USA. Each stand was sampled using four 10-m radius plots for the tree stratum, and 20 1-m² quadrats for shrub and herb strata. Changes in canopy cover by species, origin (native and exotic) and life form were analyzed. 2007 data were subjected to clustering to examine the divergence in species composition of the stands with time. Shrub cover increased from 3 to 31%, and exotic herbaceous cover increased from 3 to 40%. Cover of native annuals had increased from 2004 (17%) to 2005 (33%), but then dropped to 15% in 2007. Forty percent of the stands were dominated by the shrub species Ceanothus palmeri, and associated with higher pre-fire conifer cover and fire severity. More than 50% of the stands were dominated by exotic annuals and associated with lower fire severity and less steep slopes. The remaining stands (<10%) were dominated by chaparral shrubs and occurred on lower elevation, steep west-facing slopes. Species traits predict their dynamics following disturbance, as environmental conditions change. Establishment and increasing abundance of species dependent on dispersal to reach a site, including exotic and native herbaceous species, occurred in years 2–4. Differences among stands in species composition 4 years post-fire were associated with topographic and fire severity gradients.     

Increase in nonnative understorey vegetation cover after nonnative conifer removal and passive restoration

Nonnative conifers are widespread in the southern hemisphere, where their use as plantation species has led to adverse ecosystem impacts sometimes intensified by invasion. Mechanical removal is a common strategy used to reduce or eliminate the negative impacts of nonnative conifers, and encourage native regeneration. However, a variety of factors may preclude active ecological restoration following removal. As a result, passive restoration – unassisted natural vegetation regeneration – is common following conifer removal. We asked, ‘what is the response of understorey cover to removal of nonnative conifer stands followed by passive restoration?' We sampled understorey cover in three site types: two‐ to ten‐year‐old clearcuts, native forest and current plantations. We then grouped understorey species by origin (native/nonnative) and growth form, and compared proportion and per cent cover of these groups as well as of bare ground and litter between the three site types. For clearcuts, we also analysed the effect of time since clearcut on the studied variables. We found that clearcuts had a significantly higher average proportion of nonnative understorey vegetation cover than native forest sites, where nonnative vegetation was nearly absent. The understorey of clearcut sites also averaged more overall vegetation cover and more nonnative vegetation cover (in particular nonnative shrubs and herbaceous species) than either plantation or native forest sites. Notably, 99% of nonnative shrub cover in clearcuts was the invasive nonnative species Scotch broom (Cytisus scoparius). After ten years of passive recovery since clearcutting, the proportion of understorey vegetation cover that is native has not increased and remains far below the proportion observed in native forest sites. Reduced natural regeneration capacity of the native ecosystem, presence of invasive species in the surrounding landscape and land‐use legacies from plantation forestry may inhibit native vegetation recovery and benefit opportunistic invasives, limiting the effectiveness of passive restoration in this context. Abstract in Spanish is available with online material.     

Vegetation succession among and within structural layers following wildfire in managed forests

Question: How does vegetation develop during the initial period following severe wildfire in managed forests? Location: Southwestern Oregon, USA. Methods: In severely burned plantations, dynamics of (1) shrub, herbaceous, and cryptogam richness; (2) cover; (3) topographic, overstory, and site influences were characterized on two contrasting aspects 2 to 4 years following fire. Analysis of variance was used to examine change in structural layer richness and cover over time. Non‐metric multidimensional scaling, multi‐response permutation procedure, and indicator species analysis were used to evaluate changes in community composition over time. Results: Vegetation established rapidly following wildfire in burned plantations, following an initial floristics model of succession among structural layers. Succession within structural layers followed a combination of initial and relay floristic models. Succession occurred simultaneously within and among structural layers following wildfire, but at different rates and with different drivers. Stochastic (fire severity and site history) and deterministic (species life history traits, topography, and pre‐disturbance plant community) factors determined starting points of succession. Multiple successional trajectories were evident in early succession. Conclusions: Mixed conifer forests are resilient to interacting effects of natural and human‐caused disturbances. Predicting the development of vegetation communities following disturbances requires an understanding of the various successional components, such as succession among and within structural layers, and the fire regime. Succession among and within structural layers can follow different successional models and trajectories, occurs at different rates, and is affected by multiple interacting factors.     

