Early post-fire succession in California chaparral: Changes in diversity, density, cover and biomass

For four consecutive years, following the fires in November 1993, temporal variations in species richness, cover and biomass of component plant groups in early post-fire chaparral succession were monitored on different aspects at the Stunt Ranch Santa Monica Mountains Reserve, southern California. Plant groups were categorized based on growth form, life form, ability to fix nitrogen, geographic origin and regeneration strategies. North-facing slopes exhibited higher species richness, higher species turnover rate over time and faster vegetation recovery in terms of biomass accumulation and return to pre-fire species composition. This was probably due to higher species richness and biomass of nitrogen-fixing species found on north-facing slopes in comparison to south-facing slopes. On both north- and south-facing slopes, annuals had the highest species turnover rate, followed by herbaceous perennials and shrubs. In the first four post-fire years, annual species were the largest floristic group, but herbaceous perennials and shrubs were the major contributors to community biomass. Nitrogen-fixing species and exotics contributed significantly to early post-fire community structure. Although the general trends in post-fire succession are clear in terms of temporal changes in the relative proportions of different plant groups, environmental variation and the nature of plant life histories of component species, especially dominant species, could alter such trends significantly.     

Early post-fire plant succession in Peninsula Sandstone Fynbos: The first three years after disturbance

The early post-fire plant succession in fynbos vegetation in the Mediterranean-type climate area of South Africa was studied. Relatively little has been published on this early stage of plant succession in fynbos. Annual sampling over the first three post-fire years confirmed a steady, but relatively slow increase in plant canopy cover of shrubs and graminoids (mainly Restionaceae), whereas cover of geophytes and other herbs peaked in the first year and declined significantly, thereafter. Cover of annual plants increased each year, which may relate to the persistence of a relatively open vegetation cover by the third year. The responses of reseeder and resprouter species of the Restionaceae to the post-fire environment appeared to be habitat dependent. Cover of the reseeders increased rapidly in seep areas, but their recovery was distinctly delayed in dryland areas outside the seeps. Re-establishment of the many reseeder Erica species appeared to be delayed until the second post-fire year. Seed banks of these species were possibly negatively impacted by the fire, and required dispersal of seed from unburnt areas for recruitment. In contrast to some current generalisations, species richness appeared to increase after the fire; less certainly from the first to the second year, but more certainly from the second to the third year. Therefore, this study does not support a short-term monotonic decline in species richness after fire in fynbos.     

Influence of fire on critically endangered Swartland Shale Renosterveld in the Cape Floristic Region

Questions

The degree to which renosterveld shrublands are fire‐dependent is currently unclear. To address this issue, the following questions were asked: (1) does smoke stimulate germination of soil‐stored seeds in renosterveld; (2) does recently‐burned renosterveld display changed composition and higher diversity than unburned vegetation; and (3) how do the species compositions of renosterveld soil seed banks and standing vegetation compare?

Location

Swartland, Cape Floristic Region, South Africa.

Methods

Soil seed bank samples from a north‐ and south‐facing slope were smoke‐treated and germinated to test for smoke‐stimulated germination. Burned standing vegetation was surveyed 16 months post‐fire, as was unburned vegetation on the same slopes. Seed bank species richness and density were compared between smoke‐treated and untreated samples within and between slopes. Burned and unburned standing vegetation were compared within and between slopes in terms of species richness, abundance and aerial cover. Compositional similarity of the seed banks and standing vegetation was assessed.

Results

Seed banks were dominated by annuals and graminoids. Smoke treatment had no effect, except for driving significantly higher species richness and seedling density in south‐facing slope perennial shrubs. Species richness and seedling density were significantly higher in seed banks on the south‐facing slope compared to the north‐facing slope. Burned standing vegetation exhibited significantly higher diversity than unburned vegetation. Annuals and graminoids displayed significantly higher species richness and aerial cover in burned renosterveld. The north‐facing slope contained less than half the number of species/m² compared to the south‐facing slope. The seed banks and standing vegetation showed low to intermediate similarity (Sørensen = 31%–53%), but grouped close together on an NMDS plot, suggesting intermediate similarity overall.

