Comparison of three seed trap types in a chalk grassland: toward a standardised protocol

In a chalk grassland in north-western France, vegetation and seed rain were studied along two transects. We compared the efficiency of three types of seed traps (funnel traps, sticky traps, pots exposed in the field) to estimate the annual seed rain and to reveal various processes involved in plant community regeneration. Data were analysed to compare seed density, species richness and composition across seed traps and vegetation. Geostatistical analyses (semivariograms. were used to detect possible autocorrelation and to examine patterns of spatial variation in seed rain and vegetation. The results show that (1) funnel traps are most efficient (56 species, 8079 seeds.m^–2). (2) Each type of seed traps brings different information about the processes involved in seed dispersal and seedling establishment. Sticky traps help to understand import and export of seeds in a community through anemochory. Funnel traps may provide a measure of the auto-regeneration capacity of the vegetation through its local seed production. Pot traps integrate various processes involved between seed dispersal and seedling emergence. (3.) The exposition to environmental conditions generates marked spatial patterns in species richness measured in pots and in vegetation, selecting species able to survive. Pot traps indicate that numerous species producing seeds in standing vegetation are unable to establish in the field through seedling recruitment. Our study emphasizes the need of using different types of seed traps and spatial sampling configurations to assess the regenerative potential of standing vegetation and the various mechanisms involved in seed dispersal. A standardised protocol for seed rain sampling was finally developed.     

Productivity and species richness across an environmental gradient in a fire-dependent ecosystem

The fire-dependent longleaf pine-wiregrass (Pinus palustris Mill.-Aristida beyrichiana Trin. & Rupr.) savannas of the southeastern United States provide a unique opportunity to examine the relationship between productivity and species richness in a natural ecosystem because of the extremely high number of species and their range across a wide ecological amplitude (sandhills to edges of wetlands). We used a natural gradient to examine how plant species richness and plant community structure vary with standing crop biomass (which in this system is proportional to annual net productivity) as a function of soil moisture and nitrogen mineralization rates in a frequently burned longleaf pine-wiregrass savanna. Highest ground cover biomass and highest species richness were found at the same position along the gradient, the wet-mesic sites. Relative differences in species richness among site types were independent of scale, ranging from 0.01 m(2) to 100 m(2). Nitrogen availability was negatively correlated with species richness. Dominance of wiregrass (in terms of biomass) was consistent across the gradient and not correlated with species richness. Regardless of site type, the community structure of the savannas was characterized by many perennial species with infrequent occurrences, a factor in the low temporal heterogeneity (percent similarity between seasons and years) and high within-site spatial heterogeneity (percent dissimilarity of vegetation composition). The coexistence of numerous species is likely due to the high frequency of fire that removes competing hardwood vegetation and litter and to the suite of fire-adapted perennial species that, once established, are able to persist. Our results suggest that soil moisture is an important factor regulating both the number of species present and community production within the defined gradient of this study.     

Relationship between grass species richness and ecosystem stability in Kruger National Park, South Africa

The relationship between grass species richness and ecosystem stability was investigated in the Kruger National Park. A total of 135 489 individual grasses were identified from 189 sites spread across 19 485 km² of savanna biome, making this one of the largest studies of its kind. At each site, grass percentage abundance and standing crop were measured at one year intervals to provide an index of ecosystem function. A severe drought that affected the region between 1991 and 1993 provided a convenient perturbation. At the height of the drought, mean species richness declined by 12.7% while standing crop declined by 38.1%, from 3199 to 1979 kg ha^?1. Percentage grass abundance declined to 87.5% of its pre‐drought value. After the drought had passed species richness, standing crop and percentage abundance recovered to 92.1%, 113.8% and 92.8% of their pre‐perturbation values, respectively. Statistical analysis of these responses revealed that grass assemblages of low species richness were more resistant to drought than assemblages of high species richness. Species‐poor sites also showed better recovery from perturbation after the drought had passed. These findings suggest that ecosystem stability may be negatively related to grass species richness in South African savanna grasslands.     

