Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin,USA

As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether “nurse plants” will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220–350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25–33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.     

Effect of grazing on plant patterns in arid ecosystems of Patagonian Monte

Our objective was to assess the relationship between the spatial patterning of perennial grasses (total, grazed, and non‐grazed) and shrub patches in rangelands under different grazing pressures of the Patagonian Monte. We selected three grazed paddocks with the usual stocking rate for the area, where previous studies showed that a piosphere formation is common. At each paddock, we analysed the grain of heterogeneity at sites located at two distances from the single watering point (near, far), using high‐resolution aerial photographs. At these sites, we also assessed in the field the density, size, cover, and spatial patterning of grazed and non‐grazed perennial grasses and shrub patches. The grain of heterogeneity of shrub patches was coarser in sites near the watering point than in those distant from it, as a consequence of the increase in size of both, bare soil and shrub patches. Field sampling showed that a coarse grain of heterogeneity relative to fine‐grained sites resulted from changes in species composition, increased bare soil areas and reduced perennial grass cover. In coarse‐grained sites, lower perennial grass cover resulted from lower density and/or smaller size of grass bunches than in fine‐grained sites. We did not find significant differences among sites in the proportion of perennial grazed grasses. Since the density and cover of perennial grasses was higher in fine‐ than in coarse‐grained sites, we suggested that fine‐grained sites are more important as feeding stations than coarse‐grained sites. The consequences of this differential use could lead to degradation of fine‐grained sites and to higher homogeneity in spatial plant structure and floristic composition within paddocks with respect to the condition observed at present, increasing the size of the highly degraded zone within the piosphere. At the patch level, we found that at about one third of the sampled transects, both total and non‐grazed perennial grasses were spatially aggregated with shrub patches. However, in most transects grazed perennial grasses were indifferently distributed in relation with shrub patches, showing that grazers display high selectivity of foraging sites at macro level (i.e. high and low grazing pressure sites at the paddock level), but random occupancy of vegetation units (randomness in the distribution of grazed perennial grasses at the patch level). The intensity of the positive association between non‐grazed grasses and shrub patches was higher in fine‐grained than in coarse‐grained sites and may be attributed to higher protection against herbivores associated to denser shrub patches in fine‐ relative to coarse‐grained sites. We concluded that a feedback exists between the spatial distribution of species preferred by grazers and the spatial patterning of use of these species.     

Seasonal variation in the seed bank of a Patagonian grassland in relation to grazing and topography

Abstract. The germinable seed bank of Festuca spp., Carex patagonica, and other annuals and perennials in a semiarid Patagonian grassland was analyzed every three months. The effects of grazing, topography and microsites with respect to established grass tussocks on the germinable-seed bank were also analyzed. The total germinable-seed bank was larger in summer after seed rain. At this time of the year most of the seeds were from annuals and perennial dicots. Seeds of the dominant Festuca spp. were the main components of the graminoid seed bank, which was homogeneously distributed in patches of bare soil. In spring, i.e. some months after the seed rain, the germinable-seed bank of most of the species was greatly reduced, while the seed bank of Carex patagonica did not change significantly. At this time of the year, the graminoid seed bank was heterogeneously distributed in space, with the seeds accumulating in wind-protected microsites. In the grazing treatments, the seed bank of the perennial grasses was reduced proportionally to the decrease of the plant cover. On the other hand, the germinable-seed bank of C. patagonica increased with the grazing treatments, in relation to the increase of vegetation cover. Topographical position had two types of effect on the seed bank of perennial grasses. One was the increase of the germinable-seed bank, after the seed rain, on the relatively warmer northeast facing slopes, which was related to a higher seed rain in these places. The other was an increased seed loss on slopes. The different seed-bank strategies observed in this grassland are compared and discussed in relation to strategies described for other grasslands.     