Conclusions

Elevated germination of perennial shrubs in smoke‐treated seed bank samples and increased diversity of post‐fire standing vegetation suggest the renosterveld in this study shows elements of a fire‐driven system. Certain species only recruited in burned sites, suggesting fire‐stimulated germination. Aspect had a major influence on plant community composition, with the mesic south‐facing slope being more diverse than the xeric north‐facing slope. The similarity between the seed banks and standing vegetation was higher than previously shown for renosterveld, and appears to be higher than for fynbos.     

Local lithological drivers of post-fire vegetation recovery and implications for fire-prone regions

Local ecosystem resilience to fire disturbance can be influenced by multiple factors, from topography and climate, to fire history and pre-fire structure of biotic communities. Here we investigated the factors affecting post-fire recovery of scrub vegetation in areas under Mediterranean climate affected by frequent fires. We hypothesized that, under comparable climatic and topographic conditions, geological factors (with bedrock type as a proxy) would be at least as important as fire history in explaining patterns of post-fire recovery. We surveyed scrub vegetation in a mountain study area in Portugal, using a stratified random sampling scheme, with fire frequency, time since last fire, and bedrock type (granite vs. schist) as stratifying layers. Based on vegetation and plant community data from 40 plots, we analyzed total species richness and composition, and the relative abundance of functional groups defined on the basis of general (non fire-specific) life-history traits. We found that, at a local scale, lithology can override fire history in determining post-fire recovery. Vegetation plots on granite exhibited a considerable development of tall scrubs and higher values of total species richness. They also hosted higher numbers of animal-dispersed woody species, of trees and tall scrubs, of woody deciduous species, and of forest, edge and tall scrub species. Differences in the post-fire development of scrub vegetation and in the functional profile of plant communities highlight the need to consider local geological diversity when establishing priorities for post-fire active restoration under scenarios of limited resources.     

Direct and indirect responses of tallgrass prairie butterflies to prescribed burning

Fire is an important tool in the conservation and restoration of tallgrass prairie ecosystems. We investigated how both the vegetation composition and butterfly community of tallgrass prairie remnants changed in relation to the elapsed time (in months) since prescribed fire. Butterfly richness and butterfly abundance were positively correlated with the time since burn. Habitat-specialist butterfly richness recovery time was greater than 70 months post-fire and habitat-specialist butterfly abundance recovery time was approximately 50 months post-fire. Thus, recovery times for butterfly populations after prescribed fires in our study were potentially longer than those previously reported. We used Path Analysis to evaluate the relative contributions of the direct effect of time since fire and the indirect effects of time since fire through changes in vegetation composition on butterfly abundance. Path models highlighted the importance of the indirect effects of fire on habitat features, such as increases in the cover of bare ground. Because fire return intervals on managed prairie remnants are often less than 5 years, information on recovery times for habitat-specialist insect species are of great importance.     

Soil seed bank dynamics in post-fire heathland succession in south-eastern Australia

Soil seed banks can exert a strong influence on the path of vegetation succession following fire, with species varying in their capacity to persist in the seed bank over time, leading to changes in seed bank composition and propagules available for post-fire colonisation. This study examined the effect of time since fire on soil seed bank dynamics in a chronosequence of seven sites spanning 26 years in a south-eastern Australian sand heathland. No significant change was evident in the species richness and density of the germinable soil seed bank, but species composition differed significantly among young (0–6 years since fire), intermediate (10–17 years since fire) and old-aged (24–26 years since fire) sites (using presence/absence data). No significant trend was observed in the similarity between the extant vegetation and the soil seed bank with time since fire. A total of 32% of the species recorded in the soil seed bank were not present in the above-ground vegetation at the same site, which suggests that species requiring fire for germination may be present in the seed bank. Most species present in the extant vegetation were not recorded (63%) or were in very low abundances in the soil seed bank (29%). The mode of regeneration appears to be the major determinant of species absence in the soil seed bank, as 66% of species occurring in the extant vegetation but not in the seed bank have the capacity to regenerate by resprouting. This study shows that a major shift in the successional pathway after fire due to altered seed bank composition is unlikely in this vegetation; most species not recorded in the seed bank are either resprouters (obligate or facultative) or serotinous, suggesting that they will readily regenerate following fire. Unless fire frequencies are high and kill fire-sensitive obligate seeders before they reach maturity, the chance that the soil seed bank could substantially alter vegetation composition within the study area after fire is low. However, it is unclear how successional pathways may alter in response to severe fires with the potential to kill both seeders and resprouters.     

