Postwildfire seeding to restore native vegetation and limit exotic annuals: an evaluation in juniper‐dominated sagebrush steppe

Reestablishment of perennial vegetation is often needed after wildfires to limit exotic species and restore ecosystem services. However, there is a growing body of evidence that questions if seeding after wildfires increases perennial vegetation and reduces exotic plants. The concern that seeding may not meet restoration goals is even more prevalent when native perennial vegetation is seeded after fire. We evaluated vegetation cover and density responses to broadcast seeding native perennial grasses and mountain big sagebrush (Artemisia tridentata Nutt. spp. vaseyana [Rydb.] Beetle) after wildfires in the western United States in six juniper (Juniperus occidentalis ssp. occidentalis Hook)‐dominated mountain big sagebrush communities for 3 years postfire. Seeding native perennial species compared to not seeding increased perennial grass and sagebrush cover and density. Perennial grass cover was 4.3 times greater in seeded compared to nonseeded areas. Sagebrush cover averaged 24 and less than 0.1% in seeded and nonseeded areas at the conclusion of the study, respectively. Seeding perennial species reduced exotic annual grass and annual forb cover and density. Exotic annual grass cover was 8.6 times greater in nonseeded compared to seeded areas 3 years postfire. Exotic annual grass cover increased over time in nonseeded areas but decreased in seeded areas by the third‐year postfire. Seeded areas were perennial‐dominated and nonseeded areas were annual‐dominated at the end of the study. Establishing perennial vegetation may be critical after wildfires in juniper‐dominated sagebrush steppe to prevent the development of annual‐dominated communities. Postwildfire seeding increased perennial vegetation and reduced exotic plants and justifies its use.     

Are two strategies better than one? Manipulation of seed density and soil community in an experimental prairie restoration

Restoration practitioners have a variety of practices to choose from when designing a restoration, and different strategies may address different goals. Knowledge of how to best use multiple strategies could improve restoration outcomes. Here, we examine two commonly suggested strategies in a single tallgrass prairie restoration experiment: increased forb sowing density and prairie soil inoculation. We designed a study with two different forb seeding densities. Within these densities, we transplanted seedlings into 1‐m² plots that had been grown in either a whole prairie soil inoculum or sterilized prairie soil. After 4 years, we found positive effects of both high forb sowing density and inoculation treatments on the ratio of seeded to nonseeded plant cover in these plots, and negative effects of both treatments on nonseeded plant diversity. No effects of either treatment were seen on seeded plant diversity. Each strategy also affected the plant community in different ways: high forb sowing density increased seeded forb richness and decreased native nonseeded plant cover, while inoculation decreased non‐native cover, and tended to increase average successional stage of the community. These effects on restoration outcome were typically independent of each other, with the result that plots with both manipulations had the most positive effects on restoration outcomes. We thus advocate the combined use of these restoration strategies, and further studies which focus on both seeding and soil community manipulation in restoration.     

Fire severity and seedling establishment in Pinus halepensis woodlands, eastern Iberian Peninsula

Given the observed heterogeneity in fire severity produced within wildfires, we asked to what extent this heterogeneity might affect post-fire regeneration. For this purpose, we studied the post-fire dynamics of Pinus halepensis (Aleppo pine) in the eastern Iberian Peninsula. Sampling was stratified on the basis of fire severity. We defined three fire severity classes based on the degree of consumption of the pine canopy. The results suggested that there is no clear relationship between seedling density and fire severity; however, mortality was lower and growth (height, shoot biomass and root biomass) was higher in the high severity class. These results can be explained by soil processes: Sites in the high fire severity class may have sustained higher fire intensities, resulting in higher soil organic matter mineralisation and higher ash deposition, and thus in higher post-fire soil fertility. This higher fertility would produce faster growth in pine seedlings. Independent of the severity class, seedling mortality was higher in quadrats (50 × 50 cm) with higher cover of the perennial grass Brachypodium retusum (Poaceae), suggesting a possible competitive effect. For all plots in all 3 severity classes, spatial analysis suggests an aggregate seedlings pattern, but with independence from the position of the adult (source) trees. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Determinants of coastal treeline and the role of abiotic and biotic interactions

