Resilience of Mediterranean shrubland to a severe drought episode: the role of seed bank and seedling emergence

Extreme climate events, such as severe drought episodes, may induce changes in vegetation if they induce species‐specific adult mortality and changes in the seedling recruitment pattern. In 2005 a severe drought occurred in Doñana National Park (south Spain) causing extensive shrubland mortality. Over the following years we monitored the soil seed bank and seedling emergence via a gradient of canopy dieback induced by the drought episode. The canopy dieback corresponded to an increase in emergence of seedlings of woody species in 2007, probably because of the reduced competition induced by canopy loss. The soil seed bank of woody species sampled in 2008 was less abundant on plots with a higher proportion of dead vegetation, probably because of depletion of the seed bank as a result of the increased germination in the previous year and also as a result of a reduction in seed supply in these sites. Accordingly, in 2009 we detected reduced emergence of woody species on plots that had suffered the greatest shrub mortality. We failed to find any significant changes in patterns of the soil seed bank and seedling emergence of short‐lived herbaceous species, indicating greater resilience in these types of species. This study highlights the resilience of Mediterranean shrublands to climate fluctuations at one extreme of the variability characteristic of these ecosystems. An increase in the frequency of severe drought episodes – increasingly probable under the new climate conditions – does have the potential, however, to induce changes in vegetation, especially in woody communities that need more time to replenish their seed banks.     

Similarity between seed bank and vegetation in Mediterranean grassland: a predictive model

Abstract. The similarity in species composition between seed bank and vegetation was analysed in Mediterranean grasslands in relation to altitude, topography and grazing. Soil samples were collected in permanent plots in autumn at the end of the summer drought period and in spring, before the new seed fall and after the natural winter seed stratification. The seed bank composition was determined by greenhouse germination over a nine-month period. Presence/absence of species in the standing vegetation throughout the complete annual cycle, and the percentage area of bare ground in October, were recorded in the same plots. The species composition of the standing vegetation is clearly determined by altitude, topography and grazing, while the floristic composition of the seed banks is only related to altitude and topography in the case of autumn seed bank and with any of the three factors in the spring seed bank. Relative abundances of grasses, legumes and forbs also show different patterns in vegetation and seed bank data. Sørensen similarity between the autumn seed bank and the vegetation declines as altitude rises, but there are no significant differences for topography and grazing. This similarity decreases in the case of the spring seed bank and does not show any significant relationship with any of the factors. The perennial/ annual ratio and the proportion of bare soil in October are proposed as explanatory variables in a predictive model of similarity between the seed composition of the seed bank and vegetation.     

Year-to-year Oscillations in Demography of the Strictly Biennial <Emphasis Type="Italic">Pedicularis sylvatica</Emphasis> and Effects of Experimental Disturbances

In 1998–2001, I studied disturbance effects on the population structure and dynamics of a grassland strict biennial Pedicularis sylvatica, and on the species demography (monthly dynamics of seedling recruitment in 1998 and within- and between-year survival in 1998–2000). In two Czech populations, I established three experimental disturbance regimes: (1) a gap treatment, that simulated grazing by clipping vegetation and creating small gaps, (2) a mowing treatment, where I clipped the vegetation, and (3) a no management treatment, where I left the vegetation untreated. The number of recruiting seedlings varied greatly by year, and demographic structure of populations showed significant year-to-year oscillations in mean seedling numbers, from low (3 ± 0.7 s.e. per 0.25 m² plot) to high (103 ± 20). Inversely in the same years and plots, mean adult numbers in populations oscillated from high (12 ± 2) to low (0.7 ± 0.3). Disturbance effects were only important for seedling recruitment in early census dates in all years. In 1998, most seedlings recruited in April–May in gaps in both sites, but most died before winter. Within- and between-year survival was not affected by disturbance regimes but fluctuated significantly among years. Between-year survival increased with increasing size of the overwintering bud and was higher in disturbance treatments. Since the oscillations in population structure did not significantly vary in response to experimental disturbances, population dynamics may be driven endogenously rather than by disturbance events. The weak disturbance effects on species demography may also indicate population resilience to changes in habitat quality. However, since disturbances promoted seedling recruitment, grazing or mowing regimes are strongly recommended, as they create regeneration opportunities and maintain habitat quality, meeting the species long-term conservation goals.     

