Journal of Vegetation Science

Published by Wiley
Online ISSN: 1654-1103
Print ISSN: 1100-9233
Publications
Questions: 1) What is the effect of functional type on species’ distribution model performance, and which plant species’ traits explain the most variation? 2) Does model performance vary more as a result of functional type, individual species characteristics, or modeling method?; Location: California’s Southwest Ecoregion, USA; Methods: We developed species distribution models (SDMs) for 45 plant species using four modeling methods (GLMs, GAMs, classification trees, and Random Forests). Using AUC as a performance measure of prediction accuracy, we carried out regression analyses to compare the effects of functional type, longevity, dispersal mechanism, range size, cover, species prevalence, and model type. Results: Functional type explained more variation in model performance than any other variable, but other species traits and biogeographical factors were also significant. Differences in prediction accuracy reflected variation in species’ life history, disturbance response, and rarity. AUC was significantly higher for species that are longer-lived, found at intermediate levels of abundance, and have smaller range sizes. Models also performed better for shrubs than for subshrubs and perennial herbs. The disturbance response functional type with the highest SDM accuracy was obligate seeding shrubs with ballistic dispersal that regenerate via fire-cued germination from a dormant seed bank. Conclusions: The effect of species characteristics on model performance overrides any differences in modeling technique. Prediction accuracy may be related to the way a suite of species characteristics covary along environmental gradients, and including disturbance response was important because SDMs predict the realized niche. Classification of plant species into functional types, particularly in disturbance-prone ecosystems, may provide a strong framework for evaluating performance of SDMs.
 
Questions: How did an initial tree harvest in 1894 influence the spatial and temporal patterns of Pinus ponderosa recruitment? How do these patterns compare to our understanding of P. ponderosa stand dynamics prior to Euro-American settlement? How might spatial pattern information, particularly with respect to patch characteristics, inform current restoration and management practices?Location: A 2.59-ha permanent sample plot in the Fort Valley Experimental Forest, Flagstaff, Arizona. The plot was selectively harvested in 1894 and measured in 1909 and 2002.Methods: We used historical stem-map and ledger data, contemporary data, and dendrochronological techniques to reconstruct stand structure (tree size, age, location) in three scenarios: (1) unharvested (1909), (2) harvested (1909), and (3) contemporary (2002). We used Clark and Evans' R, Ripley's K(t) univariate analysis, and correlogram analysis to assess the spatial pattern in each scenario. We also used Ripley's K12(t) bivariate analysis and tree age data to examine spatial and temporal recruitment patterns as observed in the contemporary scenario.Results and Conclusions: The unharvested stand was aggregated at scales up to 28 m. The selective harvest accentuated the spatial patchiness of the stand in 1909 and changed spatial patterns by homogenizing tree size within patches. By 2002, the stand was a single patch dominated by small trees. Post-harvest recruitment patterns were not spatially random; Pinus seedlings initially established in natural grass openings and then proceeded to fill-in stump patches created by harvesting. Knowledge of spatial pattern should be explicitly incorporated into restoration activities in these forests.
 
The first broad-based, paleoecological analysis of a sedimentary sequence on the British chalk, dating to the terminal Pleistocene, reveals a history of climatic, vegetational and faunal change. The past co-occurrence of currently allopatric species among molluscs, beetles and plants supports hypotheses of the impermanence of communities. Modern pollen rain data are utilized to refine the ecological interpretation of the fossil pollen data.The presence of the Windermere Interstadial (Allerod) and the Loch Lomond re-advance (Younger Dryas) are represented by decreased abundances of arboreal taxa and increased representation of cold grassland elements. Open grassland habitats appear to have been a continuous landscape element, at least locally, since the late-glacial period on the northern British chal Mands, although their species composition has changed greatly in the last 11400 yr.
 
Species and site scores for the first and second axis of the DCA based on basal area of trees (eigenvalues were 0.731 and 0.053 respectively and the variance explained by the first two axes was 82.7%). q = oak forest and s = pine forest (labels indicating the sampling year and 'b' for before the storm and 'a' for after). The third axis (not shown; eigenvalue = 0.009) did not reveal interpretable patterns for species or sites.
Logistic regression for mortality. Species are entered as dummy variables and are compared to all unspecified species. Positive values for parameter estimates indicate increased susceptiblity with increases in that parameter, whereas negative values indicate decreased susceptibility.
Logistic regression for mortality. Positive values for parameter estimates indicate increased susceptiblity with in- creases in that parameter, whereas negative values indicate decreased susceptibility. Damage types are entered as dummy variables and are compared to undamaged trees. N
Logistic regression for damage. Positive values for parameter estimates indicate increased susceptiblity with increases in that parameter, whereas negative values indicate decreased susceptibility. All forms of tree damage are considered. Species are entered as dummy variables and are compared to all unspecified species. N
We measured tree damage and mortality following a catastrophic windthrow in permanent plots in an oak forest and a pine forest in central Minnesota. We monitored changes in forest structure and composition over the next 14 years. Prior to the storm, the oak forest was dominated by Quercus ellipsoidalis, and the pine forest by Pinus strobus. The immediate impacts of the storm were to differentially damage and kill large, early-successional hardwoods and pines. Subsequent recovery was characterized by the growth of late-successional hardwoods. In both forests the disturbance acted to accelerate succession. Ordination of tree species composition confirmed the trend of accelerated succession, and suggested a convergence of composition between the two forests.
 
We studied the effects of windthrow on the understory plant species composition of a pine forest (dominated by Pinus strobus) and an oak forest (dominated by Quercus ellipsoidalis). We recorded the presence of vascular plant species in randomly located quadrats in the two forests, and in three microsite types associated with tipup mounds (pit, old soil and new soil) in the pine forest at irregular intervals over the course of 14 years. The understories of the two forests remained distinct throughout the study. The frequency of occurrence of a number of forest floor species considerably increased; few species decreased. The disturbance specialists Rubus idaeus and Polygonum cilinode increased in frequency throughout the study in the pine forest, but are beginning to decline in the less disturbed oak forest. Annuals and biennials preferentially colonized the disturbed soil of microsites on tipups, and declined in frequency after about 7 yr. Both forests have increased in understory species richness, but have not changed substantially in the distribution of growth forms. Despite early differences in species composition, microsite types associated with tipup mounds became more similar through time. Although small in magnitude, there was a directional change in understory composition at both forests, with no apparent sign of a return to pre-disturbance conditions.
 
Question: Are seed size and plant size linked to species responses to inter-annual variations in rainfall and rainfall distribution during the growing season in annual grasslands? Location: A 16-year data set on species abundance in permanent plots 15 km north of Madrid in a Quercus ilex subsp. ballota dehesa. Methods: At species level, a GLM was used to analyse the effects of various rainfall indices (total autumn rainfall, early autumn rainfall and spring drought) on species abundance residuals with respect to time and topography. We also assessed the importance of seed size and plant size in the species responses at community level using species as data points. Seed mass and maximum stem length were used as surrogates for seed size and plant size, respectively. Results: Seed mass and plant size may explain some of the fluctuations in the floristic composition of annual species associated with autumn rainfall patterns. Species that are more abundant in dry autumns have greater seed mass than those species that are more abundant in wet autumns. Early autumn rainfall seems to favour larger plants. Conclusions: Our empirical results support the hypothesis that autumn rainfall patterns affect the relative establishment capacity of small and large seedlings in annual species.
 