Recent die-off of coastal forests has been attributed primarily to the effects of sea level rise by correlation with tide-gage records. Due to the temporal and spatial scales involved, direct monitoring of sea level rise impacts is challenging and its attribution can be confounded by both land-use history and species interactions. Here we present experimental evidence for a micro-tidal, oligohaline estuarine system that the location of coastal treeline is determined by both environmental controls and positive and negative species interactions. We conducted field surveys and a transplant experiment to determine the controls on pine seedling establishment and survival along a salinity and flooding gradient. Using a two-way changepoint model we determined that sawgrass cover (Cladium jamaicense) and salinity concentrations interact to control natural pine regeneration (Pinus taeda). The long-term removal of sawgrass resulted in increased soil salinity and high rates of (planted) pine seedling mortality. In contrast, pine seedlings planted directly under sawgrass were able to survive at the same level as upland forest plots because of reduced salinity levels. This research provides evidence that sawgrass can facilitate pine seedling survival, but also suggests that either competitive exclusion by sawgrass or dispersal limitations prevent initial pine seedling regeneration. We propose that forest dynamics are closely coupled to fire, which consumes sawgrass biomass and reduces competition between pine seedlings and grass. Following fire, pine seedling establishment and the regrowth of sawgrass facilitates long-term pine seedling survival. Under this scenario, recent marsh invasion into coastal forests may not necessarily represent a permanent state change in locations where abiotic stress is not the only determinant for community composition.     

Seeding tallgrass prairie in monospecific patches promotes native species establishment and cover

In tallgrass prairie reconstruction, the way desired seeds are arranged on the landscape may affect species establishment, species persistence, and the establishment and persistence of undesired (nonseeded) species from the local propagule pool. To test effects of species seeding pattern on how grasslands develop spatially, we seeded 20—4 × 4–m bare soil plots with 16 tallgrass prairie species. Treatment plots were divided into 16—1 × 1–m subplots, 64—0.5 × 0.5–m subplots, 256—0.25 × 0.25–m subplots, or 1,024—0.125 × 0.125–m subplots. Each species was hand broadcast into separate subplots (1 m² total area/species) within each plot. An additional treatment included uniformly mixing and broadcasting all seeds across a plot. We recorded species cover at the 0.125 × 0.125–m scale within each plot at the beginning of the second and third growing seasons. While species persistence was greatest within plots seeded with larger subplots, plots with smaller subplots were more spatially diverse and less occupied by nonseeded species over time than larger subplot and mixed plots. As is common in reconstruction efforts, establishment was variable among species and seeding with monospecific subplots enhanced colonization of desired rhizomatous species (e.g., Heliopsis helianthoides, Monarda fistulosa, Elymus virginicus) into unoccupied locations at the expense of species from the local propagule pool. Results suggest that seeding species in smaller, monospecific patches could result in grasslands with a more balanced native species composition than those established with a seed mixture approach.     

Invasion resistance and persistence: established plants win,even with disturbance and high propagule pressure

