The effect of shrub clearing and grazing on the composition of a Mediterranean plant community: functional groups versus species

Abstract. Fuel-breaks to impede the spread of fires in Mediterranean woody vegetation are created by clearing of shrubs, followed by very intensive cattle grazing before the fire season. The present research analysed the effects of these two disturbance types on herbaceous community composition at the functional group and the species level, by using canonical discriminant ordination analysis and by categorical modeling methods. Grazing caused an increase in the abundance of small species, geophytes, and species with an early flowering period that ends before the grazing period. The abundance of many different species increased as a result of clearing. However, no association was found between positive or negative response to clearing and any species attribute tested. Several a priori functional groups defined by life form and family showed responses to either grazing or clearing. Links between these responses and individual species attributes are discussed. The results emphasize the different nature of the two disturbances: grazing as a selective agent, and shrub clearing as a generalized one. It also stresses the importance of the plant architecture and the morphological and phenological niche in determining community composition under extremely intensive grazing conditions.     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

Vegetation-environment relationships in a Negev Desert erosion cirque

The relationship of desert vegetation to topographic and edaphic factors in Makhtesh Ramon, an erosional cirque in the Negev Desert of Israel, was analyzed using redundancy analysis ordination. Altitude and geological substrate had the most significant correlations with vegetation patterns. In particular, significant and monotonic increases in both total species richness and the percentage of annual species were found with increasing altitude. A comparison was made of the effectiveness of Whittaker's diversity sampling plots (Shmida 1984) versus the log-series survey method of McAuliffe (1990). We concluded that at small sample sizes, the Mc Auliffe plots were more likely to yield interpretable ordination results, even though they included only the perennial flora.     

Secondary succession after fire in a Mediterranean dwarf-shrub community

Abstract. Changes in vegetation composition after fire in a Sarcopoterium spinosum dwarf-shrub community on phosphorus-deficient terra rossa soil overlying limestone were studied over a 10-yr period. Herbicide application and phosphate fertilizer were combined to provide a range of initial conditions for recovery. The site was undisturbed during the growing season in winter but was grazed by cattle in summer. Encroachment rate of S. spinosum was mainly a function of the density of viable buds on plants that survived the fire and/or the shrub control treatments. This, together with the very low rate of shrub establishment from seed set a short-to-medium term limit to the attainable cover of S. spinosum. The leguminous shrub Calycotome villosa appeared only in the third year after the fire, apparently from seed. After shrub control, the encroachment rate of C. villosa was low, even after 9 yr. On the other hand, where no herbicide was applied, and with the cessation of goat grazing, C. villosa became increasingly prominent, to the extent that by the 8th year after the fire, it inhibited the continued expansion of S. spinosum. Phosphorus nutrition tended to reduce the cover of S. spinosum because of its stimulating effect on the predominantly leguminous herbaceous vegetation. C. villosa also appeared to benefit from greater phosphorus availability, despite greater competition from the annual vegetation. These findings show that, depending on the context, secondary succession after fire can take different courses. Whereas the combined effect of herbicide control and P-enrichment cannot prevent shrub encroachment, it can slow down the rate of succession for more than a decade, despite the presence of ample shrub seed sources in the area.     

Stimulation of seed germination in an abandoned hay meadow

Abstract. In order to stimulate germination from the soil seed bank in a formerly species-rich hay meadow for conservation purposes, eight gaps were made in each of four habitat types: (1) Dry/open, (2) Dry/shaded, (3) Moist/open and (4) Moist/ shaded; the meadow was abandoned 12 yr before the experiment started. The mean tree cover of the gaps was reduced from 50 % to 22 %. Light was estimated as distance to nearest tree towards south and shade as tree cover. Soil moisture content was estimated from topography and measured from soil samples. There were no significant differences in total herb number between strata before the experiment started. The number of the target group herb indicators of old, traditional management, was however higher in dry, open areas than in shaded areas whether moist or dry. After the treatment, 15 herbs appeared anew in the gaps, nine of these were indicators of traditional management. 80 % of the indicators present after treatment were either new or had increased in frequency in the gaps. More indicators of traditional management appeared in gaps in the open, on dry soils in slopes than in shaded gaps, on moist soils and flat surface. In order to stimulate germination from the seed bank of as many herb indicators of traditional management as possible in abandoned hay meadows of the region, dry, sloping areas, situated as far as possible from large trees towards south and in the open, should be chosen for making gaps for germination.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

Do rivers function as corridors for plant dispersal?