Inter-relationships between standing crop, biodiversity and trait attributes of hydrophytic vegetation in artificial waterways

1. The extensive British canal system potentially provides a favourable habitat for aquatic plants and, because of its uniformity, it is ideal for studying relationships between disturbance intensity (largely from boat traffic) and communities of aquatic macrophytes. 2. The standing crop and species composition of aquatic plants were measured in summer at 396 sites distributed randomly over the canal network. We also quantified the number of rare species and ‘attribute groups’ (groups of species sharing similar suites of biological traits). These data were analysed in relation to standing crop, assumed to indicate a disturbance gradient. 3. Consistent with unimodal models and the Intermediate Disturbance Hypothesis, maximum hydrophyte species richness occurred at an intermediate biomass (50–200 g DWm^?2). This corresponded to a low frequency of low magnitude disturbance (light boat traffic) on navigable sections, or occasional high magnitude disturbance (the post‐interventionist phase) on sections currently unnavigable and subject to active vegetation management. The frequency of rare species was also the greatest under these conditions, reflecting the availability of regeneration niches. 4. Sorting of species into attribute groups revealed that the overall relationship between species richness and standing crop comprised of closely overlapping unimodal responses of nine attribute groups, superimposed on a core vegetation of Potamogeton pectinatus, together with greater representation of filamentous algae, lemnids and elodeids with increasing standing crop (i.e. assumed low disturbance). High species richness was associated with the overlap of functionally different groups of species, rather than with disturbance‐mediated coexistence of functionally similar plants. 5. The analysis of a matrix of sites and the representation of plant traits weighted by the biomass of species that displayed them, in relation to different aspects of disturbance, highlighted a shift from traits associated with resilience (turions, unanchored floating or submerged leaves, low body flexibility, budding, small body size), or competitiveness (entire leaves, low reproductive output, high biomass density, large body size) at high standing crop, through to attributes more associated with resistance to disturbance (rhizomes, tubers, streamlining of foliage, low biomass density) at low standing crop. 6. Comparison with a stochastic null model of change in species number along a constrained gradient, after correction for variation in sampling effort, indicated that sites towards the tails of the gradient (excluding those with extremely low biomass) supported more species than might be expected from chance alone, while the most species‐rich sites in mid‐gradient generally supported many fewer species than expected. 7. We suggest that a disturbance regime that maintains intermediate standing crops would be appropriate for the conservation of species‐rich aquatic vegetation. Precise definition of this regime, under a range of circumstances, requires the study of temporal change at representative sites.     

Richness of species and growth-forms within sclerophyll and mesophyll vegetation in eastern Australia

Abstract The patterns in total species richness and in the richness of the dominant growth-forms of vegetation communities of coastal sclerophyll and mesophyll vegetation in eastern Australia are examined. Plant species richness data were obtained from two 500 m² quadrats from 50 sites within a single geographical region north of Sydney, New South Wales. Concentrically nested subquadrats within each quadrat enabled the determination of species-area relationships for total species richness and its components. Three growth-forms were examined (trees, shrubs and ground cover) and patterns in the richness of these components were compared to those exhibited by total species richness. Total species richness was higher in sclerophyll communities on Hawkesbury Sandstone soils than in adjacent mesophyll communities on Narrabeen shales and sandstones. Significant patterns in total species richness within the two soil types were also found. Shrub and ground cover species richness showed strong correlations with total species richness, with higher richness in the sclerophyll communities. However, tree species richness contributed little to the patterns in total species richness. The results of this study suggest that differential patterns in the components of total species richness must be taken into account for effective modelling of natural areas based on species richness and diversity parameters.     

Effects of fire regime on plant species richness and composition differ among forest,woodland and heath vegetation

Question

Do the effects of fire regimes on plant species richness and composition differ among floristically similar vegetation types?

Location

Booderee National Park, south‐eastern Australia.