Small mammal abundance in Mediterranean post-fire habitats: a role for predators?

We studied patterns of small mammal abundance and species richness in post-fire habitats by sampling 33 plots (225 m² each) representing different stages of vegetation recovery after fire. Small mammal abundance was estimated by live trapping during early spring 1999 and vegetation structure was sampled by visual estimation at the same plots. Recently–burnt areas were characterised by shrubby and herbaceous vegetation with low structural variability, and unburnt areas were characterised by well developed forest cover with high structural complexity. Small mammal abundance and species richness decreased with time elapsed since the last fire (from 5 to at least 50 years), and these differences were associated to the decreasing cover of short shrubs as the post-fire succession of plant communities advanced. However, relationships between vegetation structure and small mammals differed among areas burned in different times, with weak or negative relationship in recently burnt areas and positive and stronger relationship in unburnt areas. Furthermore, the abundance of small mammals was larger than expected from vegetation structure in plots burned recently whereas the contrary pattern was found in unburned areas. We hypothesised that the pattern observed could be related to the responses of small mammal predators to changes in vegetation and landscape structure promoted by fire. Fire-related fragmentation could have promoted the isolation of forest predators (owls and carnivores) in unburned forest patches, a fact that could have produced a higher predation pressure for small mammals. Conversely, small mammal populations would have been enhanced in early post-fire stages by lower predator numbers combined with better predator protection in areas covered by resprouting woody vegetation.     

Vegetation succession after fire in sclerophyll woodland communities in south-eastern Australia

The dynamics of vegetation regeneration after burning were examined in three dry sclerophyll communities near Canberra, in south-eastern Australia. Changes in seedling and regrowth populations were followed in permanent quadrats during the first two years after burning, compared with both the preburn vegetation and population changes over the same period in adjacent, unburnt plots. All species represented either by living plants in the tree and shrub strata and/or by seed in the soil and litter prior to burning regenerated during the first year after the fire treatments. No new species invaded the areas after burning. Species varied in their regenerative strategy and recovered after the fires either by germination of seed residual in the soil and ash or released from trees after burning, by regrowth from surviving vegetative organs, or by a combination of germination and regrowth. Both seedling input and the vegetative recovery of populations were higher during the first than second year after burning. The vegetative multiplication and seed gertnination of many species were stimulated by fire. It was concluded that the regeneration of the communities studied, as well as the post-fire recovery of numerous different communities reported in the literature, closely resembled an initial floristic composition model. It is considered that the process of vegetation redevelopment after a disturbance (i.e. secondary succession) will be influenced greatly by the species composition at the time of disturbance, and by the type of disturbing agent. A single model would not be expected to adequately describe secondary succession following disturbance by agents imposing different stress conditions on a community.     

Does helimulching after severe wildfire affect vegetation recovery in a coastal area of Northwest Spain?

Helimulching is commonly applied after high-severity wildfires in North America because of its effectiveness in reducing post-fire runoff and erosion. However, its use in other parts of the world is still very limited and information about its effects in different environments is scarce. In this study, the effects of helimulching on vegetation recovery and species composition were assessed in 70 experimental plots (80 m² each) established in five shrubland areas in Northwest Spain affected by wildfire in summer 2013. The effects on shrub, forb, fern and grass cover, and on total vegetation cover, as well as on species richness, Shannon diversity index and heterogeneity were studied over the 2 years following the fire. The impacts of soil burn severity and mulch depth on these variables were also considered. Overall, the mulching treatment had little effect on the cover variables. Although it had a positive effect on forb cover, these species represented only a small portion of the total vegetation cover. Soil burn severity was not a significant factor in explaining the variation in the variables under study. The treatment had a low impact on species composition. In the mulched plots only three non-native species were recorded and these displayed a limited capacity to act as invasive species as they were absent at the end of the period of study. The results indicate that helimulching is a feasible soil stabilization treatment with neutral effects on vegetation cover and the composition of shrubland in coastal areas of Northwest Spain.