A scheme of different equalizations of the size of site groups. Each line represents distribution of a species in the original ( a ) and equalized ( b - d ) data sets, each data set with four site groups. Segments are site groups labelled A, B, C and D, and segment lengths correspond to the number of sites in each group. The thick parts of each segment represent sites with occurrence of the given species, and the thin parts represent sites where the species is absent. In ( b ) to ( d ), the four lines represent in turn equalizations used for calculation of species fidelity to the target site groups A, B, C and D. 
Relative frequencies of species occurrences within (vertical axis) and outside (horizontal axis) the target group of 204 rock-outcrop dry grassland sites, compared with (a) 298 sites of other types of rock-outcrop dry grasslands and (b) 15 487 sites of all other types of grasslands. Each dot is a species. Curves represent the value of Φ = 0.5 for cases when the size of the target group is equalized to 50%, 10% and 2% of the entire data set, respectively. Φ -values > 0.5 are above these curves, 
Comparison of some properties of different fidelity measures with properties of the phi coefficient applied to the site groups of equalized size. P/A = presence/absence. *Chi-square enables distinguishing between positive and negative fidelity if supple- mented by additional information on whether the observed occurrence frequency of the species in the target site group is higher or lower, respectively, than the expected frequency.
Aim: Concentration of species occurrences in groups of classified sites can be quantified with statistical measures of fidelity, which can be used for the determination of diagnostic species. However, for most available measures fidelity depends on the number of sites within individual groups. As the classified data sets typically contain site groups of unequal size, such measures do not enable a comparison of numerical fidelity values of species between different site groups. We therefore propose a new method of measuring fidelity with presence/absence data after equalization of the size of the site groups. We compare the properties of this new method with other measures of statistical fidelity, in particular with the Dufrêne-Legendre Indicator Value (IndVal) index.
 
Questions: What are the relationships between weed species traits and their change in distribution over a 30-year period? What does it tell us about factors that have driven shifts in the composition of weed communities? Location: France. Methods: We analysed the links between change in the status of weed species in sunflower crops (decreasing or increasing) and a set of 17 traits using data sets collected in the 1970s and the 2000s, respectively. We analysed the contribution of traits to explain changes in the status of species both individually and in a multivariate way by mean of a clustering of species into functional groups. Results: 69% of the most widespread species had significantly changed their frequency rank status over the last 30 years. Nearly two thirds of the increasing species belonged to a single functional group, out of the five groups identified in this analysis. Overall, the weed flora occurring in sunflower crops has specialised since the 1970s in favour of ‘sunflower mimicking’ functional groups: increasing species were more nitrophilous, more heliophilous, less sensitive to sunflower herbicides and shared a rapid summer life cycle. Conclusions: The individual trait approach gave some indication as to the environmental factors likely to have caused the shift in sunflower weed communities. The functional group approach seemed to outperform direct trait comparisons as it accounted for major traits combinations i.e. cases where a species has a number of favourable traits, but is severely disadvantaged by the possession of one or a few deleterious traits.
 
Patterns of plant succession were studied in areas of scorched and blown-down forest resulting from the 1980 eruption of Mount St. Helens, Washington. Changes in species abundance were observed for 7 years in permanent sample plots representing four post-disturbance habitats, or site types. Total plant cover and species richness increased with time on all site types. In blown-down forests supporting snowpack at the time of eruption, understory recovery was dominated by the vegetative regeneration of species persisting through disturbance. In forests without snowpacks, plant survival was poorer. Increases in cover and diversity were dominated first by introduced grasses, then by colonizing forbs characteristic of early successional sites. Epilo-bium angustifolium and Anaphalis margaritacea showed widespread recruitment and clonal expansion throughout the devastated area. As a result, species composition on previously forested sites converged toward that on formerly clearcut sites, where early serai forbs resprouted vigorously from beneath the tephra.Total plant cover and species diversity were poorly correlated with post-disturbance habitat and general site characteristics (e.g. distance from the crater, elevation, slope, and aspect). However, distributions of several life-forms (e.g. low sub-shrubs and tall shrubs) were strongly correlated with depth of burial by tephra and with cover of tree rootwads. Thus, early community recovery may reflect microsite variation or chance survival and recruitment rather than broad-scale gradients in environment or disturbance.Recovery of pre-disturbance composition and structure will undoubtedly be much slower than after other types of catastrophic disturbance. The rate and direction of community recovery will largely depend on the degree to which original understory species survived the eruption.
 
Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage.
 
Trends in mean annual temperature and precipitation from 1965 to 2007 in (a, b) eastern France and (c, d) western Switzerland: (a, c) Yearly mean surface temperature anomalies (overall mean of temperature as baseline) and (b, d) annual sum of precipitation anomalies (overall mean of annual sum of precipitation as baseline) averaged for (a, b) 10 elevation sites (ranging from 236 to 1,120 m a.s.l.) in the French part of the Jura range and for (c, d) three elevation sites (ranging from 316 m to 1,073 m a.s.l.) in the Swiss part of the Jura range. Filled dark grey bars are positive anomalies, whereas empty bars are negative anomalies. Filled light grey bars are 1989 and 2007 sampling times. The dark curves have been smoothed using a 10-year filter. Data were gathered from the French National Climatic Network (Météo-France) and the Swiss Climatic Network (Météo-Suisse).
Relationship between species mean abundance–dominance in a relevé (cover abundance scale ranging from 1 to 6) (Braun-Blanquet 1932) and species mean rank of their recording sequence in a relevé in 2007. Each cross represents a species. Average values are computed for herb and fern species only and for the 154 re-surveyed relevés only.
Scatter diagrams of relevé scores in 1989 and 2007: (a, b) along the first and (c, d) second CA89 axes, using both (a, c) the truncated and (b, d) complete 2007 species lists. The first axis corresponds to a gradient of altitude (from lowland to mountain) and the second axis represents a gradient of water availability (from dry to wet). Relevés in 1989 were used as active variables, whereas relevés in 2007 were included in the analyses only as supplementary variables. Dashed bisecting lines represent absence of differences between 1989 and 2007 plot scores (no shift).
Changes in (a) 1st, (b) 5th and (c) 9th decile positions of species altitudinal ranges for both lowland and mountain species between 1989 and 2007, using the truncated 2007 species lists. L89 (grey bars) and L07 (dark bars) refer to lowland species (n=31) in 1989 and 2007, respectively. M89 (grey bars) and M07 (dark bars) refer to mountain species (n=29) in 1989 and 2007, respectively. Means are shown with standard error. Bars not sharing a common letter differ significantly between the two geographic species groups (Student's two-sample t-test between lowland and mountain species, P<0.05). Bars not sharing a common number differ significantly between periods (Student's paired sample t-test between 1989 and 2007, P<0.05).
Question: How strong are climate warming-driven changes within mid-elevation forest communities? Observations of plant community change within temperate mountain forest ecosystems in response to recent warming are scarce in comparison to high-elevation alpine and nival ecosystems, perhaps reflecting the confounding influence of forest stand dynamics.Location: Jura Mountains (France and Switzerland).Methods: We assessed changes in plant community composition by surveying 154 Abies alba forest vegetation relevés (550-1,350 m a.s.l.) in 1989 and 2007. Over this period, temperatures increased while precipitation did not change. Correspondence analysis (CA) and ecological indicator values were used to measure changes in plant community composition. Relevés in even- and uneven-aged stands were analysed separately to determine the influence of forest stand dynamics. We also analysed changes in species distribution to detect shifts along the elevation gradient by focusing on the lowest, central and highest positions of lowland and mountain species altitudinal ranges.Results: We found significant shifts along the first CA axis, which reflected a change in plant community composition towards a greater frequency of lowland species. Analyses of ecological indicator values indicated increases in temperature and light availability in A. alba stands, particularly in even-aged stands. However, no major changes in overall species distribution were found.Conclusions: The community-level changes are consistent with effects of climate warming and local stand dynamics. Changes in species distribution were small in comparison to observed local temperature increases, perhaps reflecting dispersal limitation, phenotypic plasticity or microclimatic buffering by the tree canopy. Causality cannot rigorously be inferred from such a descriptive study; however, we suggest that recent warming is now driving plant community change in the climatically more moderate mid-elevation forest setting.
 
