Soil seed bank and species turnover in a limestone grassland

Abstract. The vegetation and soil seed bank in a limestone grassland in southern Sweden were studied in permanent 1-m² subplots which had been either grazed or not grazed for 17 yr. Of the 92 species recorded, 18 were present only in the seed bank, 28 were more frequent and 24 were less frequent in the seed bank than in the vegetation and 22 were not detected in the seed bank. Among the species present in the seed bank, therophytes were over-represented. Species turnover in the vegetation was estimated from presence/absence data collected in 1980, 1986 and 1990. Turnover was high, but there were no differences between grazed and ungrazed subplots. The turnover for individual species was also high in many cases. There was no clear relationship between the turnover of a species and its presence in the persistent seed bank.     

Community assembly and biomass production in regularly and never weeded experimental grasslands

We studied the natural colonisation of new species in experimental grasslands varying in plant species richness (from 1 to 60) and plant functional group richness (from 1 to 4) in either regularly or never weeded subplots during the first 3 years after establishment. Sown species established successfully, with no differences in species richness or their relative abundances between the regularly and never weeded subplots during the study period. Aboveground biomass of sown species increased with increasing sown species richness in both treatments. While a positive relationship between sown species richness and total aboveground biomass (including colonising species) existed in the 2nd year after sowing in the regularly and never weeded subplots, this positive relationship decayed in the 3rd year in the never weeded subplots because of a higher biomass of colonising species in species-poor mixtures. Total aboveground biomass varied independently of total species richness 3 years after sowing in both treatments. Jaccard similarity of coloniser species composition between regularly and never weeded subplots decreased from the 2nd to the 3rd year, indicating a divergence in coloniser species composition. Coloniser immigration and turnover rates were higher in regularly weeded subplots, confirming that weeding counteracts species saturation and increases the chance that new colonisers would establish. Although our study shows that low diversity plant communities are unstable and converge to higher levels of biodiversity, the effects of initially sown species on community composition persisted 3 years after sowing even when allowing for succession, suggesting that colonising species mainly filled empty niche space.     

Tree species richness and turnover throughout New Zealand forests

Abstract. Patterns of mortality, recruitment, and forest turnover were investigated using permanent plot data from temperate forests in 14 localities throughout New Zealand. Tree mortality and recruitment rates were calculated from tagged trees ≥ 10 cm diameter at 1.4 m on individual 400 m² plots, and turnover rates were calculated as the mean of mortality and recruitment rates. Turnover rates (1.4% per year) were very similar to those recorded for tropical forests (i.e. 1.5% per year). As was shown in tropical forests, we also found significant relationships between forest turnover and species richness. In New Zealand forests there was also a decrease in species richness and turnover rates with increasing latitude. Although species richness is well known to decline with latitude, our study provides support for a possible link between seasonality and disturbance with tree turnover and species diversity. While tree mortality and recruitment rates were approximately in balance at some localities, in others there were imbalances between mortality and recruitment rates.     

Annual Glyphosate Treatments Alter Growth of Unaffected Bentgrass (Agrostis) Weeds and Plant Community Composition

Herbicide resistance is becoming more common in weed ecotypes and crop species including turfgrasses, but current gaps in knowledge limit predictive ecological risk assessments and risk management plans. This project examined the effect of annual glyphosate applications on the vegetative growth and reproductive potential of two weedy bentgrasses, creeping bentgrass (CB) and redtop (RT), where the glyphosate resistance (GR) trait was mimicked by covering the bentgrass plants during glyphosate application. Five field plots were studied in habitats commonly inhabited by weedy bentgrasses including an agricultural hayfield, natural meadow, and wasteland. Results showed that annual glyphosate treatment improved bentgrass survivorship, vegetative growth, and reproductive potential compared with bentgrass in unsprayed subplots. In the second year of growth, RT plants had an 86-fold increase in flower number in glyphosate-treated subplots versus controls, while CB plants had a 20-fold increase. At the end of the three year study, plant community composition had changed in glyphosate-treated subplots in hayfield and meadow plots compared to controls. Soils in subplots receiving glyphosate had higher nitrate concentrations than controls. This is the first study to mimic the GR trait in bentgrass plants with the goal of quantifying bentgrass response to glyphosate selection pressure and understanding the impacts on surrounding plant communities.     

