Spatial patterns and dynamics of woody vegetation in an arid savanna

The spatial distribution of woody plants was studied in an arid savanna in Botswana. The study included stands of mixed species and sizes as well as monospecific even-sized stands of different size classes of the tree Acacia erioloba and the shrub Acacia mellifera. In the case of A. mellifera both dense stands on overgrazed land and more open stands were included. The analysis used all plant-to-plant distances, and individuals were represented with a realistic canopy extension. The mixed stands showed aggregated distribution of individuals, mainly caused by strong clumping of small shrubs. In A. erioloba saplings were aggregated, small trees were randomly or regularly distributed and large trees were randomly spaced. In open stands of A. mellifera aggregation increased with size of the shrubs, while in dense stands with overgrazing aggregation decreased with increasing size. The different patterns are discussed in relation to the relative importance of inter- and intraspecific competition for water and of disturbance by fire as regulatory mechanisms for total amount and spatial distribution of woody plants in this savanna.     

When do nurse plants stop nursing? Temporal changes in water stress levels in Austrocedrus chilensis growing within and outside shrubs

Question: Does the proximity of shrubs affect seasonal water stress of young Austrocedrus chilensis trees (a native conifer of the Austral Temperate Forest of South America) in xeric sites? Location: A. chilensis xeric forest in northwest Patagonia, Argentina. Methods: We examined the dependence of predawn twig water potential on tree development (seedling to adult) and proximity to nurse shrubs during spring and summer. We analysed spatial associations of seedlings, saplings and adult trees with nurse shrubs, and also evaluated if trees affected shrub canopy vitality. Results: Water stress in Austrocedrus trees was affected by shrub presence. Small trees (i.e.<0.5 m in height) growing in the open were most stressed, particularly in summer. Small trees growing within a shrub canopy had low water stress and little change between spring and summer. The opposite trend, however, was true for the medium‐height category (i.e. 0.5‐1.5 m in height); trees in this size category were more stressed when growing within the shrub canopy than in the open. Larger Austrocedrus trees (i.e.>2 m in height) were not affected by shrub presence. Austrocedrus trees were spatially associated with shrubs in all height classes; however, the percentage of living shrub canopy decreased with tree height. Conclusions: In xeric areas of northwest Patagonia, the strength and direction of interactions between A. chilensis and shrubs, in terms of tree water stress, are dynamic and modulated by tree size and environmental conditions. Overall, positive effects of shrubs on early developmental stages appear to be more important than subsequent negative interactions, since nursing effects could generate a spatial association of shrubs and Austrocedrus trees that persists through later successional stages. These findings shed light on mechanisms behind successional changes, and have important conservation and management implications.     

The small scale spatial pattern of C3 and C4 grasses depends on shrub distribution

At micro‐site scale, the spatial pattern of a plant species depends on several factors including interactions with neighbours. It has been seen that unfavourable effects generate a negative association between plants, while beneficial effects generate a positive association. In grasslands, the presence of shrubby species promotes a particular microenvironment beneath their canopy that could affect differently the spatial distribution of plants with different tolerance to abiotic conditions. We measured photosynthetic active radiation, air temperature and wind speed under shrub canopies and in adjacent open sites and analysed the spatial distribution of four grass species (two C₃ and two C₄) in relation to shrub canopy in a grazed sub‐humid natural grassland in southern Uruguay. Radiation, air temperature and wind speed were lower under shrubs than in adjacent open sites. The spatial distribution of grasses relative to the shrub canopy varied depending on the photosynthetic metabolism of grasses. C₄ grasses showed a negative association or no correlation with the shrubs, whereas C₃ grasses showed a positive association. Our results highlight the importance of the photosynthetic metabolism of the grasses in the final outcome of interactions between grasses and shrubs. Micro‐environmental conditions generated underneath shrubs create a more suitable site for the establishment of C₃ than for C₄ grasses. These results show that facilitation could be more important than previously thought in sub‐humid grasslands.     