Successional patterns on tropical inselbergs: A case study on the Nouragues inselberg (French Guiana)

A number of plant communities have been described on tropical inselbergs, known as hot spots of plant and animal biodiversity. However, few studies tried to question what drives seral processes in these harsh environments, submitted to natural hazards (violent storms, intense runoff and lightning strikes) which may destroy the vegetation cover and its accumulated organic matter. We analysed quantitative data from the granitic Nouragues inselberg (French Guiana) in order to discern how successional processes featured their variety. We showed that the transition from herbaceous carpets (bromeliaceous mats and grassy meadows) to woody vegetation (shrub thickets) was not conditioned by slope, but was truly successional. We also showed that there was a cycle of change in shrub thickets, reinitiated by the destruction of scrub vegetation by fire (lightnings), wood-destroying fungi and termites.     

Species composition and invasion in NW Argentinian secondary forests: Effects of land use history,environment and landscape

Abstract. Successional patches are a large component of forest ecosystems throughout the world and their vegetation composition is conditioned by multiple factors such as land use history, disturbances, environmental conditions and landscape context. We investigated the relative contribution of historical, environmental, biotic and spatial factors in determining vegetation composition and invasion by exotic species in secondary forest patches of Sierra de San Javier, Tucumán, Argentina. We estimated canopy cover for shrub, vine and tree species distributed over 51 patches with known land use history. We also recorded environmental, historical and spatial variables and used multivariate techniques to explore the relationship between forest composition and explanatory variables. Land use, time since abandonment, altitude, slope and cover of different strata were related to the vegetation pattern in the study site, and they were all significantly structured over space. Exotic species appeared to differ from natives in their response to explanatory variables. Overall, exotic species were dominant on the edges of young patches originated from herbaceous crops, but the total number of exotic species was related to the distance to urban areas and small farms identified as potential sources of exotic propagules. Vegetation composition of secondary forests in NW Argentina was related to historical and environmental factors, but spatial variables strongly influenced vegetation composition as well as the variation in explanatory variables.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Roller Chopping Effectively Reduces Shrub Cover and Density in Pine Flatwoods

Changes to fire regimes have resulted in excessive shrub growth and declines in the species rich herbaceous ground layer of pyric savanna and grassland systems worldwide, including the pine flatwoods of the Southern Coastal Plain of the United States. Prescribed burning and roller chopping during growing (April–October) and dormant (November–March) seasons are management practices promoted to reduce shrub invasion and increase herbaceous plant growth in flatwoods. However, relatively little is known about the seasonal effects these activities have on shrubs and herbaceous vegetation. We assessed the effects of prescribed burning and roller chopping on herbaceous and shrub characteristics in pine flatwoods and explored how grazing may mediate these treatments. We used a paired design, with comparison made between sampling locations randomly located within treated (e.g. burned) and adjacent untreated areas. Growing season burning was more effective at reducing shrub cover and height than dormant season burning. However, shrub re‐growth occurred the second year post‐burn. Roller chopping and roller chopping/burning combinations led to decreases in shrub cover and height for 2 years post‐treatment. Decreases in shrub density were seen on sites subject to growing season roller chopping and grazing. Decreases in herbaceous vegetation were observed following all treatments, possibly the result of grazing. If reductions in shrub density are required, growing season roller chopping in combination with grazing may be the only effective treatment. However, initial deferment from grazing following burning and roller chopping treatments may be necessary to permit re‐establishment and growth of forbs and graminoids.     

Plant population and community responses to removal of dominant species in the shortgrass steppe

Question: What are the plant population‐ and community‐level effects of removal of dominant plant species in the shortgrass steppe? Location: The Shortgrass Steppe Long‐Term Ecological Research site in northern Colorado, USA. Methods: We annually measured plant cover and density by species for 10 years after a one‐time aboveground removal of the dominant perennial grass, Bouteloua gracilis. Removal and control plots (3 m × 3 m) were within grazed and ungrazed locations to assess the influence of grazing on recovery dynamics. Our analyses examined plant species, functional type, and community responses to removal, paying special attention to the dynamics of subdominant and rare species. Results: Basal cover of B. gracilis increased by an average of 1% per year, but there was significantly less plant cover in treatment compared to control plots for 5 years following removal. In contrast to the lower cover in treatment plots, the plant density (number of plants m^?2) of certain subdominant perennial grasses, herbaceous perennial and annual forbs, a dwarf shrub, and cactus increased after removal of the dominant species, with no major change in species richness (number of species per 1 m × 1 m) or diversity. Subdominant species were more similar between years than rare species, but dominant removal resulted in significantly lower similarity of the subdominant species in the short term and increased the similarity of rare species in the long term. Conclusions: Removal of B. gracilis, the dominant perennial grass in the shortgrass steppe, increased the absolute density of subdominant plants, but caused little compensation of plant cover by other plants in the community and changes in species diversity.     