Question: What are the trends and patterns in the application of ordination techniques in vegetation science since 1990?Location: Worldwide literature analysis.Methods: Evaluation of five major journals of vegetation science; search of all ISI-listed ecological journals. Data were analysed with ANCOVAs, Spearman rank correlations, GLMs, biodiversity indices and simple graphs.Results: The ISI search retrieved fewer papers that used ordinations than the manual evaluation of five selected journals. Both retrieval methods revealed a clear trend in increasing frequency of ordination applications from 1990 to the present. Canonical Correspondence Analysis was far more frequently detected by the ISI search than any other method. Applications such as Correspondence Analysis/Reciprocal Averaging and Detrended Correspondence Analysis have increasingly been used in studies published in “applied” journals, while Canonical Correspondence Analysis, Redundancy Analysis and Non-Metric Multidimensional Scaling were more frequently used in journals focusing on more “basic” research. Overall, Detrended Correspondence Analysis was the most commonly applied method within the five major journals, although the number of publications slightly decreased over time. Use of Non-Metric Multidimensional Scaling has increased over the last 10 years.Conclusion: The availability of suitable software packages has facilitated the application of certain techniques such as Non-Metric Multidimensional Scaling. However, choices of ordination techniques are currently less driven by the constraints imposed by the software; there is also limited evidence that the choice of methods follows social considerations such as the need to use fashionable methods. Methodological diversity has been maintained or has even increased over time and reflects the researcher's need for diverse analytical tools suitable to address a wide range of questions.
 
Monthly precipitation (mm) at Quebrada El Cobre, Reserva Nacional Las Chinchillas, Chile. 
We followed plant cover and soil seed density of shrubs and herbs in two markedly contrasting years with regard to annual rainfall in two opposite-facing slopes (mesic vs xeric) and an intervening, relatively wet, ravine of a typical creek in semi-arid Chile. During the ENSO year 1997 cover of ephemerals increased in all three sites; 43% vs 8% on the xeric slope 75% vs 26% on the mesic slope and 57% vs 32% in the ravine. The number of species was almost three times higher on the xeric slope (27 vs 10 species), increased by 47% on the mesic slope (28 vs 19) and by 14% in the ravine (24 vs 21). Cover of shrubs plus perennial grasses increased from 52% in 1996 to 59% in 1997 on the xeric slope, but no significant cover changes were found on the mesic slope or the ravine. On the xeric slope peak density of seeds was 4500/m2 in 1996, while during the ENSO year it was 24000/m2. On the mesic slope equivalent values were 3000 and 17000 seeds/m2 while in the ravine figures were 8500 and 27000 seeds/m2. The strong responses of ephemerals and seed bank to the ENSO-driven wet year of 1997 demonstrate the importance of this phenomenon in replenishing, recovering and probably maintaining ephemeral vegetation in this region.
 
Question: How do aggregate trait values and functional diversity of leaf traits linked to palatability and decomposability change during a woody post-cultural succession spanning 200 years?Location: Coastal Marlborough, South Island, New Zealand.Methods: The biomass of all woody species was determined in 32 20-m × 20-m plots ranging from 10 to 200 years in time since last disturbance. Species abundances were combined with data on leaf nutrient, secondary metabolite and structural carbohydrate content to calculate biomass-weighted trait means (i.e. aggregate trait values) and functional diversity index values for each plot. Aggregate trait values and functional diversity were regressed on successional age and total live above-ground carbon content to examine functional shifts with succession and one consequence of succession – increasing above-ground carbon.Results: Almost all significant regressions between aggregate trait values and both successional age and above-ground carbon indicated a shift toward increased leaf palatability and decomposability during succession. The relationships were all non-linear, with aggregate trait value shifts occurring relatively early in the successional sequence. There was weak evidence for an increase in functional richness with succession, but this was a secondary effect relative to the shifts in aggregate trait values.Conclusions: These results are in direct contrast with studies of the early stages of herbaceous post-cultural successions from grasslands to shrublands, which have found a shift towards communities of decreasing palatability and decomposability, suggesting that functional shifts in woody succession may be fundamentally different.
 
Question: Habitats are characterized by their plant species composition. Therefore, climate change impacts on habitats can be assessed by two complementary statistical approaches: either directly by modelling the climate envelope of the habitat, or indirectly by modelling the habitat in terms of its plant species. How do these approaches differ in their projected habitat distribution? What are the consequences for nature conservation? Location: Europe. Methods: Potential climate change impacts on the distribution of European protected Natura 2000 sites were modelled for five natural and semi-natural grassland habitat types, defined by the EU Habitats Directive, using data from the Atlas Florae Europaeae and reports on Natura 2000 sites. We used random forests (RF) and logistic regression (GLM) to model current and potential future distributions for 2050. Results: All habitats are projected to lose between 22% and 93% of their range in the ‘no dispersal’ scenario. In the ‘unrestricted dispersal’ scenario, almost all habitats gain suitable climate space, between 5% and 100% of their current range. In the direct habitat approach, both model algorithms have high discriminatory performance on test data and are well calibrated. In the indirect species approach, only GLM shows high model performance; RF models are overfitted. Projections of occurrence probabilities differ more strongly between model approaches (‘direct’ versus ‘indirect’) than between model algorithms (GLM versus RF). Conclusions: Habitats are complex entities. Because of their dynamic nature, particularly in the face of climate change, we suggest modelling the future distribution of habitat types not exclusively based on their current definitions and mapped distributions, but also based on their constituent elements, and in particular their characteristic plant species.
 
Comparisons among three analytical software packages.
CCA triplot of Boyd Tract data, as produced by SYN-TAX 2000 . 
Personal computers of ever-increasing speed have motivated programmers of multivariate software to adapt their programs to be run in Microsoft Windows and Macintosh platforms. Updated versions of these multivariate programs appear more and more frequently and are marketed intensively. In this review we provide a comparative analysis of the most recent versions of three analytical software packages –Canoco for Windows 4.5, PC-ORD version 4 and SYN-TAX 2000. The three packages share two characteristics. First, the most recent versions are now compatible with the most recent Windows platforms and should therefore be accessible for use by virtually all vegetation scientists. Second, they have capabilities for numerous multivariate techniques, although each package has some unique techniques. Thus, any one of the packages will have much to offer the user.
 
