Changes in Mediterranean plant succession: old-fields revisited

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

Patterns of species diversity on an oceanic versus a continental island mountain: a hypothesis on species diversification1

Abstract. The vegetation on a wet mountain slope on Haleakala (an oceanic island) is compared with that on Kinabalu (a continental island) to examine relationships between regional floristic richness and α- and β-diversities. The two mountains are similar in their constant tropical climate, generic and family-level floristic elements and geological age of the summit regions, but different in regional floristic richness (rich on Kinabalu vs. poor on Haleakala). α-diversity of canopy and subcanopy tree species was much higher on Kinabalu than in comparable zones on Haleakala. Average turnover rate of species (as logarithmic community similarity) on the slope was one order of magnitude greater on Kinabalu than on Haleakala (0.127 vs. 0.017 per 100 m alt.). While there were genera with wide altitudinal ranges on both mountains, a large proportion of the genera was differentiated into parapatric altitudinal congeners on Kinabalu. By contrast, most genera are altitudinally monotypic on Haleakala. The number of sympatric congeners per genus, and the frequency of multi-specific genera per plot were high on lower slopes but decreased with increasing altitude on Kinabalu, whereas the values were low across all altitudes on Haleakala. These patterns suggest that sympatric and parapatric species radiation was less on Haleakala than on Kinabalu. This may be related to Haleakala's initially poor and disharmonic flora.     

Effects of disturbance intensity and frequency on early old‐field succession

Abstract. Early old‐field succession provides a model system for examining vegetation response to disturbance frequency and intensity within a manageable time scale. Disturbance frequency and intensity can interact with colonization and competition to influence relative abundance of earlier and later successional species and determine, respectively, how often and how far succession can be reset. We tested the joint effects of disturbance frequency and intensity on vegetation response (species richness, abundance, canopy structure) during the first six years of succession by clipping the dominant species (D) or all species (T) in spring and fall of each year (S), once per year in summer (Y1), each two years in summer (Y2), or each four years in summer (Y4). Vegetation response reflected disturbance effects on expansion of a later monospecific dominant perennial herb, Solidago altissima, and persistence of the early, richer flora of annuals. A more abundant and taller top Solidago canopy developed on plots clipped each 2 yr or less frequently. Plots clipped yearly or seasonally were richer, but had less abundant, shorter, and differently stratified canopy. Disturbance mediated the relative abundance of early and later successional species; however, frequency and intensity effects were not completely congruent. Persistence of a richer early successional flora increased through the most frequent disturbance (S), and was magnified by disturbance intensity. Disturbance as extreme as clipping all vegetation twice yearly did not cause a drop in species richness, but maintained the early successional community over the first six years of succession. We conclude that clipping disturbance influenced the rate of succession, but the early community could rebound through the range of disturbance frequency and intensity tested.     

Vegetation analysis of the Mediterranean region of Nile Delta

Thirty-four vegetation clusters identified in the present study, after the application of TWINSPAN and DCA multivariate techniques, were assigned into 8 vegetation types, each of definite vegetation and habitat characters. The suggested vegetation types are well segregated along the DCA axis one which reflects soil moisture, salinity (as indicated by EC values), fertility (as indicated by the organic matter and nitrogen contents) and species diversity gradients. In general, soil moisture and soil fertility increase and species diversity decreases with the following sequence of vegetation types:Echinops spinosissimus-Ononis serrata on inland sand dunes,Pancratium maritimum on coastal sand dunes,Halocnemum strobilaceum-Salsola kali in saline sand deposits,Atriplex halimus-Chenopodium murale along the terraces and slopes of drains,Arthrocnemum glaucum-Tamarix nilotica in salt marshes,Chenopodium murale along the slopes of drains,Phragmites australis along the littoral zones of drains, andLemna gibba-Potamogeton crispus in the water zone. This sequence reflects also a gradient of human interference, starting with the vegetation of the less disturbed habitats (sand dunes and saline sand deposits) and ending with the fully man-made habitats (drain zones).     