Direct versus indirect facilitation (herbivore mediated) among woody plants in a semiarid Chaco forest: A spatial association approach

In arid environments, direct facilitation (microhabitat amelioration) and indirect facilitation (‘associational resistance’ via protection from herbivory) among plants of different species may act simultaneously. Little is known about their relative effects. One way to disentangle the effects is by evaluating spatial associations. We examined the relative importance of these two mechanisms of facilitation in the semiarid Chaco vegetation of north‐central Argentina, through an eight‐way observational study in which we quantified the degree of spatial association between saplings of each of two key tree species, Schinopsis lorentzii (Anacardiaceae) and Aspidosperma quebracho‐blanco (Apocynaceae), with shrub neighbours either possessing spines or without spines and in both an ungrazed site and a site with a long history of cattle grazing. We analysed data across 400 subparcels at each site with spatial analysis by distance indices. Saplings of both tree species showed positive spatial associations with spiny shrubs in the grazed site but not in the ungrazed site, and never with non‐spiny shrubs. This result suggests that spiny shrubs may indeed provide associational resistance for saplings of key tree species in grazed habitats in these dry subtropical forests, that is, that indirect facilitation may predominate over direct facilitation. If confirmed by experimental studies, this result can have implications for the silvopastoral management of rapidly expanding ranches in the semiarid Chaco, where current practice includes the near elimination of native shrubs.     

Can isotropy vs. anisotropy in the spatial association of plant species reveal physical vs. biotic facilitation?

In dryland ecosystems and other harsh environments, a large part of the vegetation is often clustered, appearing as ‘islands’. If ‘independent’ species, usually colonizers, can be distinguished from species which are ‘dependent’ on the presence of the colonizing species for successful establishment and/or persistence, the type of spatial pattern of the association ‐ isotropic (spatially symmetric) or anisotropic (spatially asymmetric) ‐ can give information on the underlying environmental factors driving the process of association. Modified spatial pattern analysis based on Ripley's K‐function can be applied to bivariate clustered patterns by cardinal direction in order to detect possible anisotropy in the pattern of association. The method was applied to mapped distribution patterns of two types of semi‐arid shrubland in southeastern Spain. In shrubland of Retama sphaerocarpa, low shrubs of Artemisia barrelieri were significantly clustered under the canopy of the Retama shrubs in all four cardinal directions, suggesting an isotropic facilitation effect. In low shrubland dominated by Anthyllis cytisoides and Artemisia barrelieri, Anthyllis shrubs occurred more frequently than expected on the eastern side (and downslope) of an Artemisia shrub. The possible environmental factors driving the two association patterns are discussed and recommendations for further applications of the analytical method are given.     

Long‐term mortality patterns of the deep‐rooted Acacia erioloba: The middle class shall die!

Question: Is there a relationship between size and death in the long‐lived, deep‐rooted tree, Acacia erioloba, in a semi‐arid savanna? What is the size‐class distribution of A. erioloba mortality? Does the mortality distribution differ from total tree size distribution? Does A. erioloba mortality distribution match the mortality distributions recorded thus far in other environments? Location: Dronfield Ranch, near Kimberley, Kalahari, South Africa. Methods: A combination of aerial photographs and a satellite image covering 61 year was used to provide long‐term spatial data on mortality. We used aerial photographs of the study area from 1940, 1964, 1984, 1993 and a satellite image from 2001 to follow three plots covering 510 ha. We were able to identify and individually follow ca. 3000 individual trees from 1940 till 2001. Results: The total number of trees increased over time. No relationship between total number of trees and mean tree size was detected. There were no trends over time in total number of deaths per plot or in size distributions of dead trees. Kolmogorov‐Smirnov tests showed no differences in size class distributions for living trees through time. The size distribution of dead trees was significantly different from the size distribution of all trees present on the plots. Overall, the number of dead trees was low in small size classes, reached a peak value when canopy area was 20 ‐ 30 m², and declined in larger size‐classes. Mortality as a ratio of dead vs. total trees peaked at intermediate canopy sizes too. Conclusion: A. erioloba mortality was size‐dependent, peaking at intermediate sizes. The mortality distribution differs from all other tree mortality distributions recorded thus far. We suggest that a possible mechanism for this unusual mortality distribution is intraspecific competition for water in this semi‐arid environment.     