Question: Following a volcanic eruption of ca. 232 AD, known as the Taupo eruption, the emergent conifer Libocedrus bidwillii expanded on Mt. Hauhungatahi, upwards above the current tree-line, and downwards into the mixed montane forest. We ask: (1) if current age-structures at different altitudes support the patterns predicted by the temporal stand replacement model, with cohort senescence and progressively depleting recruitment at ca. 600 year intervals (average cohort age) since the eruption: and (2) if the case history of the population sheds light on the persistence of mixed conifer-hardwood forests in general. Location: Mt. Hauhungatahi, Tongariro National Park, New Zealand. Methods: The species composition and structure of seven stands covering the altitudinal range of Libocedrus bidwillii, were quantified. Libocedrus trees were cored, and regression equations used to predict ages. Cohorts were identified. Results: Libocedrus densities and basal areas, and the abundance of seedlings and saplings, peaked at different altitudes. At the species’lower limits there has been no recruitment for ca. 550 years, and the angiosperm Weinmannia racemosa has gained dominance. In the tree line and sub-alpine forest stands, a low level of continuous regeneration has been boosted by periodic cohort recruitment following exogenous disturbances. Conclusions: In the montane zone, the Libocedrus age structure, and its replacement by Weinmannia, are consistent with a model of depleting cohorts separated by ca. 600 years since the Taupo eruption. At higher altitudes more frequent disturbances and reduced competition have allowed Libocedrus persistence. Comparison with other studies suggests long-term relationships between gymnosperms and angiosperms are mediated by the scale and frequency of disturbance.
 
We document post-fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade-intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade-intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post-fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.
 
We present results from repeated analyses (1962, 1993) of a permanent plot established in 1947, combined with retrospective stand age structure data, in an old Pinus sylvestris stand in Muddus National Park, northern Sweden. The study points towards a successional pathway governed by concurrent disturbance effects of climate variability, reindeer grazing and fire. This is intermediate to the two often advocated ideas on dynamics in boreal forests, that is, one of disturbance-related tree regeneration/mortality and one of continuous regeneration.
 
Questions: Most modern fire-prone landscapes have experienced disruptions of their historic fire regimes. Are the primary tallgrass prairies of the Flint Hills reflective of a history of continuous fire occurrence? Did fire frequency, severity, size and seasonality change in connection with changes in land use? Has fire occurrence been related to drought conditions? Location: Edges of Cross Timbers forest stands at the Tallgrass Prairie Preserve (TGPP) in the Flint Hills of Osage County, Oklahoma, USA. Methods: Cross-sections of 76 Quercus stellata were collected from Cross Timbers stands at or near the grassland edge in the TGPP. Dendrochronological methods were used to identify years of formation for tree rings and fire scars. Superposed epoch analysis was used to evaluate the effect of drought conditions on fire occurrence. Results: Fires were recorded in 46.6% of the years between 1729 and 2005. In 41 cross-sections at one site, the mean fire interval between 1759 and 2003 was 2.59 years, with fire interval decreasing from a mean fire interval of 3.76 years in the early part of the record to 2.13 years in modern times. No extended periods without fire were recorded in the study area. Drought conditions had no significant effect on fire occurrence. Conclusions: In contrast with many fire-prone landscapes worldwide, the prairies of the Flint Hills have experienced no recent fire suppression or exclusion. Changes in fire frequency mark transitions in land use, primarily from being traditionally used by Native Americans to being managed for cattle production.
 
Questions: (1) Is climate a strong driver of vegetation dynamics, including interannual variation, in a range margin steppic community? (2) Are there long-term trends in cover and species richness in this community, and are these consistent across species groups and species within groups? (3) Can long-term trends in plant community data be related to variation in local climate over the last three decades? Location: A range margin steppic grassland community in central Germany. Methods: Cover, number and size of all individuals of all plant species present in three permanent 1-m2 plots were recorded in spring for 26 years (1980–2005). Climatic data for the study area were used to determine the best climatic predictor for each plant community, functional group and species variable (annual data and interannual variation) using best subsets regression. Results: April and autumn temperature showed the highest correlation with total cover and species richness and with interannual variations of cover and richness. However, key climate drivers differed between the five most abundant species. Similarly, total cover and number and cover of perennials significantly decreased over time, while no trend was found for the cover and number of annuals. However, within functional groups there were also contrasting species-specific responses. Long-term temperature increases and high interannual variability in both temperature and precipitation were strongly related to long-term trends and interannual variations in plant community data. Conclusions: Temporal trends in vegetation were strongly associated with temporal trends in climate at the study site, with key roles for autumn and spring temperature and precipitation. Dynamics of functional groups and species within groups and their relationships to changes in temperature and precipitation reveal complex long-term and interannual patterns that cannot be inferred from short-term studies with only one or a few individual species. Our results also highlight that responses detected at the functional group level may mask contrasting responses within functional groups. We discuss the implications of these findings for attempts to predict the future response of biodiversity to climate change.
 
Location of fossil midden study site (darkened circle). Present day lakes Pyramid Lake and Walker Lake are represented by a darker pattern. 
Plant macrofossils extracted from fossil woodrat (Neotoma spp.) middens at a single locale in the northwestern Great Basin were used to examine vegetation dynamics during the last 30 000 yr. Although the modern assemblage of xeric species at the study site is a recent occurrence, a large proportion of the modern plant taxa near the study locale were also found 12 000 - 30 000 yr BP. The persistence of extant species through time was likely facilitated by within-species genetic diversity and the formation of coenospecies. The diverse topographic and microhabitat features in the northwestern Great Basin also allowed different species to coexist during glacial periods. Changes in species composition occurred during two time intervals: 20 000 - 30 000 and 10 000 - 12 000 yr BP. Vegetation changes during 20 000 - 30 000 yr BP were cyclic; community composition oscillated between two groups of taxa. Vegetation changes between 10 000 - 12 000 yr BP occurred during the Pleistocene-Holocene transition and were largely directional from the Pleistocene assemblages through two transition assemblages to a Holocene assemblage. These changes in species composition generally reflect changes in climate. The presence of relatively mesic species during 10 000 - 30 000 yr BP is consistent with the regional late-Pleistocene climate, and the gradual loss of relatively mesic species during the Holocene parallels the change to a more xeric climate. Contrasted with other areas of North America and Europe, the magnitude of vegetation changes at our study area were relatively small. Furthermore, the persistence of many species through time at this site in the northwestern Great Basin also differs from results at other study sites in North America and Europe. These differences are probably related to land form characteristics and genetic diversity within species.
 
Of 34 species of vascular plants recorded in an unimproved calcareous grassland community in 1960 and subsequently buried for 32 yr, just six spp. were recovered as viable seeds from the original land surface, following excavation of an experimental earthwork. Five of the six are known or suspected to form persistent seed banks; the status of Carex flacca as a persistent seed is now established. The exceptionally low density of viable seeds at 35 seeds/m2 and the paucity of other plant remains confirm that these calcareous soils have supported rapid decomposition. The complete absence of viable seeds known to be short-lived indicated that the macrofauna has played no significant role in the introduction of viable seeds from the modern-day surface down to depths in excess of 1.6 m, at least not in recent years. Because of the precise dates of burial and excavation, the results offer a rare opportunity to confirm laboratory and other indirect observations of long-term seed persistence following burial in temperate climate calcareous soils.
 