     

Secondary succession in coastal dune fynbos: variation due to site and disturbance

Succession after fire and bushcutting in coastal dune fynbos was monitored for two and a half years and comparisons were made with adjacent, mature (13 year-old) fynbos. Sixty-two to 68% of pre-disturbance species, including all the dominants, were found in the successional communities 1.5 yr after disturbance: the patterns thus fitted the initial floristic composition model. On the more mesic south facing slope, post-fire succession differed from the north-facing (burnt) and bushcut sites in that ordinations showed a clear separation between the mature and successional communities. This difference was due to the post-fire abundance on the southfacing slope site of short-and medium-lived species not present in the mature fynbos. One and a half years after disturbance, species richness and equitability had increased relative to mature vegetation. This increase was greatest for the south facing slope where short-and medium-lived species and juveniles of pre-disturbance dominants co-occurred. In general, successional patterns were consistent with those described for other fynbos and fire-prone mediterranean shrublands.     

Post-fire regeneration in a Mediterranean pine forest with historically low fire frequency

Species of Mediterranean vegetation are known to regenerate directly after fire. The phenomenon of autosuccession (direct regeneration) has been found to be often combined with an increase of species richness during the first years after fire due to the high abundance of short-lived herbaceous plants facilitated by plentiful nutrients and light. The high degree of vegetation resilience, which is expressed in terms of autosuccession, has been explained by the selective pressure of fire in historic times. According to existing palaeoecological data, however, the Pinus halepensis forests in the Ricote Mountains (Province of Murcia, SE Spain) did not experience substantial fire impact before the presence of man nor are they especially fire-prone today. Therefore, we studied post-fire regeneration to find out if direct succession is present or if species from pre-fire vegetation are absent during the post-fire regeneration stages. Patterns of succession were deduced from observations made in sample plots on sites of a known regeneration age as well as in adjacent unburnt areas. The results of the vegetation analyses, including a Detrended Correspondence Analysis, indicate that Pinus halepensis forest regeneration after fire resembles autosuccession. As regards the presence of woody species, there is a high percentage similarity on north (83%) and south (70%) facing slopes during the first year after fire vs. reference areas which is due, for example, to direct regeneration of the resprouting Quercus coccifera or seeders like Pinus halepensis or Fumana laevipes. However, if herbaceous species are included in the comparison, the similarity on north-facing sites decreases (to 53%) with the presence of additional species, mainly ruderals like Anagallis arvensis or Reseda phyteuma, and even woody species on the burnt plots. This effect indicates “enhanced autosuccession”, which was not found on south-facing sites where overall species richness was very high irrespective of the impact of fire. Locally we found limited regeneration of some species, for example Pinus halepensis at high altitudes (1000 m), even 22 years after fire. As we assume that historical fires did not play an important role in the area and direct succession is present nevertheless, our results support the theory that autosuccession is not a process restricted to fire-prone areas. Fire has been only one of several selective forces since human settlement that probably led to a set of species pre-adapted against recurrent disturbance.     

Fire-mediated dieback and compositional cascade in an Amazonian forest

The only fully coupled land-atmosphere global climate model predicts a widespread dieback of Amazonian forest cover through reduced precipitation. Although these predictions are controversial, the structural and compositional resilience of Amazonian forests may also have been overestimated, as current vegetation models fail to consider the potential role of fire in the degradation of forest ecosystems. We examine forest structure and composition in the Arapiuns River basin in the central Brazilian Amazon, evaluating post-fire forest recovery and the consequences of recurrent fires for the patterns of dominance of tree species. We surveyed tree plots in unburned and once-burned forests examined 1, 3 and 9 years after an unprecedented fire event, in twice-burned forests examined 3 and 9 years after fire and in thrice-burned forests examined 5 years after the most recent fire event. The number of trees recorded in unburned primary forest control plots was stable over time. However, in both once- and twice-burned forest plots, there was a marked recruitment into the 10-20cm diameter at breast height tree size classes between 3 and 9 years post-fire. Considering tree assemblage composition 9 years after the first fire contact, we observed (i) a clear pattern of community turnover among small trees and the most abundant shrubs and saplings, and (ii) that species that were common in any of the four burn treatments (unburned, once-, twice- and thrice-burned) were often rare or entirely absent in other burn treatments. We conclude that episodic wildfires can lead to drastic changes in forest structure and composition, with cascading shifts in forest composition following each additional fire event. Finally, we use these results to evaluate the validity of the savannization paradigm.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Community dynamics in relation to management of heathland vegetation in Scotland