Spatial association of Aspidosperma quebracho‐blanco juveniles with shrubs and conspecific adults in the Arid Chaco,Argentina

Abstract The survival of Aspidosperma quebracho‐blanco juveniles in the Arid Chaco is facilitated under the canopy of nurse plants. The possible effects of nurse plants were studied at intra‐ and interspecific levels by analysing the spatial distribution of juveniles and adults of A. quebracho‐blanco, of the main shrubs Larrea divaricata and Mimozyganthus carinatus, and of the group of deciduous and evergreen shrub species, and their pair associations. Data were analysed using the SADIE (Spatial Analysis by Distance Indices) software. A. quebracho‐blanco seedling abundance followed the distribution pattern of the main shade‐providing species: an aggregated spatial distribution pattern in most of the categories studied. The seedling bank of A. quebracho‐blanco also showed an aggregated pattern and was spatially associated with shrubs and adults of its own species. The intensity of the association depended on the functional types: deciduous Fabaceae, deciduous non‐Fabaceae, evergreen and conspecific adults, each of which provides a different canopy structure and therefore different amounts of shade. The spatial association was significant with the evergreen group, and less significant with the deciduous Fabaceae group. There was no positive association with deciduous non‐Fabaceae, or with gaps (open sky). The differences generated by canopy cover may influence the nurse effect, as observed in the intensity of association of A. quebracho‐blanco with shrubs and conspecific adults.     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Effects of pioneer shrubs on the recruitment of the fleshy‐fruited tree Olea europaea ssp. cuspidata in Afromontane savanna

Question: Is seedling recruitment of a fleshy‐fruited tree in degraded Afromontane savanna dependent on shelter from pioneer shrubs, and is shelter availability related to shrub traits? Location: Degraded montane savanna in northern Ethiopia (13°36′N, 39°21′E). Method: Nurse plants of Olea europaea ssp. cuspidata seedlings were recorded using T‐square plotless sampling and clustered according to shrub traits, using Ward's method after Principal Components Analysis. Facilitation was further examined through experimental planting and Kaplan‐Meier survival analysis. Results: Both in grazed and protected areas, Olea recruits were found exclusively under shrubs, primarily under Euclea racemosa although Acacia etbaica was more abundant. Olea recruitment is distributed randomly at landscape scale, but depends on shelter at patch scale. Shelter ability is related to shrub shape and species identity. Dense multi‐stemmed shrubs with a wide base and crown on a mulch‐rich mound are key recruitment foci. Euclea shrubs have these favoured traits and probably act as preferential perching sites for avian seed dispersers. Soil and organic matter accumulation under Euclea shrubs may also create favourable conditions for Olea germination and survival. Experimentally planted seedlings had a better chance for survival under Euclea. Conclusions: Olea regeneration is probably subject to both passive (disperser‐mediated) and active facilitation. Small changes of shrub traits can alter the suitability of a patch for Olea recruitment. Protection of shrubs can increase facilitation for seedlings, while pruning may reduce competition for saplings and thus enhance forest succession. Planting of raised Olea seedlings under Euclea shrubs in years with a good rainy season may further assist forest restoration.     

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

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

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Restoration of Midwestern U.S. Savannas: One Size Does Not Fit All

Lowland savannas are a rare variant of Midwestern United States savanna occurring on alluvial soils, for which reference information is sparse. To evaluate the appropriateness of using upland savanna as a surrogate source of reference information for lowland savanna, we studied a pre‐Euro‐American lowland savanna using original U.S. Public Land Survey data and other historical records. Historical vegetation was reconstructed and compared among upland savannas, lowland savannas, and lowland forests; we also evaluated potential disturbance dynamics maintaining these systems. We found that all three communities were dominated by members of the genus Quercus but also had extensive representation by many other tree species, especially notable for savannas in this region. There were no clear size–density relationships for species in the genus Quercus, indicating that these historical savannas were not characterized exclusively by large, scattered oak trees but rather by trees of many oak species and nonoak species in a wide range of size classes. Both upland and lowland savannas also contained a substantial shrub component. We found no evidence that lowland savannas were maintained by flooding, although the uneven‐aged canopy structure suggested that periodic disturbance occurred. Restoration of lowland savanna in this region should include provisions for maintaining nonoak species and shrubs, with disturbance timed to maintain an uneven‐aged canopy structure. Although the appropriateness of historical data in the face of climate change may be questionable, in this region, a warmer climate may actually help promote the “oak parkland” that was present from 8,000 BP up to Euro‐American settlement.     

