Spatial relationships between the standing vegetation and the soil seed bank in a fire-prone encroached dehesa in Central Spain

Postfire vegetation regeneration in many fire-prone ecosystems is soil seed bank dependent. Although vegetation and seed bank may be spatially structured, the role of prefire vegetation patterns and fire in determining postfire vegetation patterns is poorly known. Here, we investigated the spatial patterning of species abundance and richness in the vegetation and seed bank of a Mediterranean encroached dehesa in Central Spain. The seed bank was studied with and without a heat shock simulating a spatially homogeneous fire. Semivariograms and cross-semivariograms showed that species richness in the vegetation was aggregated in patches, mainly of herbs, with highest values corresponding to high herb cover and low tree cover. Species richness in the seed bank was also structured in patches, but the spatial pattern was weak. Seedling density of germinates in the seed bank also showed weak spatial pattern. Heating increased overall germination and species richness, and the intensity of the spatial pattern of species richness, particularly of herbaceous species. However, seed bank density patterns disappeared after heat shock because of increased germination of shrubs without spatial pattern. Our results document that the spatial structure of plant richness in the vegetation may persist after fire due to the spatial patterns of herbaceous species in the seed bank, and that postfire species richness patterns can arise independently of fire intensity patterns. However, the spatial structure of the vegetation after fire can be altered by the feedback between shrub encroachment and an eventual fire because of the ubiquitous germination of shrubs.     

Time of germination and establishment success after fire of three obligate seeders in a Mediterranean shrubland of central Spain

Aim Determining how differences in time of germination can affect plant establishment in plant communities that, after a disturbance, must reestablish from seeds under climatic conditions subject to extremes, such as the Mediterranean. Although early germination may be beneficial for survival in summer, when drought is severe, this may expose the seedlings to winter extremes, thus to higher mortality. Understanding how sensitive is the establishment of different species to temporal patterns of germination will help to understand the factors that control species distribution and community stability in disturbance‐prone environments, as well as its sensitivity to changes in weather patterns as climate changes. Methods An experimental fire was made in early fall in an old Cistus–Erica shrubland in Toledo (central Spain). After fire, germination, survival and growth of the three dominant seeder species (Cistus ladanifer, Erica umbellata and Rosmarinus officinalis) were monitored during the first 3 years after fire. Seedlings were tagged to identify their time of emergence, and divided into cohorts according to their month of germination. Differences in survival of the various cohorts were evaluated by means of a Wilcoxon (Gehan) statistic. Height of surviving, tagged plants was compared among cohorts by means of a Kolmogorov–Smirnov test. Results The year following fire was one of the driest on record, while the next one was one of the wettest. Germination was more abundant during the first than during the second year. Establishment was mainly from first‐year germination, as the majority of second‐year germinated seedlings died. Temporal patterns of germination within a year and between years varied between species. Seedling mortality was highest immediately following germination, not in summer. Mortality was related to time of germination: during a given period of time, the mortality of younger seedlings was higher than that of older ones. However, survival was not highest for the first cohorts. In general, the earlier the seedlings germinated the more vigourous they became, more clearly so for Cistus than for Rosmarinus, but differences tended to disappear with time. Overall, time of germination varied between species and affected differently seedling survival and vigour of the various species. Rosmarinus and Cistus had sufficient survivors to reestablish the initial population. Erica, despite abundant germination, suffered a strong population reduction. Main conclusions Mediterranean shrub species differ in their temporal patterns of germination and survival after fire. The effect of time of germination is complex: germinating early is advantageous since old seedlings fared better than younger ones when confronted with the same rigours. However, germinating early might expose the seedlings to greater hazards and the first cohort might not survive best. The temporal window for establishment is narrow and mainly restricted to the first year after fire. Second year seedlings, irrespective of most favourable conditions, survived very little. Missing the window of establishment might virtually lead to a population collapse, despite having very high germination, as found for Erica.     

Delayed post-fire seedling emergence linked to season: a case study with Leucopogon species (Epacridaceae)

In many fire-prone habitats around the world, natural fire regimes have shaped the evolution of the associated flora. Critical life history stages are often linked to fire in species that occur in these fire-prone habitats but many species are unstudied. We investigated seedling emergence patterns over time, after several fires in south-eastern Australia, for three obligate seeders in the Genus Leucopogon (Family Epacridaceae): L. setiger, L. esquamatus and L. exolasius. Fixed quadrats were monitored for 12 to 30 months for newly emerged seedlings, both after fire and in unburnt L. exolasius and L. esquamatus habitats. There was a flush of seedling emergence in the first year after fire for all three Leucopogon species, with smaller pulses recorded in subsequent years. Time elapsed between fire and the onset of emergence differed between fires, but not between Leucopogon species. Whatever the timing of the fire, seedling emergence was restricted to the late autumn and winter periods, coinciding with emergence in unburnt habitat. This contrasts with patterns of emergence previously reported for other taxa, and also in this study, where emergence of the Leucopogon species after fire was delayed compared with co-occurring species in other genera. Our results suggest that seasonal factors are important to the germination ecology of these species and that combinations of fire-related and seasonal factors are necessary to maximise germination. Rainfall has a non-seasonal pattern in the study region and seasonal emergence post-fire has not previously been recorded. The magnitude of delay to emergence of species with seasonal emergence patterns will be determined by the season of fire but not by a seasonal pattern of rainfall in the post-fire year. A shift of the peak fire season could increase this delay, possibly affecting plant population recovery. This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatial concordance at a regional scale in the regeneration process of a circum-Mediterranean relict (Buxus balearica): connecting seed dispersal to seedling establishment