Disturbance: a double-edged sword for restoration in a changing climate

Ecological restoration often relies on disturbance as a tool for establishing target plant communities, but disturbance can be a double-edged sword, at times initiating invasion and unintended outcomes. Here we test how fire disturbance, designed to enhance restoration seeding success, combines with climate and initial vegetation conditions to shift perennial versus annual grass dominance and overall community diversity in Pacific Northwest grasslands. We seeded both native and introduced perennial grasses and native forbs in paired, replicated burned-unburned plots in three sites along a latitudinal climate gradient from southern Oregon to central-western Washington. Past restoration and climate manipulations at each site had increased the variation of starting conditions between plots. Burning promoted the expansion of extant forbs and perennial grasses across all sites. Burning also enhanced the seeding success of native perennial grass and native forbs at the northern and central site, and the success of introduced perennial grasses across all three sites. Annual grass dominance was driven more by latitude than burning, with annuals maintaining their dominance in the south and perennials in the north. At the same time, unrestored grasslands surrounding all sites remained dominated by perennial grasses, suggesting that initial plot clearing may have allowed for annual grass invasion in the southern site. When paired with disturbance, further warming may increase the risk of annual grass dominance, a potentially persistent state.     

Continuous grazing disrupts desert grass-soil seed bank composition under variable rainfall

Long-term field studies help unveil mechanisms of grass soil seed bank (SSB) persistence and resilience. We wonder whether grazing is a disruptive force that changes grass SSB composition and size, while heavy rainfall is a positive bottom-up force that restores the grass SSB in grazed areas of the Monte desert, Argentina. We used core sampling to study the grass SSB in cattle-free (17 year) and grazed (10 year) grasslands. The SSB size in cattle-free sites usually responded in a continuous positive fashion to rainfall, and the proportion of palatable grasses in the SSB was similar among years. Most grass species formed a transient SSB, but the presence of a stand of perennial plants would prevent their local extinction. In contrast, the SSB size in the grazed habitats showed a threshold response to rainfall (i.e. it did not increase under low-moderate rainfall), the seeds of palatable grasses were always scarce, and the proportion of seeds of less palatable grasses increased under extreme rainy conditions. Seed production from grasses that grew tillers from axillary buds during a drought was zero, suggesting that the bank of plants could have a role in the replenishment of the grass SSB only in mesic years. Within the time span assessed here, continuous heavy grazing together with scarce as well as heavy rainfall were disruptive forces that reduced the number and proportion of seeds from palatable grasses in the SSB, which are also the species preferred by seed-eating animals.

     

Sheep Grazing Decreases Organic Carbon and Nitrogen Pools in the Patagonian Steppe: Combination of Direct and Indirect Effects

We explored the net effects of grazing on soil C and N pools in a Patagonian shrub–grass steppe (temperate South America). Net effects result from the combination of direct impacts of grazing on biogeochemical characteristics of microsites with indirect effects on relative cover of vegetated and unvegetated microsites. Within five independent areas, we sampled surface soils in sites subjected to three grazing intensities: (1) ungrazed sites inside grazing exclosures, (2) moderately grazed sites adjacent to them, and (3) intensely grazed sites within the same paddock. Grazing significantly reduced soil C and N pools, although this pattern was clearest in intensely grazed sites. This net effect was due to the combination of a direct reduction of soil N content in bare soil patches, and indirect effects mediated by the increase of the cover of bare soil microsites, with lower C and N content than either grass or shrub microsites. This increase in bare soil cover was accompanied by a reduction in cover of preferred grass species and standing dead material. Finally, stable isotope signatures varied significantly among grazed and ungrazed sites, with δ¹⁵N and δ¹³C significantly depleted in intensely grazed sites, suggesting reduced mineralization with increased grazing intensity. In the Patagonian steppe, grazing appears to exert a negative effect on soil C and N cycles; sound management practices must incorporate the importance of species shifts within life form, and the critical role of standing dead material in maintaining soil C and N stocks and biogeochemical processes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  RAG designed study, performed research, analyzed data, wrote the paper; ATA designed study, wrote the paper; CGGM designed study, performed research, analyzed data; MGP performed research; OES designed study; RBJ designed study, contributed new methods.     