Disturbances and propagule pressure are key mechanisms in plant community resistance to invasion, as well as persistence of invasions. Few studies, however, have experimentally tested the interaction of these two mechanisms. We initiated a study in a southwestern ponderosa pine (Pinus ponderosa Laws.)/bunch grass system to determine the susceptibility of remnant native plant communities to cheatgrass (Bromus tectorum L.) invasion, and persistence of cheatgrass in invaded areas. We used a 2 × 2 factorial design consisting of two levels of aboveground biomass removal and two levels of reciprocal seeding. We seeded cheatgrass seeds in native plots and a native seed mixture in cheatgrass plots. Two biomass removal disturbances and sowing seeds over 3 years did not reverse cheatgrass dominance in invaded plots or native grass dominance in non-invaded native plots. Our results suggest that two factors dictated the persistence of the resident communities. First, bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey) was the dominant native herbaceous species on the study site. This species is typically a poor competitor with cheatgrass as a seedling, but is a strong competitor when mature. Second, differences in pretreatment levels of plant-available soil nitrogen and phosphorus may have favored the dominant species in each community. Annual species typically require higher levels of plant-available soil nutrients than perennial plants. This trend was observed in the annual cheatgrass community and perennial native community. Our study shows that established plants and soil properties can buffer the influences of disturbance and elevated propagule pressure on cheatgrass invasion.     

Endophytes inconsistently affect plant communities across Schedonorus arundinaceus hosts

Fungal endophytes in cool-season grasses may affect communities at multiple trophic levels. However, it is unclear whether community-scale endophyte effects arise due to the endophyte itself or as a result of unique, endophyte–host interactions. We used a long-term field experiment to test whether common-toxic (CT) and non-ergot alkaloid-producing (novel) endophytes in Schedonorus arundinaceus (tall fescue) forage cultivars consistently affect communities across tall fescue hosts. Tilled plots (2 × 2 m; Guelph, ON) were seeded with Georgia 5 and Jesup cultivars containing either the CT or AR542 (novel) endophyte and allowed to be re-colonized by plant species from the local propagule pool. Non-seeded control plots were included to assess effects of seeding the non-native grass. We assessed plant, invertebrate, soil moisture, and soil nutrient responses to the endophyte–cultivar treatments after four growing seasons. Seeding tall fescue affected plant species abundances, but not richness, and did not consistently alter soil moisture and nutrient pools. Endophyte identity in the tall fescue cultivars affected the communities, but effects were not consistent between cultivars. Within Georgia 5, the AR542 endophyte reduced tall fescue abundance and altered the invertebrate community relative to CT plots. Within Jesup, the AR542 endophyte reduced species evenness and decreased soil moisture during dry periods relative to CT plots. Endophyte effects were not consistent between cultivars, and it is probable that the community-scale effects of endophyte infection in tall fescue cultivars arise due to unique interactions between cultivar and endophyte.     

Can Carbon and Phosphorous Amendments Increase Native Forbs in a Restoration Process? A Case Study in the Northern Tall‐grass Prairie (U.S.A.)

In the northern Great Plains (United States), sites with less than 20% of native species are difficult to restore. We have experimented with a restoration method that shows some promise. It consists of systematically installing simulated small‐scale patches (8.0 m² in size) over 25% of an old field and then seeding these patches with native species. The working hypothesis is that these patches will generate a constant source of propagules which in time will lead to increases in native species diversity within the surrounding grass matrix. The objective of this paper was to determine whether soil amendments should be used to facilitate the establishment and persistence of native species (primarily forbs) within these patches. We seeded the patches with a mixture of native grass and forb species and applied four soil treatments: P fertilization, C additions, C + P, and a control (no amendments). Results for the first 5 years were as follows: (1) seeded forb richness was mostly unaffected by soil amendments; (2) seeded and nonseeded forb biomass and density were substantially reduced by C additions, whereas they were unaffected or increased under P additions; (3) both seeded and non‐native grass biomass substantially increased with C additions; and (4) there was an inverse relationship between native seeded forbs and non‐native grass biomass. Our conclusions are that: (1) P amendments are a potential tool for enhancing native seeded forb biomass in simulated small‐scale disturbance patches; and (2) C additions, although enhancing seeded grass biomass do not reduce the biomass of non‐native grasses.     