Methods

We completed floristic surveys of 87 sites in Sydney Coastal dry sclerophyll vegetation, where fire history records have been maintained for over 55 years. We tested for associations between different aspects of the recent fire history and plant species richness and composition, and whether these relationships were consistent among structurally defined forest, woodland and heath vegetation types.

Results

The relationship between fire regime variables and plant species richness and composition differed among vegetation types, despite the three vegetation types having similar species pools. Fire frequency was positively related to species richness in woodland, negatively related to species richness in heath, and unrelated to species richness in forest. These different relationships were explained by differences in the associations between fire history and species traits among vegetation types. The negative relationship between fire frequency and species richness in heath vegetation was underpinned by reduced occurrence of resprouting species at high fire frequency sites (more than four fires in 55 years). However, in forest and woodland vegetation, resprouting species were not negatively associated with fire frequency.

Conclusions

We hypothesize that differing relationships among vegetation types were underpinned by differences in fire behaviour, and/or biotic and abiotic conditions, leading to differences in plant species mortality and post‐fire recovery among vegetation types. Our findings suggest that even when there is a high proportion of shared species between vegetation types, fires can have very different effects on vegetation communities, depending on the structural vegetation type. Both research and management of fire regimes may therefore benefit from considering vegetation types as separate management units.     

Relationship between light availability and species richness during fen grassland succession

The complex relation between standing crop and species richness in herbaceous plant communities is still one of the most intensively discussed topics in vegetation ecology. In this study, we focus on the extent to which light availability is able to explain this relationship in fen grasslands. We analyse changes of light availability (measured as relative irradiance, RI), standing crop and species richness during fen grassland succession. Our study include communities representing differences in drainage intensity and nutrient status (‘land use intensification sere’) and successional stages developing after abandonment (‘abandonment seres’). Both in the land use intensification sere and in the abandonment seres, we recorded an increase in standing crop reaching highest values of approx. 800 g m^‐2. RI, species richness and number of small‐growing species declined in the successional seres. RI was highest in stands of low‐productive communities, where light compensation points (5% RI) were already attained at soil surface and light saturation points (30% RI) at soil surface or at 15 cm height. Horizontal heterogeneity of RI at soil surface and at 30 cm height decreased in the abandonment seres, but not at 30 cm height in the land use intensification sere. Furthermore, we detected a signiñcant negative relation between standing crop and RI. Species richness declined with increasing standing crop and thus consequently also with decreasing RI. This result points out that light competition might be of importance for this pattern. The number of small‐growing species decreased exponentially with an increase in standing crop (and decrease in RI). It can be assumed that low light availability negatively affects small‐growing species at standing crop values higher than 400–500 g m². As the maintenance of small‐growing species and the improvement of habitat conditions for their establishment are important aims of nature conservation in fen grasslands, management strategies should be orientated at maintaining standing crops that are not higher than these values.     

Effect of local and regional processes on plant species richness in tallgrass prairie

Historically, diversity in a community was often believed to result primarily from local processes, but recent evidence suggests that regional diversity may strongly influence local diversity as well. We used experimental and observational vegetation data from Konza Prairie, Kansas, USA, to determine if: (1) there is a relationship between local and regional richness in tallgrass prairie vegetation; (2) local dominance reduces local species richness; and (3) reducing local dominance increases local and regional species richness. We found a positive relationship between regional and local richness; however, this relationship varied with grazing, topography and fire frequency. The decline in variance explained in the grazed vegetation, in particular, suggested that local processes associated with grazing pressure on the dominant grasses strongly influenced local species richness. Experimental removal of one of the dominant grasses, Andropogon scoparius , from replicate plots resulted in a significant increase in local species richness compared to adjacent reference plots. Overall all sites, species richness was higher in grazed (192 spp.) compared to ungrazed (158 spp.) areas. Across the Konza Prairie landscape, however, there were no significant differences in the frequency distribution of species occurrences, or in the relationship between the number of sites occupied and average abundance in grazed compared to ungrazed areas. Thus, local processes strongly influenced local richness in this tallgrass prairie, but local processes did not produce different landscape-scale patterns in species distribution and abundance. Because richness was enhanced at all spatial scales by reducing the abundance of dominant species, we suggest that species richness in tallgrass prairie results from feedbacks between, and interactions among, processes operating at multiple scales in space and time.     