Effects of Caragana microphylla patch and its canopy size on “islands of fertility” in a Mongolian grassland ecosystem

In arid and semiarid regions, variations in “islands of fertility” accompanied by discontinuous vegetation is frequently observed. However, the effects of vegetation patches on soil, including the influence of canopy size, are not fully understood, particularly under conditions of severe grazing. We examined the effects of patches of mound-forming shrub, Caragana microphylla, and the plant’s canopy size on these islands of fertility in a heavily grazed Mongolian grassland. In 11 patches with various canopy sizes (32.5–180 cm in diameter), we compared the chemical properties of soils among three microsites: Mound, Below, and Around, which were inside, below, and outside of C. microphylla mounds, respectively. Total carbon (C) and most essential elements for the plants were more concentrated in Mound, but total nitrogen (N) and nonlimiting elements, such as exchangeable sodium (Na), did not significantly differ among microsites. Larger canopies more strongly affected the enrichment of total C and most essential elements, including total N, in Mound. These results suggest that C. microphylla patches substantially enrich total C and most essential elements and that the extent of enrichment was intensified with canopy size. However, under severe grazing, total N may be relatively more affected by the redistribution of resources through grazing, particularly when the canopy size is small.     

Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0–10 cm² basal area), medium (10–30 cm²), and large ( > 30 cm²) clones were initially mapped in five 0.91-m² quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.     

Post‐fire shrub recruitment in a semi‐arid grassland: the role of microsites

Question: (1) Which factors regulate post‐fire recruitment and spread of the shrub Senecio bracteolatus in Patagonian grasslands? (2) What is the role of the grass Stipa speciosa on S. bracteolatus establishment in the post‐fire succession? Location: Northwest Patagonia, Argentina. Methods: We studied the effect of fire on S. bracteolatus recruitment and density by comparing these variables between burned and unburned grasslands. In burned areas, we compared abiotic characteristics and seedling establishment under the canopy of grasses (S. speciosa) and in gaps (inter‐tussock areas). Post‐fire interactions between S. bracteolatus seedlings and S. speciosa were studied using field and greenhouse experiments. Results: Density of S. bracteolatus was higher in burned than in unburned areas. In burned sites, seedlings were more abundant under tussock grasses, whereas juveniles were more abundant in gaps. Tussocks generated more attenuated micro‐environmental conditions than gaps during stressful summers. Gaps were more abundant in burned sites, while “under tussock” microsites were more frequent in unburned sites. In burned areas, tussocks allowed higher establishment of seedlings (facilitation), but gaps allowed more seedling growth and higher persistence of juveniles. Conclusions: Fire promoted S. bracteolatus recruitment in Patagonian grasslands by increasing the availability of favourable gap microsites. Grass protection for shrub seedlings became negative with time, probably due to competition with grasses. Gaps led to better performance and persistence of shrub plants. Six years after fire, higher shrub recruitment and adult density (observed as a trend) in burned grassland provides an opportunity for potential S. bracteolatus invasion.     

Past and present vegetation in relation to solifluction on Ben Arkle,Sutherland

Background: Forest succession in tropical pastures usually starts from woody vegetation patches. Patches may arise within the grass matrix at microsites with favourable soil conditions or through facilitation by established nurse plants.

Aims: We report the formation of woody vegetation patches in tropical pastures after investigating whether patch formation was associated with micro-scale terrain features and whether facilitation was important for patch initiation.

Methods: The study was conducted in three pasture sites in the Atlantic forest domain of Brazil. We compared soil, terrain and species abundance patterns among pairs of woody patch and open pasture plots.

Results: The effect of variation in soil physical and chemical attributes was limited. Some species were able to establish in the grass matrix and survive disturbance from grazing and fire, while other species only established in patches, under other already established trees or shrubs. Some of these species were exotics, which are commonly eliminated in restoration efforts.

Conclusions: Allowing the establishment of species capable of withstanding pasture environments, including exotics, can accelerate succession. Furthermore, the abilities to endure competition from grasses and survive fire are key features of species suitable for the initial stages of forest restoration in tropical pastures.     

