Community changes following shrub invasion of grassland

Summary We studied the development of the shrub Baccharis pilularis ssp consanguinea and its effects on herbs of the annual grassland in Northern California. A series of shands of Baccharis was sampled of ages ranging from 1 yr to>9 yr, representing most of the life cycle of the shrub. In each stand we examined shrub biomass, structure and litterfall. We also determined cover and biomass of all herbaceous species and estimated seed production, seed rain and storage of seed in the soil. Abundances of all herbaceous species declined greatly after Baccharis formed a closed canopy at 2–3 yr, and little seed of herbaceous species was either dispersed into shrub stands or stored in the soil. Exclosures suggested that herbivory by small mammals in the closed shrub stands may be important in reducing the abundance of herbaceous species following shrub invasion of grassland.     

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.     

Effects of invasive plants on fire regimes and postfire vegetation diversity in an arid ecosystem

We assessed the impacts of co‐occurring invasive plant species on fire regimes and postfire native communities in the Mojave Desert, western USA. We analyzed the distribution and co‐occurrence patterns of three invasive annual grasses (Bromus rubens, Bromus tectorum, and Schismus spp.) known to alter fuel conditions and community structure, and an invasive forb (Erodium cicutarium) which dominates postfire sites. We developed species distribution models (SDMs) for each of the four taxa and analyzed field plot data to assess the relationship between invasives and fire frequency, years postfire, and the impacts on postfire native herbaceous diversity. Most of the Mojave Desert is highly suitable for at least one of the four invasive species, and 76% of the ecoregion is predicted to have high or very high suitability for the joint occurrence of B. rubens and B. tectorum and 42% high or very high suitability for the joint occurrence of the two Bromus species and E. cicutarium. Analysis of cover from plot data indicated two or more of the species occurred in 77% of the plots, with their cover doubling with each additional species. We found invasive cover in burned plots increased for the first 20 years postfire and recorded two to five times more cover in burned than unburned plots. Analysis also indicated that native species diversity and evenness as negatively associated with higher levels of relative cover of the four invasive taxa. Our findings revealed overlapping distributions of the four invasives; a strong relationship between the invasives and fire frequency; and significant negative impacts of invasives on native herbaceous diversity in the Mojave. This suggests predicting the distributions of co‐occurring invasive species, especially transformer species, will provide a better understanding of where native‐dominated communities are most vulnerable to transformations following fire or other disturbances.     

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.     

Effects of Prescribed Burning on Herbaceous and Woody Vegetation in Northern Lowland Meadows

Abstract Spring burning of sedge‐grass meadows in the Slave River Lowlands (SRL), Northwest Territories, Canada was applied between 1992 and 1998 to reduce shrub encroachment and enhance Bison bison (bison) habitat, although the impact of fire on preferred bison forage was unknown before management. In the summer of 1998 we conducted a study in the Hook Lake area of the SRL to test the effect of burn frequency (unburned, burned once, or burned three times since 1992) on herbaceous plant community composition and Salix spp. L. (willow) shrub vigor. Plant species abundance, litter biomass, soil pH, and depth of the organic soil horizon were measured in 300 1‐m² quadrats nested within 30 1,000‐m² plots in both burned and unburned dry meadows. To test the relationship between frequency and willow vigor, all willow shrubs within the plots were assigned a vigor score from I (dead) to IV (flourishing). The spring burns appear to have reduced willow vigor; however, shrub survival remained high (76%) on the most frequently burned meadows. Ordination plots resulting from canonical correspondence analysis suggest that multiple spring burns influenced plant community composition in dry meadow areas and that less palatable bison forage species (e.g., Carex aenea Fern. and Juncus balticus L.) were correlated with a regime of three spring burns. Our results suggest that frequent spring fires in the Hook Lake area have only a small negative effect on willow cover but may reduce the abundance of primary bison forage plants compared with less frequently burned meadows.     