Assessing the level of plant invasion: A multi-scale approach based on vegetation plots

Recent studies highlight the importance of selecting the appropriate scale and indices of invasion level for evaluating the abundance and impact of alien plants. Our survey considers the use of vegetation plot databases compared with floristic checklists to address invasion patterns regarding alien–native relationships across vegetation types by means of a multi-scale approach. We analysed the alien–native richness relationship in 1077 vegetation plots from the Basque Country (N. Spain) at ecosystem level and phytosociological class and alliance levels. According to our results, the alien species richness (Alo)–native species richness (Nat) relationship is variable and depends not only on the scale but also on the vegetation type. In contrast with other multi-scale approaches, no negative correlation has been detected at any studied level. The strong correlation existing between plot number and cumulative Alo and cumulative Nat highlights the constraints of using checklists to generalize invasion patterns. Our results demonstrate that the combined use of both relative alien species richness and relative alien species cover facilitates the understanding of invasion patterns across plant communities at different scales. In addition to climate, disturbance and propagule pressure, habitat type proved to be an important filter for alien species, capable of explaining such patterns.     

雅鲁藏布江河岸植物物种丰富度分布格局及其环境解释

沿环境梯度物种多样性的分布格局及其环境影响因子的研究是生态学研究的重点内容之一。雅鲁藏布江是我国海拔最高的大河, 目前还没有针对其流域的河岸植物群落物种丰富度分布格局的系统研究。作者沿海拔梯度从雅鲁藏布江上游源头区至下游共设置了15个海拔样带, 在每个海拔样带随机取6个5 m×5 m样方, 共对90个样方的河岸植物群落物种组成和植被类型进行了调查, 共记录到238种维管束植物。DCA排序结果表明物种组成变化主要受海拔和经度控制。CCA排序区分出5种大的植被类型, 即高寒草甸或高寒草原、高山草原、两种亚高山灌丛草原和林缘灌丛植被。各植被类型的分布和变化主要受海拔和经度影响。雅鲁藏布江河岸植物物种丰富度沿海拔和经度梯度呈浅“U”形分布格局, 物种丰富度与年均降水量和植被盖度成显著正相关。     

Fine‐scale vascular plant species richness in different alpine vegetation types: relationships with biomass and cover

Abstract. Vascular plant species richness was related to biomass and vegetation cover in nine different alpine vegetation types on the Hardangervidda mountain plateau, western Norway. Each vegetation type was sampled within an 8m × 6m area, and the species‐richness pattern analysed. Evidence for a unimodal relationship between species richness and both biomass and cover was found at the within‐vegetation type scale. Cover was a better predictor for species richness than biomass, suggesting that light may be an important factor influencing species richness at this scale in alpine vegetation. The possibility that the results are an artefact of small grain size is also discussed, and several arguments for an ecological explanation of the humpback relationship between species richness and cover are discussed.     

Standard survey designs drive bias in the mapping of upland swamp communities

Vegetation maps are critical biodiversity planning instruments, but the classification of vegetation for mapping can be strongly biased by survey design. Standardization of survey design across different vegetation types is therefore increasingly recommended for vegetation mapping programs. However, some vegetation types have complex small‐scale vegetation patterns that are important in characterizing these vegetation types, and standard designs will often not capture these patterns. The objective of this paper was to investigate the magnitude of potential map bias that results from survey design standardization and recommend approaches to deal with this bias. We surveyed upland swamps of the Greater Blue Mountains World Heritage Area Australia using two contrasting survey designs, including the standard 400 m² single quadrat design recommended and used by authorities. We then derived a classification for these swamps and tested the effect of survey design on this classification, species richness and the type of species detected (obligate or facultative swamp species). Species richness and species type were not significantly different among survey techniques. However, more than 40% of swamps clustered differently among survey designs. Thus, one of the 10 derived communities (which is floristically consistent with a previously mapped endangered community) was indistinct, and some individual swamps misclassified using the standard survey design. An effect of landscape position on swamp floristic patterns and a significant trend for high similarity scores among swamps surveyed with multiple small quadrats compared to the standard survey design was also determined. Australian upland swamps are classified at the global scale as shrub‐dominated wetlands, and complex floristic patterns have been recorded in shrub‐dominated wetlands in both northern and southern hemispheres. We therefore advocate either multiple survey designs or different survey standards for upland swamp communities and other vegetation types that have complex floristic patterns at small scales.     