内蒙古典型草原小叶锦鸡儿灌丛化对水分再分配和利用的影响

灌丛化是全球草原地区存在的主要环境问题。通过对内蒙古典型草原区小叶锦鸡儿灌丛和草地斑块冠层降雨再分配、地表径流、土壤含水量的对比观测,研究了小叶锦鸡儿灌丛化对该区水分再分配和利用的影响。结果表明,灌丛和草地斑块的冠层截留量分别占降雨量的20.86%和7.88%,灌丛和草地斑块的平均地表径流系数分别为5.95%和17.19%。土壤含水量观测结果显示,0—60 cm土层中,降雨事件过程中,灌丛斑块较草地斑块能捕获更多水分,灌丛斑块植被冠层下方土壤含水量高于草地斑块;而在雨后无有效降水补充土壤水分的前提下,0—60 cm土层中,灌丛斑块土壤水分蒸散发量高于草地斑块,其中0—10cm土层中灌丛斑块土壤水分蒸散发速率低于草地斑块,10—60 cm土层中灌丛斑块土壤水分蒸散发速率高于草地斑块。研究认为,在水分为关键性限制因子的干旱半干旱区,小叶锦鸡儿灌丛化过程增加草原生态系统中水分分布的空间异质性,灌丛斑块能捕获、利用更多水分以维持更多的生物量。     

Seed distribution of four co-occurring grasses around Artemisia halodendron shrubs in a sandy habitat

In a natural population of the perennial semi-shrub Artemisia halodendron in a shifting sandy habitat in the Horqin Desert of eastern Inner Mongolia, six isolated adult A. halodendron individuals of similar canopy size were chosen as target plants. The density of seeds in the top 5 cm soil depth around shrubs was measured using transects aligned to the four main wind directions and at different distances from the shrub base on both the windward and leeward sides. The effects of shrub presence on seed distribution of four co-occurring grasses were examined by linking seed distribution to seed traits. Of the four species, Setaris viridis and Eragrostis pilosa had small but similar seed mass, while Chloris virgata and Aristida adscensionis had large but similar seed mass. The species were grouped into two cohorts: small-seeded vs. large-seeded cohorts, and shrub presence effects on seed distribution of both cohorts were examined. We found marked difference in the seed distribution pattern among species, especially between the small-seeded and large-seeded cohorts. The small-seeded cohort had significantly higher seed accumulation on the windward than the leeward sides in the most and least prevailing wind directions and much higher seed accumulation on the leeward than the windward sides in the second and third most prevailing wind directions, while opposite patterns occurred in the large-seeded cohort. Four species also showed marked variation in the seed distribution pattern among transects and between windward and leeward sides of each transect. This study provided further evidence that shrubs embedded in a matrix of herbaceous plants is a key cause of spatial heterogeneity in seed availability of herbaceous species. However, seed distribution responses to the presence of shrubs will vary with species as well as with wind direction, sampling position (windward vs. leeward sides of the shrub) and distance from the shrub.     

Regional productivity mediates the effects of grazing disturbance on plant cover and patch‐size distribution in arid and semi‐arid communities

Patch‐size distribution and plant cover are strongly associated to arid ecosystem functioning and may be a warning signal for the onset of desertification under changes in disturbance regimes. However, the interaction between regional productivity level and human‐induced disturbance regime as drivers for vegetation structure and dynamics remain poorly studied. We studied grazing disturbance effects on plant cover and patchiness in three plant communities located along a regional productivity gradient in Patagonia (Argentina): a semi‐desert (low‐productivity community), a shrub‐grass steppe (intermediate‐productivity community) and a grass steppe (high‐productivity community). We sampled paddocks with different sheep grazing pressure (continuous disturbance gradients) in all three communities. In each paddock, the presence or absence of perennial vegetation was recorded every 10 cm along a 50 m transect. Grazing effects on vegetation structure depended on the community and its association to the regional productivity. Grazing decreased total plant cover while increasing both the frequency of small patches and the inter‐patch distance in all communities. However, the size of these effects was the greatest in the high‐productivity community. Dominant species responses to grazing explained vegetation patch‐ and inter‐patch‐size distribution patterns. As productivity decreases, dominant species showed a higher degree of grazing resistance, probably because traits of species adapted to high aridity allow them to resist herbivore disturbance. In conclusion, our findings suggest that regional productivity mediates grazing disturbance impacts on vegetation mosaic. The changes within the same range of grazing pressure have higher effects on communities found in environments with higher productivity, markedly promoting their desertification. Understanding the complex interactions between environmental aridity and human‐induced disturbances is a key aspect for maintaining patchiness structure and dynamics, which has important implications for drylands management.     