Small-scale spatial dynamics of plant species in a grassland community over six years

Abstract. In a species-rich mountain grassland in the Krkonosse Mts., Czechoslovakia, data from four permanent plots of 50 cm x 50 cm were recorded annually from 1985 to 1990 to study the spatial dynamics of the species. Plots were divided into 15 x 15 subplots and the number of vegetative units of all plants within each subplot was determined. There was not much net change at the plot level, but the subplots were very dynamic. Two aspects of the spatial dynamics of the species were followed: (1) persistence, i.e. the tendency of the species to remain in the same subplot, and (2) long-distance spreading, i.e. movement to subplots beyond the immediate neighbourhood. Species differed widely in their persistence and longdistance spreading and were classified into mobility types: long-range guerrilla, short-range guerrilla, phalanx and 'sitting’. The mobility types were, to a certain extent, correlated with the growth form of plants, but some species of one growth form showed different types of small-scale dynamics and some species with different growth forms had the same spatial dynamics.     

High species mobility in species-rich plant communities: An intercontinental comparison

Using results from a long-term study of fine-scale dynamics in grasslands in four widely separated study areas from two continents, we provide further evidence to support the idea of the carousel model as an aid to describe the high fine-scale temporal and spatial species mobility found in grassland communities. Cumulative species numbers on small subplots in plots situated in stable plant communities, determined as the sum of species appearing in these subplots in one or more years over a period of time, are very high. In floristically different species-rich grasslands, varying from moist pine savannas in North Carolina and Mississippi, to humid chalk grassland in the Netherlands and seasonally dry limestone grassland in Sweden, average species numbers on subplots of 0.01 m² in plots of 2.5 m² over the period 1985–1989 were similar, most plots falling in the range 10.8–13.2. The total cumulative species numbers were similar as well, most plots falling in the range 17.4 and 20.9. Yearly average species numbers remained relatively constant. Considerable species turnover is occurring in all these communities; on average three species appear and three disappear each year in each 0.01 m² subplot. Total species accumulation on 0.01 m² subplots over the period 1985–1989 varied considerably, from 4.1 to 11.6, and is correlated with the cumulative species total on the plot, the latter figure being considered as correlated with the size of the species pool.     

哀牢山中山湿性常绿阔叶林林窗更新研究

对云南哀牢山中山湿性常绿阔叶林林窗更新状况的调查结果表明,林内优势树种在林窗阶段不占优势,其在林窗内的重要值显著低于成熟林分;林窗内外的乔木幼苗在组成上存在较大差异,林窗内的幼苗多样性(H=2．73,D=0．89)高于林冠下(H=2．25,D=0．82),二者的Jaccard相似系数只有0．6．许多树种的幼苗只出现在林窗内;林窗内的乔木幼苗密度(38636株·hm²)远远高于林冠下(17323株·hm²)．林窗内和林冠下的乔木幼苗(H≤50cm)均以高度为11～20cm幼苗的在数量上占优势,但林窗内高度超过20cm的幼苗比例明显高于林冠下;林窗大小对乔木幼苗更新有重要影响,不同树种的幼苗密度高峰出现在不同大小级别的林窗内．     

Temporal and spatial patterns in emergence and early survival of perennial plants in the Sonoran Desert

Seedling emergence and survival of 15 perennial species were studied for six years in a 557-m² permanent plot at Tumamoc Hill, Arizona, USA, an ungrazed site in the northern Sonoran Desert. The minimum rain required for germination and emergence ranged from 17.5 to 35.6 mm. Few species emerged in every year of the study. First-year survival averaged across all 15 species was 3.7%; only 0.1% of seedlings lived as long as four years. The odds of survival in the first year improved with increased rain. About three times as many seedlings died from predation as desiccation. In 2-m² subplots, mortality of three woody species in the first 30 days after emergence appeared to be independent of seedling density. Short-, moderate-, and long-lived species displayed distinct survival strategies. Long-lived species compensated for generally poor seedling survival by frequent germination and emergence. Moderate-lived species exhibited highly episodic germination and emergence, a potentially risky behavior that might have been offset to some extent by relatively good long-term survival. Short-lived species had the highest seedling survival. Because these species can bloom in their first year, good early survival meant that some individuals were able to reproduce before they died.     

