Plant strategies and secondary succession on Brittany heathlands after severe fire

Abstract. Plant succession on heathlands subjected to major fire disturbance and humus burn in 1976 was studied over twelve years following burning. Life history strategies of principal heathland species are described with reference to concepts outlined by Grime (1979) and Whittaker & Goodman (1979). Heathlands and closely related communities are characterised by dominance of speciestolerant of physical stress (‘S’ strategists) whereas species which colonise disturbed sites are closer to rude-rals (‘R’ strategists). After severe burning three main successional patterns were identified. They depend on water and nutrient availability relative to temporal population dynamics. Recovery of heathland is often retarded due to monospecific dominance, e.g. of Polytrichum commune, Molinia caerulea and Betula pubescens. These patterns of secondary succession illustrate the inhibition model advanced by Connell & Slatyer (1977).     

Spatio-temporal patterns in post-burn succession on Brittany heathlands

Heathlands in the Paimpont area, Brittany, were ravaged by numerous fires in the summer of 1976. The period in which the fires occurred, and their intensity had important consequences for recolonization processes. In the tall Ulex europaeus heathland described in this paper, fire occurred relatively late (mid-July), and was of high intensity. Colonisation was mainly from propagules, often from species foreign to the original heathland. Plant succession together with vertical and horizontal structure was studied in permanent plots over 13 yr. Three stages were distinguished. Vertical structure varied greatly among the three stages. It was very simple in the colonisation phase, became more complicated during the competition phase, but became simpler again in the stabilisation phase. This was particularly evident for the moss layer, which, 13 yr after the fire, was made up of Polytrichum commune alone. Mosses showed interspecific competition with size as the decisive factor: the large mosses gradually eliminated the smaller ones. Horizontal structure was studied using spatial autocorrelation, which provided some indication of the distinctive -ness and scale of spatial pattern. Some species were weakly aggregated such as Ceratodon purpureus, Polytrichum pili-ferum, and P. juniperinum, whileothers, like the P. formosum-P. commune complex, Agrostis curtisii and Ulex minor were strongly aggregated. In the current community, 13 yr after the fire, a mosaic can be distinguished with three components: (1) a moss layer of Polytrichum commune on which a herbaceous layer, dominated by Agrostis curtisii has developed; (2) a birch wood (both Betulapéndula and B. pubescens), which established on the Polytrichum - Agrostis combination; (3) patches of Ulex minor heathland, with a few Frica and Calluna plants. Extreme simplicity and stability are the two prominent features of the present vegetation state.     

Vegetation dynamics in Brittany heathlands after fire

The vegetation dynamics of heathlands in Brittany have been followed for three years in areas subject to fire in August 1976. The pre-fire vegetation had been analysed and mapped before its destruction. The structure of the community and the processes taking place in it (in terms of biomass, primary productivity, phenology, mineral nutrition and food value) had been examined. The redevelopment of the canopy was studied by the point-contact method along permanent line transects. This semi-quantitative study permits calculation of the relative frequency of each species and, from this, its cover. The growth form of each species and the stratification of the community are also indicated by this method. Permanent plots were also used to record changes in the vegetation, by means of a census of individuals and records of the development and growth strategy of each species. The plots were located in homogeneous areas, or on bare soil around seed parents in order to examine seed dispersal and seedling establishment. These two methods yielded detailed information on the nature of the secondary successions following fire in the heathlands of Brittany.

     

Colonization and early succession on anthropogenic soils

Abstract. This paper reports on vegetation development on permanent experimental plots during five years of succession. Nine (1 m²) plots were filled with three typical substrates from man-made habitats of urban and industrial areas in the region of Berlin. The three substrates (a commercial ‘topsoil’, a ruderal ‘landfill’ soil and a sandy soil), differ in organic matter and nutrient contents. Relevés of species composition and percent cover of each species present were made monthly during the growing season from the start of vegetation development. This paper describes the different successional pathways on topsoil and ruderal soil and the colonization process on sandy soil. On topsoil, ruderal annuals are dominant in the first year and are replaced by short-lived perennials from the second year. Those species were replaced by long-lived perennial herbs (Ballota nigra or Urtica dioica) from the third year of succession onwards. On the ruderal land-fill soil the early successional stages are less sharp and the perennial Solidago canadensis is able to dominate within one year after the succession was initiated. On sandy soil there is still an ongoing colonization process, where pioneer tree species like Betula pendula and Populus nigra play a main role. The importance of ‘initial floristic composition’, the role of substrate for community structure and the peculiarities of successional sequences on anthropogenic soils in the context of primary and secondary successions are discussed.     