Influence of grazing and fire on breeding birds and perennial plants in Cyprus scrub and forest systems

Grazing and browsing by sheep and goats has been an important anthropogenic influence on ecosystems in the Mediterranean region for centuries. This influence has changed significantly in recent decades, with a general shift from range grazing to the penning of animals. The intermediate disturbance hypothesis (IDH) proposes that perturbation - including anthropogenic disturbance - is the norm for Mediterranean ecosystems, and that higher species diversity is found under conditions of continuous, moderate disturbance. Here we test the IDH as it relates to grazing of scrub and open forest habitats in Cyprus, while also testing for the effects of fire. We carried out surveys of breeding birds and vegetation at 48 study sites in scrub and open woodland across Cyprus. We estimated relative grazing pressure (past and present) and fire history at these sites, and tested for associations between these factors, breeding bird species richness and perennial vegetation diversity. We found moderate current grazing reduces density of lower and middle level scrub, and a higher level of diversity of perennial vegetation at moderately over intensively grazed sites. We found that moderate historical grazing pressure had a positive influence on richness of breeding bird species in lower scrub habitat such as phrygana, but that this effect was weaker the taller and denser the habitat. By contrast, current grazing pressure had a negative effect on species richness in lower scrub, but species richness in grazed habitats was higher in denser, taller scrub habitat such as garrigue and maquis. Our study suggests grazing plays an important role in maintaining habitat heterogeneity but the impact on avian species richness depends on the density and height of the vegetation.     

Long-term changes in a Wisconsin Fagus-Acer forest in relation to glaze storm disturbance

Changes in the composition of a Fagus-Acer (Beech-Sugar maple) forest in southeastern Wisconsin over a 16-yr period from 1971 to 1987 are analyzed in relation to a severe glaze (ice) storm disturbance occurring within the census period. Landscape topography created ‘windward’ and ‘leeward’ forest aspects with respect to storm severity, which resulted in greater canopy opening on the windward aspect. In the tree stratum, most species remained stable in density and most of the common species increased in basal area into larger size classes. However, Fagus grandifolia, Ulmus rubra, and the small tree Ostrya virginiana suffered net losses that suggest synergistic effects between glaze storm disturbance and other factors upon tree mortality. In the sapling stratum, canopy opening strongly promoted release of shade-to levant Acer sac-charum. On the windward forest aspect, sapling densities of less shade-tolerant species also increased, in contrast to the absence of such increases on the leeward forest aspect. In the shrub (regeneration) stratum, species responses were heterogeneous. Regeneration of most species increased over the 16-yr period, and some less shade-tolerant species showed increased regeneration differentially on the windward forest aspect. Overall, disturbance appears to have accelerated forest succession toward increased dominance by A. saccharum and persistence of both Fagus and Tilia americana through their capacities for root sprouting. However, forest succession was retarded somewhat on the windward aspect through increased recruitment of less-shade tolerant species. These results parallel those of other studies of glaze storm disturbance, and they illustrate how spatially heterogeneous disturbance intensity may contribute to maintenance of forest diversity.     

Limiting mechanisms in the regeneration of the Chilean matorral – Experiments on seedling establishment in burned and cleared mesic sites

The Chilean matorral is characterized by multispecific shrub clumps in dry areas but has a continuous canopy in wetter sites. It has been hypothesized that this difference is due to easier recolonization of open patches by shrub seedlings under more mesic conditions. Within the mesic range of the matorral we designed a field experiment to compare shrub seedling emergence, growth, and survival under the closed canopy of a secondary forest versus three types of open patches: burned, cleared of shrubs but with a herbaceous layer present, and clear without a herbaceous layer. After the first summer, survival of Quillaja saponaria seedlings was 75% in the burned site, 30% in the cleared patch without herbaceous vegetation, and 15% with herbaceous vegetation present, whereas there was 0% survival under the secondary forest canopy. After eight years, the percentages had dropped to: 22%, 12% and 3%, respectively. These results contrast strongly with the seedling establishment patterns in drier areas of the matorral where early seedling survival is higher under the shade of large shrubs. In the experimental mesic sites, seedlings did best on the burned site, not only in terms of survival, but also in terms of growth. After one year, seedling mean height was 10.2 cm in the burned site, whereas 3.8 cm and 5.3 cm in the cleared patches without and with herbaceous respectively. After eight years, mean height differences between treatments had increased further: 147.7 cm in the burned site, 40.3 cm in the cleared patch without herbaceous cover and 13 cm in the cleared patch with herbs. Our results indicate that the facilitative effect of nurse shrubs on seedling establishment found in dry ranges of the matorral is less important in more mesic sites. This difference may explain the continuous shrub cover in relatively mesic areas as opposed to the characteristic patchy structure of the matorral in its drier range.     

Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland

Facilitative or positive interactions among species are driven mainly by the environmental amelioration or protection from grazing provided by nurse plants. Some studies have suggested that protection from grazing is inconsequential in water-limited environments because of low herbivore densities and their grazing effects. Others, however, argue that herbivores have a major effect on semi-arid plant communities, and that protection from grazing is a significant factor driving positive plant–plant interactions in such environments. We identified a gradient in grazing pressure in a semi-arid shrubland in south-eastern Australia along which we compared soil condition, incident radiation and plant composition beneath two nurse shrub species with open (shrub-free) interspaces. Our aim was to assess the degree of microclimatic amelioration provided by both shrubs, and changes in the interactions (intensity, importance and frequency) between both nurse shrubs and their understorey species, and their effects on species richness at the community level. Both the relative interaction intensity (RII) and interaction importance (I_imp) indices of plant–plant interactions were generally positive and independent of grazing pressure. Soil beneath both nurse plants had significantly greater indices of nutrient cycling and infiltration, and contained more C and N than soil in the open. Almost twice as many species occurred under the canopies of both shrubs (44 species) than in the open (23 species), and the composition of species differed significantly among microsites. Fifty-four percent of all perennial plant species occurred exclusively under shrubs. Our results suggest that environmental amelioration is a stronger driver of the facilitatory effect of shrubs on their understorey species than protection from grazing. Our conclusions are based on the fact that the substantial effect of plant–plant interactions on plant species richness was largely independent of grazing pressure. Irrespective of the underlying mechanism for this effect, our study illustrates the ecological role of shrubs as refugia for understorey plants in semi-arid environments and cautions against management practices aimed at reducing shrub populations.     

Leaf litterfall, fine-root production, and decomposition in shrublands with different canopy structure induced by grazing in the Patagonian Monte, Argentina

Selective sheep grazing in the Patagonian Monte induces the reduction of total and perennial grass cover, species replacement within life forms, and the increase in dominance of long-lived evergreen woody plants with slow growth rates and high concentration of secondary compounds in leaves. We hypothesized that these changes in the canopy structure induced by sheep grazing will affect the mass, chemistry and decomposability of leaf litter and fine roots. We selected two sites in the Patagonian Monte, representative of ungrazed and grazed vegetation states. At each site, we assessed canopy structure (total cover and absolute and relative grass and shrub cover), monthly leaf litterfall, and fine-root biomass and production in the upper soil (15 cm). We also estimated the rates of mass, C, soluble phenolics, lignin and N decay in litterbags containing both leaf litter and fine roots of each site under field conditions during two consecutive years. The ungrazed site exhibited higher total plant cover, absolute and relative grass- and shrub-cover than the grazed one. Leaf litterfall was lower at the grazed site than at the ungrazed site. Fine-root production did not vary between sites. Leaf litter and fine root tissues had higher concentration of secondary compounds at the grazed than at the ungrazed site. However, fine roots showed lower mass and C decay than leaf litter, attributable to the predominant secondary compound (lignin and soluble phenolics, respectively). Leaf litter decomposed slower but released more N during decay at the ungrazed than at the grazed site, probably due to its low concentration of secondary compounds. We concluded that changes in canopy structure induced by grazing disturbance such as those explored in our study could reduce leaf litterfall mass and increase the concentration of secondary compounds of both leaf litter and fine roots leading to slow N release to soil during decay.