Extreme drought stress shifts net facilitation to neutral interactions between shrubs and sub‐canopy plants in an arid desert

The stress gradient hypothesis (SGH) predicts that the importance or intensity of competition and facilitation will change inversely along abiotic stress gradients. It was originally postulated that increasing environmental stress can induce a monotonic increase in facilitation. However, more recent models predicted that the relationship between severity and interaction exhibits a hump‐shaped pattern, in which positive interactions prevail under moderate stress but decline at the extreme ends of stress gradients. In the present study, we conducted a field experiment along a temporal rainfall gradient for five consecutive years, in order to investigate interactions in a shrub‐herbaceous plant community at the southern edge of the Badain Jaran Desert, and, more specifically, investigated the effects of Calligonum mongolicum, a dominant shrub species, on both abiotic environmental variables and the performance of sub‐canopy plant species. We found that shrubs can improve sub‐canopy water regimes, soil properties, plant biomass, density, cover, and richness and, more importantly, that the positive effect of shrubs on sub‐canopy soil moisture during the summer diminishes as rainfall decreases, a pattern that partly explains the collapse of the positive interaction between shrubs and their understory plants. These results provide empirical evidence that the positive effect of shrubs on understory plant communities in extreme arid environments may decline and become neutral with increasing drought stress.     

Nurse plants,tree saplings and grazing pressure: changes in facilitation along a biotic environmental gradient

Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a field experiment in which tree saplings of four different species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures differing in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the different tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not differ between species. The relative facilitation of sapling browsing and sapling survival – i.e. the difference between saplings inside and outside the shrub canopy – decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these findings do not agree with current conceptual facilitation-stress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.     

Non‐destructive dry matter estimation of Alhagi sparsifolia vegetation in a desert oasis of Northwest China

Question: Can above‐ground biomass of naturally growing Alhagi sparsifolia shrubs be estimated non‐destructively? Location: Qira oasis (37° 01′N, 80° 48′E, 1365 ma.s.l.) at the southern fringe of the Taklamakan desert, Xinjiang, NW China. Methods: Two methods were compared to estimate above‐ground biomass (AGB) of Alhagi. At first shrub AGB was estimated by manual ground measurements (called ‘allometric approach’) of length, width and height of 50 individuals. Subsequently regression equations were established between calculated shrub canopy volume and shrub AGB (r²= 0.96). These equations were used to calculate AGB from manual ground measurements in 20 sample plots within the Alhagi field. Secondly, kite‐based colour aerial photography coupled with the use of a Geographic Information System (called ‘GIS approach’) was tested. First and second order polynomial regressions between AGB data of the 50 individual shrubs and their respective canopy area allowed to automatically calculate the AGB of all remaining shrubs covered by the photograph (r²= 0.92 to 0.96). The use of non‐linear AGB regression equations required an automatised separation of shrubs growing solitary or in clumps. Separation criteria were the size and shape of shrub canopies. Results: The allometric approach was more reliable but also more time‐consuming than the GIS‐based approach. The latter led to an overestimation of Alhagi dry matter in densely vegetated areas. However, this systematic error decreased with increasing size of the surveyed area. Future research in this field should focus on improvements of AGB estimates in areas of high shrub density.     