Early secondary succession in a southeastern U.S. alluvial floodplain

Abstract. Species number, turnover and early successional patterns were examined over the first five years of old-field succession in a former bottomland hardwood forest. Number of species, measured by walk-through surveys, increased up to Year 3, but decreased in Years 4 and 5. Species turnover in 20 quadrats indicated that species’invasions remained high throughout the study, while losses of species were initially high but declined. Detrended Correspondence Analysis (DCA) on species cover in each year showed the existence of a compositional gradient that could be significantly related to relative elevation, an indirect measure of hydrology, in Years 2, 3, 4, and 5. DCA of the pooled five-year species cover data showed greater compositional changes in lower elevation quadrats, and those higher elevation quadrats dominated by the woody vine, Campsis radicans, remained similar over time.     

Spatial distribution patterns of Rhynchostegium megapolitanum at the landscape scale ‐ an expanding species?

Question: Is Rhynchostegium megapolitanum an expanding species? Location: Viennese Basin (120‐220 m a.s.1.), Austria. Methods: 121 dry grasslands, were investigated for the occurrence of R. megapolitanum. Nineteen environmental variables at 50 randomly selected sites, species composition at sites with and without R. megapolitanum and the spatial patterns of distribution of the species at the landscape scale were analysed. We compared actual distribution data of three rare species (Didymodon acutus, Pleurochaete squarrosa R. megapolitanum) and a common one (Brachythecium rutabulum) with the distribution obtained by vouchers that were collected between 1860 and 1940 in the investigated area. We calculated a GIS based model pattern and compared it with the actual distribution. Results: R. megapolitanum was detected in 28 of these sites, almost 50 % of its populations produced sporophytes. We found significant differences between sites with and without R. megapolitanum with regard to grassland size, the percentage of silt and of sand in the soil. There were fewer occurrences of historic herbarium vouchers of R. megapolitanum than our current field survey discovered. The GIS based analyses of distribution patterns at the landscape scale showed a clustering of sites in which R. megapolitanum was present or absent. Simulations with a spatially realistic expansion model showed high similarities to the actual distribution of the species. Conclusions: All these analyses suggest that R. megapolitanum has been expanding in the investigated area. A significant increase in temperature and nitrogen deposition within the last hundred years might be the underlying cause for the species' spread.     

A model of alpine species distribution in relation to snowmelt time and altitude

Abstract. The focus of this study is the response of species to time of snowmelt and altitude in alpine areas and an examination of changes in species response to snowmelt as altitude increases and temperature decreases. Transects (n= 43) were placed evenly along an altitudinal gradient at Finse, Hardanger‐vidda, western Norway, from ridges to late snowbeds. These gradients were systematically sampled (‘Repeated Gradient Analysis, RGA’) and an adjusted F‐test was used to determine repeated trends in species distribution along the transects. Of the 41 taxa analysed 22 showed a significant change in expected occurrence in response to time of snowmelt (when a site becomes free of snow) as altitude increased. Three types of response were observed: (1) no change in response: (2) increased occurrence as altitude increases, i.e. the taxon invades snow‐free sites as altitude increases, and (3) decreased occurrence as altitude increases, i.e. the taxon retreats from snow covered areas. It is suggested that the changes in response are due to both environmental factors (temperature related) and biological interactions. Decreases in expected occurrence are probably due to increased environmental severity as altitude increases (temperature related decreases). These species are represented by taxa preferring intermediate cover of snow. The invasion of earlier snow‐free sites is probably due to reduced competition from lee‐side taxa as altitude increases. A predictive model based on the species‐environmental relationships suggests that a 1°K temperature increase changes the limits of occurrence in response to time of snowmelt from 3 to 20 days for the different taxa.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

The relationship between the soil seed bank and above-ground vegetation of a coastal barrier island

Abstract. The germinable soil seed bank is described from a coastal barrier island off the northwest coast of Florida, USA. Soil samples collected from seven vegetation types, recently deposited dredge spoil and unvegetated areas in autumn 1990 and spring 1991 were placed out in greenhouse trays. 110 taxa germinated from the samples with the largest number (41) being C₃ perennial dicots. The largest number of taxa germinated from dry (57) and wet (54) swales, the fewest (one species: Heterotheca subaxillaris) from strand. Similarity of seed bank densities to above-ground species cover was low (Jaccard's Index = 0.36), not different between vegetation types, but higher in the autumn than in the following spring. Compositional gradients in the seed bank and above-ground vegetation determined using DCA ordination were highly correlated and related to distance from mean high water, and plot elevation. At the landscape scale, the seed bank provided an equally clear delineation of vegetation types to that based upon the above-ground vegetation. The seed bank of low disturbance, late-succession vegetation types (wooded dunes, swales, marshes) was well developed (high species richness, emergent density, and percentage annual species) with the exception that the large-seeded woody species (i.e. Quercus spp.) were absent from the wooded dune seed bank. By contrast, a poorly developed and transient seed bank occurred in more frequently disturbed (extensive sand movement, salt spray), early successional dredge spoil, unvegetated areas and strand. These contrasts support a general pattern of increasing seed bank development and a persistent rather than transient seed bank with decreasing disturbance frequency, increasing time since disturbance and successional maturity.     

