Directed dispersal and decomposition drive cyclic succession in arid subtropical thicket

The influence of animal-mediated processes on the purported cyclical succession model of vegetation dynamics in arid subtropical thicket of southern South Africa was examined. We used spatially explicit sampling methods to evaluate the influence of dispersal and decomposition on plant community structure. The harvester ant Messor capensis was the only ant species recorded harvesting Portulacaria afra fruit and the elaiosome bearing seed of the tree Gymnosporia polyacantha from experimental piles. Propagules were dispersed to nest mounds positioned beneath woody shrubs. Vertebrates dispersed fleshy-fruited tree species to regeneration sites beneath P. afra clumps. The minute seeds of succulent shrubs were secondarily dispersed from beneath shrubs to ant nest sites located in the open. The results from two experiments indicate that soil nutrients and decomposition activity are spatially localised. The rate of below-ground decomposition, and the incidence of an invertebrate detritivore assemblage, was high beneath P. afra clumps compared with exposed nutrient-enriched and grassy sites. Animal dispersal vectors are integral to the regeneration of major plant groups in arid subtropical thicket by moving seeds from competitively harsh environments to safe sites beneath nurse plants. Decomposition and nutrient release are localised beneath P. afra clumps due to the unique conditions that support a suite of detritivore taxa recorded nowhere else at the study site.     

Effects of goat pastoralism on ecosystem carbon storage in semiarid thicket,Eastern Cape,South Africa

Abstract Intensive pastoralism with goats transforms semiarid thicket in the Eastern Cape, South Africa from a dense vegetation of tall shrubs to an open landscape dominated by ephemeral grasses and forbs. Approx. 800 000 ha of thicket (which prior to the introduction of goats had a closed canopy and a Portulacaria afra Jacq. component) have been transformed in this manner. Ecosystem C storage in intact thicket and loss of C due to transformation were quantified. Carbon storage in intact thicket was surprisingly high for a semiarid region, with an average of 76 t C ha^?1 in living biomass and surface litter and 133 t C ha^?1 in soils to a depth of 30 cm. Exceptional C accumulation in thicket may be a result of P. afra dominance. This succulent shrub switches between C₃ and CAM photosynthesis, produces large quantities of leaf litter (approx. 450 g m^?2 year^?1) and shades the soil densely. Transformed thicket had approx. 35% less soil C to a depth of 10 cm and approx. 75% less biomass C than intact thicket. Restoration of transformed thicket landscapes could consequently recoup more than 80 t C ha^?1.     

Spontaneous Return of Biodiversity in Restored Subtropical Thicket: Portulacaria afra as an Ecosystem Engineer

An accepted criterion for measuring the success of ecosystem restoration is the return of biodiversity relative to intact reference ecosystems. The emerging global carbon economy has made landscape‐scale restoration of severely degraded Portulacaria afra (spekboom)‐dominated subtropical thicket, by planting multiple rows of spekboom truncheons, a viable land‐use option. Although large amounts of carbon are sequestered when planting a monoculture of spekboom, it is unknown whether this is associated with the return of other thicket biodiversity components. We used available carbon stock data from degraded, restored, and intact stands at one site, and sampled carbon stocks at restored stands at another site in the same plant community. We also sampled plant community composition at both sites. The total carbon stock of the oldest (50 years) post‐restoration stand (250.8 ± 14 t C ha^?1) approximated that of intact stands (245 t C ha^?1) and we observed a general increase in carbon content with restoration age (71.4 ± 24 t C ha^?1 after 35 and 167.9 ± 20 t C ha^?1 after 50 years). A multiple correspondence analysis separated degraded stands from stands under restoration based on ground cover, floristic composition, and total carbon stock. Older post‐restoration and intact stands were clustered according to woody canopy recruit abundance. Our results suggest that spekboom is an ecosystem engineer that promotes spontaneous return of canopy species and other components of thicket biodiversity. The spekboom canopy creates a cooler micro‐climate and a dense litter layer, both likely to favor the recruitment of other canopy species.     