Short‐term effects of herbicides and a prescribed fire on restoration of a shrub‐encroached pine savanna

Shrub encroachment occurring worldwide in savannas and grasslands has commonly been hypothesized to result from anthropogenically altered environments. Two disturbance‐based approaches to restoration have involved: (1) application of selective herbicides to reduce density/cover of shrubs; (2) reinstatement of natural fire regimes to generate environmental conditions favoring herbaceous species. We studied short‐term responses of native shrubs, vines, and grasses in a Louisiana pine savanna to herbicides followed by a prescribed fire and fire alone. Plots established in the summer, 2013, were hand‐sprayed in the fall with Imazapyr and Triclopyr, Triclopyr alone, or no herbicide, then prescribed burned the following spring. Numbers of species of shrubs and vines at scales of 1 and 100 m², numbers of stems and regrowth of stems produced by six common species of shrubs, and the number of flowering culms of perennial C₄ grasses were assessed postfire in 2014. Compared with fire alone, herbicides followed by fire resulted in (1) small reductions in species richness of shrubs and no effects on vines, (2) fewer stems comprising shrub genets, but similar postfire regrowth of resprouting shrub stems, and (3) fewer flowering culms of C₄ grasses. Underground storage organs of savanna shrubs and vines survived both aboveground disturbances. Thus, single applications of herbicides followed by fires reduced, but did not reverse shrub encroachment and negatively affected grasses. Because effects of herbicides overrode those of prescribed fires, these disturbances did not act synergistically, suggesting that reinstating natural fire regimes should be a priority in restoration of savannas and grasslands.     

Gopher Tortoise Response to Habitat Management by Prescribed Burning

ABSTRACT As quality of forested habitat declines from altered fire regimes, gopher tortoises (Gopherus polyphemus) often move into ruderal areas to the detriment of the animal and land manager. We evaluated effects of a dormant-and-growing-season prescribed fire on habitat and gopher tortoise use of degraded longleaf pine (Pinus palustris) forests surrounding military training areas. We burned 4 of 8 sites in winter 2001–2002 and again in April 2003. Changes in vegetation measured during 2001–2004 indicated that burn treatments did not increase herbaceous vegetation. Similarly, movement patterns, burrow usage, and home range of tortoises radiotracked from 2002–2004 did not differ between treatments. Woody cover initially was reduced in the forests postburn, and we found more new burrows in burned forest sites. Once shrub cover was reduced, tortoises started using forested habitat that had become overgrown. However, shrub reduction may be temporary, as woody stem densities increased postburn. Thus, the one-time use of fire to manage tortoise habitat may not rapidly restore the open canopy, sparse woody midstory, and abundant herbaceous vegetation that this species requires. Repeated prescribed fires or additional management techniques may be needed for complete restoration.     

Barriers to shrub reestablishment following fire in the seasonal submontane zone of Hawai'i

Introduced grass species have invaded extensive areas of Hawaii Volcanoes National Park and increased the size and frequency of fire. Following fire, grass cover is enhanced while native shrub cover is reduced; the reduction in most shrubs persists for at least 20 years even in the absence of fire. Shrub seedlings were planted in burned and unburned plots with and without grass cover. Biomass of 14 month old shrub seedlings was generally highest in recently burned/grass removed plots, intermediate in old burn/grass removed plots, and lowest in unburned/grass removed plots. In contrast, shrub biomass in plots with grass cover was low and did not differ significantly among burn treatments. Light competition is likely to be responsible for differences in shrub growth rates; grass cover reduced light to 1–10% of background levels. In addition, pool sizes of available soil N were highest in recently burned, intermediate in old burn, and lowest in unburned areas.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Effects of Prescribed Fire on Shinnery Oak ( Quercus havardii) Plant Communities in Western Oklahoma