Effects of cutting and nitrogen deposition on biodiversity in Cantabrian heathlands

Questions: Does the diversity of heathland vegetation change when subjected to experimental disturbances such as cutting and nitrogen fertilization? Do changes in the vegetation structure negatively affect the regeneration of the dominant species Calluna vulgaris? Is cutting an alternative method of conserving the diversity and maintaining the structure of heathlands in the Cantabrian Mountains? Location: Calluna vulgaris heathlands on the southern slopes of the Cantabrian Mountain range, NW Spain. Methods: A total of 60 plots were treated with different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (56 kg‐N.ha^?1.yr^?1). The changes in the cover values of the species present were monitored over a five year study period. The cover values were used to calculate abundance and species richness. Results: Fertilizing with nitrogen allows biodiversity to increase over time. However, the greatest biodiversity is associated with the cutting plus fertilization treatment, since cutting allows gaps to be opened that are easily colonized by pioneer annual species, while fertilization mainly favours an increase in the mean number of perennial herbs (graminoids and forbs). Increased perennial herb richness also corresponds to a rise in their cover values. The recovery of the dominant woody species in these communities, Calluna vulgaris, is not impeded by the increase in perennial herbs species' cover values. Conclusions: In the Calluna vulgaris heathlands studied, cutting plus fertilization allowed an increase in biodiversity over time. No displacement of the dominant woody species, Calluna vulgaris, is observed due to the presence of the perennial herbs. Cutting patches of heathland is recommended as a mechanism for maintaining high vegetation diversity, when grazing is not possible.     

Influence of a developing tree canopy on the yield of Pennisetum pedicellatum sown on a mine spoil

Abstract. Shoot and root biomass yield of a sown grass, Pennisetum pedicellatum, were measured at below-canopy, canopy edge and open locations in young monoculture stands of eight tree species planted on a coalmine spoil. Incident light as percentage of full sunlight decreased from open to canopy edge to below-canopy locations. The shoot and root weights of Pennisetum in different tree stands for each of the three locations were significantly different and were significantly related to each other, and to percentage sunlight across all tree species plots and locations. The gradient of incident light was the principal factor governing the gradient of grass biomass under developing canopies of tree plantations on the mine spoil.     

Effects of long‐term grazing management on sand dune vegetation of high conservation interest

Question: Can long‐term grazing management maintain and restore species‐rich sand dune plant communities within a sand dune site of high conservation interest? Location: Newborough Warren, North Wales, UK. Methods: Vegetation changes that occurred between 1987 and 2003, subsequent to grazing by domestic livestock being introduced to the site after decades with little or no stock grazing, were analysed using data collected from permanent monitoring quadrats over a 16‐year period. Results: At the plant community level, grazing brought about a shift from a tall‐grass dominated, species‐poor community to a more species‐rich community in the dry dunes, but did not change community type in dune slacks. However, at the species level, grazing enhanced the abundance of some desired perennial, annual and biennial species, graminoids and bryophytes in both habitat types. The increased frequency of positive indicator species for habitat condition suggests that grazing was beneficial for species of conservation interest. Ellenberg nitrogen (N) values decreased after grazing in dry habitats but showed no long‐term change independent of grazing, suggesting no increase in site fertility over the study period. Surprisingly, light (L) values also decreased in the dry dunes after grazing. Conclusions: Long‐term grazing management can play an important role for the conservation of dune communities and associated species. Because of its positive effects on species diversity, plant communities and habitat condition in sand dunes, livestock grazing is recommended for conservation management.     