Demography of the long‐lived conifer Agathis ovata in maquis and rainforest,New Caledonia

Abstract. The endemic New Caledonian conifer Agathis ovata occurs as an emergent tree in fire‐prone shrublands (maquis), and fire‐sensitive rainforest. Growth, survivorship and recruitment over 5 yr were compared for populations from forest and maquis on ultramafic substrates in New Caledonia to investigate whether demographic behaviour varied in response to the strongly contrasting forest and shrubland environments. Growth of seedlings and of small (30–100 cm height) and large (100 cm height; 5 cm DBH) saplings was slow, but varied significantly among stages, site types and years. The greatest difference in growth rates was among stages, seedlings growing 0.34 cm.yr^?1, small saplings 1.06 cm.yr^?1 and large saplings 2.13 cm.yr^?1. Tree DBH increased by only 0.05 cm.yr^?1 and, based on these rates, individuals with DBH of 30 cm are estimated to be more than 700 yr old. Few trees (3.5%) produced cones in any year and seedling recruitment was low, but some recruitment was recorded each year in both maquis and forest. Rates of recruitment per parent were highest in forest (1.28.yr^?1, cf 0.78.yr^?1), but the higher density of trees in maquis meant that overall recruitment was greater there (92 ha^?1.yr^?1, cf 56 ha^?1.yr^?1). Seedling mortality ranged from 0.9 to 2.9% among years with no significant difference between maquis and forest. No sapling mortality was recorded, but annual tree mortality ranged from 0 to 1.4%. Evidence from a recently burned site indicated that while trees may survive fire, seedlings and saplings do not. Post‐fire seedling recruitment per ha from surviving trees was four times lower than in unburned sites, but growth rates were four times higher. Similar demographic attributes, including high survivorship, low growth rate and low rates of recruitment over a long reproductive life, characterize Agathis ovata populations in both maquis and rainforest in New Caledonia and are indicative of a broad tolerance of light environments that is unusual among tree species. These demographic attributes help to explain the long‐term persistence of the species in these strongly contrasting habitats.     

SEED BANK OF A FRESHWATER TIDAL WETLAND: TURNOVER AND RELATIONSHIP TO VEGETATION CHANGE

Depletion during spring germination (turnover), longevity, and successional relationships were studied at High Marsh (HM), Cattail (CT), and Shrub Forest (SF) sites in a freshwater tidal wetland over three years. There was significant seasonal reduction in size and composition of seed banks from all sites. Turnover was greatest in HM surface (0–2 cm) samples where 29 x more seeds germinated in March than in June. In CT and SF samples turnover was considerably less. Magnitude (34–97%) was related to species composition and factors affecting field germination. Decrease in density with depth (0–10 cm) was log-linear in March samples. Except for SF 30–32 cm, few seeds and species were found at 8–10, 15–17, or 30–32 cm. Three seed bank strategies were distinguished: (a) complete turnover (Type II, sensu Thompson and Grime 1979), (b) high turnover with some reserve (Type III), and (c) large long-term seed reserve (Type IV). Longevity of many species appeared to be restricted; 31–56% at each site were present only in surface samples, and 29–52% germinated only in March samples. Although some species were important at all three sites, the seed bank composition of each was distinct, and was related to vegetation composition at each site. Because the bulk of the seed bank at each site resembles its site vegetation more than that of another site, it is not possible to predict succession.     

Strategies for Mixed-Grass Prairie Restoration: Herbicide, Tilling, and Nitrogen Manipulation

Large areas of North American prairie have been planted with grasses introduced from Eurasia. We examined three strategies (herbicide, tilling, and nitrogen manipulation) for enhancing the establishment of seedlings of native species and suppressing the introduced grasses Agropyron cristatum (crested wheat grass) and Bromus inermis (smooth brome). Plots (5 × 15 m) were subjected to one of three levels of tilling (none, intermediate, complete) and four levels of nitrogen (none, intermediate, high, and sawdust added to immobilize nitrogen). Treatments were applied in a factorial design with twelve treatments and ten replicates. Seeds of 41 native species were drilled into the plots in May 1992. Following the failure of seeds to establish in 1992, a subplot (5 × 13 m) within each main plot was sprayed with the herbicide glyphosate in April 1993. The nitrogen treatments were repeated in Spring 1993. In August 1993, the density of native seedlings in sprayed subplots was 20 times that in unsprayed subplots. Within sprayed subplots, native seedling density and the cover of bare ground decreased significantly with increasing nitrogen availability. Plots receiving sawdust had significantly higher mean cover of bare ground and significantly lower concentrations of soil available nitrogen. Native seedling density was significantly higher in plots receiving the highest intensity of tilling. The responses of native seedlings to all these factors point to the importance of neighbor-free establishment sites as a prerequisite for prairie restoration.     