Grassland responses to multiple disturbances on the New England Tablelands in NSW,Australia

Disturbances have played a significant role in shaping vegetation patterns following European settlement and agricultural development in Australia, particularly over the last 100 years. However, little is known about the temporal dynamics of plant communities in relation to disturbances and their interactions. In this study we examined the response and recovery of temperate grassland communities to disturbance, using a multi-factorial experiment involving grazing exclusion (absent and present), fire (absent and present), soil cultivation (none, 5 and 20 cm) and soil amendment (none, fertiliser and fertiliser plus clover seeds) on the New England Tablelands in NSW, Australia. Temporal patterns of plant communities were analysed using detrended correspondence analysis for eight surveys over 24 months. Vegetation patterns at each survey were analysed using principal components analysis. The effects of treatments on malleability (Bray–Curtis dissimilarity) of plant communities were analysed using a linear mixed model, and the relationships between malleability and plant species groups were depicted using a generalised additive model and further analysed using a linear model. Perennial native grasses and a non-native forb (Hypochaeris radicata) initially dominated the vegetation, but after the disturbances H. radicata and other non-native species dominated. Compositional changes from the initial states were greatest in the first spring (7 months after treatment application), and then the vegetation tended to recover towards its original state. Soil cultivation resulted in the greatest deviation in community composition, followed by soil amendment, fire, with grazing exclusion the least. The recovery process and rate of recovery varied with treatment reflecting the dominance of soil cultivation and its interactions with other forms of disturbances. Soil amendment and grazing exclusion tended to reduce the effects of soil cultivation. Malleability was negatively related to perennial native grasses; positively to other non-native species, annual native grasses and perennial native sedges/rushes; and negatively to H. radicata when its cover was below 18%, but positively when above 18%. The degree of malleability reflected the high resilience of the vegetation to disturbance, and was mainly due to the recovery of perennial native grasses and H. radicata. This resilience demonstrated that the small-scale disturbances did not cause vegetation to cross an ecological threshold and that the present vegetation is resilient to common disturbances occurring at small scales. The results also suggest (1) that the present vegetation has developed mechanisms to adapt to these disturbances, (2) the importance of disturbance scale or (3) that the ecological threshold had already been crossed and the present vegetation is in a degraded state compared with its original state before the end of the 19th century.     

Differential Indirect Effects of Excluding Livestock and Rabbits from Chalk Heath on the Associated Leafhopper (Hemiptera: Auchenorrhyncha) Fauna

Preliminary results are presented of sampling the leafhopper assemblages on a field experiment designed to examine the differential effects of rabbits and livestock (mainly sheep) on the vegetation of chalk heath in southern England. Experimental plots that excluded livestock either allowed entry by rabbits or excluded them. Results were compared with those from plots grazed by both livestock and rabbits. After 7 years, exclusion of grazing herbivores had resulted in predictable increases in vegetation height, but no major changes were detected in the species composition of the vegetation. As expected, ungrazed plots had higher species richness and greater abundances of several individual leafhopper species. However, plots grazed only by rabbits had a leafhopper assemblage that was distinct from either ungrazed or mixed grazing plots. It is suggested that rabbit grazing may have subtle effects on grassland invertebrate assemblages that are not necessarily predictable from an examination of the species composition of the vegetation. Chalk heath vegetation contains an unusual mixture of calcicole and calcifuge plant species, but the leafhopper assemblage included a restricted number of calcareous grassland specialist species and only one species strongly associated with acidic grasslands; most leafhoppers recorded were generalist grassland species.     