Disturbance regimes and vegetation dynamics: role of floods in riverine wetlands

Abstract. This study tested whether the frequency of flood disturbances was able to slow down or stabilize vegetation succession in former braided channels over a decade. According to the Patch Dynamics Concept and to succession theory, species richness and diversity should be high but stable in the frequently (40 days/year) flooded channel, and should change over time in the infrequently (1 day/year) flooded one. Within the frequently disturbed channel, composition of vegetation as well as species richness and diversity appeared stable through dynamic equilibrium over the decade. Only one zone, because of particular geomorphological features that decreased disturbance intensity, developed highest diversity and richness as expected from the Intermediate Disturbance Hypothesis. The highest disturbance effect decreased species richness and was related to a higher spatial heterogeneity of the substrate (number of grain-size classes). In the other zones, richness and diversity appeared to be lowest where disturbance frequency was lowest or disturbance intensity was highest. From 1981 to 1987, the infrequently flooded channel underwent succession, and species richness increased in the major part of the channel, whereas diversity increased only in its extreme parts.     

Species response curves along environmental gradients. A case study from SE Norwegian swamp forests

Abstract. Vegetation science has relied on untested paradigms relating to the shape of species response curves along environmental gradients. To advance in this field, we used the HOF approach to model response curves for 112 plant species along six environmental gradients and three ecoclines (as represented by DCA ordination axes) in SE Norwegian swamp forests. Response curve properties were summarized in three binary response variables: (1) model unimodal or monotonous (determinate) vs. indeterminate; (2) for determinate models, unimodal vs. monotonous and (3) for unimodal models, skewed vs. symmetric. We used logistic regression to test the influence, singly and jointly, of seven predictor variables on each of three response variables. Predictor variables included gradient type (environmental or ecocline) and length (compositional turnover); species category (vascular plant, moss, Sphagnum or hepatic), species frequency and richness, tolerance (the fraction of the gradient along which the species occurs) and position of species along each gradient. The probability for fitting a determinate model increased as the main occurrence of species approached gradient extremes and with increasing species tolerance and frequency and gradient length. Appearance of unimodal models was favoured by low species tolerance and disfavoured by closeness of species to gradient extremes. Appearance of skewed models was weakly related to predictors but was slightly favoured by species optima near gradient extremes. Contrary to the results of previous studies, species category, gradient type and variation in species richness along gradients did not contribute independently to model prediction. The overall best predictors of response curve shape were position along the gradient (relative to extremes) and tolerance; the latter also expressing gradient length in units of compositional turnover. This helps predicting species responses to gradients from gradient specific species properties. The low proportion of skewed response curves and the large variation of species response curves along all gradients indicate that skewed response curves is a smaller problem for the performance of ordination methods than often claimed. We find no evidence that DCA ordination increases the unimodality, or symmetry, of species response curves more than expected from the higher compositional turnover along ordination axes. Thus ordination axes may be appropriate proxies for ecoclines, applicable for use in species response modelling.     

Management of three endangered plant species in dynamic Baltic seashore meadows

Question: Arctophila fulva var. pendulina, Primula nutans var. jokelae and Puccinellia phryganodes are threatened early successional species growing in the seashore meadows of the northern Baltic Sea. Patches formed by these species are destined to be replaced by other species during primary succession and in order to persist in the area they have to continuously colonize new areas. We studied whether the displacement of the species could be slowed down and their sexual and/or vegetative reproduction enhanced by management targeted to surrounding vegetation. Location: Bothnian Bay, Baltic Sea, W Finland. Methods: Vegetation surrounding patches of all study species was mown in four successional growing seasons. Moreover, the impact of additional soil turning on creating new favourable growing sites was tested for A. fulva. Results: Deterioration of suitable habitats of A. fulva and P. nutans was markedly slowed down by management and the vegetative and/or sexual reproduction of these species was enhanced. In the case of P. phryganodes, however, no positive response to management was obtained. Conclusions: In order to improve the long‐term persistence of these three species successional vegetation changes should be slowed down and their dispersal and colonization success improved by continuous management of the populations. We further suggest that the colonization of new areas should be aided by transplantations to the non‐vegetated islets, which have recently risen from the sea and cannot be reached by means of dispersal.     