Seaweed succession on artificial reefs on different bottom substrata

Artificial rest reefs were set on sandy and rocky bottoms at 5–10 m depth along the coast of southern Japan. Mature thalli ofSargassum, Gelidium and other seaweds were transported from other coastal areas, packed in mesh bags and attached to the reefs to start the beds. After one year, the seaweed flora on the reef on a sandy bottom consisted of more than 20 species, includingSargassum spp. andGelidium amansii, which are important animal food species. Coralline algae were the dominants on the rocky bottom reefs. The lower biomass on reefs on the rocky bottom was due to grazing by urchins. The same number of species was present in the first and second years on reefs on sandy bottoms, but there were moreSargassum thalli the second year.Maximum algal biomass of the artificial reef in May of the second year was 9998 g wet wt m^–2 in sandy areas, 441 g wet wt m^–2 in boulder areas and 228 g wet wt m^–2 in rocky areas. Reefs on rocky bottoms continued to be covered by coralline algae and several species ofCodium andDictyota.     

Conservation of heathland by sheep grazing in Brittany (France): Importance of grazing period on dry and mesophilous heathlands

Heathlands are characteristic semi-natural ecosystems of western Europe. Nowadays they are threatened because of their lack of agricultural value. Management of the remaining heathland areas necessitates precise knowledge about different management tools that can be used, including grazing. The effects of sheep grazing on dry and mesophilous heathlands of Brittany were studied in relation to grazing season and duration. Management aims were to control shrubs and to maintain the dominance of heathers. The grazing pressures applied here were quite high, about 10–15 sheep ha⁻¹ yr⁻¹. Erica ciliaris appeared to be very sensitive to grazing at the end of summer. This implicates special care in the management of mesophilous heathland. In dry heathland, Erica cinerea did not show this sensitivity. Control of shrubs was achieved by most grazing regimes which led to a decrease of 11–22 cm in gorse height, except summer grazing which seemed to stimulate their growth.This study underlined the relevance of the use of sheep grazing as a heathland management tool but also the necessity of prior experiments to adapt these operations mainly in terms of grazing period to the type of heathland and the objectives put forward.     

Benthic colonization and succession on temperate sublittoral rocky cliffs

Patterns of benthic colonization and succession were investigated on a temperate rocky cliff (Aegean Sea, Eastern Mediterranean). Cement and ceramic panels deployed on the rocky substratum at 25-30 m depth were sampled every 3 months over a 2-year period yielding 28 floral and 156 animal species. Diversity, cover and abundance had low values at short immersion periods and increased at the long ones. The vagile fauna responded to the increasing habitat complexity offered by the development of algal turfs and sessile species, and the community structure remained highly dynamic. Unlike the duration of immersion, the type of substratum didn't affect species colonization except for decapods that showed a preference for cement panels. Three to four stages of succession were recorded over immersion periods, according to vagile and sessile biota, respectively, during which species richness and abundance increased. The structure of the developed communities on both artificial materials differed from the natural algal-dominated benthic community of the same area, suggesting that the recovery of rocky shore communities on temperate cliffs is time consuming.     

Diatom flora of a kettle-hole bog in relation to hydrarch succession zones

Diatom assemblages were compared to the vegetational seres and to physical and chemical parameters of a kettle-hole bog which had distinct hydrarch successional zones and was in turn surrounded by a cattail marsh and a willow swamp. Multivariate analysis of variance showed the water parameters (temperature, pH, conductivity, color, free calcium) among the zones were significantly different (p < 0.0001). Diatoms were enumerated using the first 500 valves counting method and results showed each zone had a distinct flora. Cluster analysis of the diatom data by both species and zones grouped the bog successional zones into one major group and the marsh/swamp zones into the second major group. The occurrence of well defined seral stages of vegetation and diatoms suggests microorganisms are undergoing ecological succession in kettle-hole bogs.     

Soil nitrogen mineralization in a coastal fynbos succession

Patterns of net nitrogen mineralization and nitrification in 0–7.5 cm deep mineral soils of different stages (seral ages 1, 6 and 20 years) of a post-fire coastal fynbos succession were assayed using laboratory andin situ incubations. No evidence of increasing allelopathic inhibition of nitrification with successional development was found as NO₃−N was the predominant product at all seral stages and the NO₃−N∶NH₄−N ratio remained constant. Rather the results of field incubations of soils beneathProtea repens stands of different successional ages showed that increased mineralization and nitrification appeared to be associated with increased soil total N content rather than with successional age. Further, the incubation of soilsin situ during the dry summer months showed that NO₃−N production appears to be closely related to temperature and soil moisture content, both of which are variables that vary throughout succession due to the changing structure of the vegetation.     