Regenerative role of seed banks following an intense soil disturbance

Our main aim was to determine the contribution of the seed bank to vegetation regeneration following a disturbance consisting in a deep ploughing and a thorough homogenisation of a perennial grassland. In the seed bank prior to disturbance, seed distribution through the vertical soil profile was evaluated to determine the initial seed species structure. Then, several characteristics of the shallow seed bank and the extant vegetation were evaluated prior and following field disturbance: seed species composition and abundance, and species composition of the aboveground vegetation. The contribution of seed rain versus seed bank was evaluated by means of the comparison of the vegetation developed in plots filled with sterilised soil (seed bank removal) and the vegetation developed in non-sterilised plots in the field. The distribution of seeds through the profile indicated a sharp decline in abundance with depth, and it was probably linked to propagule morphology, with small and rounded seeds proner to being buried deeper than larger seeds. In the grassland prior to disturbance, the aboveground vegetation and seed bank species composition showed very low similarity index, most likely because during the 5 years following field abandonment, sheep pressure had caused a faster change in aboveground vegetation species composition than in seed bank species composition. Ploughing and homogenisation of the grassland led to low seed abundance in the shallow soil layer caused by dilution of the seed bank. Regardless of impoverishment in seed abundance and species richness, comparison between sterilised and non-sterilised plots showed that the seed bank acted as an effective source of colonising species and determined the aboveground species composition. To summarise, this study outlines the importance of considering several characteristics of the seed bank, such as species composition and seed abundance, in the understanding of the function of seed bank and dynamics of the vegetation following a deep ploughing and homogenisation treatment.     

The role of drought,disturbance, and seed dispersal in dominance shifts in a temperate grassland

Aims

More intense and frequent droughts predicted for the future will heavily affect grasslands worldwide and, along with various other disturbances, possibly trigger major vegetation shifts. Therefore, documenting these changes and understanding the mechanisms behind them is essential. Our study aimed to investigate how dominant species in a semiarid grassland respond to a combination of small-scale experimental disturbances and naturally occurring droughts and uncover the underlying mechanisms.

Location

Central Hungary.

Methods

We conducted a small-scale disturbance experiment in a semiarid temperate grassland and followed recovery for 18 years, including severe drought events. In 1999, we established 16 sites, eight dominated by Festuca vaginata and another eight by Stipa borysthenica, the two dominant species of sand grasslands in the region. At each site, three permanent plots were marked that received either a cutting or digging treatment or remained as controls. We monitored the cover and density changes of Festuca and Stipa annually.

Results

In the early years following the disturbance, Festuca recovered at a similar rate under both disturbance treatments, while Stipa recovered faster in cut than in dug plots. When natural drought events caused major diebacks of both species, Stipa recovered very quickly and regained dominance in initially Stipa-dominated plots, and it also took over in initially Festuca-dominated control and cut plots. However, digging at Festuca-dominated sites delayed drought-induced Stipa colonisation and thus favoured Festuca recovery. We found that the poor performance of Stipa in dug plots was related to sharply reduced seedling establishment, which resulted from the low number of seeds captured by the bare soil surface after digging.

Conclusions

Our results demonstrate that recurring drought events may induce dominance shifts in temperate grasslands, but small-scale disturbances can modulate vegetation responses. Our findings emphasise the importance of post-disturbance regeneration patterns in drought-induced vegetation shifts and show that seed dispersal strategy may have a major effect on vegetation dynamics.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

Role of soil seed bank along a disturbance gradient in an alpine meadow on the Tibet plateau