The reaction of different species of gapmakers (DBH > 20 cm) to the natural disturbances in 81 gaps (0.6822 ha) in Fiby urskog sampled from autumn 1988 to spring 1989.
Gap area estimated by plots in the spruce forest of Fiby urskog, autumn 1988.
Number of seedlings and saplings in different height classes (cm) in the biggest gap (0.29 ha) created in the winter of 1982/1983 in Fiby urskog.
We investigated patterns of disturbance and recovery in Fiby urskog, a primeval spruce (Picea abies) forest, situated south of the border between the Boreo-nemoral and Boreal regions in East-central Sweden. The main types of disturbances are storm damage, fungal infection and insect attacks. The response of the different tree species varied and the mode of tree-fall depended on the different combinations of disturbance agents. The DBH distributions of gap creators and gap-border trees were almost the same. There was a high age diversity (100–240 yr) among the fallen trees. We concluded that all canopy trees (DBH > 20 cm) had the same probability of being felled by storms, irrespective of their age and DBH.According to an estimate along transect lines, gaps made up 31% of the spruce forest area. Individual gap sizes ranged from 9 m2 to 360 m2, but 83% of the gaps were < 150 m2. The varied age structure of logs in individual gaps indicated that gap enlargements were common. 96 tree-falls were observed on four days with an hourly mean wind speed > 12.0 m/s; all trees fell in the direction of the wind. However, when we consult the 30-yrrecord(l 959–1989)ofthemeanhourly wind speed >12.0 m/s, it is clear that the pattern of storm-directions does not match the pattern of orientation of fallen logs.The present disturbance regime and the predominance of small gaps were more favourable for the regeneration oí Picea abies than of light-demanding tree species. In one large, 2900 m2 gap, not crossed by the transects, all the major tree species had established within 7 yr, suggesting that classical succession in the sense of complete species replacement or ‘relay floristics’ didnot occur. Our observations seem rather to fit the ‘initial floristic’ model. Estimates of turnover time ranged from 170 to 228 yr, depending on the method used.
 
Species native to the southwest of Western Australia, representing a range of plant families, life-history strategies, fire-response syndromes, seed-store types and seed weights, were tested for viability using tetrazolium chloride and for germination under combinations of constant temperatures of 15 °C or 23 °C, constantly dark or 12 h diurnal whitelight conditions, and with, or without, addition of gibberellic acid (GA3, 50 mg/l). Species previously known to require a heat-shock treatment to overcome dormancy due to an impervious testa were pre-treated prior to imposition of temperature, light and GA3 conditions. The test environmental conditions related to differences between winter and autumn temperatures and surface and buried seed germination positions of post-fire habitats.
 
Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long-term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species-rich plots were more stable in terms of species abundance and composition compared with species-poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small-scale species turnover was negatively related to species diversity, indicating more stability in species-rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.
 
There is growing evidence that plant and animal species are arranged in hierarchies of relative competitive performance. More work is needed to determine which plant traits best predict relative competitive performance. We therefore measured relative competitive performance of 63 terrestrial herbaceous plant species using Trichostema brachiatum as a reference species (that is, phytometer or target species). The neighbour species came from a wide array of terrestrial vegetation types (e.g. rock barrens, alvars, old fields), and represented a wide array of growth forms (e.g. small rosette species such as Saxifraga virginiensis and large clonal graminoids such as Agropyron repens). The experiment was repeated with two pot sizes: large (control) and small (stress treatment). Relative competitive performance in large pots (controls) was highly correlated with that in small pots (stress treatment) (r= 0.90, p < 0.001). The hierarchy of relative competitive performance in the large pots was also highly correlated with the hierarchy in the small (stressed) pots (rs= 0.91, p < 0.001). Principal components analysis and multiple linear regression showed that plant size (measured by total biomass, above-ground biomass, below-ground biomass, canopy area, height and leaf area index) and leaf shape (measured as length to width ratio, length, width) were the two characteristics that best predicted relative competitive performance (large pots, r2= 0.55; small pots, r2= 0.48).
 
Question: What changes in species composition and cover have occurred in chaparral as a function of fire history across an ecoregion? Location: San Diego County, California, USA. Methods: Stands in which 40 mid-elevation chaparral vegetation plots (each 400 m2 in area) were located in the 1930s were resurveyed in 2001. We stratified the stands into Infrequently versus Frequently burned (0–1 versus 2 or more fires recorded in the 91-yr period), and Immature versus Mature (31 yr versus >31 yr since last fire), resulting in four groups. Ten stands were randomly selected from each of these groups for survey. Results: There were no major shifts in life form composition, e.g., live oak trees were not invading chaparral that had experienced little or no fire, nor were subshrubs or herbaceous species replacing shrubs in areas that had experienced more frequent fires. However, there was a notable increase in the frequency of the subshrub Eriogonum fasciculatum across all fire history groups. In the mature stands with infrequent fire, average cover of resprouting shrubs increased (from 72 to 91%) and cover of obligate seeding shrubs (species with fire-cued germination) decreased (from 21 to 6%) significantly. Mature stands with frequent fire showed a significant decrease in resprouter cover (from 87 to 80%) and increase in obligate seeders (from 10 to 16%). Conclusions: While the tremendous changes in land use in southern California have been predicted to cause shifts in chaparral composition, these shifts are difficult to detect because species longevity and fire cycles are on the order of decades to a century. In this study, the expected trends could only be detected in groups that were mature at the time of the second survey.
 
Location of soil profiles along the altitudinal transect from Saint-Michel-de-Maurienne, Savoy, France. The number of levels with charcoal over the number of levels sampled per profile is shown in parentheses. Vertical dashed lines are the present-day limits of vegetation belts. The current ranges of the main modern canopy tree species are displayed by continuous black (abundant) and dashed grey (not abundant) lines. 'Maur' corresponds to the profiles names of the main transect on the Saint-Michel-de-Maurienne catchment area, and 'Breq' corresponds to the names of two additional profiles on the Mont-Brequin alongside the main transect on the same catchment area.  
Absolute taxonomic contribution to the first two axes of correspondence analysis. Dots indicate the taxa which have a significant contribution to the corresponding axis.  
Shrub, tree and total mean (± SE) taxonomic richness for the main type of forest communities characterised by the dominating canopy species according to mid-Holocene charcoal assemblages (A) and modern phytosociological relevés (B).  
Ordination of charcoal assemblages and main contributing taxa (underlined) on the first two axes of correspondence analysis. Arrows show the direction of the axis of contribution of the main contributing taxa. First CA axis is plotted against the altitudes of assemblages. Linear regression; r = 0.72; P < 0.0001. Name of taxa: ABAL = Abies alba; ACOP = Acer cf. opalus; ACPL = Acer cf. platanoides; ACPS = Acer pseudoplatanus; ALGI = Alnus glutinosa/A. incana; ALVI = Alnus viridis; ARUU = Arctostaphylos uva-ursi; BEPP = Betula spec.; CFLA = cf. Larix decidua; CFON = cf. Ononis; COAV = Corylus avellana; ERIC = Ericaceae; FREX = Fraxinus excelsior; JUSP = Juniperus spec.; LAPI = type Larix-Picea; PICE = Pinus cembra; PISU = Pinus sylvestris/ uncinata; POSP = Populus spec.; PRSP = Prunus spec.; RODO = cf. Rhododendron; ROMA = Rosaceae maloideae; SARA = Sambucus racemosa; SASP = Salix spec.; VAMY = Vaccinium cf. myrtillus; VAUL = Vaccinium cf. uliginosum; VAVI = Vaccinium cf. vitis-idaea.  
Current land-use a bandonment and the current rise in temperature in the Alps both suggest that tree limits may change. When it is assumed that the climate of the early mid-Holocene between 8000 and 5000 yr before present is analogous to that of the predicted climate of the late 21st century, palaeo-ecological studies of the early Holocene may provide data for the prediction of the vegetation pattern in a century from now. It appears that mid-Holocene charcoal assemblages can be used to reconstruct the spatial patterns of the vegetation before, or during, the practice of slash-and-burn.
 