Invasion and persistence of bird-dispersed,subtropical thicket and forest species in fire-prone coastal fynbos

Abstract. Seedling abundance at four microsites (open fynbos, beneath emergent fynbos shrubs, beneath thicket, and beneath forest) was determined at three coastal dune landscapes, located along a gradient of increasing summer rainfall and where fire-dependent fynbos was the predominant vegetation. At all sites thicket seedlings were most common beneath emergent fynbos shrubs and under thicket clumps; seedlings of forest species were most abundant at forest microsites although some individuals were recorded beneath thicket. Very few thicket seedlings were observed in open fynbos. Birds play a keystone role in facilitating establishment of the fleshy fruit-bearing thicket flora. Seedling abundance at microsites of different thicket and forest species was generally unrelated to fruit abundance. Germination success of most species was highest under shaded conditions; soil organic content had no effect on germination. Removal of pulp and birdingestion enhanced the germination, relative to untreated controls, of two out of three species tested. A simple Markov model predicted a gradual increase in cover of the thicket and forest component and a gradual decline in fynbos under a ‘normal’ (20-yr interval) fire regime simulated over 10 cycles. Although inter-fire seedling establishment under emergent fynbos shrubs is important in the initial colonisation of fynbos by obligate resprouting thicket shrubs, these species persist and expand by vegetative recruitment after and between fires, respectively. In the prolonged absence of fire, the endemic-rich and fire-dependent fynbos flora would be replaced by species-poor forest and thicket.     

Structure and dynamics of a tropical dry forest in Ghana

Forest under low rainfall (averaging 745 mm yr^-1) on the Shai Hills in S.E. Ghana has redeveloped following cessation of farming in the 1890s. Forest stature is low, with a canopy at about 11 m, principally of three species, Diospyros abyssinica, D. mespiliformis and Millettia thonningii. Drypetes parvifolia and Vepris heterophylla are common understorey trees. Twelve species of woody liane were recorded. Species of thicket vegetation in the area were also present at low density. Most species are evergreen.Tree mortality averaged 2.3% yr^-1 and exceeded recruitment (1.5% yr^-1). Differences between species in mortality and recruitment were pronounced: canopy species showed a small decline in density; understorey species increased markedly and the thicket species declined. Seed production was very variable, but seedling establishment was very poor for all species. Seedling mortality was high (11% yr^-1) especially for small seedlings. These population trends probably represent the latter stages of succession of forest regrowth after farming about 100 years ago.Compared with tropical rain forest, Shai Hills forest has similar relative tree diameter growth (1–3.5% yr^-1), mortality and recruitment rates, and small-litter fall (5.52 t ha^-1 yr^-1).Shai Hills forest differs from rain forest by its short stature, relatively few (evergreen) tree species, poor regeneration from seed, high soil nutrient status and low rainfall. Similar forests have been reported in east Africa and in parts of New Guinea.Abbreviations dbh diameter at breast height (1.3 m) - gbh girth at breast height died May 1984     

Spatial/temporal variations in shrub thicket soil seed banks on an Atlantic Coast barrier island

Potential species replacement within low-diversity shrub thicket communities was investigated for a Virginia barrier island. Seed bank species composition was quantified in a glasshouse study using soil samples collected beneath closed Myrica cerifera thickets, as well as from thicket gaps. Samples were collected from productive and aging thickets, corresponding to differences in soil age. These data were compared to species presently occurring within the thickets and gaps. Seedbank species composition was not indicative of current community composition for either the intact thickets or the gaps. Seed banks resembled a more pioneer community. Thirteen families, 23 genera, and 25 species were identified from the seed bank beneath the M. cerifera thickets. Four species were woody. The within-gap seed bank included 19 families, 30 genera, and 34 species. Eight species were woody. The current community included 21 families, 33 genera, and 36 species beneath the intact thickets as well as within the thicket gaps. Eighteen species were woody. The species richness of gaps was more than three times that of intact thickets. For low-diversity shrub thickets, gaps enhance species richness.     