Short‐Term and Long‐Term Effects of Soil Ripping,Seeding, and Fertilization on the Restoration of a Tropical Rangeland

Rangeland degradation is a serious problem in semiarid Africa. Extensive areas of bare, compacted, nutrient‐poor soils limit the productivity and biodiversity of many areas. We conducted a set of restoration experiments in which all eight combinations of soil tilling, fertilization, and seeding with native perennial grasses were carried out in replicated plots. After 6 months, little aboveground biomass was produced in plots without tilling, regardless of seeding or fertilization. Tilling alone tripled plant biomass, mostly of herbaceous forbs and annual grasses. Perennial grasses were essentially limited to plots that were both tilled and seeded. The addition of fertilizer had no significant additional effects. After 7 years, vegetation had declined, but there were still large differences among treatments. After 10 years, one tilled (and seeded) plot had reverted to bare ground, but the other tilled plots still had substantial vegetation. Only one seeded grass (Cenchrus ciliaris) was still a contributor to total cover after 10 years. We suggest that restoration efforts on these soils be directed first to breaking up the surface crust, and second to the addition of desirable seed. A simple ripping trial inspired by this experiment showed considerable promise as a low‐cost restoration technique.     

Does Restoration Enhance Regeneration of Seasonal Deciduous Forests in Pastures in Central Brazil?

The goal of restoration is to accelerate ecosystem recovery, but in ecosystems that naturally regenerate rapidly restoration techniques need to be selected carefully to facilitate rather than impede natural recovery. We compared the effects of five restoration techniques, such as plowing the soil, removing grasses, adding forest litter, seeding, and planting nursery‐growing seedlings, on the regeneration of seasonal deciduous forest trees in four abandoned pastures in central Brazil. We monitored all woody stems immediately prior to treatments and again 14 months after the treatments. We recorded an average of 16,663 tree stems per hectare and a total of 83 species before implementing treatments. Planting strongly increased species richness; adding litter and seeding had weaker positive effects on richness; and plowing and grass removal had no effect. Plowing substantially reduced the density of naturally established stems. Despite the high survival of planted seedlings, stem density in planting treatments did not change because the tractor and digging holes to plant seedlings caused mortality of naturally regenerating seedlings. Tree stems grew more in the grass release plots than in the control plots. Our results suggest that early succession of seasonal deciduous forest in pastures in the region studied does not need to be stimulated once the perturbation is stopped and that intensive restoration efforts may actually slow recovery. We recommend only enrichment planting of seedlings that are not able to resprout.     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Effects of long-term fire exclusion on tree species composition and stand structure in an old-growth Pinus palustris (Longleaf pine) forest

Frequent fire is an integral component of longleaf pine ecosystems, creating environmental conditions favoring survival and growth of juvenile pines. This study examined stand structure, species composition, and longleaf pine regeneration in an old-growth tract of longleaf pine forest (Boyd Tract) experiencing long-term (>80 yr) fire exclusion in the Sandhills of North Carolina. Sampling of woody stems (i.e., 2.5 cm diameter at breast height) and tallies of longleaf pine seedlings were carried out in plots established randomly on upland, mesic areas and lowland, xeric areas within the Boyd Tract. Dominant woody species in mesic plots were black oak, hickories, and large, sparse longleaf pines. Xeric plots had high densities of turkey oak with the large longleaf pines, as well as higher frequencies of smaller longleaf stems. These differences between areas were associated with higher clay content of upland soils and higher sand content of lowland soils. Age-class frequency distributions for fire-suppressed longleaf pine following the last wildfire at the Boyd Tract approximately 80 yr ago contrasted sharply with data from an old-growth longleaf tract in southern Georgia (Wade Tract) that has been under a long-term frequent fire regime. Post-burn recruitment for the Boyd Tract wildfire appears to have been initially high on both site types. Longleaf pine recruitment diminished sharply on the mesic site, but remained high for 60 yr on the xeric site. Currently, longleaf pine regeneration is minimal on both site types; several plots contained no seedlings. Sharp contrasts in longleaf pine dominance and stand structure between the Boyd and Wade Tracts demonstrate the importance of large-scale disturbance, especially hurricanes and fire, in shaping the structure and function of longleaf pine ecosystems of the southeastern United States. In particular, long-term exclusion of fire on the Boyd Tract has altered stand structure dramatically by permitting hardwoods to occupy at high densities the characteristically large gaps between longleaf stems that are maintained by fire and other disturbances.     