Species richness and phenology of vegetation along irrigation canals and drains in the Nile Delta,Egypt

This study evaluates the species richness, phenology and effect of environmental factors on the composition of canal and drain vegetation in the Nile Delta, Egypt. Altogether, 365 stands were sampled along the terraces, slopes, littoral zones and water zones of 28 canals and 10 drains. Smooth species compositional changes were found in the first three zones. The slope plant community has the highest total number of species and species richness and a medium value of species turnover. The water zone has the lowest values of these variables.The number of species with seedlings and vegetative plants had two peaks: one in winter and the second in summer, which was related to the contrasted behaviour of the winter and summer weeds. The number of species with vegetative and fruiting plants was relatively higher, while the number with dormant plants was relatively lower in the canal vegetation when compared to that of the drains. The number of species with vegetative, flowering and fruiting plants increased with the decrease of canal-drain width.     

Two bromeliad species with contrasting functional traits partition the understory space in a South American xerophytic forest: correlative evidence of environmental control and limited dispersal

We examined the spatial distribution of two bromeliad species with contrasting functional traits in the understory of a xerophytic South American Chaco forest. Aechmea distichantha is a facultative terrestrial species with well-developed phytotelma and short rhizomes, whereas Bromelia serra is a strictly terrestrial species with soil-exploring roots and long rhizomes. Both bromeliads develop colonies on relatively elevated patches in Schinopsis balansae forests. We evaluated the roles of environmental controls, limited dispersal, and interspecific competition as drivers of the different distribution of these bromeliads. We mapped the overstory, understory and topography of 16 forest plots with bromeliads (400 m² each, subdivided in 100 4-m² subplots). We sampled soil characteristics on sectors dominated by each bromeliad species. We used structural equation modeling to assess direct and indirect associations of each bromeliad species cover with environmental conditions, abundance of conspecifics in the vicinity, and local abundance of the other species. A. distichantha cover increased on elevated subplots with high tree/shrub basal area, whereas B. serra cover showed the opposite pattern. In addition, A. distichantha cover was negatively associated with B. serra cover, but not vice versa, and cover of both species increased with the abundance of nearby conspecifics, suggesting that limited vegetative dispersal partly accounted for their distribution. Sectors dominated by A. distichantha had lower soil bulk density and higher organic matter content than those dominated by B. serra. According to our model, influences of competition and limited vegetative dispersal reinforce the association between distribution of these bromeliads and environmental heterogeneity of the forest understory.     

The relative contribution of elastic and osmotic adjustments to turgor maintenance of woody species.

To determine how tissue water relations vary and contribute to turgor maintenance in species from contrasting ecological zones, seedlings of jack pine ( Pinus banksiana Lamb.), black spruce ( Picea mariana [Mill] B.S.P.) and flooded gum ( Eucalyptus grandis W. Hill ex Maiden) were subjected to an 8 day drought stress by water withholding with and without prior mild water stress conditioning. Jack pine, a deep-rooted species from dry, sandy boreal sites, lost turgor at the lowest relative water content (75–65%) and water potential, and had lowest maximum bulk elastic modulus (E_max of 5.2–5.8 MPa). Although this suggests a high inherent dehydration tolerance, jack pine did not further adjust its elasticity when repeatedly stressed. Black spruce, a shallow-rooted species from predominantly moist sites in the boreal region, lost turgor at intermediate relative water content (86–76%) and water potential, but could adjust its elasticity to maintain turgor in repeatedly stressed tissues. Flooded gum, a deep-rooted species from moist, warm temperate-subtropical regions, had a low inherent drought tolerance since it lost turgor at higher relative water content (88–84%) and water potential, but was capable of some adjustment when the stress was repeated. Elastic adjustment (<3.7 MPa) was more important for turgor maintenance than osmotic adjustment (<0.13 MPa), which was statistically nonsignificant. Maximum bulk modulus of elasticity, but not osmotic potentials at full turgor, was significantly correlated with the relative water content and water potential at zero turgor in droughted seedlings. These results highlight the importance of tissue shrinkage for dehydration tolerance. Both the inherent capacity for turgor maintenance of a species under drought and its ability to adjust to repeated drought should be considered in genetic selections for drought tolerance.     