How Fast Can Carbon Be Sequestered When Restoring Degraded Subtropical Thicket?

Is the maximum rate of carbon sequestration reported for the CAM‐C₃ plant Portulacaria afra (spekboom), viz. 15.4 t CO₂ ha^?1 yr^?1, unusual in comparison with other plants with similar physioliogies, or could such rates be expected routinely in restoration with P. afra? Private sector investors in thicket restoration need an answer to this question in order to assess the feasibility of using carbon finance as the main income stream from their investments. A literature review showed that 15.4 t CO₂ ha^?1 yr^?1 is not an unusual rate of carbon sequestration for CAM plants in arid and semi‐arid environments, which suggests that investors in thicket restoration should not consider this an outlier value. The results also suggest that carbon finance could be used to fund restoration using other CAM plants in degraded xeric thickets in countries such as Argentina, Chile, Mexico, and Madagascar.     

Portulacaria afra is constrained under extreme soil conditions in the Fish River Reserve, Eastern Cape, South Africa

Restoration of degraded thicket landscapes can be achieved by planting Portulacaria afra (spekboom) cuttings. The factors determining P. afra abundance in thicket types not dominated by P. afra are of interest to restoration practitioners because they could influence restoration protocols using P. afra cuttings in these landscapes. We consequently investigated the relationship between P. afra cover and soil properties, namely pH, EC, organic C, particle size distribution and total content of 26 elements at 78 sites in the Fish River Reserve, Eastern Cape province, South Africa. P. afra cover showed a consistent pattern of constraint at extreme levels (both high and low) of subsoil EC, organic C, sand content, Ca, Zn and Al. The results suggest that P. afra is most competitive in intermediate edaphic environments in the Fish River Reserve. It remains to be investigated in restored landscapes whether P. afra cutting survivorship and growth are greater in such soils.     

An exotic invasive shrub has greater recruitment than native shrub species within a large undisturbed wetland

The idea that invasive species have higher recruitment and tolerate a wider range of conditions than native species requires more rigorous examination across a range of community types. We aimed to compare the recruitment and distribution patterns of adults and seedlings of an exotic invasive plant, glossy buckthorn (Frangula alnus), with four co-occurring native shrub species within a heterogeneous Wisconsin wetland. Detailed vegetation survey data were analyzed for spatial and compositional patterns of shrub distributions. In adult plant frequency and cover, buckthorn was not significantly different from the native winterberry. However, the number of glossy buckthorn seedlings exceeded by more than seven times the combined number of seedlings of the four native species. Sample units containing buckthorn adults were also much more likely to contain seedlings than for native shrubs. However, native seedlings were not more likely to occur at sites lacking adults, suggesting no greater dependence on recruitment away from adults in native species. Buckthorn, winterberry, poison sumac, and dogwood all showed preference for sites with higher tree densities and lower predominance of obligate wetland species in an ordination of 114 species. Glossy buckthorn adults and seedlings and winterberry seedlings were more widely distributed across seven community types than adults and seedlings of the other native species, suggesting broad tolerance to the conditions in different community types. High recruitment is the key factor that may allow glossy buckthorn to overcome community resistance and spread.     