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

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

Pine weevil damage to Norway spruce seedlings: effects of nutrient‐loading,soil inversion and physical protection during seedling establishment

• 1 The pine weevil Hylobius abietis (L.) feeds on the bark of young conifer seedlings and is one of the most economically important forest pests in Europe.
• 2 In a field experiment, we examined the combined effects of the treatments: nutrient‐loading of seedlings, planting in scarified plots and protection of seedlings against pine weevil damage for either half a season or a full season.
• 3 Nutrient loading had no significant effect on the amount of pine weevil feeding.
• 4 Seedling mortality was significantly reduced when seedlings were protected from pine weevil feeding during establishment. This occurred even though the debarked area of protected seedlings 5 weeks after the shields had been removed was similar to that of the unprotected seedlings. This indicates that initial protection rendered the seedlings more tolerant to later wounding by pine weevil.
• 5 Planting in soil inversion significantly reduced feeding compared with planting in humus.
• 6 We conclude that nutrient‐loading of seedlings in the autumn before planting would not increase pine weevil feeding after planting. Mortality could be reduced by treatments that postpone the start of pine weevil feeding on seedlings. Such treatments, combined with planting in soil inversion, would result in increased seedling growth, vitality and tolerance to pine weevil attack.

     

Mycorrhiza development of loblolly pine seedlings in relation to soil reaction and the supply of nitrate

Summary A study was made of the relative influence of nitrate and soil pH on mycorrhiza development in seedlings of loblolly pine (Pinus taeda L.). The investigation was conducted in the greenhouse, using as the growth medium topsoil collected from a young pine plantation.Lime-induced Fe-deficiency occurred at pH 7.5, resulting in chlorotic seedlings with few mycorrhizas. Chlorosis was corrected, and normal mycorrhiza development restored, by adding Fe-EDTA without altering soil pH.Application of 18 1/2, 37, and 74 pounds of N per acre as NaNO₃ reduced mycorrhiza development at age 21 weeks, but had no effect at age 45 weeks. At 21 weeks, the degree of infection varied inversely as the percentage total N in seedling roots. The effect of NaNO₃ was due to the nitrate ion, since Na₂CO₃ did not reduce mycorrhiza development even though it raised soil pH.Alkalinityper se did not affect mycorrhiza formation in loblolly pine seedlings, but only indirectly through its influence on host nutrition. Normal mycorrhiza development was possible at pH values of 7.2 and 7.5, provided Fe deficiency was corrected, and soil nitrate level was kept low. The results may be interpreted in terms of the carbohydrate — nitrogen balance in the root tissues.     

Seedling emergence after sod cutting in grass heath

Abstract. Experimental plots in grass heath, previously subjected to various sod cutting treatments, contained a seed store of ericoid dwarf shrubs, but seeds of grasses of the former grass heath were not found. The density of emerging dwarf-shrub seedlings in the plots was determined mainly by soil moisture, but with density of seeds stored in the soil and soil acidity as modifying factors. It is concluded that the restoration of a dwarf-shrub heathby sod cutting is relatively successful, as compared to other measures, because the grass seeds become obsolete.     