We examined the role of the soil seed bank along a grazing disturbance gradient and its relationship with the vegetation of alpine meadows on the Tibet plateau, and discussed the implications for restoration. The seed bank had a high potential for restoration of species-rich vegetation; 62 species were identified in the vegetation and 87 in the seed bank, 39 species being common to both. Mean seed density was 3069–6105 viable seeds m⁻². The density of buried seeds increased significantly with increasing disturbance, indicating that restoration of disturbed areas is not seed limited. Seed density and species richness decreased with depth. The proportion of perennial species decreased with decrease in disturbance both in seed bank and in vegetation. A large portion of species with persistent seeds in the disturbed areas indicate that this seed type can be regarded a strategy of adaptation to current disturbances. Detrended correspondence analysis (DCA) showed significant differences of species composition between seed bank and vegetation, except for the seriously disturbed site. Our results suggest that the establishment of new species in severely disturbed areas is more dependent on the seed bank. By contrast, the restoration in less-disturbed and mature meadows does not rely on seed banks, and the establishment of the vegetation in these communities is more likely to rely on seed dispersal from the standing vegetation and on species with vegetative reproduction.     

Vole mound effects and disturbance rate in a mediterranean plant community under different grazing and irrigation regimes

A factorial field experiment was used to assess the influence of soil-disturber mammals in the structure of a 9-year-old Mediterranean annual plant community subjected to different sheep grazing and irrigation regimes. We estimated the disturbance rate (mound building activity) by Mediterranean voles, their effects on vegetation and the mechanisms of these effects during a period of vole outbreak. The effects on vegetation were analysed at the levels of species, functional groups and plant community. Disturbance rate was high and voles can disturb the entire soil surface once every four or five years. The availability of certain trophic resources (perennial plants) appeared to drive vole expansion in the experimental plots and it was independent of the irrigation and grazing treatments. Mound building activities largely affected vegetation but conserved plot differences. Total vegetation cover, absolute cover of all functional groups, mean vegetation height and species richness were less on mounds than on undisturbed ground. These effects did not change the relative abundance of annuals, perennials, grasses and forbs. Only the relative abundance of small-seeded species decreased on mounds. As the proportion of these seeds was similar in both types of patches, we suggest that small-seeded species had more difficulties for germinating or emerging when they are buried during mound formation. Irrigation and sheep grazing promoted large changes in the vegetation parameters but these effects were, in general, similar on mounds and undisturbed ground. Our results show that the availability of germinable seeds may be the major limitation for mound revegetation, probably due to the scarcity of seeds existing at the depths from which soils are excavated. Our results also suggested a resource limitation on mounds. The results provide additional evidence that soil disturbances by small herbivore mammals exert relevant ecological effects on abandoned Mediterranean croplands. We discuss the ecological implications of vole mound-building activities for plant succession, plant species conservation and forage resource availability for livestock. This revised version was published online in June 2006 with corrections to the Cover Date.     

Dynamics of grazing lawn formation: an experimental test of the role of scale‐dependent processes

Grazing lawns are characteristic for African savanna grasslands, standing out as intensely grazed patches of stoloniferous grazing‐tolerant grass species. Grazing lawn development has been associated with grazing and increased nutrient input by large migratory herds. However, we argue that in systems without mass migrations disturbances, other than direct grazing, drive lawn development. Such disturbances, e.g. termite activity or megaherbivore middens, also increase nutrient input and keep the bunch vegetation down for a prolonged time period. However, field observations show that not all such disturbances lead to grazing lawns. We hypothesize that the initial disturbance has to be of a minimal threshold spatial scale, for grazing intensity to be high enough to induce lawn formation. We experimentally tested this idea in natural tall savanna grassland. We mowed different‐sized plots to simulate initial disturbances of different scales (six times during one year) and applied fertilizer to half of the plots during two years to simulate increased nutrient input by herbivores or termite activity. Allowing grazing by naturally occurring herbivores, we followed the vegetation development over more than three years. Grazing kept bunch grass short in coarser, fertilized plots, while grasses grew out toward their initial height in fine‐scale and unfertilized plots. Moreover, lawn grasses strongly increased in cover in plots with an increased nutrient input but only after coarser scale disturbance. These results support our hypothesis that an increased nutrient input in combination with grazing indeed induces grazing lawn formation, but only above a threshold scale of the initial disturbance. Our results provide an alternative mechanism for the development of grazing lawns in systems that lack mass migrating herds. Moreover, it gives a new spatial dimension to the processes behind grazing lawn development, and hence help to understand how herbivores might create and maintain spatial heterogeneity in grassland systems.     