Cross-Scale Patterns in Shrub Thicket Dynamics in the Virginia Barrier Complex

Abstract To interpret broad-scale erosion and accretion patterns and the expansion and contraction of shrub thickets in response to sea level rise for a coastal barrier system, we examined the fine-scale processes of shrub recruitment and mortality within the context of the influence of ocean current and sediment transport processes on variations in island size and location. We focused on Myrica cerifera shrub thickets, the dominant woody community on most barrier islands along the coastline of the southeastern USA. Observations suggest that M. cerifera, a salt-intolerant species, is increasing in cover throughout the Virginia barrier islands, yet rising sea level in response to climate change is increasing erosion and reducing island area. Our objective was to explain this apparent paradox using pattern–process relationships across a range of scales with a focus on ocean currents and sediment transport interacting with island characteristics at intermediate scales. Multi-decadal comparisons across scales showed a complex pattern. At the scale of the entire Virginia barrier complex, modest decreases in upland area were accompanied by large increases in shrub area. Responses were more variable for individual islands, reflecting inter-island variations in erosion and accretion due to differences in sediment transport via ocean currents. Several islands underwent dramatic shrub expansion. Only for within-island responses were there similarities in the pattern of change, with a lag-phase after initial shrub colonization followed by development of linear, closed canopy thickets. Understanding the fine-scale processes of shrub seedling establishment and thicket development, in conjunction with the influence of ocean currents and sediment transport, provides a framework for interpreting island accretion and erosion patterns and subsequent effects on shrub thicket expansion or contraction across scales of time and space.     

Spatial and temporal growth dynamics of Barrier Island shrub thickets

Spatial and temporal dynamics of barrier island shrub thickets were investigated on Hog Island, a barrier island along the Eastern Shore peninsula of Virginia. The island thickets are dominated by the actinorhizal shrub, Myrica cerifera. On the accreting, northern half of the island, thickets expanded as swale area increased. Analysis of aerial photographs revealed that total thicket area increased by nearly 400% from 1949 to 1989, paralleling a substantial increase in area for the northern portion of the island. Thicket characteristics varied with the age and position of the three field sites on Hog Island. At the youngest site examined (20 yr), recruitment and shrub growth were high. Shrub growth remained high and recruitment continued in the stable central portions of the island (50 yr). However, at the oldest site near a bay side salt marsh (≈130 yr), recruitment was not evident, shrub growth was reduced, and mortality was most apparent. Several “life stages” for shrub thickets were identified relative to island accretion. After the stabilization of new land, seedlings establish and grow rapidly to form a thicket. Once established, a thicket rapidly expands in the relatively protected, mesic swale environment. Thicket decline follows, characterized by an increase in vine growth on and around the shrubs and the formation of gaps as the thicket breaks up. Spatial variation in M. cerifera thickets on Hog Island is related to length of time soils have been stabilized and, especially, to time since thicket establishment.     

Changes in Spatial Patterns of Caragana stenophylla along a Climatic Drought Gradient on the Inner Mongolian Plateau

Few studies have investigated the influence of water availability on plant population spatial patterns. We studied changes in the spatial patterns of Caragana stenophylla along a climatic drought gradient within the Inner Mongolian Plateau, China. We examined spatial patterns, seed density, “nurse effects” of shrubs on seedlings, transpiration rates and water use efficiency (WUE) of C. stenophylla across semi-arid, arid, and intensively arid zones. Our results showed that patches of C. stenophylla populations shifted from a random to a clumped spatial pattern towards drier environments. Seed density and seedling survival rate of C. stenophylla decreased from the semi-arid zone to the intensively arid zone. Across the three zones, there were more C. stenophylla seeds and seedlings underneath shrub canopies than outside shrub canopies; and in the intensively arid zone, there were almost no seeds or seedlings outside shrub canopies. Transpiration rates of outer-canopy leaves and WUE of both outer-canopy and inner-canopy leaves increased from the semi-arid zone to the intensively arid zone. In the intensively arid zone, transpiration rates and WUE of inner-canopy leaves were significantly lower and higher, respectively, than those of outer-canopy leaves. We conclude that, as drought stress increased, seed density decreased, seed proportions inside shrubs increased, and “nurse effects” of shrubs on seedlings became more important. These factors, combined with water-saving characteristics associated with clumped spatial patterns, are likely driving the changes in C. stenophylla spatial patterns.     