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.     

A multifaceted view on the impacts of shrub encroachment

Shrub encroachment, a global phenomenon with management implications, is examined in two papers in the current issue of Applied Vegetation Science. Barbosa da Silva et al. show that encroachment simplifies the herbaceous community, and Pittarello et al. illustrate how pastoral practices can restore encroached grasslands. While detrimental effects of shrub encroachment on grassland vegetation are often reported, we argue for a more holistic view when assessing this land‐cover change.     

Is prescribed fire a suitable management tool to reduce shrub encroachment in palm savannas?

Shrub encroachment in grasslands is a worldwide problem that has many ecological consequences, transforming previously open environments into dense forests. Disruption of natural fire regimes is one of the main causes of shrub encroachment, and the use of prescribed fire is a common strategy used to restore these ecosystems. In this study, we provide information about how a palm tree savanna under a process of shrub encroachment responds to the reintroduction of fire. We describe the effects of a first fire event on vegetation composition and structure using an experimental approach. We examine a species‐specific response to the fire. After one prescribed fire event applied to four study areas of 16 ha each, we analyzed the change in vegetation physiognomy and composition in burned and control plots for 1 year. Low‐intensity prescribed fire decreased height and cover of most shrub species and increased herbaceous vegetation cover over time. We classified shrub and herbaceous species response to fire according to the time they became present and their phenological characteristics. Our results can help stakeholders to determine if prescribed fire is helpful at reducing shrub encroachment in short term in similar ecosystems, considering how plant community responds to the reintroduction of fire after decades of fire suppression.     

川西亚高山针叶林土壤微生物及酶对林下植物去除的响应

生物多样性与生态系统功能的关系是生态学领域研究的热点与难点。但因受研究手段的限制,有关森林物种组成及其多样性变化对土壤微生物数量和酶活性影响的研究少有报道。采用人工去除灌草层的实验方法,研究了川西亚高山针叶林灌草层丧失对土壤微生物数量和酶活性的影响。结果表明:1)灌草层去除后,土壤细菌和真菌数量以CK(对照)RH(除草)RS(除灌),而土壤放线菌数量则以RHCKRS;2)灌草层去除后,土壤微生物群落构成发生改变,真菌比例有所下降;3)灌草层去除后,土壤酶活性随之发生变化,各种酶活性均以CKRHRS。表明林下灌草层去除,尤其是灌木层去除,导致土壤微生物数量下降、群落构成发生变化以及土壤酶活性下降,从而在一定程度上影响到森林生态系统的物质循环功能。     

Longleaf Pine (Pinus palustris P. Mill.) Restoration Using Herbicides: Overstory and Understory Vegetation Responses on a Coastal Plain Flatwoods Site in Florida,U.S.A.

A study was conducted on a Coastal Plain flatwoods site in Florida to determine the effects of common forestry herbicides on Longleaf pine seedling survival and growth and on the understory vegetation. Following removal of the overstory slash pine, five low‐rate herbicide treatments were applied over the top of planted Longleaf pine seedlings to provide short‐term understory vegetation control and accelerate seedling growth. The objective was to increase Longleaf pine growth by reducing the shrub competition while increasing the herbaceous ground cover. Despite causing reduction in seedling survival over the control treatment, imazapyr (0.21 ae kg/ha) resulted in the highest seedling growth (height and volume). The significant reduction of shrub cover, density, and height by imazapyr was believed to be responsible for the improved seedling growth in this treatment. Both hexazinone (0.56 ai kg/ha) and sulfometuron methyl (0.26 ai kg/ha) + hexazinone (0.56 ai kg/ha) treatments also reduced cover of Runner oak, a major shrub species, but the response was evident only 8 months after treatment. Although sulfometuron methyl (0.26 ai kg/ha) and sulfometuron methyl + hexazinone treatments did not result in any significant change in overall grass, forb, and shrub cover, both treatments resulted in greater Longleaf pine growth compared to the control. None of the herbicides significantly affected the major understory grasses and forbs. Overall, imazapyr provided the best desired results with significant increase in seedling growth and better control of shrub species with no significant effects on grass and other herbaceous species cover.     

