Herbivory by subterranean termite colonies and the development of fairy circles in SW Namibia

1. The growth pattern of Namibian fairy circles was examined in relation to environmental, termite, and plant variables to provide support for the sand termite (Psammotermes allocerus Silvestri) hypothesis of circle origin. 2. New and young circles were associated with the highest number of sand termites and their foraging grass sheetings which were both considerably lower in mature and senescent circles. 3. Circles increased in size with age, and apart from the senescent stage had higher soil moisture levels than the matrix. 4. In laboratory trials sand termites browsed seedling roots, negatively impacting root and leaf number. 5. This provides a potential mechanism for circle formation through central‐based foraging by P. allocerus which eliminates Stipagrostis grass clumps around its nest system. 6. The resulting bare disc is postulated to be maintained through a combination of a depleted seed bank, termite herbivory on seedlings, and excavations by pugnacious ants (Anoplolepis steingroeveri Forel).     

Adopting a spatially explicit perspective to study the mysterious fairy circles of Namibia

The mysterious ‘fairy circles’ are vegetation‐free discs that cover vast areas along the pro‐Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25‐ha aerial images of north‐west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self‐organized vegetation pattern. Finally, we test if an existing partial‐differential‐equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key‐processes in arid areas such as plant competition for water and local resource‐biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas‐leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self‐organized vegetation patterns that emerge from positive biomass‐water feedbacks involving water transport by extended root systems and soil‐water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self‐organization to match the birth‐ and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.     

Are Namibian “Fairy Circles” the Consequence of Self-Organizing Spatial Vegetation Patterning?

Causes of over-dispersed barren “fairy circles” that are often surrounded by ca. 0.5 m tall peripheral grasses in a matrix of shorter (ca. 0.2 m tall) grasses in Namibian grasslands remain mysterious. It was hypothesized that the fairy circles are the consequence of self-organizing spatial vegetation patterning arising from resource competition and facilitation. We examined the edaphic properties of fairy circles and variation in fairy circle size, density and landscape occupancy (% land surface) with edaphic properties and water availability at a local scale (<50 km) and with climate and vegetation characteristics at a regional scale. Soil moisture in the barren fairy circles declines from the center towards the periphery and is inversely correlated with soil organic carbon, possibly indicating that the peripheral grass roots access soil moisture that persists into the dry season within fairy circles. Fairy circle landscape occupancy is negatively correlated with precipitation and soil [N], consistent with fairy circles being the product of resource-competition. Regional fairy circle presence/absence is highly predictable using an empirical model that includes narrow ranges of vegetation biomass, precipitation and temperature seasonality as predictor variables, indicating that fairy circles are likely a climate-dependent emergent phenomenon. This dependence of fairy circle occurrence on climate explains why fairy circles in some locations may appear and disappear over time. Fairy circles are only over-dispersed at high landscape occupancies, indicating that inter-circle competition may determine their spacing. We conclude that fairy circles are likely to be an emergent arid-grassland phenomenon that forms as a consequence of peripheral grass resource-competition and that the consequent barren circle may provide a resource-reservoir essential for the survival of the larger peripheral grasses and provides a habitat for fossicking fauna.     

Ants and the enigmatic Namibian fairy circles – cause and effect?

1. Parts of the Namibian landscape show extensive surface perturbation in the form of long‐lived, yet dynamic ‘fairy circles'. While exerting profound ecological effects on 7.3% of the land surface, the origin and nature of these large bare discs embedded in an arid grassland matrix remains unresolved. 2. We found no evidence to support the current hypothesis of a termite origin for fairy circles but instead observed a strong spatial association between fairy circles and large nests of the ant Black pugnacious ant Anoplolepis steingroeveri Forel, with much higher ant abundances on the circles compared with the matrix. 3. Aggression trials showed that different colonies of A. steingroeveri were located on different circles, and that the species was polydomous. 4. Fairy circles and Pogonomyrmex ant nests both have a bare disc surrounding the nest, are overdispersed (evenly spaced), and are associated with elevated soil moisture. Fairy circle soils exhibited a five‐fold increase in soil moisture when compared with the matrix. 5. Senescent Stipagrostis obtusa (Delile) Nees seedlings were only observed on the circles and not in the matrix, and were found to have a reduction in both root length and number of roots. 6. Anoplolepis steingroeveri excavated the root system of both S. obtusa seedlings on the disc and Stipagrostis ciliata (Desf.) de Winter grasses on the perimeter of the circles, where they tended honeydew‐secreting Meenoplidae bugs that fed on grass roots and culms. The bugs occurred almost exclusively on grasses associated with the circles. This ant–bug interaction is a possible mechanism for the observed reduction in root length and number of senescent grass seedlings on the circles.     