Changes in structural and compositional attributes of shinnery oak (Quercus havardii Rydb.) plant communities have occurred in the twentieth century. These changes may in part relate to altered fire regimes. Our objective was to document effects of prescribed fire in fall (October), winter (February), and spring (April) on plant composition. Three study sites were located in western Oklahoma; each contained 12, 60 × 30‐m plots that were designated, within site, to be seasonally burned, annually burned, or left unburned. Growing season canopy cover for herbaceous and woody species was estimated in 1997–1998 (post‐treatment). At one year post‐fire, burning in any season reduced shrub cover, and spring burns reduced cover most. Winter and annual fires increased cover of rhizomatous tallgrasses, whereas burning in any season decreased little bluestem cover. Perennial forbs increased with fall and winter fire. Shrub stem density increased with fire in any season. Communities returned rapidly to pre‐burn composition with increasing time since fire. Fire effects on herbaceous vegetation appear to be manifested through increases in bare ground and reduction of overstory shrub dominance. Prescribed fire can be used as a tool in restoration efforts to increase or maintain within and between community plant diversity. Our data suggest that some plant species may require or benefit from fire in specific seasons. Additional research is needed to determine the long‐term effects of repeated fire over time.     

Seed dispersal by cattle may cause shrub encroachment of Grewia flava on southern Kalahari rangelands

Abstract. Shrub encroachment, i.e. the increase in woody plant cover, is a major concern for livestock farming in southern Kalahari savannas. We developed a grid‐based computer model simulating the population dynamics of Grewia flava, a common, fleshy‐fruited encroaching shrub. In the absence of large herbivores, seeds of Grewia are largely deposited in the sub‐canopy of Acacia erioloba. Cattle negate this dispersal limitation by browsing on the foliage of Grewia and dispersing seeds into the grassland matrix. In this study we first show that model predictions of Grewia cover dynamics are realistic by comparing model output with shrub cover estimates obtained from a time series of aerial photographs. Subsequently, we apply a realistic range of intensity of cattle‐induced seed dispersal combined with potential precipitation and fire scenarios. Based on the simulation results we suggest that cattle may facilitate shrub encroachment of Grewia. The results show that the severity of shrub encroachment is governed by the intensity of seed dispersal. For a high seed dispersal intensity without fire (equivalent to a high stocking rate) the model predicts 56% shrub cover and 85% cell cover after 100 yr. With fire both recruitment and shrub cover are reduced, which may, under moderate intensities, prevent shrub encroachment. Climate change scenarios with two‐fold higher frequencies of drought and wet years intensified shrub encroachment rates, although long‐term mean of precipitation remained constant. As a management recommendation we suggest that shrub encroachment on rangelands may be counteracted by frequent fires and controlling cattle movements to areas with a high proportion of fruiting Grewia shrubs.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Return of Fire as a Restoration Tool: Long‐Term Effects of Burn Severity on Habitat Use by Mexican Fox Squirrels

After decades of suppression, fire is returning to forests of the western United States through wildfires and prescribed burns. These fires may aid restoration of vegetation structure and processes, which could improve conditions for wildlife species and reduce severe wildfire risk. Understanding response of wildlife species to fires is essential to forest restoration because contemporary fires may not have the same effects as historical fires. Recent fires in the Chiricahua Mountains of southeastern Arizona provided opportunity to investigate long‐term effects of burn severity on habitat selection of a native wildlife species. We surveyed burned forest for squirrel feeding sign and related vegetation characteristics to frequency of feeding sign occurrence. We used radio‐telemetry within fire‐influenced forest to determine home ranges of Mexican fox squirrels, Sciurus nayaritensis chiricahuae, and compared vegetation characteristics within home ranges to random areas available to squirrels throughout burned conifer forest. Squirrels fed in forest with open understory and closed canopy cover. Vegetation within home ranges was characterized by lower understory density, consistent with the effects of low‐severity fire, and larger trees than random locations. Our results suggest that return of low‐severity fire can help restore habitat for Mexican fox squirrels and other native wildlife species with similar habitat affiliations in forests with a historical regime of frequent, low‐severity fire. Our study contributes to an understanding of the role and impact of fire in forest ecosystems and the implications for forest restoration as fire returns to the region.     

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract. In the mesic grasslands of the central United States, the shrub Cornus drummondii has undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused by C. drummondii expansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C₄ grass Andropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency in A. gerardii and lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain in A. gerardii in the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state of C. drummondii dominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect of C. drummondii dominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability. C. drummondii removal facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.     

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.     