Long-Term Effects of Reclamation Treatments on Plant Succession in Iceland

The long‐term effects (20–45 years) of reclamation treatments on plant succession are examined at two localities in Iceland that were fertilized and seeded from 1954 to 1979 with perennial grasses or annual grasses, or left untreated. The areas that underwent reclamation treatments had significantly higher total plant cover (7–100%) than the untreated control plots (<5%), and floristic composition was usually significantly different between treated and untreated plots. Dwarf‐shrubs (Calluna vulgaris and Empetrum nigrum), bryophytes, biological soil crust, grasses, and shrubs characterized the vegetation in the treated plots, but low‐growing herbs that have negligible effects on the environment, such as Cardaminopsis petraea and Minuartia rubella, and grasses characterized the control plots. The seeded grass species had declined (<10%, the perennials) or disappeared (the annuals) but acted as nurse species that facilitated the colonization of native plants. It seems that by seeding, some factors that limit plant colonization were overcome. Soil nutrients, vegetation cover, litter, and biological soil crust were greater in the treated areas than the control plots. This may have enhanced colonization through an increase in soil stability and fertility, increased availability of safe microsites, increased moisture, and the capture of wind‐blown seeds. This study demonstrates the importance of looking at the long‐term effects of reclamation treatments to understand their impact on vegetation succession.     

Adaptation of plant‐mycorrhizal interactions to moisture availability in prairie restoration

The strength and direction of plant response to inoculation with arbuscular mycorrhizal fungi (AM fungi) is dependent on both abiotic and biotic contexts, often generating patterns of AM fungal mediation of plant adaptation. However, knowledge of plant‐community level effects of these interactions in grassland restoration is limited. We conducted a field inoculation experiment by inoculating five plant species native to a drier prairie and five plant species native to a moister prairie with mycorrhizal fungal communities from each prairie type. Species were paired by genus or family to account for phylogenetic effects. The inoculated plants were transplanted to study plots seeded with a restoration seed mix. Plots were manipulated to create either moister or drier conditions similar to environments of the plant species and mycorrhizal communities. In both transplanted and seeded plant species, we found that only drier prairie‐range species benefited from moisture‐regime matched AM fungal inoculum. Other seeded prairie plant species demonstrated a negative response to inoculation, likely due to the earlier successional stage of these species. Additionally, nonseeded plants benefited from inoculation in different ways: native nonseeded plants had highest cover with drier prairie inoculum in drier conditions, while nonnative plants had highest cover with moister prairie‐origin inoculum. These results suggest that use of local AM fungi may be particularly important in restorations at drier sites, even at relatively small differences in moisture availability. Further, specific knowledge of relative responsiveness of seeded plant species and nonseeded plant species to AM fungal inoculation will be useful in planning restorations.     

Tree dieback and regeneration in secondary Acacia zanzibarica woodlands on an abandoned cattle ranch in coastal Tanzania

Questions: Bush encroachment is a major problem when African savanna ecosystems are used for cattle ranching. How do secondary woodlands develop after ranching is abandoned? What are the patterns and rates of tree mortality and regeneration? Location: Mkwaja Ranch (now part of Saadani National Park) in coastal Tanzania. Methods: Ninety‐seven circular plots (4‐m radius) were set in secondary Acacia zanzibarica woodland along a gradient of tree density. Variables relating to tree and grass layers and soil characteristics of plots were recorded. Seedlings were counted twice in the wet seasons, and resprouts once. Tree flowering and pod production were assessed during the fruiting season, while survival of trees initially present was recorded after 12 and 32 months. Interrelationships among variables were investigated using multiple linear regression, binary logistic regression and mixed effects models. Results: After 32 months, over one‐third of trees in plots had died. Most died after fire, especially on heavy soils, and mortality was significantly related to the tree live biomass ratio and soil conditions. Seed production was very low, especially in denser stands. Numbers of seedlings correlated with soil and grass variables but not with seed rain. Half of trees killed above‐ground produced coppice shoots from the base; in contrast, root suckering was independent of topkill. By the end of the study, no seedlings survived and no resprouts emerged above the grass layer. Conclusions: A. zanzibarica woodlands at Mkwaja Ranch were able to develop because of ranching, and can only persist under intensive grazing. The woodlands do not represent a successional stage towards forest and will probably revert to predominantly grassland vegetation within 10–20 years unless grazing pressure from wild ungulates increases considerably and/or fire regimes change.