Edaphic properties enable facilitative and competitive interactions resulting in fairy circle formation

Millions of generally regularly spaced, roughly circular barren patches called fairy circles occur in a narrow band ca 100 km inland of the south‐west African coast. These generally have conspicuously taller peripheral grasses in a shorter grass matrix. The origins of these fairy circles are controversial, but one possibility is that they are self‐organizing emergent vegetation patterns that are the consequence of interplay between positive (facilitative) and negative (competitive) interactions between grasses. We hypothesized that the coarse textured sand on which fairy circles occur creates a hydraulically and nutritionally connected landscape, in which neighbouring fairy circles competitively influence each other over several metres, while providing opportunity for focusing of resources around the peripheral grasses. To test our hypotheses we conducted three main groups of analyses: 1) we measured grass biomass to assess facilitative and competitive effects of the component grasses; 2) across a region with fairy circles we measured the size and density of fairy circles and correlated that with water infiltration rates into soil; 3) we measured the capacity of soil to conduct water pulses and ¹⁵N tracers. We found evidence of facilitative interactions in the periphery of the fairy circles and competitive suppression of the matrix grass proximal to the periphery. Across the region, fairy circle size was positively correlated with soil infiltration rates and negatively with precipitation. This suggests that fairy circles emerge in soils with high capacity for water flux that enables landscape hydraulic connectivity. Water‐ and ¹⁵N‐pulse experiments showed that edaphic resources were highly mobile, moving up to 7.5 m over a period of 1–3 weeks. We concluded that the evidence is consistent with an emergent vegetation pattern explanation for the origins of fairy circles and that the circles are more closely associated with a highly connective edaphic environment, rather than with particular biota.     

Effects of plant-soil feedback on tree seedling growth under arid conditions

Aims Plants are able to influence their growing environment by changing biotic and abiotic soil conditions. These soil conditions in turn can influence plant growth conditions, which is called plant–soil feedback. Plant–soil feedback is known to be operative in a wide variety of ecosystems ranging from temperate grasslands to tropical rain forests. However, little is known about how it operates in arid environments. We examined the role of plant–soil feedbacks on tree seedling growth in relation to water availability as occurring in arid ecosystems along the west coast of South America.Methods In a two-phased greenhouse experiment, we compared plant–soil feedback effects under three water levels (no water, 10% gravimetric moisture and 15% gravimetric moisture). We used sterilized soil inoculated with soil collected from northwest Peru (Prosopis pallida forests) and from two sites in north-central Chile (Prosopis chilensis forest and scrublands without P. chilensis).Important findings Plant–soil feedbacks differed between plant species and soil origins, but water availability did not influence the feedback effects. Plant–soil feedbacks differed in direction and strength in the three soil origins studied. Plant–soil feedbacks of plants grown in Peruvian forest soil were negative for leaf biomass and positive for root length. In contrast, feedbacks were neutral for plants growing in Chilean scrubland soil and positive for leaf biomass for those growing in Chilean forest soil. Our results show that under arid conditions, effects of plant–soil feedback depend upon context. Moreover, the results suggest that plant–soil feedback can influence trade-offs between root growth and leaf biomass investment and as such that feedback interactions between plants and soil biota can make plants either more tolerant or vulnerable to droughts. Based on dissecting plant–soil feedbacks into aboveground and belowground tissue responses, we conclude that plant–soil feedback can enhance plant colonization in some arid ecosystems by promoting root growth.     

Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

Forest fire may disrupt plant–microbial feedbacks

Plant–microbial feedbacks are important drivers of plant community structure and dynamics. These feedbacks are driven by the variable modification of soil microbial communities by different plant species. However, other factors besides plant species can influence soil communities and potentially interact with plant–microbial feedbacks. We tested for plant–microbial feedbacks in two Eucalyptus species, E. globulus and E. obliqua, and the influence of forest fire on these feedbacks. We collected soils from beneath mature trees of both species within native forest stands on the Forestier Peninsula, Tasmania, Australia, that had or had not been burnt by a recent forest fire. These soils were subsequently used to inoculate seedlings of both species in a glasshouse experiment. We hypothesized that (i) eucalypt seedlings would respond differently to inoculation with conspecific versus heterospecific soils (i.e., exhibit plant–microbial feedbacks) and (ii) these feedbacks would be removed by forest fire. For each species, linear mixed effects models tested for differences in seedling survival and biomass in response to inoculation with conspecific versus heterospecific soils that had been collected from either unburnt or burnt stands. Eucalyptus globulus displayed a response consistent with a positive plant–microbial feedback, where seedlings performed better when inoculated with conspecific versus heterospecific soils. However, this effect was only present when seedlings were inoculated with unburnt soils, suggesting that fire removed the positive effect of E. globulus inoculum. These findings show that external environmental factors can interact with plant–microbial feedbacks, with possible implications for plant community structure and dynamics.     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

Novel technique shows different hydrophobic chemical signatures of exotic and indigenous plant soils with similar effects of extracts on indigenous species seedling growth

Changes to ecosystem abiotic parameters are regarded as possible mechanisms facilitating plant invasion and community composition shifts. This study compared the hydrophobic chemical signatures of soil from exotic bitou bush (Chrysanthemoides monilifera spp. rotundata) invaded, indigenous acacia (Acacia longifolia var. sophorae) dominated and bare sand (unvegetated) habitats using a novel, rapid, capturing technique which utilised Amberlite® XAD4 resin filled bags that were placed in situ. The hydrophobic chemical signature of the bitou bush soil extract was significantly different to the acacia soil and bare sand extracts. High concentrations of 18 sesquiterpenes dominated the hydrophobic signature of the bitou bush extract. Low concentrations of all three extracts did not significantly affect the seedling growth of three indigenous test species under laboratory conditions, however, at higher concentrations, the extracts from soil inhabited by plants, whether exotic or indigenous, similarly inhibited the seedling growth of two species, while seedling growth of the third species was inhibited by extracts from all three soil types. These results do not support the hypothesis that exotic invasive species are more likely to exhibit allelopathic effects than indigenous plant species.     

Soil microbial communities from an elevational cline differ in their effect on conifer seedling growth

Sub-alpine environments consist of altitudinal gradients associated with dramatic changes in plant growth and community composition, but the role of soil feedbacks and microbe interactions is largely unknown. Here, we examine the influence of the overall soil microbial community, with a focus on ectomycorrhizal and dark septate endophytic root colonizing fungi, from low, mid, and high elevations on the growth of Pinus contorta and Picea glauca × engelmannii. The influence of the soil microbial community was tested on seedlings from the same three elevations in order to determine ‘home’ versus ‘away’ effects on conspecifics of differing elevations. The low elevation soil was the most fertile and harbored a soil microbial community with an overall negative effect on seedling growth. In contrast, the high elevation soil was the least fertile and had a microbial community that enhanced seedling growth. However, only the soil microbial community in the highest elevation soil resulted in a stronger influence on the native P. contorta seedlings than seedlings originating from lower elevations. Despite the overall influence of the soil microbial community, ectomycorrhizal colonization was significantly correlated with P. glauca × engelmannii growth rates, but colonization by dark septate endophytes showed no relationship with seedling growth. The results provide evidence that plant—soil microbial community relationships are dependent on soil environment. Moreover, our results provide further support for the importance of soil microbes in facilitating seedling growth toward the edge of their elevational range.     