This paper is a tribute to A.S. Watt who published his ‘Pattern and process in the plant community’ almost 50 years ago. Watt's interpretation of the plant community “as a working mechanism, which maintains and regenerates itself” is still highly relevant, although the keywords have changed. ‘Process’ in Watt's view involves both upgrade and downgrade aspects, whereas ‘Pattern’ was not specified, neither quantified. Nowadays, process is mainly approached as ‘disturbance’, that is natural disturbance and ‘pattern’ as patch structure. Together they make up the ‘patch dynamics’ of the community. Some implications of patch dynamics for phytosociology are discussed. A ‘Wattian’ concept of the plant community combines the Gleasonian idea of individualistic behaviour of species with the Clementsian (or rather Braun-Blanquetian) notion of community dynamics. Later work by Harper (demography), Grubb (regeneration niche) and earlier work of Sernander (forest gap dynamics) is significant for the understanding of the patch-dynamic nature of the community.Recent interest in plant species mobility can easily be linked to the concept of patch dynamics. Examples of mobility in a limestone grassland are given and a system of mobility types is proposed.Some perspectives for the study of patch dynamics are mentioned. Numerical pattern analysis should have a more prominent place in this type of study; the significance of the study of small permanent plots in a stand is emphasized, and unprejudiced demographic studies, as well as experimental studies of small-scale species replacement are recommended.
 
Vegetation development in 68 abandoned gravel pits and their surroundings in five regions in Sweden is described by means of classification and ordination. Most pits were less than 10 yr old, but some pits in central Sweden were up to 100 yr old. For 26 pits in central Sweden the results of mechanical and chemical soil analyses are presented.Vegetation in young pits is more homogeneous than the vegetation in the surroundings. The surrounding vegetation type strongly affects pit vegetation. There are regional differences in pit vegetation between pits in coniferous forests in different parts of Sweden. Old gravel pits are clearly distinguished from young ones and show more resemblance to the surrounding vegetation. Tree layer cover and nitrogen content increase, whereas C/N ratio and pH decrease with age. Species richness of the gravel pits was not related to their size.
 
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.
 
Previous studies on secondary succession in abandoned agricultural land in the Mediterranean area were carried out by the chronosequence method, including data from different sites. A unique opportunity to study secondary succession arose from a situation in which different parts of one homogeneous East-Mediterranean vineyard were abandoned for 5, 8, 15 and 35 yr, and did not suffer from any disturbance subsequently. Most of the perennial species that colonized the abandoned vineyard were fleshy fruited species, which apparently were dispersed by birds from the surrounding maquis into the vineyard. These bird-dispersed species were the first to be established, and were the dominant plant group according to dispersal modes. The abandoned vine plants and their supporting columns provided the birds with perching and feeding sites, enhancing the arrival of bird-dispersed species regardless of their life forms. Under these conditions the most important attribute that affected vegetation dynamics was seed dispersal mode. Trees were among the first to colonize in the vineyard, implying that no facilitation was needed for their establishment. Annual plant species were the only species to disappear during succession. Almost all perennial species which had arrived persisted in the vineyard, and no replacement of perennial species was found. The rate of succession was rapid, as expressed by the short time (8–15 yr) needed for the stabilization of species composition, for growth to average height of late succession trees, and for reaching high cover of the invading perennial species in the abandoned vineyard. The secondary succession described above differs from that in the western Mediterranean by the absence of perennial species replacement and its rapid rate. The possible causes are discussed.
 
Recruitment and regeneration: Significant interrelationships between plot-scale variables as indicated from MLR analyses. Positive relationships are drawn in white, negative in black lines. The thickness of lines indicates the relative weight of the variables as predictors (additive adj. R2 at model inclusion step; indicated in relation to the 100% scale). Arrows show logical predictive pathways; if no arrows are indicated, the direction of the correlation is unclear. FCI is shown as clay variable.
Frequency distribution and fitted log-normal curves of bole diameter (at 20-cm above ground) of the 783 living trees plus the 64 dead trees measured in June 2002 on the 97 study plots (A1), the 698 living trees in June 2003 (A3), the 542 living trees in February 2005 (A5), dead trees in June 2002 (D2), dead trees from June 2002 until June 2003 (D3), dead trees from June 2003 until February 2005 (D5), and the total of dead trees in February 2005 (DT). A t-test matrix table for a cross-comparison of the different categories of tree bole diameters (log-transformed) is also shown.
Questions: Bush encroachment is a major problem when African savanna ecosystems are used for cattle ranching. How do secondary woodlands develop after ranching is abandoned? What are the patterns and rates of tree mortality and regeneration? Location: Mkwaja Ranch (now part of Saadani National Park) in coastal Tanzania. Methods: Ninety-seven circular plots (4-m radius) were set in secondary Acacia zanzibarica woodland along a gradient of tree density. Variables relating to tree and grass layers and soil characteristics of plots were recorded. Seedlings were counted twice in the wet seasons, and resprouts once. Tree flowering and pod production were assessed during the fruiting season, while survival of trees initially present was recorded after 12 and 32 months. Interrelationships among variables were investigated using multiple linear regression, binary logistic regression and mixed effects models. Results: After 32 months, over one-third of trees in plots had died. Most died after fire, especially on heavy soils, and mortality was significantly related to the tree live biomass ratio and soil conditions. Seed production was very low, especially in denser stands. Numbers of seedlings correlated with soil and grass variables but not with seed rain. Half of trees killed above-ground produced coppice shoots from the base; in contrast, root suckering was independent of topkill. By the end of the study, no seedlings survived and no resprouts emerged above the grass layer. Conclusions: A. zanzibarica woodlands at Mkwaja Ranch were able to develop because of ranching, and can only persist under intensive grazing. The woodlands do not represent a successional stage towards forest and will probably revert to predominantly grassland vegetation within 10–20 years unless grazing pressure from wild ungulates increases considerably and/or fire regimes change.
 
As a consequence of socio-economic changes, many Castanea sativa coppices have been abandoned and are now developing past their usual rotation length. Do we have to expect changes in stand structure and composition of abandoned Castanea sativa coppice invaded by other species? Is a tree ring-based approach adequate to early recognise changes in inter-specific competitive interaction? Lowest alpine forest belt of the southern Swiss Alps. We selected a 60-year old abandoned Castanea sativa coppice stand with sporadic Fagus sylvatica and Quercus cerris mixed in. Using tree-ring based indices we analysed differences in the species-specific response to competition. Analyses were performed by comparing how subject dominant trees (10 Castanea, 5 Fagus, 5 Quercus) have differently faced competition from their immediate Castanea coppice neighbourhood, taking into account the changes over time and space. Although no species appears yet to have made a difference in the surrounding coppice mortality, there are species-specific differences in growth dominance, which indicate potential successional processes. Castanea sativa growth dominated in the early stages of stand development. However, after approximately 30–35 yr Fagus sylvatica and Quercus cerris became much more dominant, indicating a change in competitive potential that does not favour Castanea sativa. Without interventions this coppice will develop into a mixed stand. A tree-ring based approach allows an early recognition of forthcoming changes in stand composition and structure and is likely to be an important tool for forest landscape management.
 
Salt marshes along the Atlantic coast of France have been converted into solar salt pans since the 7th century. Salt production declined strongly from the 18th century onward, leading to the abandonment of many of these salt pans. High soil salinity is a residue from the original salt production and varies among salt pans according to time since abandonment, the current flooding regime and the effect of drainage measures. The relationships between the plant communities and seven environmental variables were investigated by Detrended Correspondence Analyses (DCA). Duration of flooding, electric conductivity and sodium saturation were most strongly related to variation in vegetation. The Heleo-chareto-Hippuridetum vulgaris and the Caricetum ripariae occur in lagoons with slightly saline soil that are flooded for the longest time period; the Scirpetum maritimi compacti occurs in salt pans with saline soil flooded for a long period; the Alopecuro-Juncetum gerardii occurs on saline soil where flooding is of short duration whereas the Carici-Lolietum perennis is never flooded and occurs on only slightly saline soil.Soil salinity and duration of flooding provide a satisfactory explanation of the variation in species composition in abandoned salt pans but land-use practices, especially grazing, have to be taken into account to fully understand their floristic composition.
 