The paper describes studies of post-fire succession in heathland vegetation in N.E. Scotland, dominated by Calluna vulgaris. A preliminary model (Legg, 1978) suggested good agreement between simulation of succession on the basis of a Markov chain and observations of stands at different stages of development after burning, at least in the earlier stages. Vegetation transitions are currently being recorded in permanent plots on burnt areas. First results confirm the view that (a) the post-fire succession has the properties of a Markov process, (b) this type of model remains valid when constructed from records of actual transitions, rather than data obtained by inference from evidence of transition. Comparing successional events in stands where, at the time of burning, the Calluna population was in pioneer-, building-, mature-and degenerate phases, shows that transition matrices generally agree with the Markov hypothesis, but not in the case of stands where Calluna was degenerate when burnt. The composition of establishing vegetation 1 year after fire is not confined to species normally associated with the early stages of succession, but reflects the composition of the stand before burning. Redevelopment after fire is described in terms of an initial floristic composition of species with strategies permitting early re-establishment, selected by the recurrence of the fire factor. Subsequent transitions represent changes in their relative abundance due to differing growth properties and competitive interactions. This interpretation applies only under conditions of recurrent incidence of fire (normally once in 10–15 yr). If fire does not recur, Calluna stands pass into the degenerate phase, where changes in the nature of relay floristics may come into play (e.g. with tree colonization).Nomenelature follows Clapham, Tutin & Warburg (1962) for vascular plants; Smith (1978) for bryophytes.     

Species–area relationships in the Hantam-Tanqua-Roggeveld,Succulent Karoo,South Africa

The Hantam-Tanqua-Roggeveld subregion is part of the Succulent Karoo hotspot of biodiversity which stretches along the southwestern side of South Africa and Namibia. Forty Whittaker plots were surveyed in the spring of 2005, in eight vegetation associations, to gather diversity data for the Hantam, Tanqua Karoo and Roggeveld areas. Seven plot sizes were used to construct species–area curves using three different models namely: the untransformed linear function, the power function and the exponential function. In general, the power and exponential functions produced a more significant fit to the data than the untransformed linear function. Linear regressions using environmental parameters indicated that altitude, mean annual precipitation and mean annual temperature were significant predictors of species richness at the 1, 10, 100 and 1000 m² scales. To illustrate the variation in species–area curves and species richness across the landscape, a transect through the study area is discussed. The transect stretches eastwards from the Tanqua Karoo across the escarpment into the Roggeveld and crosses five different vegetation associations. Differences between associations were found in species richness in the 1000 m² plots. Each association also produced species–area curves with their own characteristics. Slope values for the samples within an association did not differ significantly, although the intercept value often did. Comparisons between associations along the transect revealed significant differences in the slope value between the associations, except for the Dicerothamnus rhinocerotis Mountain Renosterveld which did not differ significantly from the associations bordering it on either side.     

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

Question

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

Location

Booderee National Park, south‐eastern Australia.

Methods

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

Results

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

Conclusions

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

Vegetation establishment in post-fire Adenostoma chaparral in relation to fine-scale pattern in fire intensity and soil nutrients

At fine spatial scales (0.1–10m), chaparral communities have been shown to be strongly influenced by canopy-gap patterns, leading to periodicities in vegetation at 4–5 m spatial scales. Fine-scale variations in fire behavior and post-fire erosion can lead to changes in the patterning of viable seeds and nutrients and may alter the spatial patterning of post-fire chaparral communities. This study deals with the relationship among fire behavior, post-fire nutrient availabilities and vegetation patterns in a 1-yr old, post-fire Adenostoma fasciculatum chaparral community in the Sierra Nevada Mountains, California, USA. Variations in mineral soil exchangeable cations (Ca, Mg) and extractable phosphorus (P0₄-P) were correlated with ash distribution. Cations and measures of ammonium and nitrate were also correlated with fire intensity, measured by the diameter of the smallest remaining A. fasciculatum twigs following fire. Fire intensity was correlated with the pattern of post-fire vegetation establishment based on first axis DC A scores. However, ash PO4-P was more highly correlated with sample DCA scores, local species richness and total cover (p < 0.01), suggesting that small-scale variations in PO4-P which correlate with ash distributions may be important in structuring this community. Two- and three-term local variance analysis revealed a maximum of pattern intensity in DCA first axis scores at 4–5 m intervals that likely corresponds to pre-fire canopy-gap patterns. However, total cover showed pattern at spatial scales of 8–10 m, and was correlated at this scale with patterns of ash distribution and fire intensity. Microtopographic patterns also occur at similar spatial scales. Microtopographic patterns appear important in determining post-fire plant nutrient and water distributions and, thereby, patterns of plant establishment. Thus, the scale and intensity of post-fire vegetation pattern may differ considerably from pre-fire conditions.     