Composition and seasonal flux of the soil seed bank of species-rich Themeda triandra grasslands in relation to burning history

Abstract. Most species-rich grasslands dominated by Themeda triandra in southeastern Australia have been ungrazed and frequently burnt for decades. The seedling emergence technique was used to determine the size and taxonomic composition of the soil seed bank of five grasslands that had different fire histories (i.e. burnt at 1 yr, 3 yr and > 10 yr intervals) and this was compared to the standing vegetation at each site. A nested sampling design (subplot, plot, site) was used to determine the effect of spatial scale on the patterns observed in both the vegetation and the seed bank. Temporal variation in the seed bank was assessed by repeated soil sampling over a two year period. 61 native and 30 exotic species were recorded in the vegetation. Richness varied more between sites than within sites. Sites were therefore internally homogeneous for species richness. However, no correlation between burning frequency and richness was found. DCA ordination separated the sites into distinct groups, but sites with similar fire history did not necessarily group closely. 60 taxa germinated from the soil seed bank, comprising 32 native and 28 exotic species; 11 species, mostly therophytes, were restricted to the seed bank. The richness of the seed bank was significantly lower than the vegetation at all spatial scales. No correlation between seed bank richness and fire history was found. The seed bank of species-rich grasslands is dominated by a limited number of widespread, highly clumped, annual, native and exotic monocots. Most native hemicryptophytes, and perennials in general, were represented in the soil by a transient seed bank. Only 12 % of study species, all therophytes, were considered to form large, persistent seed banks, the size of which was greater in unburnt grasslands at all times of the year. The distinct floristic patterns observed in the vegetation are less clearly represented in the seed bank. The seed bank represents a floristically distinct (and less variable) component of the vegetation when compared to the standing flora. The size of the long-term soil seed bank suggests that it has little functional importance for many native species and probably contributes little to seedling regeneration processes following disturbance. Altering established fire regimes is likely to only change the composition and small-scale richness of the existing site vegetation and will not re-integrate species previously lost from the vegetation due to past management. It is suggested that the maintenance of vegetation remnants and processes that encompass a range of long-term burning histories will be necessary if the flora is to be conserved in situ. Restoration of degraded grasslands cannot rely on the soil seed bank but rather, will be dependent on the reintroduction of propagules.     

Aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis>) stands and their contribution to plant diversity in a semiarid coniferous landscape

We conducted a field study to determine the relative contributions of aspen (Populus tremuloides), meadow, and conifer communities to local and landscape-level plant species diversity in the Sierra Nevada and southern Cascade Range, northeastern California, USA. We surveyed plant assemblages at 30 sites that included adjacent aspen, conifer, and meadow communities across a 10,000-km² region. We statistically investigated patterns in local and landscape-scale plant diversity within and among the three vegetation types. Summing across sites, aspen stands supported more plant species overall and more unique plant species than either meadow or conifer communities. Local richness and diversity did not differ between aspen and meadow plots; conifer forest plots were significantly lower in both measures. Heterogeneity in species composition was higher for aspen forest than for meadows or conifer forest, both within sites and between sites. Plant communities in aspen stands shared less than 25% of their species with adjacent vegetation in conifer and meadow plots. Within aspen forest, we found a negative relationship between total canopy cover and plant diversity. Our results strongly support the idea that plant communities of aspen stands are compositionally distinct from adjacent meadows and conifer forest, and that aspen forests are a major contributor to plant species diversity in the study region. Current patterns of aspen stand succession to conifer forest on many sites in the semiarid western US are likely to reduce local and landscape-level plant species diversity, and may also have negative effects on other ecosystem functions and services provided by aspen forest.     