Question: In the Northern Hemisphere, species with dispersal limitations are typically absent from secondary forests. In Australia, little is known about dispersal mechanisms and other traits that drive species composition within post-agricultural, secondary forest. We asked whether mode of seed dispersal, nutrient uptake strategy, fire response, and life form in extant vegetation differ according to land-use history. We also asked whether functional traits of Australian species that confer tolerance to grazing and re-colonisation potential differ from those in the Northern Hemisphere.Location: Delatite Peninsula, NE Victoria, Australia.Methods: The vegetation of primary and secondary forests was surveyed using a paired-plot design. Eight traits were measured for all species recorded. ANOSIM tests and Non-metric Multi-dimensional Scaling were used to test differences in the abundance of plant attributes between land-use types.Results: Land-use history had a significant effect on vegetation composition. Specific leaf area (SLA) proved to be the best predictor of response to land-use change. Primary forest species were typically myrmecochorous phanerophytes with low SLA. In the secondary forest, species were typically therophytes with epizoochorous dispersal and high SLA.Conclusions: The attributes of species in secondary forests provide tolerance to grazing suggesting that disturbance caused by past grazing activity determined the composition of these forests. Myrmecochores were rare in secondary forests, suggesting that species had failed to re-colonise due to dispersal limitations. Functional traits that resulted in species loss through disturbance and prevented re-colonisation were different to those in the Northern Hemisphere and were attributable to the sclerophyllous nature of the primary forest.
 
Question: How do various plant functional groups and types are related to regeneration of limestone grasslands and to temporal changes in cover of trees and shrubs? Which mechanisms are responsible for changes in differently treated grasslands? Location: Southern Poland, 50° 01'N, 19° 50'E. Methods: A 12-year restoration experiment after the clearing of a 35-yr-old secondary pine wood developed on limestone grassland, and a parallel observation of succession in neighbouring abandoned grassland were carried out. Changes in cover of species with different functional traits and established strategies in relation to time and cover of shrub layer were analysed using multiple linear regression. Results: Over 12 years the composition and cover of species changed in the studied grasslands, but significant differences still occurred between the old grassland and the grassland restored in former wood gaps and that developed in former closed wood. Despite the very close proximity of the old grassland, the composition of restored grasslands was still much determined by initial conditions. In these grasslands, the cover of Festuco-Brometea species was significantly lower than in the old grassland. In all sites, the cover of species with large leaves and with leaves distributed regularly along the stem, and the cover of CS strategists increased. With increasing shrub cover in restored grasslands, the cover of annuals and biennials, tall species, and R and CS strategists decreased. Conclusions: Limited availability of seeds seems the principal reason for the weak regeneration of xerothermic limestone grasslands. Trees and shrubs inhibit the developed of both early successional species and perennials occurring in established communities. They significantly hinder regeneration of grasslands and should be cut every five or six years.
 
In an abandoned field in the sandy desert of Kuwait annual plants were less numerous in stands dominated by Tribulus terrestris than in adjacent stands dominated by other species.In the Tribulus stands, annuals were smaller in phytomass and stature. Possible physical and biotic factors were investigated in both types of stands in order to determine the causative agent(s) for the lesser density and development of annuals in the Tribulus stands. Physical conditions of moisture, light and soil characteristics did not appear to be limiting to annuals in the Tribulus stands.Experimental evidence is presented for the leaching from T. terrestris shoots of water-soluble substances which strongly inhibit the germination and radicle elongation of most of the associated annual species. The effect of these substances is fairly specific. Preliminary analysis indicates the presence of phenolic compounds in the leachate of T. terrestris, which are believed to play a significant role in the growth inhibition. The possible role of chemical inhibition on certain annuals in stands of T. terrestris is discussed.
 
When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi-natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long-term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass-dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.
 
Development of planted seedlings of four canopy tree species in recently abandoned pastures (mown and unmown) and in ca. 40-yr old secondary dry forest in Guana-caste National Park, Costa Rica, was studied from July 1989–June 1992. The species were the light-demanding Cedrela odorata and Swietenia macrophylla, and the shade-tolerant Hymenaea courbaril and Manilkara chicle. Seedling mortality was high and primarily correlated with dry season drought. After the first dry season, M. chicle and C. odorata showed < 5 % survival in the pasture and 20 % in the forest, but after 3 yr survival had decreased to 3% in all plots. Survival of S. macrophylla was highest in the mown pasture (45 % after 3 yr) and lowest in the forest (10 %). For H. courbaril, survival was 40–55 % under all three conditions.Growth rates were low, with a height increment of < 15 cm/yr, possibly due to low precipitation in 1990 and 1991. All species grew taller in the pasture than in the forest.Differences in soil depth, texture and drainage appeared to contribute to variation in the results both within and between plots. Herbivory contributed to the lower survival of C. odorata and S. macrophylla in the forest.
 
Questions: Did the forest area in the Swiss Alps increase between 1985 and 1997? Does the forest expansion near the tree line represent an invasion into abandoned grasslands (ingrowth) or a true upward shift of the local tree line? What land cover / land use classes did primarily regenerate to forest, and what forest structural types did primarily regenerate? And, what are possible drivers of forest regeneration in the tree line ecotone, climate and/or land use change?Location: Swiss Alps.Methods: Forest expansion was quantified using data from the repeated Swiss land use statistics GEOSTAT. A moving window algorithm was developed to distinguish between forest ingrowth and upward shift. To test a possible climate change influence, the resulting upward shifts were compared to a potential regional tree line.Results: A significant increase of forest cover was found between 1650 m and 2450 m. Above 1650 m, 10% of the new forest areas were identified as true upward shifts whereas 90% represented ingrowth, and we identified both land use and climate change as likely drivers. Most upward shift activities were found to occur within a band of 300 m below the potential regional tree line, indicating land use as the most likely driver. Only 4% of the upward shifts were identified to rise above the potential regional tree line, thus indicating climate change.Conclusions: Land abandonment was the most dominant driver for the establishment of new forest areas, even at the tree line ecotone. However, a small fraction of upwards shift can be attributed to the recent climate warming, a fraction that is likely to increase further if climate continues to warm, and with a longer time-span between warming and measurement of forest cover.
 
Question: Predicting the impact of land-use change on vegetation is vital to understanding how biodiversity and ecosystem function may respond. Is it correct to assume that abandonment is an extreme form of grazing reduction? Location: Borders and central Scotland. Methods: The analysis used data sets from two identical experiments where the impacts of two unfertilized, extensively grazed treatments and one unfertilized abandoned treatment were compared against the species dynamics of a pasture subject to normal, productive grazing management over a 16-year period. Initial multivariate analysis using Principal Response Curves was used to assess if particular traits were associated with either extensive or abandoned treatments, and was checked using univariate tests of individual traits. RLQ analysis followed by clustering into response groups was used to assess if species behaved in a similar manner between sites. Results: For many traits/attributes the shift in value or proportion was approximately linear across the extensification treatments as grazing was removed. However, certain traits showed step changes and quadratic responses. Leaf dry matter content, an important effect trait, was in the latter group. Most traits/attributes and species behaved similarly at the two sites. However, traits such as regenerative strategy, seed length, longevity and mass and seed bank type behaved differently, indicating that they are not predictable response traits. Conclusion: The results indicate that responses to grazing removal during extensification are largely straightforward and largely independent of species pool. However, there are discrepancies that suggest that simple analyses of the impacts of land-use changes such as grazing reduction may hide more complex responses.
 