Re-establishment of Calluna vulgaris (L.) Hull in an eight-year grazing experiment on upland acid grassland

Upland heathland is an internationally important habitat but a large area in the UK has been degraded to acid grassland by intensive livestock grazing. Re-establishment of dwarf shrubs, particularly Calluna vulgaris (L.) Hull, is a key objective for restoring heathland on these sites. A replicated plot-scale experiment was set up to examine effects of disturbance and seed addition on C. vulgaris establishment in a Nardus stricta L. grassland under three grazing regimes: sheep only (1.5 ewes ha^?1 for 10 months per year); cattle only (0.5 heifers ha^?1 in summer only); and, the cattle regime combined with sheep (1.0 ewes ha^?1 for 10 months per year). Early results of the experiment have been reported previously but it was not known if these results were an indication of the longer-term restoration success. Here we evaluate the success of the restoration methods (disturbance, seeding treatments and grazing regime) eight years after the treatments began. In seeded plots, young C. vulgaris plants had greatest above-ground height, dry weight and shoot length if grazing was excluded or the cattle-only regime was applied. C. vulgaris cover was greatest, and increased most, in plots that had been disturbed, seeded and ungrazed or subjected to the cattle-only regime. The vegetation in these plots also became more similar to reference sites with 50% or more cover of C. vulgaris. The invasive Juncus effusus L. was more frequent in disturbed and grazed plots but less frequent in plots with C. vulgaris established from added seed. Previous results that showed the benefits of disturbance and seeding treatments were still valid but changes in the vegetation composition were still occurring and longer-term studies will be needed to determine when grazing regimes including sheep might be reintroduced.     

Effect of sheep grazing on seed circulation on the Loess Plateau

In grazing ecosystems, mature seeds fall directly to the soil to form the soil seed bank (SSB), or are ingested by grazing livestock to become part of the dung seed bank (DSB; i.e., seed circulation). Both the SSB and DSB form the basis for the natural regeneration of vegetation. However, little is known about the relationships between the SSB, DSB, and aboveground vegetation (AGV) community under different stocking rates (SRs). This study investigated the relationships between the SSB, seeds in Tan sheep (Ovis aries) dung, and AGV at different SRs (0, 2.7, 5.3, and 8.7 sheep ha^–1) in a semiarid region of the Loess Plateau in China. We found that Tan sheep grazing increased the species richness heterogeneity of grassland vegetation, and negatively influenced the density of AGV. Under natural conditions, 17 species from soil‐borne seeds and 10 species from Tan sheep dung germinated. There was low species similarity between the soil and DSBs and AGV. Sheep SR and the seed banks (soil and dung) were negatively correlated with AGV. Seeds are cycled from herbage to livestock to soil during cold season grazing; the seasonal nature of this seed dispersal is an adaptation to harsh, semiarid environments. Increased seed bank diversity under sheep grazing facilitates grassland regeneration on the Loess Plateau, similarly to other semiarid regions globally.     

Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NW Greece)

We investigate the persistent soil seed bank composition and its relation to the above-ground flora of grazed and non-grazed sub-Mediterranean deciduous oak forests of NW Greece. Twenty-eight taxa were recorded in the soil seed bank and 83 taxa (70 taxa in plots of seed bank sampling) in the above-ground vegetation. The dominant tree species and many woodland species found in the above-ground vegetation were absent from the soil seed bank. Similarity between the soil seed bank and the above-ground vegetation decreased with grazing, and grazing led to a decrease of species richness in above-ground vegetation and soil seed bank. Beta diversity of vegetation among grazed and among non-grazed plots did not differ, but was significantly higher between grazed and non-grazed areas. Beta diversity of the soil seed bank declined with grazing. When applying classification tree and logistic regression analyses, non-grazed forest sites are clearly differentiated by the presence of Phillyrea latifolia, Euphorbia amygdaloides and Brachypodium sylvaticum. PCA ordination of above-ground species composition reflected a gradient from sites grazed by ruminants to non-grazed sites, but no clear structure was detected in the seed bank.     

Plant traits and regeneration of urban plant communities after disturbance: Does the bud bank play any role?

Questions: What is the relative role of the bud bank, seed and various species traits in the regeneration of urban plant communities after severe disturbances? Do invasive and exotic species, highly abundant in disturbed communities, regenerate better than native species after disturbance? Methods: Hand tilling was applied to three urban plant communities with and without additional herbicide treatment to exclude regeneration from the bud bank. Plant traits were determined from the literature and databases. Species responses to the treatments were evaluated with RDA analyses in CANOCO. Linear models were applied to identify traits that could predict the responses of species to disturbance. Results: The bud bank played a key role in regeneration in the plots without herbicide. In the plots with herbicide treatment, the seed bank was important in re‐establishing vegetation after disturbance. Exclusion of the bud bank by using herbicide allowed the establishment of small annuals, whereas biennials and perennials were successful in plots where the bud bank was not inhibited by herbicide. Exotic species with a long residence time in the local flora were successful in plots where regeneration from the bud bank was excluded, whereas species with short residence times or that were invasive were suppressed by both types of disturbance. Conclusion: In response to various types of disturbance, species with different regeneration strategies (either seeds or bud bank) were promoted. Exotic species were suppressed primarily by disturbance, which suggests that factors other than just regenerative capability contributed to the high abundance of exotics in urban communities.     

The positive and negative conservation impacts of sheep grazing and other disturbances on the vascular plant species and vegetation of lowland subhumid Tasmania

Summary An important conservation question for grazed areas of lowland subhumid Tasmania is ‘what effects do different, practical disturbance regimes have on native vegetation?’ An experiment designed to determine the single and interactive effects of fire and sheep grazing was established at four sites with distinct vegetation types. There were significant interactive effects of fire and sheep grazing on vegetation attributes at all sites. An analysis of published and new data indicated that there were several vascular plant species that appeared dependent on sheep grazing for their persistence in the present landscape, while there were others that were intolerant of this disturbance but required other types of disturbance, such as mowing. However, most native species appeared to survive in a wide variety of disturbance regimes short of ploughing and fertilization. The implications of these results are that a variety of disturbance regimes is necessary to maintain biological diversity in this environment, and that the naturalness of the regime is not necessarily relevant to its use for conservation.     

Do plant-community-based grazing regimes lead to epizoochorous dispersal of high proportions of target species?

The contribution of large-herbivore epizoochory to the transfer of seeds within and between areas is thought to be significant. But often seeds of ubiquitous species are dispersed, which may enhance ruderalization processes. In order to study the dispersal of target species by sheep, we employed a community-based grazing approach followed by intra- and inter-area sheep transfers (maximum transfer distance 3 km). In case of inter-area transfers, well-developed target communities of an open inland sand ecosystem are used as “source”, linked to less-developed sand habitats (“sink”) via sheep. Also other factors determining which species become dispersed under field conditions were tested: seed surface structure, seed mass, plant seed-releasing height and animal behaviour. Finally the influence of animal movement on seed detachment and the actual arrival of seeds within a “sink” were studied.