Tropical forest structure and diversity in relation to altitude and disturbance in a Biosphere Reserve in central India

Questions: Is species diversity affected in protected areas where human activities are permitted or tolerated? On plots of a fixed size, does stem density alone predict number of species? Are differences in density related to disturbance and altitude? Location: Achanakmar‐Amarkantak Biosphere Reserve, central India. Methods: 42 sites, each with three replicate 10‐m radius plots, were examined. All trees (≥ 30 cm GBH) in each plot were measured for girth at breast height. α‐diversity, species richness and evenness were calculated for each site. The sites were ordinated by Nonmetric Multidimensional Scaling (NMS) using relative importance values of component species. Correspondence Analysis was used to broadly delineate communities. Anthropogenic disturbances were recorded in terms of percentage of trees lopped, scale of lopping, number of domestic livestock dung piles and foot trails (both livestock and people) for each plot. Results: The NMS analysis exhibited a near linear arrangement of sites with no evidence of discrete vegetation zones. NMS axes were significantly related to altitude and disturbance scores. With increasing elevation, basal area increased but number of species, α‐diversity and its components declined monotonically. The number of species and indices of species diversity were positively associated with tree lopping and also with total disturbance. Number of species was controlled by stem density only in plots not dominated by Shorea robusta. Conclusions: Recent levels of human disturbance are associated with higher species diversity in this biosphere reserve. There is some evidence that stands at all altitudes follow the same successional pattern to dominance by Shorea, a successional pattern that also results in decreased diversity without disturbance.     

An island biogeographical view of the successional pathway in wet dune slacks

Abstract. Wet dune slacks occur as small, naturally fragmented systems in the dune landscape, isolated from other slacks. We studied the effects of slack isolation and area on the rate and direction of primary succession in a chronosequence of dune slacks. The results indicate that important changes occur in community characteristics over a period of 50 yr. Total cover and number of species increase as a result of the endogenous succession process, during which organic matter and nutrients accumulate. Consequently, competitive interactions shift from competition for nutrients to competition for light. Local factors thus determine, at least partially, the community composition in the slack. However, differences in community composition with increasing age are smaller when the slack is more isolated or smaller, suggesting a slower succession rate and biomass accumulation. Together with a lower contribution of slow dispersing species in more isolated slacks, this indicates that species accumulation is dispersal limited and thus influenced by regional factors. The stochastic variation resulting from this dispersal limited species accumulation causes a divergent successional pathway.     

Community structure of endemic Mimosa species and environmental heterogeneity in a semi‐arid Mexican valley

Abstract. In this paper we analyse six communities with seven Mimosa species in the Tehuacán‐Cuicatlán valley, Mexico. All species are endemic to Mexico and four are endemic to the valley. Mimosa species are found in (1) the ‘matorral xerófilo’ (arid tropical scrub): Mimosa calcicola, M. lacerata, M. luisana, M. polyantha and M. purpusii, and (2) the ‘selva baja caducifolia’ (tropical deciduous forest): M. adenantheroides, Mimosa texana var. filipes. Most of them occur in similar soil environments, while M. polyantha and M. calcicola establish in particular soil conditions and only M. luisana establishes in two different sites showing a wider range of adaptation to soil characteristics. The communities studied include 24 plant families, 51 genera and ca. 70 species (5% of the total flora estimated in the valley). Heterogeneity was found among the communities. Our results point to the replacement of ‘matorral xerófilo’ and ‘selva baja caducifolia’ by ‘matorral espinoso’ (thorny scrub). Thorny species (e.g. Acacia cochliacantha, Mimosa spp.) are becoming the dominant/codominant elements in the communities. Within the communities, Mimosa species have a significant influence on soil pH, organic matter and electrical conductivity values. These species contribute to ameliorate soil nutrient conditions (OM, N_tot, P, Ca, Mg, Na and K contents) as well as other environmental factors (e.g. temperature, shade) under their canopy in degraded plant communities and may serve as ‘resource islands’.     

Soil pH and species diversity in coastal dunes

Soil pH was measured at two different spatial scales in coastal dunes on Norderney, North Sea, and in Mecklenburg-Vorpommern, Baltic Sea, Germany. Relationships between the variability in soil pH, species richness and species diversity are presented. Species richness and diversity were highest in grey dunes, where soil pH was at intermediate levels; both variables were lower in yellow and brown dunes. The variability in pH increased with increasing species diversity and also with scale. Overall, soil pH variability decreased with increasing vegetation cover. The lowest pH heterogeneity was found in heath dominated by Empetrum nigrum L. and grey dunes dominated by Campylopus introflexus (Hedw.) Brid. Increasing abundance of dominant species and decreasing species diversity of vegetation apparently reduces soil heterogeneity. Decreasing species diversity of vegetation is likely to explain decreasing variability in soil pH.