Temporal and micro-spatial patterning of seedling establishment. Consequences for patch dynamics in the southern Monte,Argentina

We compared the temporal and micro-spatial patterning of seedling emergence and establishment of two cohorts of perennial grasses and shrubs in the southern Monte, Argentina. Samplings were carried out in two contrasting areas (grazed and non-grazed) during four years. We found lower densities of emerged and established seedlings of perennial grasses in the grazed relative to the non-grazed area. Conversely, emerged seedlings of shrubs did not vary between the grazed and the non-grazed area and densities of established shrub seedlings were higher in the grazed than in the non-grazed area. We only found differences between cohorts in seedling emergence of perennial grasses. These differences could be associated with the amount of precipitation in the year previous to the emergence. Both in the grazed and non-grazed area, seedlings of perennial grasses were concentrated at the periphery of plant patches. We defined a plant patch as a discrete unit of the spatial pattern of vegetation surrounded by, at least, 20 cm of bare soil from the nearest neighbour patch. Emergence in perennial grasses was more frequent at the southern/western patch-periphery than at other patch-periphery locations. Established seedlings of perennial grasses, however, were homogeneously distributed throughout patch periphery. Emergence in shrubs was more frequent at the centre and periphery of patches than at inter-patch microsites. In contrast, established seedlings of shrubs were homogeneously distributed among microsites. Our results suggests that differential seedling establishment between life forms is the outcome of complex biotic and abiotic interactions and feedbacks at the patch level between seedlings and established plants. Both life forms appear to have a different role in the origin, dynamics, and maintenance of spotting vegetation. Because of the ability to establish both at inter-patch and patch microsites, shrubs could be identified as colonizers or initiators of small plant patches in bare soil or they may contribute to increase the cover and size of pre-existing plant patches. Both processes would be promoted in grazed areas. Due to the ability to establish at patch peripheries, perennial grasses would contribute to the isodiametric growth of pre-existing patches, preferentially in non-grazed areas. This revised version was published online in June 2006 with corrections to the Cover Date.     

Plant‐soil feedbacks from 30‐year family‐specific soil cultures: phylogeny,soil chemistry and plant life stage

Intraspecific negative feedback effects, where performance is reduced on soils conditioned by conspecifics, are widely documented in plant communities. However, interspecific feedbacks are less well studied, and their direction, strength, causes, and consequences are poorly understood. If more closely related species share pathogens, or have similar soil resource requirements, plants may perform better on soils conditioned by more distant phylogenetic relatives. There have been few empirical tests of this prediction across plant life stages, and none of which attempt to account for soil chemistry. Here, we test the utility of phylogeny for predicting soil feedback effects on plant survival and performance (germination, seedling survival, growth rate, biomass). We implement a full factorial experiment growing species representing five families on five plant family‐specific soil sources. Our experiments exploit soils that have been cultured for over 30 years in plant family‐specific beds at Oxford University Botanic Gardens. Plant responses to soil source were idiosyncratic, and species did not perform better on soils cultured by phylogenetically more distant relatives. The magnitude and sign of feedback effects could, however, be explained by differences in the chemical properties of “home” and “away” soils. Furthermore, the direction of soil chemistry‐related plant‐soil feedbacks was dependent on plant life stage, with the effects of soil chemistry on germination success and accumulation of biomass inversely related. Our results (1) suggest that the phylogenetic distance between plant families cannot predict plant–soil feedbacks across multiple life stages, and (2) highlight the need to consider changes in soil chemistry as an important driver of population responses. The contrasting responses at plant life stages suggest that studies focusing on brief phases in plant demography (e.g., germination success) may not give a full picture of plant–soil feedback effects.     