Response of Betula glandulosa seedlings to simulated increases in nutrient availability,temperature and precipitation in a lichen woodland at the forest–tundra ecotone

Over the last few decades, shrub species have expanded rapidly in open tundra environments due to climate change. Previous experimental studies in this environment have shown that nutrient addition and, to a lesser extent, warming, had positive effects on shrub growth. However, the response of shrub species in open forested ecosystems such as lichen woodland is still largely unknown. The main objective of this study was to evaluate the performance of Betula glandulosa (Michx., dwarf birch) seedlings subjected to direct (warmer temperature, increased precipitation) and indirect (increased nutrient availability) effects of climate change in a lichen woodland (25 % tree cover). The study took place 10 km south of the subarctic treeline in western Québec (Canada). At the end of the second growth season, seedling leaf, woody stem and root biomass along with leaf area had increased significantly in response to nutrient addition. Moreover, seedlings exposed to nutrient addition had greater nitrogen, phosphorus and potassium concentrations in their leaves. Warming treatment also had a significant but weaker impact on leaf and woody stem biomass, while increase in precipitation had only a slight impact on seedling root biomass. Our results indicate that B. glandulosa response to simulated changes in the abiotic environment is similar to that observed in open tundra, suggesting that this species could also become more widespread in the forested ecosystems of the forest–tundra ecotone.     

Relationships between environment and mountain riparian plant communities associated with two rare tertiary-relict tree species, Euptelea pleiospermum (Eupteleaceae) and Cercidiphyllum japonicum (Cercidiphyllaceae)

Euptelea pleiospermum and Cercidiphyllum japonicum are dominant relictual tree species of the riparian plant communities in the subtropical region of Shennongjia Mountains, central China. Community survey and multivariate analysis were carried out to characterize their ecological ranges and regeneration characteristics, and further to explore the factors determining their present distribution boundaries and the barriers restricting their range expansion. We recorded 303 woody species, among which there were 14 relictual, rare and/or endemic woody species. Results obtained from cluster analysis corroborated the results from non-metric multidimensional scaling (NMS) ordination, classifying the 42 plots into three well-defined associations. Correlation analysis revealed that variation in species composition among associations was partly explained by some physical (altitude, slope, soil temperature) and chemical (available K, NH₄, pH) properties. Multiple regression analyses with data of adults and seedlings with physical and chemical variables revealed contrasting patterns of the two dominant relictual species. Both taxa have a high resprouting rate that keeps the stands stable irrespective of scarcity of seedlings. However, expansion of the distribution range is difficult in such a situation. E. pleiospermum prefers riparian habitats with poor nutrients in the south-facing slope of the Shennongjia Mountains, which may indicate that this species is light-demanding at some life stages and well adapted to a nutrient-poor habitat where competition from other species is limited. The Cercidiphyllum japonicum population was more scattered and fragmented, and preferred riparian habitats on gentle slopes. Both species are favored by the mild and moist microclimate, but the current conditions in these subtropical mountain riparian habitats probably do not really provide optimal ecological conditions for the two relictual species.     

Rate of Carbon Sequestration at Two Thicket Restoration Sites in the Eastern Cape, South Africa