POST-FIRE RECOVERY OF FLORIDA LAKE WALES RIDGE VEGETATION

Community responses to fire of five major vegetation associations of the Lake Wales Ridge were investigated during a 5-yr post-fire period. Vegetation of southern ridge sandhills, sand pine scrub, scrubby flatwoods, flatwoods, and swales was sampled using permanent line transects and quadrats in both burned and unburned (control) areas. Fire passed through the mosaic of vegetation leaving a pattern of unburned and lightly to severely burned areas. There was little change in soil chemistry (pH, K, P, Mg) with the exception of a short-lived increase in Ca. Vegetation recovery rates, measured by percentage cover and Horn's Index, were rapid. Typically, less than 2 yr was needed for recovery of poorly drained flatwoods and swale associations and 1 to 4 years for better drained scrubby flatwoods and southern ridge sandhills. Species diversity (H′) increased significantly following fire at poorly drained sites due to increased evenness, but was largely unchanged at better drained sites. There was virtually no change in vascular plant species composition for the sprouting associations. Fire is not a succession-initiating disturbance in the Clementsian sense. The species present prior to burning either resprout soon after fire or resist fire, thus rapidly restoring the preburn conditions. Limited data suggest fire in the associations dominated by seeding species likewise does not initiate a relay type succession. The ridge vegetation exhibits marked resiliency to fire as a result of an evolutionary history of a stressful environment including winter droughts, acidic, nutrient-poor sand substrates, and frequent lightning-caused low intensity burns.     

Post-fire regeneration in a Mediterranean pine forest with historically low fire frequency

Species of Mediterranean vegetation are known to regenerate directly after fire. The phenomenon of autosuccession (direct regeneration) has been found to be often combined with an increase of species richness during the first years after fire due to the high abundance of short-lived herbaceous plants facilitated by plentiful nutrients and light. The high degree of vegetation resilience, which is expressed in terms of autosuccession, has been explained by the selective pressure of fire in historic times. According to existing palaeoecological data, however, the Pinus halepensis forests in the Ricote Mountains (Province of Murcia, SE Spain) did not experience substantial fire impact before the presence of man nor are they especially fire-prone today. Therefore, we studied post-fire regeneration to find out if direct succession is present or if species from pre-fire vegetation are absent during the post-fire regeneration stages. Patterns of succession were deduced from observations made in sample plots on sites of a known regeneration age as well as in adjacent unburnt areas. The results of the vegetation analyses, including a Detrended Correspondence Analysis, indicate that Pinus halepensis forest regeneration after fire resembles autosuccession. As regards the presence of woody species, there is a high percentage similarity on north (83%) and south (70%) facing slopes during the first year after fire vs. reference areas which is due, for example, to direct regeneration of the resprouting Quercus coccifera or seeders like Pinus halepensis or Fumana laevipes. However, if herbaceous species are included in the comparison, the similarity on north-facing sites decreases (to 53%) with the presence of additional species, mainly ruderals like Anagallis arvensis or Reseda phyteuma, and even woody species on the burnt plots. This effect indicates “enhanced autosuccession”, which was not found on south-facing sites where overall species richness was very high irrespective of the impact of fire. Locally we found limited regeneration of some species, for example Pinus halepensis at high altitudes (1000 m), even 22 years after fire. As we assume that historical fires did not play an important role in the area and direct succession is present nevertheless, our results support the theory that autosuccession is not a process restricted to fire-prone areas. Fire has been only one of several selective forces since human settlement that probably led to a set of species pre-adapted against recurrent disturbance.