Effects of ice duration on plankton succession during spring in a shallow polymictic lake

1. Long-term records of air temperature and ice phenology (ice duration), and phyto- and zooplankton time series (1979–1997) were used to study the effects of ice duration on the successional pattern within plankton communities during spring in a shallow polymictic lake. 2. Water temperature in March was significantly lower after cold winters when compared to average or mild winters. Mean water temperature in April was not significantly different after mild, average or cold winters, but showed an overall significant negative correlation with ice duration. 3. Ice duration affected the timing and the magnitude of the peak abundance of diatoms, rotifers and daphnids during spring, but had no direct effects on the timing and maximum of chlorophytes, cryptophytes, cyanobacteria, bosminids and cyclopoid copepods. 4. Plankton groups which appeared first in the seasonal succession (i.e. diatoms, rotifers and daphnids) reached maximum abundance earlier after mild and average winters. The peak abundance of diatoms was negatively correlated with ice duration, whereas that of rotifers and daphnids was independent of the conditions during the preceding winter. 5. Temperature alone was generally a poor predictor of the timing and magnitude of both phyto- and zooplankton maxima. Turbulence may be important in the timing and the magnitude of peaks in diatoms, while total algal biomass was the most important determinant for the timing of the rotifer maximum. The magnitude of the daphnid maxima were significantly influenced by water temperature in March and April, and by rotifer abundance. The magnitude of the bosminid maximum was correlated with food availability and predation, whereas the timing of the maximum was more closely related to water temperature in May. 6. We conclude that, as a result of the low heat storage capacity of shallow lakes, the effects of winter on planktonic communities are short lived, and soon overtaken by the prevailing weather and by biotic interactions.     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Dynamics of nitrogen and phosphorus retention during wetland ecosystem succession

We compared the mechanisms of nitrogen (N) and phosphorus (P) removal in four young (<15 years old) constructed estuarine marshes with paired mature natural marshes to determine how nutrient retention changes during wetland ecosystem succession. In constructed wetlands, N retention begins as soon as emergent vegetation becomes established and soil organic matter starts to accumulate, which is usually within the first 1–3 years. Accumulation of organic carbon in the soil sets the stage for denitrification which, after 5–10 years, removes approximately the same amount of N as accumulating organic matter, 5–10 g/m²/yr each, under conditions of low N loadings. Under high N loadings, the amount of N stored in accumulating organic matter doubles while N removal from denitrification may increase by an order of magnitude or more. Both organic N accumulation and denitrification provide for long-term reliable N removal regardless of N loading rates. Phosphorus removal, on the other hand, is greatest during the first 1–3 years of succession when sediment deposition and sorption/precipitation of P are greatest. During this time, constructed marshes may retain from 3 g P/m²/yr under low P loadings to as much as 30 g P/m²/yr under high loadings. However, as sedimentation decreases and sorption sites become saturated, P retention decreases to levels supported by organic P accumulation (1–2 g P/m²/yr) and sorption/precipitation with incoming aqueous and particulate Fe, Al and Ca. Phosphorus cycling in wetlands differs from forest and other terrestrial ecosystems in that conservation of P is greatest during the early years of succession, not during the middle or late stages. Conservation of P by wetlands is largely regulated by geochemical processes (sorption, precipitation) which operate independently of succession. In contrast, the conservation of N is controlled by biological processes (organic matter accumulation, denitrification) that change as succession proceeds.     

Phytoplankton community succession in a lake subjected to artificial circulation

East Sidney Lake, a small, eutrophic bottom release impoundment in NY, has undergone artificial circulation for three seasons. The artificial circulation system resulted in an overall reduction in the physical stability of the water column, making the lake subject to alternating periods of weak chemical stratification and mixing. Phytoplankton community succession exhibited a high degree of regularity from year to year, culminating in mid summer dominance by heterocystous cyanophytes in all years. Changes in the physical structure of the water column, with attendant changes in Z _eu :Z _mix , were not important determinants of phytoplankton community makeup in East Sidney Lake. Seasonal patterns and community characteristics were not affected by artificially induced alterations in stability, but instead were most sensitive to surface temperatures, flushing rate and TN:TP. The timing of cyanophyte blooms was not affected by artificial circulation, nor was maximum seasonal phytoplankton biomass reduced.     