Fire Season Effects on Flowering Characteristics and Germination of Longleaf Pine (Pinus palustris) Savanna Grasses

Pinus palustris (longleaf pine) savannas depend on contiguous grass cover to facilitate frequent surface fires. Enhanced flowering, seed production, and germination may be linked to season of fires. We assessed the effect of month of prescribed fire (February, April, May, and July) on percentage of plants with flowering culms (FCs%), FC/area, FC/plant, seed production, and germination for five warm seasons, fall‐flowering grasses. Multivariate analysis indicated the response of flowering and fruiting to burn month varied among the grass species. The dominant species, Sporobolus junceus and Schizachyrium scoparium var. stoloniferum, had greater numbers of most flowering characteristics when burnt during April, May, and July. Aristida purpurascens had increased FC/plant after May and July burns. In contrast, Ar. mohrii had the fewest FC/plant and seeds/FC when burnt in July. Germination was greatest (26–60%) for Ar. purpurascens. Seeds collected following July burns for Ar. purpurascens and Ar. ternarius and after May burns for S. junceus were within the highest germination values recorded. Germination of Sc. scoparium var. stoloniferum was very low after February and July burns (≤5%). With Ar. mohrii, only seed collected following February (2%) and April burns (3%) germinated. April, May, and July fires increased seed production of dominant matrix grasses, thus facilitating the potential for recruitment of these species and facilitating seed collection from potential donor sites for ground‐layer restoration projects. Varying prescribed fire burn month captured variation in flowering characteristics among these grasses.     

Uprooting and Burial of Invasive Alien Plants: A New Tool in Coastal Restoration?

Invasive alien plants are a problem for conservation management, and control of these species can be combined with habitat restoration. Subsoil burial of uprooted plants is a new method of mechanical control, which might be suitable in disturbed habitats. The method was tested in Rosa rugosa (Japanese Rose), an invasive shrub in north‐western Europe with negative effects on coastal biodiversity. Two months after uprooting and burial in dunes of north‐eastern Denmark, 89% of the 58 shrubs resprouted from roots and rhizomes; on average 41 resprouts per shrub. Resprout density was twice as high at former shrub margins compared with the center; resprouts were taller and originated from more superficial soil layers at the margin than in the center. Resprouting was negatively correlated with fragment depth, and no resprouts were observed from greater than 15 cm depth. The number of resprouts increased with fragment dry mass (0.5–168.5 g). After 18 months with harrowing the species was still resprouting, flowering, and fruiting, albeit with no difference between shrub margin and center. Resprouts were taller (26 cm) and coverage was higher (0–4%) after two compared with three times harrowing, whereas no difference was found in cover of native dune species (1–5%). The results show that even small fragments of R. rugosa resprout, and that resprouting persists despite repeated harrowing. Thus, careful subsoil burial of all fragments is necessary, special attention should be paid to the shrub margin, and follow‐up treatments are needed. The effectiveness of the burial method is discussed for restoration of coastal dunes.     

Effects of prescribed burning on avian nest survival in the southern Great Plains

Shrubs, such as mesquite (Prosopis spp.) and cholla (Opuntia spp.), now dominate fire-suppressed grasslands in southwestern North America. Responses of birds to prescribed burning of the shortgrass prairie in this region are poorly understood. We examined daily survival rates of mourning dove (Zenaida macroura) and lark sparrow (Chondestes grammacus) nests in an experimental landscape (4,811 ha) of spatially replicated, inter-annual fire frequencies (burning every 2 yr, 4 yr, or 10 yr) near Amarillo, Texas. Herbaceous habitat structure was most developed in infrequently burned plots, but shrub densities were less variable among the burn treatments. We modeled daily nest survival (DSR) against burn frequency, shrub density at nest sites, and nest stage (incubation or nestling). Daily survival of mourning dove nests was not well-related to any measured covariate, but lark sparrow DSR was negatively related to the duration of inter-annual burn frequency. In semiarid grasslands heavily inundated with shrubs, prescribed burning may positively influence the nest success of some bird species. © 2011 The Wildlife Society.