Germination, survival and growth of three vascular plants on biological soil crusts from a Mexican tropical desert

Information about the effects of biological soil crusts (BSC) on germination, seedling survival and growth of vascular plants is controversial because they can have positive, neutral or negative effects. This controversy may be because most studies conducted until now have just analysed one or two recruitment stages independently. To understand the BSC effects on vascular plants, it is necessary to consider each stage of the recruitment process and synthesise all this information. The goal of this study was twofold. First, we analyse germination, seedling survival and growth of three vascular plants (Agave marmorata, Prosopis laevigata and Neobuxbaumia tetetzo) on BSC (cyanobacteria and mixed crust) from a tropical desert region of south-central México. Second, we synthesise the information to determine the total effect of BSC on plant species performance. We conducted experiments under controlled conditions to evaluate the proportion of germinated seeds, proportion of surviving seedlings and seedling dry weight in BSC and bare soil. Results showed that BSC have different effects on germination, seedling survival and growth of plant species. Plant species performance was qualitatively higher on BSC than bare soil. The highest performance of A. marmorata and P. laevigata was observed on cyanobacteria and mixed crusts, respectively. The highest performance of N. tetetzo was on both crust types.     

Indirect vegetation–soil moisture feedback with application to Holocene North Africa climate1

Using a fully coupled climate–terrestrial ecosystem model, we demonstrate explicitly that an initial perturbation on vegetation induces not only a direct positive vegetation feedback, but also a significant indirect vegetation–soil moisture feedback. The indirect feedback is generated through either fractional cover change or soil moisture depletion. Both indirect feedback mechanisms are triggered by a vegetation perturbation, but involve subsequent effects of soil moisture and evaporation, indirectly. An increase in vegetation tends to reduce bare‐ground evaporation through either the area reduction in bare ground or the depletion of soil moisture; the reduced evaporation may then counter the initial plant transpiration, favoring a negative net vegetation feedback. Furthermore, grasses are more effective in inducing the indirect vegetation–soil feedbacks, because of their limited plant evapotranspiration and shallower roots that tend to change surface soil moisture, and, in turn, evaporation, effectively. In comparison, trees favor a direct positive vegetation feedback due to their strong plant transpiration on subsurface soil moisture as well as a lower albedo.     

Adult trees cause density‐dependent mortality in conspecific seedlings by regulating the frequency of pathogenic soil fungi

Negative density‐dependent seedling mortality has been widely detected in tropical, subtropical and temperate forests, with soil pathogens as a major driver. Here we investigated how host density affects the composition of soil pathogen communities and consequently influences the strength of plant‐soil feedbacks. In field censuses of six 1‐ha permanent plots, we found that survival was much lower for newly germinated seedlings that were surrounded by more conspecific adults. The relative abundance of pathogenic fungi in soil increased with increasing conspecific tree density for five of nine tree species; more soil pathogens accumulated around roots where adult tree density was higher, and this greater pathogen frequency was associated with lower seedling survival. Our findings show how tree density influences populations of soil pathogens, which creates plant‐soil feedbacks that contribute to community‐level and population‐level compensatory trends in seedling survival.     

Tree seedling performance and below-ground properties in stands of invasive and native tree species

The establishment and subsequent impacts of invasive plant species often involve interactions or feedbacks with the below-ground subsystem. We compared the performance of planted tree seedlings and soil communities in three ectomycorrhizal tree species at Craigieburn, Canterbury, New Zealand – two invasive species (Pseudotsuga menziesii, Douglas-fir; Pinus contorta, lodgepole pine) and one native (Nothofagus solandri var. cliffortioides, mountain beech) – in monodominant stands. We studied mechanisms likely to affect growth and survival, i.e. nutrient competition, facilitation of carbon and nutrient transfer through mycorrhizal networks, and modification of light and soil conditions by canopy trees. Seedlings were planted in plastic tubes filled with local soil, and placed in monospecific stands. Effects of root competition from trees and mycorrhizal connections on seedling performance were tested by root trenching and use of tubes with or without a fine mesh (20 μm), allowing mycorrhizal hyphae (but not roots) to pass through. Survival and growth were highest in stands of Nothofagus and lowest under Pseudotsuga. Surprisingly, root trenching and mesh treatments had no effect on seedling performance, indicating canopy tree species affected seedling performance through reduced light availability and altered soil conditions rather than below-ground suppression from root competition or mycorrhizal facilitation. Seedlings in Pseudotsuga stands had lower mycorrhizal colonisation, likely as a result of the lower light levels. Soil organic matter levels, microbial biomass, and abundance and diversity of microbe-consuming nematodes were all highest under Nothofagus, and nematode community assemblages differed strongly between native and non-native stand types. The negative effects of non-native trees on nematodes relative to Nothofagus are likely due to the lower availability of soil organic matter and microbial biomass in these stands, and therefore lower availability of resources for nematodes. This study shows that established stands of non-native invasive tree species may adversely affect tree seedlings and soil communities through modifications of the microenvironment both above and below ground. As such, invasion and domination of new landscapes by these species is likely to result in fundamental shifts in community- and ecosystem-level properties relative to those under native forest cover.     