Abstract. We evaluated the importance of dispersal for species frequencies and distribution by comparing dispersal properties of vascular plant species with their frequencies along river banks. We assumed that species with long-floating seeds would be more frequent than species with short-floating seeds. We compiled data on frequencies of vascular plants and their dispersal properties from ten rivers in northern Sweden and compared these with boreal forests and grasslands in the same region. In all rivers, but in none of the reference areas, there was a positive relationship between floating capacity and frequency of species. A comparison of floating capacity between species with and without certain dispersal devices showed that seeds of vegetatively dispersed species had higher floating capacities than other seeds. For other dispersal categories (animal and wind dispersal), floating time did not differ from contrast groups. The results indicate that water dispersal has a certain role in structuring the riparian flora, and provide a basis for explaining species distribution patterns from dispersal characteristics. They also suggest that continuous river corridors are important for maintaining regional biodiversity.     

The role of seed rain in determining the assembly of a cliff community

Abstract. Cliff-face communities of the Niagara Escarpment in Ontario, Canada, are dominated by long-lived Thuja occidentalis and a consistent assemblage of other plants. Our objective was to determine whether seed rain plays a role in determining why these species are dominant. Seed rain was collected from the cliff face and from the surrounding plateau-and talus communities at two sites over a 2-yr period in order to compare these three adjacent, but different communities. Multivariate Discriminant Analysis first separated the two sites: primarily due to the importance of Betula papyrifera at one site. The three community types were also separated, although there was still substantial overlap. There was a predictable array of species associated with each community although the seed rain on the cliff faces differed slightly. When characterized by univariate ANOVAs, seed rain in the cliff faces and plateaus had a lower species richness and lower total seed density than the talus sites. Seeds of two of the 11 species analysed individually showed an influence of habitat type on their number. Seed morphology did not influence patterns of seed rain. Finally, there was no correlation between the seed rain and above-ground vegetation in any of the communities. We conclude that the seed rain patterns that exist do not act to filter the plants that form the mature vegetation of cliffs.     

Community-wide germination strategies in an alpine meadow on the eastern Qinghai-Tibet plateau: phylogenetic and life-history correlates

In this study, we built up a database of 633 species (48 families, 205 genera) from an alpine meadow on the eastern Qinghai-Tibet plateau. Our objective was to assess the effects of phylogenetic and life-history (life form, perenniality, seed size, dispersal strategy and period) background on the community-wide germination strategies. We found that the seeds of shrubs, perennials, and well-dispersed plants, and the smaller seeds germinated more and comparatively earlier. In one-way ANOVAs, phylogenetic groups explained 12% of the variance in GT (mean germination time for all seeds germinated of each species); life-history attributes, such as seed size, dispersal strategy, perenniality and life form explained 10%, 7%, 5%, and 1% respectively, and dispersal period had no significant effect on GT. Multifactorial ANOVAs revealed that the three major factors contributing to differences in GT were phylogenetic relatedness, seed size and dispersal strategy (explained 4%, 5% and 4% of the interspecific variation independently, respectively). Thus, seeds germination strategies were significantly correlated with phylogenetic and life-history relatedness. In addition, phylogenetic relatedness had close associations and interactions with seed size and dispersal strategy. Then, we think phylogeny and life-history attributes could not be considered mutual exclusively. Seed germination, like any other trait, is shaped by the natural history of the species and by the evolutionary history of the lineage. And a large percentage of the variance remained unexplained by our model, which suggested important selective factors or parameters may have been left out of this analysis. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Plant sexual systems, dichogamy, and herkogamy in the Venezuelan Central Plain