Ecosystem carbon storage in intact thicket in the Eastern Cape, South Africa exceeds 20 kg/m², which is an unusually large amount for a semiarid ecosystem. Heavy browsing by goats transforms the thicket into an open savanna and can result in carbon losses greater than 8.5 kg/m². Restoration of thicket using cuttings of the dominant succulent shrub Portulacaria afra could return biodiversity to the transformed landscape, earn carbon credits on international markets, reduce soil erosion, increase wildlife carrying capacity, improve water infiltration and retention, and provide employment to rural communities. Carbon storage in two thicket restoration sites was investigated to determine potential rates of carbon sequestration. At the farm Krompoort, near Kirkwood, 11 kg C/m² was sequestered over 27 years (average rate of 0.42 ± 0.08 kg C m^?2 yr^?1). In the Andries Vosloo Kudu Nature Reserve, near Grahamstown, approximately 2.5 kg C/m² was sequestered over 20 years (0.12 ± 0.03 kg C m^?2 yr^?1). Slower sequestration in the Kudu Reserve was ascribed to browsing by black rhinoceros and other herbivores, a shallower soil and greater stone volumes. Planting density and P. afra genotype appeared to affect sequestration at Krompoort. Closely‐packed P. afra planting may create a positive feedback through increased infiltration of rainwater. The rate of sequestration at Krompoort is comparable to many temperate and tropical forests. Potential earnings through carbon credits are likely to rival forest‐planting schemes, but costs are likely to be less due to the ease of planting cuttings, as opposed to propagating forest saplings.     

The role of the soil seed and seedling bank in the regeneration of diverse plant communities in the subtropical Ailao Mountains,Southwest China

We compared species composition and diversity of the soil seed and seedling banks in three secondary vegetation types (shrubland, Populus bonatii forest, Lithocarpus regrowth forest) and a primary old-growth forest in the subtropical Ailao Mountains of southwestern China to clarify the importance of seed and seedling banks for forest dynamics. The average species richness was the highest in soil samples from the shrubland (26.80 ± 1.98), and the lowest from the primary forest (9.93 ± 0.50). The density of germinable tree seeds increased from the secondary vegetation to the primary forest, and the density of shrub, forb, and graminoid seeds decreased significantly. The most abundant seedlings recorded in soil samples were light-demanding species in the shrubland and Populus bonatii forest. For ground flora, the number of shrub seedlings strongly decreased with the increase in stand age, and shade-tolerant tree seedlings tended to increase. The species similarity between the seed bank and the aboveground vegetation in all sites was low (Sørensen’s index = 0.11–0.33), however, the shrubland had higher similarity compared with the other three plant communities. In the primary forest, light-demanding woody species dominated in soil seed banks, while shade-tolerant species dominated in the overstory and the forest floor. In the primary forest, seedlings of dominant tree species were rare in the understory, and no seeds of the dominant species were found in the soil. Results indicated that the early stages of vegetation recovery should take into account the possibility of recovering soil seed bank processes. However, colonization and establishment of tree seedlings will be difficult once a primary forest is destroyed.     

Testing the Stress‐Gradient Hypothesis During the Restoration of Tropical Degraded Land Using the Shrub Rhodomyrtus tomentosa as a Nurse Plant

The relative importance of facilitation and competition between pairwise plants across abiotic stress gradients as predicted by the stress‐gradient hypothesis has been confirmed in arid and temperate ecosystems, but the hypothesis has rarely been tested in tropical systems, particularly across nutrient gradients. The current research examines the interactions between a pioneer shrub Rhodomyrtus tomentosa (the nurse plant) and seedlings of a transplanted native woody Schima superba (the target species) in a tropical system in which position on a slope corresponds with a nutrient gradient; high soil nutrients at the slope bottom and relatively low soil nutrients at the slope top. In contrast, soil physical traits were more favorable for seedling growth under the shrub than in open spaces. The effect of R. tomentosa on S. superba survival was positive (facilitation) at the top of the slope, as indicated by the relative interaction index (RII), but negative in the bottom (competition). RII indicated a positive effect on seedling height at the top of the slope but was not at the bottom. Seedling survival was positively related to soil nutrient level and negatively related to soil acidity, but seedling growth of S. superba seemed to be enhanced by the shrub canopy. Thus, the results seem to support stress‐gradient hypothesis in terms of target species survival but not growth. We suggest using the shrub as a nurse plant in forest restoration in tropical degraded land with caution because not all of its effects on target species are positive .     