Question: Can cation analyses of water samples within a peatland site be used to infer solute sources to the samples, and so provide an objective criterion of ombrotrophy?Location: Mire complexes within native Pinus sylvestris forest, in Abernethy Forest, north of the Cairngorm Mountains, Scotland, UK.Methods: Chemical analyses of major cations in 200 water samples, and χ2 analysis of contingency tables relating species occurrence in the corresponding quadrat samples to the ombrotrophic-telluric division.Results and conclusions: The chemical data (especially the ratios Ca2+/Mg2+ and Na+/Mg2+) indicated a separation of about 140 essentially ombrotrophic samples from about 60 showing clear telluric influence; these conclusions are consistent with published rainwater analyses. For hydrological and meteorological reasons, a sharp separation cannot be expected, so the limit adopted here (Ca/Mg=1.0) is to some extent arbitrary, but the methods described provide a more objective criterion of ombrotrophy than any other. Contingency tables showed highly significant associations between species occurrence and the ombrotrophic-telluric division. The strongest associations included: (ombrotrophic) Eriophorum vaginatum, Odontoschisma sphagni and Sphagnum cuspidatum; (telluric) Carex panicea, Potentilla erecta, Carex echinata, Narthecium ossifragum, Sphagnum auriculatum s.l., Agrostis canina, Molinia caerulea, Eriophorum angustifolium, and Sphagnum recurvum s.l. The Abernethy mires are arguably more comparable with mire complexes on base-poor rocks in southern Scandinavia than with most ombrotrophic sites further south and west in Britain. As in Sweden, Narthecium ossifragum and Eriophorum angustifolium, generally ubiquitous on British and Irish ombrotrophic bogs, are “fen” plants at Abernethy.
 
Secondary meadows in the Krušné hory mountains, central Europe, are not colonized by woody species, although they have been abandoned for half a century and are surrounded by Picea abies (Norway spruce) forests. The causes of inhibition of establishment of Norway spruce seedlings in the meadows were tested experimentally. The experiment was started in a masting year to ensure sufficient seed input. Four treatments (vegetation cut; vegetation cut and litter removed; all above-ground biomass and topsoil removed; control) were combined in a factorial design. The effect of browsing, mainly by deer, was assessed by fencing half of the experimental plots. Seedling establishment differed among treatments (all pair-wise differences were significant) but was independent of fencing. In contrast, seedling survival was influenced both by the treatment and fencing. At the end of the second year, only the plots with all biomass and organic topsoil removed supported viable populations of spruce seedlings. Dense herb cover and a thick layer of slowly decaying litter are considered the main factors inhibiting the establishment of woody species.
 
We used dendro-ecological techniques to investigate fundamental relationships between climate and growth of Abies alba (silver fir) in eastern France. Seven Abies forests in the Trièves region of the French Alps were chosen to represent a wide range of ecological conditions based on the results of previous forest vegetation surveys. In each forest, four trees were sampled in each of five different stands with two cores per tree. These 280 cores were studied using two separate dendro-ecological methods: the pointer years method (based on extreme growth events), and correlation functions between tree ring-widths and monthly climatic data. Data from 11 meteorological stations were combined to provide a regional analysis of precipitation and minimum and maximum temperatures. The two dendro-ecological methods appear to be complementary, as the first technique emphasizes common and low intensity linear correlations between ring-widths and climatic variations, and the second method emphasizes extreme and unusual climatic events such as exceptionally cold or dry years.Across all sites, drought in the previous year was consistently correlated with a low growth rate; however, other climatic variables varied substantially among sites. For example, drought in the current year reduced growth more in the low elevation sites than in the high elevation sites and severe winter frost reduced growth the most in the high altitude sites and the driest site. Moreover, certain growth responses are better correlated with the age of the stands, the canopy closure and the floristic composition of the community than the abiotic factors, emphasizing the value of dendro-ecological sampling based on phytosociological units.
 
We studied revegetation patterns after experimental fine-scale disturbance (e.g. uprooting) in an old-growth Picea abies forest in southeastern Norway. An experimental severity gradient was established by manipulation of the depth of soil disturbance; two types of disturbed areas were used. Species recovery was recorded in the disturbed patches in three successive years after disturbance. The cover of vascular plants and, even more so the cover of bryophytes and lichens, recovered slowly after disturbance. The least severe treatments (removal of vegetation and removal of vegetation and the litter layer) was followed by the fastest recovery. The mean number of vascular plant species was usually higher three years after disturbance than before disturbance, while the opposite was true for bryophytes. Several vascular plant species that were abundant in intact forest floor vegetation (Vaccinium myrtillus, V. vitis-idaea and Deschampsia flexuosa) recovered during a three-year period primarily by resprouting from intact rhizomes and clonal in-growth. Other important recovery mechanisms included germination from a soil-buried propagule bank (e.g. Luzula pilosa, Plagiothecium laetum agg., Pohlia nutans and Polytrichum spp.) and dispersal of propagules into the disturbed patches (e.g. Betula pubescens and Picea abies). Microsite limitation seemed to occur in several species that were abundant in the soil propagule bank (e.g. the ferns Athyrium filix-femina, Gymnocarpium dryopteris and Phegopteris connectilis) but which did not appear in disturbed patches.
 
Question: What are tree mortality rates and how and why do they vary in late-successional Picea abies-dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long-term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross-dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even-aged and originated following a stand-replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter-annual variation, which we attributed to senescence of the relatively even-aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand-replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small-scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.
 
A fertilization experiment was set up in 1969 in a mature fir forest (Abies alba Mill.) in the Vosges Mountains in France, at an elevation of 800 m, on poor granites. It consists of 65 plots (250 m2), divided into 8 treatments: control, N, P, Ca, N+P, N+Ca, P+Ca, N+P+Ca. The quantities of fertilizers were: 200 kg.ha-1 N, 150 kg.ha-1 P2O5, 1500 kg.ha-1 CaO. The humus type and the ground vegetation were described in 1969, and descriptions were repeated in 1989. Statistical analyses show that: 1. Liming, and liming only, has led to considerable alterations of the flora; the control plots are now dominated by Vaccinium myrtillus, a very acidophilous species, while the limed plots are dominated by Festuca altissima and numerous nitrophilous species. 2. Similarly, the humus layer of limed plots has changed from a mor or moder type to an acid mull type. That process might reflect an improvement of the biological activity and of the mineral nutrient cycle. 3. During the past 20 years, the unfertilized ground vegetation has changed, almost as expected from a moderate liming. Two possible causes of such a phenomenon are suggested: (a) a change in the light and temperature microclimate at the ground level, as a consequence of needle losses during the recent ‘forest decline’, which in turn could have improved organic matter mineralization; and/or (b) the chronic nitrogen deposition.
 
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Marcelo Cabido
  • National University of Cordoba, Argentina
Sandra Diaz
  • National University of Cordoba, Argentina
Sandra Lavorel
Jan Leps
  • University of South Bohemia in České Budějovice
Jean-Paul Theurillat
  • University of Geneva