Sexual system, temporal variation in sex expression, and herkogamy were evaluated in 210 plant species in the Venezuelan Central Plain (VCP). This analysis was conducted considering the life forms, habitats, flowering time, pollination systems, and dispersal syndromes of the plant species. Hermaphrodite species dominated the community (75.2%; N=158), followed by monoecious taxa (17.1%; N=36), and then dioecious (7.6; N=16). Most species were adichogamous (=homogamous) (75.2%; N=155), followed by protandrous (16.5%; N=34), and protogynous (8.3%; N=17). The frequency of herkogamous species was higher (N=124; 60.2%) than the non-herkogamous (N=82; 39.8%). The three reproductive traits interacted significantly in a way that complements their effectiveness in promoting cross-pollination and avoidance of pollen–stigma interference: hermaphrodite-adichogamous species occurred in similar frequency to herkogamous and non-herkogamous species; monoecious-adichogamous species were mostly herkogamous, and all hermaphrodite, functionally dioecious-adichogamous, species were herkogamous. Sexual system, herkogamy, and dichogamy are associated with habitats in the savanna-forest mosaic of the VCP. The proportion of dioecious species decreased from trees to annual herbs and from forest to disturbed areas. Dichogamous species were most frequent for herbaceous plants, and non-herkogamous species dominated in annual herbs. The frequency of herkogamous species decreased from forest to disturbed areas. Sexual system was significantly associated with pollination system. Most dioecious and monoecious species had generalist pollination. A three-factor log linear analysis indicated that for any combination significantly. Sexual systems and herkogamy were associated with the respective seed dispersal syndrome. Most dioecious and herkogamous species were dispersed by frugivores, while monoecious plants tended to be dispersed by granivores and wind. Flowering time of dioecious species peaked during the dry–rainy transition period, whereas that of monoecious species was greatest during the rainy season. Therefore, specific biotic and abiotic conditions may enforce the occurrence of unisexuality in the community. In a similar way, dichogamous species tend to flower during the rainy season, associated with herbaceous life form and wind pollination. These results are discussed in the context of evolutionary and ecological trends.     

Recruitment limitation in secondary forests dominated by an exotic tree

Question: What factors limit woody plant recruitment in a mosaic landscape where former agricultural lands are dominated by the invasive tree Ligustrum lucidum (Oleaceae)? Location: Subtropical northwestern Argentina. Methods: In secondary forest patches, we measured (1) tree, shrub and liana abundance in different size classes; (2) seed rain of Ligustrum and two native trees and (3) topographic, soil and light variables. We used spatial autoregressive models to test for effects of Ligustrum dominance and environment on native plant abundance in each size class. We used multiple regression on resemblance matrices to quantify the relative importance of spatial (e.g. dispersal) and environmental effects on native species composition. Results: Native tree abundance in the smallest size class was unrelated to Ligustrum canopy dominance, while native tree abundance in larger size classes and native liana abundance were negatively correlated with Ligustrum dominance. Native species composition was both environmentally and spatially structured, suggesting that some species are dispersal limited. Seed rain was spatially correlated with conspecific basal area for one of two native species, but not for Ligustrum. Conclusions: Native tree recruitment appears to be limited primarily by sapling mortality in patches dominated by the invasive Ligustrum. Ligustrum does not appear to be dispersal limited in our study area and is likely to continue spreading. Invaded patches may persist for hundreds of years.     

Colonization dynamics and facilitative impacts of a nitrogen‐fixing shrub in primary succession

Abstract. Whether nitrogen‐fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen‐fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria‐induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria‐enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self‐replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre‐Coriaria stages. These soil changes resulted in three‐ to sixfold increases in growth of Griselinia when it was grown in Coriaria‐enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.     

Secondary succession in abandoned fields of dry tropical Northern Cameroon

Abstract. This study evaluates the processes of recolonization of abandoned fields by native vegetation under conditions of intensive human activity (fire, intensive grazing, firewood cutting) in a semi-arid tropical region savanna of northern Cameroon. Secondary plant succession was studied in two series of formerly cultivated fields 1–35 yr after the beginning of the fallow period. Floristic changes and the dynamics of woody plant populations were compared between areas with vertisols (clay texture) and sandy soils, as a function of length of fallows. Vegetation changed continuously during the 35 years following field abandonment. However, a very abrupt break occurred between 6 and 10 yr, due to increasingly intense human pressure during this period. Up to that point, ecological models and mechanisms of succession presented in the literature are more or less confirmed by our results. Usually, secondary succession is blocked at a stage of wooded grassland as a result of human activities.     