Seed deposition patterns and the survival of seeds and seedlings of the palm Euterpe edulis

The seed deposition pattern created by a seed disperser is one of the components of the efficiency of a species as seed disperser, and ultimately may influence the recruitment of a plant species. In this study, we used the seeds of a bird-dispersed forest palm, Euterpe edulis, to investigate the effects of two distinct seed deposition patterns created by birds that defecate (clumped pattern) and regurgitate seeds (loose-clumped pattern) on the survival of seeds experimentally set in an E. edulis-rich site, and of seedlings grown under shade-house conditions. The study was conducted in the lowland forest of Parque Estadual Intervales, SE Brazil. Clumped and loose-clumped seeds were equally preyed upon by rodents and insects. Although clumped and isolated seedlings had the same root weight after 1 year, the isolated seedlings survived better and presented more developed shoots, suggesting intraspecific competition among clumped seedlings. Our results indicate that animals that deposit E. edulis seeds in faecal clumps (e.g. cracids, tapirs) are less efficient seed dispersers than those that regurgitate seeds individually (e.g. trogons, toucans). Intraspecific competition among seedlings growing from faecal clumps is a likely process preventing the occurrence of clumps of adult palms.     

Direct and indirect effects of grazing constrain shrub encroachment in semi‐arid Patagonian steppes

Question: What are the long‐term effects of grazing exclusion on the population structure and dynamics of, and interactions among, three dominant shrub species? Location: Grass‐shrub Patagonian steppe, Chubut, Argentina. Methods: Permanent plots were established in grazed paddocks and paddocks excluded from grazing in representative Patagonian rangelands. Shrub abundance, population size‐structure, short‐term (two 3‐yr periods) and long‐term (matrix models) population dynamics, and neighborhood interactions of three native and codominant shrub species (Mulinum spinosum, Senecio filaginoides and Adesmia volckmanni) were measured and analysed using different statistical approaches. Results: The total density of shrubs was 74% higher in paddocks excluded from grazing, owing mainly to increases in Mulinum (80%) and Senecio (68%) species. However, differences in size structure between ungrazed and grazed paddocks were only detected in Mulinum. Demographic rates differed between shrub species, time‐periods and grazing conditions. In particular, recruitment in the short term (especially in wet years) and population growth rate in the long term (λ) were higher in paddocks excluded from grazing only in Mulinum populations. Senecio populations showed a marginal increase in recruitment and mortality independent of the grazing condition in the wet and dry period. Grazing exclusion modified the balance of neighborhood interactions among the three shrub species. In grazing‐exclusion paddocks, there was a balance between positive and negative interspecific interactions, while in grazed paddocks there were more negative intraspecific and interspecific interactions, resulting in a net negative balance of neighborhood interactions. Conclusions: Our understanding of woody encroachment in arid rangelands can be informed through evaluation of direct and indirect effects of grazing exclusion on the abundance and demography of dominant woody species. In Patagonian arid steppes, the occurrence of woody encroachment in rangelands excluded from grazing can be explained by altered responses in plant‐animal and plant‐plant interactions among shrub species.     

Bioassays of nutrient limitation in a tropical rain forest soil

Summary Six speices of shrubs and one large herb with contrasting life history patterns were used as bioassays of nutrient availability in a Costa Rican lowland rain forest soil. Growth responses of the herb (Phytolacca rivinoides, Phytolaccaceae) confirmed soil measurements indicating high availability of N and potentially limiting levels of P, K, Mg and Ca. Growth responses of the shrub species (Miconia spp., Melastomataceae and Piper spp., Piperaceae) to a complete nutrient fertilizer were generally less than that of Phytolacca. Lack of a strong shrub response to +P fertilization is probably due to mycorrhizal associations and slower growth rates of woody species. In general, increased growth did not occur at the expense of phenolic production in the leaves. The results emphasize that assessment of specific nutrient limitations to plant growth vary depending on species selected for the bioassay, even among species from the same community.