Positive ecosystem engineering effects of the ant Atta vollenweideri on the shrub Grabowskia duplicata

Question: How does the ant Atta vollenweideri indirectly affect the shrub Grabowskia duplicata? Does environmental modification induced by nest building affect the life history and population structure of this dominant shrub? Location: Halophytic savanna of western Uruguay, South America. Methods: We compared the density of three shrub size classes, seedlings, saplings and adults, among patches created by ant nest building and in non‐modified areas. We studied key soil properties for plant growth in both live nests and non‐modified areas. We performed a greenhouse experiment to study the effects of soils (live nest versus non‐modified area) on shrub germination probability. We also assessed vegetative propagation by measuring the incidence of root suckers and clonal offshoots in live nests and non‐modified areas through a field survey. Results: Seedling‐size class was more abundant on live nests than in non‐modified areas, suggesting that environmental conditions of this type of habitat (i.e. increased sodium and moisture content of soil, and decreased soil compaction) could facilitate shrub recruitment. We did not find any effects of soil type (live nests versus non‐modified areas) on shrub germination probability. However, the vegetative propagation incidence was higher in the environment with live nests. Conclusions: We found a strong indirect relationship between the ant A. vollenweideri and the shrub G. duplicata. This relationship seems to be mediated by an allogenic engineering process, i.e. soil bioturbation caused by nest building. The environmental conditions of live nests seem to facilitate shrub recruitment, mainly by increasing vegetative propagation.     

Response of a monsoon forest-savanna boundary to fire protection,Weipa, northern Australia

A numerical floristic analysis of samples across a monsoon forest-savanna boundary, from an area that had been actively protected from fire for 15 years, at Weipa, northern Australia, revealed three communities: (i) a monsoon forest with a low closed canopy composed mainly of tree species with extra-Australian tropical affinities and a sparse ground layer; (ii) an ecotone with a distinct closed microphyll shrub layer beneath the open canopy of savanna trees; and (iii) a savanna dominated by Eucalyptus tetrodonta. The development of the ecotone has occurred since fire protection and is of limited extent within the fire protected block. The monsoon forest occurred on soils with significantly higher concentrations of bauxitic pisoliths than the other two communities. Soils under the monsoon forest had significantly higher concentrations of total K, S, C, N, exchangeable K and Ca, and higher pH and electrical conductivity than for soils of either of the other communities. A positive relationship between woody basal area and concentrations of surface soil total P, N, C, exchangeable Ca, CEC and gravel was detected across a 20 m transect from the ecotone community into the savanna. The invasion of monsoon forest seedlings was greatest in the ecotone, with few occurring in the savanna. It appears that the expansion of the monsoon forest requires the development of a layer of shrubs. The mechanism of this facilitation is unclear, although the possible role of nutrient enrichment by the shrubs requires further investigation.     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

Seed Bank and Understorey Species Composition in a Semi-arid Environment: The Effect of Shrub Age and Rainfall

Understorey vegetation in patches of Retama sphaerocarpa shrubsin semi-arid environments is dependent on the overstorey shrublife history. Community structure changes with shrub age asa result of physical amelioration of environmental conditionsby the canopy and organic matter accumulation in the soil. Weinvestigated the effect of the canopy on understorey speciesdiversity in the field and its relationships with the soil seedbank under 50 shrubs from 5 to 25+ years old, and compared speciescomposition in the field in a wet and a dry year. Species compositionof the soil seed bank under R. sphaerocarpa shrubs did not differsignificantly with shrub age, but seed density increased asthe shrubs aged. In the field, community composition changedwith shrub age, increasing species richness in a process thatdepended on the amount of spring rainfall. Our results suggestthat the soil seed bank is rather uniform and that the shrubcanopy strongly selects which species appear in the understorey.There were seeds of many species present under both young andold shrubs but which only established under old shrubs. Thisshowed dispersal was not limiting species abundance and suggestedthat the canopy was an important sorting factor for speciespresent in the understorey. Less frequent species contributedthe most to patch diversity, and rainfall effectively controlledspecies emergence. Understorey community composition dependedon multiple interspecific interactions, such as facilitationby the shrub and competition from neighbours, as well as ondispersal processes. Facilitation in this environment is a keyfeature in the structuring of plant communities and in governingecosystem functioning. Copyright 2000 Annals of Botany Company Community structure, competition, dispersal, facilitation, species composition, rainfall variability, Retama sphaerocarpa, seed bank, semi-arid environments     

Facilitation and interference underlying the association between the woody invaders Pyracantha angustifolia and Ligustrum lucidum

Questions: 1. Is there any post‐dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage‐protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse‐plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well.