Avian Response to Mechanical Aspen Restoration in Sierra Nevada Coniferous Forest

Using mechanical treatments to mimic natural disturbances is becoming a standard management and restoration approach. In the Sierra Nevada, as throughout much of western North America, much of aspen habitat is in poor health. Because of the high ecological value of healthy aspen, and its limited extent on the landscape, restoration to reverse the decline and improve stand health has become a management priority in the region. To evaluate the ecological effects of mechanically removing competing conifers to restore aspen in the Sierra Nevada, we compared vegetation characteristics and bird abundance in treated and untreated aspen stands on the Lassen National Forest before and up to 13 years after mechanical conifer removal. Treatments reduced total canopy cover and increased herbaceous cover and the number of aspen stems, while shrub and overstory aspen covers were unchanged. Of 10 aspen focal bird species, 7 increased in abundance following treatment relative to controls, including all species associated with early seral aspen habitat and cavity nesting species; none declined. In contrast, of the six conifer focal species, the four associated with denser conifer habitat declined as a result of the treatments. The two species associated with conifer edges and understory cover increased. Our results demonstrate mechanical conifer removal treatments can provide ecologically meaningful changes in habitat for the avian community and are an effective tool for restoring ecological values of degraded aspen habitat for birds in the Sierra Nevada.     

Vegetation mapping using hierarchical object‐based image analysis applied to aerial imagery and lidar data

Aims: The primary objective of this study is to map the distribution and quantify the cover of vegetation alliances over the entirety of San Clemente Island (SCI). To this end, we develop and evaluate the mapping method of hierarchical object‐based classification with a rule‐based expert system. Location: San Clemente Island, California, USA. Methods: We developed and tested an approach based on hierarchical object‐based classification with a rule‐based expert system to effectively map vegetation communities on SCI following the Manual of California Vegetation classification system. In this mapping approach, the shrub species defining each vegetation community and non‐shrub growth forms were first mapped using aerial imagery and lidar data, then used as input in an automated mapping rule set that incorporates the percent cover rules of a field‐based mapping rule set. Results: The final vegetation map portrays the distribution of 19 vegetation communities across SCI, with the largest areas comprised of California Annual and Perennial Grassland (35%) and three types of coastal sage scrub and maritime succulent scrub, comprising a combined 53% of the area. Map accuracy was assessed to be 79% based on fuzzy methods and 61% with a traditional accuracy assessment. The accuracy of tree identification was assessed to be 81%, but species‐level tree accuracy was 45%. Conclusions: Semi‐automated approaches to vegetation community mapping can produce repeatable maps over large spatial extents that facilitate ecological management efforts. However, some low‐statured shrub community types were difficult to differentiate due to patchy canopies of co‐occurring species including abundant non‐native grasses characteristic of complex disturbance histories. Species‐level tree mapping accuracy was low due to the difficulty of identifying species within poorly illuminated canyons, resulting from sub‐optimal image acquisition timing.     

Mechanical Land Clearing to Promote Establishment of Coastal Sandplain Grassland and Shrubland Communities

The decline in grasslands and other species‐rich early successional habitats on the coastal sandplains of the northeastern United States has spurred management to increase the area of these declining plant communities. We mechanically removed overstory oak and applied seed from a nearby sandplain grassland on the island of Martha’s Vineyard, Massachusetts, to evaluate this technique for creating an open oak community able to support sandplain herbaceous species. We compared vegetation structure and composition before and after clearing in an area of total tree removal (clearcutting), an area where 85% of tree basal area was removed (savanna cutting), and in adjacent coastal oak forest. Plant responses to clearcutting and savanna cutting were similar. Sandplain herbs colonized at high frequencies after seeding and increased herbaceous cover from less than 7% before clearing to 22–38% three growing seasons later. Pennsylvania sedge (Carex pensylvanica) increased in cover approximately 6‐fold, accounting for 84–90% of the increased herbaceous cover. Other native ruderals and exotic herbs reached 2 and less than or equal to 1%, cover, respectively, after three years. Species richness across cleared treatments increased from 30 to 79 species. All forest species were retained. Forest shrubs and trees initially declined from their dominant cover but rebounded after three years. Tree clearing plus seeding appeared to be a viable management practice for increasing cover of herbaceous sandplain species while causing minimal increases in exotic herbaceous cover. The long‐term persistence of sandplain herbs may require periodic disturbances that limit woody regrowth.