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.     

Four decades of old field vegetation development and the role of Celastrus orbiculatus in the northeastern United States

Abstract. This paper describes four decades of in situ vegetation change in a post-agricultural site in southern New England where the role of the introduced vine, Celastrus orbiculatus, has drastically modified the course of vegetation development. An initial perennial forb-grassland community was replaced by two strikingly different physiognomic vegetation types-an early hardwood forest and a vine community occupying the same site 40 yr later. This ecological phenomenon was unpredictable since Celastrus was not recorded at the site until the 1970s. Within the young hardwood forest dominated by Prunus serotina and Acer rubrum, a few old field species (five) persist, whereas 17 forest herbs have become established. Typical forest shrubs are rare or absent. The Celastrus-dominated vine community has suppressed or eliminated pre-existing growth to form a relatively stable community where a forest vegetation is the regional vegetation type. Celastrus also appears to be facilitating a native vine (Vitis labrusca) by forming a ‘ladder’ for its advance. Although A. rubrum and Quercus spp. could ultimately dominate the forest community, the contiguous and aggressive nature of Celastrus makes predictions questionable. This long-term study highlights the unpredictable nature of old field succession and the role of an exotic species in threatening the natural vegetation development.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

Plastic bet-hedging in an amphicarpic annual: an integrated strategy under variable conditions

Amphicarpy is a form of diversified bet-hedging expressed mostly in annual plants, where two types of offspring are produced with two distinct ecological roles: long-range aerial dispersers and highly competitive subterranean, sedentary fruit. Emex spinosa is a semi-arid, amphicarpic annual, inhabiting habitats with different levels of environmental variation. We tested the hypothesis that, in E. spinosa, bet-hedging may be “fine-tuned” by plasticity in the phenotype ratio (aerial/subterranean fruit mass) as a function of environmental conditions. We conducted a greenhouse experiment, manipulating nutrient availability and intraspecific density, to determine the pattern of ratio shifts. In order to determine whether the integrated strategy is an adaptation to variable habitats, a similar common garden experiment was conducted, comparing two natural populations differing in environmental variability. The offspring ratio shifted in response to both nutrient availability and plant density. In pots containing single plants the ratio increased steeply with nutrient availability, while in pots containing eight plants a more moderate increase occurred. These shifts were the result of plasticity in allocation to both achene types, as well as ontogenetic effects on aerial achene production. The degree of response increased with the heterogeneity of the habitat of origin. We found evidence for an adaptive integrated strategy, with bet-hedging “fine-tuned” by phenotypic plasticity. Strenuous conditions tended to shift the offspring ratio towards securing subterranean reproductive success, while favorable conditions resulted in a shift towards dispersible achenes. The authors Asaf Sadeh and Hagai Guterman contributed equally to this study.     

Seed dispersal and seedling emergence in an old field community in central New York (USA)

Summary Seed dispersal and seedling emergence of common taxa growing in a Solidago-dominated old field in central New York (USA) were monitored from May 1982 to June 1984. Over 3.5x10⁴ seeds per m² were captured on seed traps in each of the two years, with peaks occuring in July (due to Hieracium) and in November (due to Solidago). About 4.0x10³ seedling/m² emerged beneath the intact community in each of the two years. Although seedlings emerged predominantly in the early spring, a secondary peak occurred in September and October when many seedlings of introduced grasses appeared. Two additional aspects of the reproductive biology of the major taxa were related to the seasonal timing of seed dispersal: As the date of peak seed dispersal (among taxa) became progressively later in the season, (1) the duration of dispersal increased from about one week to about one-half year, and (2) the delay between the peak of seed dispersal and the peak of seedling emergence increased from a few days to about one-half year.