Relationships of marsh seed banks to vegetation patterns along environmental gradients

• 1 The relationship of the seed bank to the vegetation of a freshwater marsh was studied along gradients of water depth and soil organic matter content. Characters examined included standing crop, seedling density, and species composition, distribution and richness.
• 2 The seed bank differed from the vegetation in that only nine of twenty-seven species were present in both, abundant seed-bank species were uncommon as adults, and adults showed different distributions along a gradient of soil organic matter content whereas their seeds were most abundant in soils with high organic matter.
• 3 The seed bank resembled the vegetation in that separate multivariate analyses of the communities revealed that variation in the species composition of each was significantly correlated with water depth and soil organic matter content. Further, species richness in both communities decreased with water depth and increased with soil organic matter content. Lastly, the standing crop of the vegetation and the number of seedlings both decreased with water depth and increased with soil organic matter.
• 4 Consideration of spatial patterns and environmental gradients revealed more similarities between vegetation and seed banks than were obtained by comparing species lists. The results suggest that artificial stimulation of seed bank germination for management purposes will not produce vegetation changes as large as those suggested by differences in species lists.

     

Diversity and productivity of plant communities across the Inland Northwest,USA

No definitive explanation for the form of the relationship between species diversity and ecosystem productivity exists nor is there agreement on the mechanisms linking diversity and productivity across scales. Here, we examine changes in the form of the diversity–productivity relationship within and across the plant communities at three observational scales: plots, alliances, and physiognomic vegetation types (PVTs). Vascular plant richness data are from 4,760 20 m² vegetation field plots. Productivity estimates in grams carbon per square meter are from annual net primary productivity (ANPP) models. Analyses with generalized linear models confirm scale dependence in the species diversity–productivity relationship. At the plot focus, the observed diversity–productivity relationship was weak. When plot data were aggregated to a focus of vegetation alliances, a hump-shaped relationship was observed. Species turnover among plots cannot explain the observed hump-shaped relationship at the alliance focus because we used mean plot richness across plots as our index of species richness for alliances and PVTs. The sorting of alliances along the productivity gradient appears to follow regional patterns of moisture availability, with alliances that occupy dry environments occurring within the increasing phase of the hump-shaped pattern, alliances that occupy mesic to hydric environments occurring near the top or in the decreasing phase of the curve, and alliances that occupy the wettest environments having the fewest species and the highest ANPP. This pattern is consistent with the intermediate productivity theory but appears to be inconsistent with the predictions of water–energy theory.     

Environmental factors covary with plant diversity–productivity relationships among Chinese grassland sites

Aim Our objective was to document the general relationship between plant species richness (SR) and above‐ground net primary productivity (ANPP) at different spatial scales and the environmental influence on this relationship. Location Temperate and alpine grasslands of China. Methods We investigated SR and ANPP at 321 field sites (1355 plots) across the widely distributed temperate and alpine grasslands of China. Ordinary least squares (OLS) regressions were used to test SR–ANPP relationships among site means. Plot‐level data of SR and ANPP were analysed with general linear models (GLMs) and the correlation between SR and ANPP was decomposed into covariance components to test the influence of climatic variables, region, vegetation type and remaining variation among sites on SR, ANPP and their relationship. Results We found positive linear relationships between SR and ANPP among sites in both the alpine and temperate grassland regions and in different grassland vegetation types of these biomes. Environmental gradients such as growing‐season precipitation affected both SR and ANPP in parallel. However, after removing the among‐site environmental variation, residual SR and ANPP were no longer correlated at the pooled within‐site level. Main conclusions The positive SR–ANPP relationship across large‐scale environmental gradients among sites was most likely the result of climatic variables influencing SR and ANPP in parallel. Our results suggest that in China's natural grasslands there is no direct relationship between SR and ANPP, presumably because the pool of available species for local community assembly is large, in contrast to experiments where species pools are artificially reduced.     