Sheep transfers resulted in the dispersal of 56 seed species, dominated by Red List (seven species) and other target species. Quantitatively, most transported seeds belonged to target species, whereas graminoid competitors were highly under-represented. Morphological traits enhance the attachment probability regardless of seed mass. But for seeds without these epizoochory-facilitating traits, mass seems to affect attachment negatively. Plant height affected the number of species present in sheep coats but not the seed quantities. Probably certain species in the vegetation produced large numbers of seeds, e.g. low-growing Medicago minima with seed surface structures and high-growing species Verbascum phlomoides without seed surface structures. Also, although transfer half-times were three times lower than grazing half-times, naturally attached Stipa capillata and Agrimonia procera seeds showed no significant detachment rates during transfer, whereas considerable losses were found during grazing. Other 3-km sheep transfers did not result in significant losses of either epizoochorously transported seed or species quantities. Our study shows that community-based grazing can lead to the dispersal of especially target species. Besides that, different habitat fragments can be connected to each other via sheep without significant seed losses along the way.     

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

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

Shrubland Restoration Following Woody Alien Invasion and Mining: Effects of Topsoil Depth, Seed Source, and Fertilizer Addition

Invasion by woody alien plants, construction, and mining operations are among the major disturbances degrading vegetation in the Cape Floristic Kingdom, South Africa. The aim of this study was to assess whether native fynbos shrubland vegetation could be restored following dense alien invasion and disturbance by mining. An area supporting dense alien trees was cleared and topsoil was stripped and stockpiled to simulate mining disturbance. A field trial investigated the effects of topsoil depth, seed mix application, and fertilizer on native species recruitment and vegetation development over a three‐year period. Soil‐stored seed banks contributed 60% of the species recruited, indicating that areas invaded for three decades have good restoration potential. The addition of a fynbos seed mix, which included serotinous overstory species, improved both the richness and structural composition of the vegetation. Most species sown in untopsoiled plots established, but survival and growth was low compared to topsoil plots. Poor growth in combination with a lack of soil seed bank species, indicate that restoring a diverse and functional cover of indigenous vegetation on subsoil is not possible in the short‐term. Soil amelioration is required to improve rooting conditions and initiate ecosystem processes. Shallow and deep topsoil treatments yielded high plant density, richness, and projected canopy cover, but canopy cover was higher in deep topsoil plots throughout the trial. Fertilizer addition increased canopy cover in untopsoiled and shallow topsoil plots via an increase in alien annual species. Fertilizer addition ultimately may lead to increased native vegetation cover in untopsoiled areas, but as it increased proteoid mortality on deep topsoil plots, it is not recommended for sites where topsoil is available. A species‐rich and structurally representative fynbos community may be restored on topsoiled areas provided that the native disturbance regime is simulated and seeds of major structural guilds not present in the soil seed bank are included in the seed mix.     

Species richness increases the resilience of wetland plant communities in a tropical floodplain

In the last two decades, the relationship between diversity and stability/ecosystem functioning has been widely discussed and has become a central issue in ecology. Here, we assessed the relationship between wetland plant diversity and community resilience after a disturbance. Our study area was located in the Upper Paraná River floodplain (Brazil). An experiment was carried out in situ (18 1 m × 1 m plots with richness varying from 1 to 18 species). In each plot, we recorded the number of species, total per cent vegetation cover and per cent age cover of each species. The above‐ground biomass of wetland plants was removed, simulating a disturbance by animal trampling or an extreme flood. The recovery of vegetation was monitored over 3 months. According to a linear regression, the recovery of wetland plants was positively correlated with diversity. Comparisons with plots containing monocultures of one of the dominant species (Polygonum stelligerum) suggested that this species did not overyield in mixed cultures. Thus, our experiments indicate that the higher resilience in richer plots after a disturbance is mainly due to the fact that species have different resource use requirements (complementarity effect) and not due to the presence of a single, more productive species. Our experiment carried out in a more real condition (in situ) showed that biodiversity is important to wetland functioning and stability, paralleling the results obtained in laboratory and mesocosms experiments. These results also suggest that the loss of plant diversity in our study area could compromise community recovery following strong disturbances.