Soil-Behaviour of Phytophthora clandestina

Investigations were undertaken to study the nature and behaviour of P. clandestina in soil. The pathogen was recovered only from soil sievings 250–499 μm and 500 μm–0.99 mm, containing small root fragments. In soil the introduced inoculum of the fungus was incapable of saprophytically and competitively colonizing the dead cotyledons of subterranean clover used as bait material. Exposure of the inoculum to increasing numbers of microbes by adding greater proportions of nonsterile fields, oil to the growth medium of the plant had no significant effect on survival rate and fresh shoot weight of subterranean clover. However, microbes present in the field soil reduced the severity of root rot of subterranean clover. P. clandestina was, able to spread between 15–30 mm through pasteurized soil within a period of 20 days.     

Patchiness and disturbance: plant community responses to porcupine diggings in the central Negev

We tested the hypothesis that diversity and productivity of herbaceous plant communities in disturbed soil are related to the physical and biological heterogeneity of the landscape Our study was earned out on vegetation responses in porcupine diggings on a rocky slope in the central Negev desert in Israel We measured aboveground bio-mass and plant density per species in 150 porcupine diggings (15 cm deep and 15 to 20 cm wide) and in equally sized adjacent control samples in the undisturbed soil matrix We calculated mean annual biomass production, plant density and species richness for 10 sample areas along the slope In addition, we divided the plants into groups according to propagule size and dispersal mode We denoted two types of landscape heterogeneity, which we called physical and biological patchiness Physical patchiness was measured as the ratio of bare rock to soil surface Biological patchiness was the area of the soil covered by shrubs with associated soil mound and under-story relative to the total soil surface We also measured disturbance density, as the long term (17 yr) average density of newly made porcupine diggings We found that 1) the physical patchiness explained 30% of the variation of biological patchiness along the slope, while 2) the patterns of disturbance intensity and biological patchiness were similar (R-=0 386) 3) Biomass, density and species richness were significantly higher in diggings than m the soil matrix 4) Plant density in the matrix, but not m the diggings, was significantly correlated with physical patchiness, 5) species richness in diggings was significantly correlated with biological patchiness, but 6) biomass production in diggings and matrix was not affected by either physical or biological patchiness of the landscape 7) Disturbance density did not affect vegetation responses in diggings and matrix 8) A shift in the plant communities in the matrix towards plants with smaller seeds was associated with increasing physical patchiness, while m diggings there was an opposite shift 9) The proportion of wind dispersers was higher in diggings than outside, while the proportion of runoff dispersers was lower, 10) the densities of runoff dispersers in diggings and matrix were positively correlated with physical and biological patchiness 11) Physical and biological patchiness formed the two major gradients of species composition, explaining 30 and 25% respectively We conclude that the network of physical and biological patchiness and soil disturbance are important in the redistribution of resources and seeds, which control plant biomass, density, species richness and diversity The bare rock surface is the main source for runoff flow with associated soil, organic matter and nutrients The understory vegetation of shrubs provides seeds for creating and maintaining diversity The soil matrix absorbs runoff flow, and disturbances absorb runoff and trap seeds Thus, differences in landscape heterogeneity and their effects on resource and seed movement interact in controlling plant community productivity and diversity in the landscape