Fine-scale patterns of vegetation and environmental factors on an ombrotrophic mire expanse: a numerical approach

The vegetation within an ombrotrophic mire expanse in SE Norway is studied in detail. Percentage cover of 45 species in 436 sample plots (16 ×16 cm), dispersed on 26 transects, are recorded. In addition, species abundance in 6976 subplots (4×4 cm) are recorded. 14 variables are recorded for each of the sample plots, while only distance to the water-table is estimated for the subplots. Spatial co-ordinates are supplied for all sample- and subplots. DCA ordination of a data-set consisting of 412 sample plots reveals two ecologically interpretable vegetational gradients: the hummock-hollow gradient (DCA 1), and a gradient associated with the peat-production of the bottom layer (DCA 2). Passive DCA of subplots is used to get an impression of within sample plot heterogeneity, and shows that the fine-scale compositional turnover may be considerable. Partitioning of the variation in species abundance data is done by use of (partial) CCA. The fraction of unexplained variation is rather large for all the tested data-sets, but within the total variation explained, both distance to the water-table and spatial structure explain large parts.     

Detection of plot-level changes in ectomycorrhizal communities across years in an old-growth mixed-conifer forest

Understanding spatial and temporal patterns present in ectomycorrhizal fungal community structure is critical to understanding both the scale and duration of the potential impact these fungi have on the plant community. While recent studies consider the spatial structure of ectomycorrhizal communities, few studies consider how this changes over time. Ectomycorrhizal root biomass and the similarity of community composition were measured at scales up to 20 cm replicated in nine plots and over 3 yr. Soil cores were additionally stratified into three depths. Annual occurrence of the dominant ectomycorrhizal species was constant at larger spatial scales but varied more across years at a fine spatial scale. Turnover of ectomycorrhizal species between years was observed frequently at scales < 20 cm. The ectomycorrhizal community within a plot was more similar across years than it was to other plots sampled in the same year. Our results demonstrate the dynamic nature of the ectomycorrhizal community even in the absence of large-scale disturbances. The potential role of root turnover and drought stress is discussed.     

The effect of defoliation intensity and history on photosynthesis, growth and carbon reserves of two conifers with contrasting leaf lifespans and growth habits

The effects of partial defoliation on photosynthesis, whole-seedling carbon allocation, partitioning and growth were studied for two species with contrasting foliar traits. Field-grown seedlings of deciduous Japanese larch ( Larix leptolepis ) and evergreen red pine ( Pinus resinosa ) were defoliated by hand in early summer for 2 consecutive years. In the first year (1990), seedlings were defoliated by removing the distal 0, 25, 50 or 75% of each needle. In the second year (1991), seedlings were defoliated either 0 or 50%, regardless of previous defoliation treatments. Defoliation had little effect on photosynthesis and starch concentration in whole seedlings of either species in the first year. In the second year, photosynthesis increased in both species in response to the 1991 defoliation treatment, and in red pine also increased in response to the 1990 defoliation treatment. Further, in 1991 both larch and pine had decreased whole-seedling total non-structural carbohydrate concentrations in all seedlings that were defoliated at least once over the 2-yr period. This decrease was noted mostly in the starch component of the non-structural carbohydrates, and was similar in both species. In 1991, biomass was similarly decreased in both species in response to 1991 defoliation. Both species showed overcompensation in total and component biomass in seedlings defoliated by 25% in 1990. Overall, the results do not support the widely held belief that evergreen trees are substantially more affected than deciduous trees by defoliation.     