Spatial Variability in Litterfall,Litter Standing Crop and Litter Quality in a Tropical Rain Forest Region

Understanding the spatial variability in plant litter processes is essential for accurate comprehension of biogeochemical cycles and ecosystem function. We assessed spatial patterns in litter processes from local to regional scales, at sites throughout the wet tropical rain forests of northern Australia. We aimed to determine the controls (e.g., climate, soil, plant community composition) on annual litter standing crop, annual litterfall rate and in situ leaf litter decomposability. The level of spatial variance in these components, and leaf litter N, P, Ca, lignin, α‐cellulose and total phenolics, was determined from within the scale of subregion, to site (1 km transects) to local/plot (~30 m²). Overall, standing crop was modeled with litterfall and its chemical composition, in situ decomposability, soil Na, and topography (r² = 0.69, 36 plots). Litterfall was most closely aligned with plant species richness and stem density (negative correlation); leaf decomposability with leaf‐P and lignin, soil Na, and dry season moisture (r² = 0.89, 40 plots). The predominant scale of variability in litterfall rates was local (plot), while litter standing crop and α‐cellulose variability was more evenly distributed across spatial scales. Litter decomposability, N, P and phenolics were more aligned with subregional differences. Leaf litter C, lignin and Ca varied most at the site level, suggesting more local controls. We show that variability in litter quality and decomposability are more easily accounted for spatially than litterfall rates, which vary widely over short distances possibly in response to idiosyncratic patterns of disturbance.     

Nested spatial patterns in seed bank and vegetation of Mediterranean old-fields

A nested sampling design was used to describe the spatial patterns for the species richness and composition in the seed bank and vegetation of three Mediterranean old-fields (1, 7, and 15 yr after the last ploughing). Three scales were examined hierarchically: sampling units within plots of 0.25 m² for the vegetation and of 0.05 m² for the seed bank, 100 m² plots within fields, and fields of 1000 m². In spite of the strong spatial variation among sampling units, species richness and composition of both seed bank and vegetation showed hierarchically structured patterns of heterogeneity, while each old-field was a homogeneous entity. These spatial patterns tended to be partially masked when the data were aggregated at the scale of the plot. Such results stress the use of a nested sampling design for studying variation in species richness and taxonomic composition in both vegetation and seed bank. This design, in combination with CCA, also showed that the vegetation showed a coarser grain than the seed bank, probably in relation to seed clumping.     

Composition and species diversity of pine-wiregrass savannas of the Green Swamp,North Carolina

Fire-maintained, species-rich pine-wiregrass savannas in the Green Swamp, North Carolina were sampled over their natural range of environmental conditions and fire frequencies. Species composition, species richness, diversity (Exp H′, 1/C), and aboveground production were documented and fertilization experiments conducted to assess possible mechanisms for the maintenance of high species diversity in these communities. Although savanna composition varies continuously, DECORANA ordination and TWINSPAN classification of 21 sites facilitated recognition of 3 community types: dry, mesic, and wet savannas. These savannas are remarkably species-rich with up to 42 species/0.25 m² and 84 species/625 m². Maximum richness occurred on mesic, annually burned sites. Aboveground production, reported as peak standing crop, was only 293 g · m^?2 on a frequently burned mesic savanna but was significantly higher (375 g · m^?2) on an infrequently burned mesic site. Production values from fertilized high and low fire frequency sites were equivalent. Monthly harvest samples showed that savanna biomass composition by species groups did not vary seasonally, but within groups the relative importance of species showed clear phenological progressions. The variation in species richness with fire frequency is consistent with non-equilibrium theories of species diversity, while phenological variation in production among similar species and the changing species composition across the moisture gradient suggest the importance of equilibrium processes for maintenance of savanna diversity.