Changes in atrazine toxicity throughout succession of stream periphyton communities

A study was made to describe atrazine toxicity and its changes throughout succession of periphyton communities of an undisturbed Mediterranean stream. Toxicity was assessed by short-term physiological tests (concentration-effect curves of photosynthesis to atrazine) in the laboratory using artificial substrates colonized in one stream site during winter, and two stream sites (one open and the other shaded) during summer. In the winter experiment, when environmental conditions were relatively steady and chlorophyll content was low, toxicity increased according to the increases in cell density and chlorophyll content throughout colonization. EC₅₀ (concentration inhibiting photosynthesis by 50%) was above 0.8 μM atrazine until day 16 and below 0.4 μM atrazine after three weeks. In the summer experiment, under more variable environmental conditions, the differences between the EC₅₀ at the beginning and the end of the colonization experiments were not significant (one factor ANOVA) at the two sites. EC₅₀ was on average 0.89 μM atrazine in the shaded site and 0.29 μM atrazine in the open site. A significant negative correlation between irradiance and EC₅₀ was observed all the experiments were considered together (r = 0.464, n = 20, p<0.05), suggesting that light history may have an important role in the response to atrazine. This investigation reveals that the response of stream periphyton to atrazine is likely to be influenced by colonization time and the corresponding changes in algal density and community composition as well as by environmental conditions (e.g. light regime) throughout succession. This revised version was published online in September 2006 with corrections to the Cover Date.     

Seed bank dynamics of two obligate seeders, <Emphasis Type="Italic">Cistus monspeliensis</Emphasis> and <Emphasis Type="Italic">Rosmarinus officinalis</Emphasis>, in relation to time since fire

Many species in Mediterranean-type ecosystems regenerate after fire by seed germination from soil seed banks. Seed bank dynamics of two of those obligate seeders, Cistus monspeliensis and Rosmarinus officinalis, were investigated in relation to stand age since fire in southwestern Portugal. Soil seed density, annual seed input, annual seed losses through germination and seed persistence were compared between species at stands differing in age since fire (5, 10 and 35 years). Soil seed density and seed input increased over the first decade after fire and were lowest at 35-year-old stands for C. monspeliensis. In R. officinalis, few seeds were produced and found in the soil at early stages, and maximum seed input and soil seed density were attained at 35-year-old stands. Soil seed density was mostly driven by seed production in both species, which is largely dependent on plant traits and population dynamics related to fire. Overall, stand age since fire had a negligible effect on seed germination, seed persistence and viability. Ten to 39% of buried seeds were not recovered after 1 year, and viability of seeds recovered was 97–100% for C. monspeliensis and only 0–3% for R. officinalis. Variation in plant traits within the seeder syndrome was evidenced by this study. R. officinalis evidenced lower seed persistence, lower proportion of viable seed produced and lower density of viable soil seed than C. monspeliensis at any stage after fire. R. officinalis is expected to depend largely on previous year seed production for population replacement after fire.     

Primary forest succession on poor sandy soilsas related to site factors

Three main succession lines are found to occur on poor sandy soils in a blown drift sand landscape in the Netherlands. Site factors – organic matter, parent material, soil moisture regime and relief – were determined. Site types, specific combinations of these site factors, are defined. For five site types succession was studied. The primary criterion for separating succession lines and for delineating the successional stages, was the site factor organic matter and the second criterion, groundwater level. Influence of organic matter on succession was significant on site types with a buried podzol. Species composition on these site types indicated richer sites than the species composition on the site types without a buried podzol. Groundwater level of about 1m with gleyzone was critical for species requiring moist sites. Within a succession line vegetation stages are characterized and the potential forest type is presumed. The stages were related to humus form and soil development. Litter of dominant species was suggested to be the main criterion for the humus form. Initial and older vegetation stages were linked to development of soil horizons. The defined site types, their soil development and relationships with the succession lines clarify spatial and temporal structure and diversity of vegetation, and herewith is the basic information for clarifying the heterogeneity and potential biodiversity of this blown drift sand landscape.     

Spring grazing by sheep: effects on seasonal changes during early old field succession

This paper describes early secondary succession on an old field on limestone released from cultivation four years previously. Seasonal changes in plant composition after spring grazing by sheep are compared with those in ungrazed controls. Grazed and ungrazed paddocks were laid out in Latin squares. Plants were sampled before and several times after grazing in April, at several spatial scales.Major changes in plant abundance and sward characters such as height and density persisted throughout the growing season. Annual herbs increased after grazing, but annual grasses declined, as did short-lived perennial herbs. Effects on perennial herbs were weak; perennial grasses usually increased but this depended on the species. This pattern confirms that sheep grazing affects the direction, as well as the rate of succession.Some effects, such as increases in biennial herbs and in species richness, were only evident at large scales of sampling, suggesting that they arose from changes in rare and widely dispersed species. Other species were affected at different spatial scales, and no one sampling method detected the full range of effects.These results indicate the potential power of manipulating grazing early in secondary succession for directing the course of community change, for conservation or other purposes.Nomenclature follows Clapham et al. (1962)This study is funded by a grant from the Nuffield Foundation to V.K.B. We thank the Wytham Management Committee, Oxford University, for permission to use the site, and the staff of the University Farm, especially Richard Bampton and Cheryl Howes, for making the grazing possible. Alan Gange, Nicola marples, Fiona Matthews, Bridget Peace, Anne Storr and others helped at various times with field recording.