Allocation tradeoffs among chaparral shrub seedlings with different life history types (Rhamnaceae)

? Premise of the Study: California chaparral shrub species have different life history types: Nonsprouters (NS) are killed by fire and persist through a fire-stimulated seed bank; facultative sprouters (FS) reestablish by a combination of vegetative sprouting and seeding; and obligate sprouters (OS) reestablish exclusively by sprouting. Nonsprouters and FS establish seedlings in open-canopy postfire environments, whereas OS establish seedlings between fires in the shady understory. We hypothesized that allocation differences among seedlings of postfire sprouters and nonsprouters and regeneration niche differences would lead to contrasting patterns in biomass accumulation (NS > FS > OS, in sun; OS > FS > NS, in shade). ? Methods: Seedlings of three species from each life history type were grown in sun and 75% shade. We measured net carbon assimilation and biomass accumulation after one year. ? Key Results: Biomass accumulation was similar in the sun except FS>OS. In the shade, NS had lower biomass than FS and OS. Assimilation rates, nitrogen relations, and allocation differences could not fully explain biomass accumulation differences. Instead, biomass accumulation was inversely related to water-stress tolerance and shade tolerance. Additionally, OS and FS differed in root/shoot allocation even though both are sprouters. ? Conclusions: Seedling growth and carbon assimilation rates were divergent among three life history types and were consistent with differences in tolerance to water stress and shade or sun regeneration niches, but not tradeoffs in sprouting-related allocation differences per se.     

Riparian Forest Restoration: Increasing Success by Reducing Plant Competition and Herbivory

The reestablishment of riparian forest is often viewed as “best management practice” for restoring stream ecosystems to a quasi‐natural state and preventing non‐point source contaminants from entering them. We experimentally assessed seedling survivorship and growth of Quercus palustris (pin oak), Q. rubra (red oak), Q. alba (white oak), Betula nigra (river birch), and Acer rubrum (red maple) in response to root‐stock type (bare root vs. containerized), herbivore protection (tree shelters), and weed control (herbicide, mowing, tree mats) over a 4‐year period at two riparian sites near the Chester River in Maryland, U.S.A. We started with tree‐stocking densities of 988/ha (400/ac) in the experimental plots and considered 50% survivorship (i.e., a density of 494/ha [200/ac] at crown closure) to be an “acceptable or minimum” target for riparian restoration. Results after four growing seasons show no significant difference in survivorship and growth between bare‐root and containerized seedlings when averaged across all species and treatments. Overall survivorship and growth was significantly higher for sheltered versus unsheltered seedlings (49% and 77.6 cm vs. 12.1% and 3.6 cm, respectively) when averaged across all species and weed control treatments. Each of the five test species exhibited significantly higher 4‐year growth with shelter protection when averaged across all other treatments, and all species but river birch had significantly higher survivorship in shelters during the period. Seedlings protected from weeds by herbicide exhibited significantly higher survivorship and growth than seedlings in all other weed‐control treatments when averaged across all species and shelter treatments. The highest 4‐year levels of survivorship/growth, when averaged across all species, was associated with seedlings protected by shelters and herbicide (88.8%/125.7cm) and by shelters and weed mats (57.5%/73.5 cm). Thus, only plots where seedlings were assisted by a combination of tree shelters and either herbicide or tree mats exhibited an “acceptable or minimum” rate of survivorship (i.e.,>50%) for riparian forest restoration in the region. Moreover, the combined growth and survivorship data suggest that crown closure over most small streams in need of restoration in the region can be achieved most rapidly (i.e., 15 years or less) by protecting seedlings with tree shelters and controlling competing vegetation with herbicides.     

Growth,flowering and recruitment of shrubs in grazed and in protected rangeland in the arid Karoo,South Africa

A three-year study involving 1300 marked shrubs of three species differing in palatability to sheep provided evidence that changes in the demographic structure of shrub populations in arid rangelands are a function of seed availability. Sheep reduced canopy size and flowering of their preferred forage species but had no effect on survivorship of seedlings or established plants during the study. Although annual turnover in all populations was low (<6%), abundant recent recruitment of non-forage species was observed on degraded rangeland. Natality of all three species was positively correlated with flower abundance. Mortality was correlated with natality because seedlings had a low probability of surviving their first year. Recruitment of both forage and non-forage species, appeared to be inhibited by competition and seedlings seldom survived near adult plants of the same growth form.