Both microhabitat and temporal scales have significant effects on the regeneration process of plant species. However, the consistency of those patterns at large scales is largely unknown, despite of being essential from both an ecological and a conservation viewpoint. In this study, we examined the spatio‐temporal variation in the process of regeneration through seeds of the relict Mediterranean shrub Buxus balearica. By means of observations and experiments, we quantified the losses at different recruitment phases (seed rain, seed predation, seedling emergence and establishment) both at a small spatial scale (distance and microhabitat effects) and at a temporal scale (four years), and compared them at a regional scale (10 different localities). We additionally studied transition probabilities between phases in two populations and measured the size structure of 18 localities encompassing all the distributional range of the species. The results show that seed shadow follows a leptokurtic pattern which is consistently maintained until the seedling phase (i.e. there is spatial concordance). Despite post‐dispersal factors are spatially uncoupled, they do not remove the gradient established in the seed shadow. Although the importance of different seed predators varied between regions, post‐dispersal seed predation (ca 80%), together with seedling emergence (usually lower than 10%, some years even nil), and seedling mortality during the first summer (ca 70%) were – consistently in all populations – the most critical bottlenecks in the regeneration of the species. Moreover, an absence of seed bank prevents the buffering of dramatic losses during such phases. Watering of sown seeds increased seedling emergence and survival, and number of leaves per seedling, and survival was consistently higher under vegetation coverage. These results suggest that hydric stress notably contributes to limit the regeneration of this once widespread species evolved under a subtropical climate, which nowadays appears in old‐structured populations throughout its distributional area.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Successional trends in standing dead biomass in Mediterranean basin species

Question: Landscape models of fire occurrence in ecosystems assume that the time since the last fire determines vegetation flammability by enabling the accumulation of dead biomass. In this study we ask if Mediterranean basin shrublands respond to these models or, on the contrary, if initial successional stages in these ecosystems could be more flammable than later stages. Location: Mediterranean shrubland in the Valencia region, eastern Spain. Methods: Using different stages of vegetation development (5, 9, 14 and 26 years since the last fire), we first study the structural comiosition of the above‐ground biomass in 375 individuals of nine woody species. Then, we measure how the standing dead biomass varies during succession, taking into account the surface cover of each species and the quantity of total dead biomass accumulated in different successional stages (3, 9, 14 and 26 years since the last fire). Results: The largest amount of standing dead biomass at the plant community level is observed in the middle stages of the succession. Early successional species, such as Cistus spp., Ulex parviflorus and Pinus halepensis, have a higher percentage of standing dead biomass at earlier stages in the succession than species typical of later successional stages, e.g. Juniperus oxycedrus, Quercus coccifera and Quercus ilex. Conclusions: The results suggest that monotonic increase in fire hazard with increasing stand age is not necessarily the rule in Mediterranean basin shrublands, since early successional species may accumulate large amounts of standing dead biomass and thus promote fire at early successional stages.     

Vegetation recovery after fuel management in Mediterranean shrublands

Question: What is the effect of fuel management practices in the recovery capacity of seeder‐dominated shrublands? Location: Ulex parviflorus shrubland localities in Mediterranean regions of eastern Spain. Methods: We applied prescribed burning and brush‐chipping as fuel management techniques in three young and three mature shrublands, and evaluated the effects in the following four years. Results: Canopy opening by the treatments allowed increasing species richness through the four years of secondary succession. The treatments produced a change in community structure and dominant species, from the woody seeder Ulex parviflorus to the resprouter grass Brachypodium retusum. Vegetation response was conditioned by both shrubland developmental stage and treatment applied. Burning resulted in more severe modification of the ecosystem, increasing bare soil cover. Four years after fuel management in different aged Mediterranean gorse shrublands, vegetation response followed a similar pattern with the exception of the young, brush‐chipped shrublands. The treatments applied for controlling Ulex parviflorus were seen to be very effective, with the exception of brush‐chipping in young shrublands. Conclusions: Selective brush‐chipping in middle‐aged or mature gorse shrublands would combine a drastic reduction in fire hazard with ecosystem conservation and regeneration.     

Spatial segregation of pines and oaks under different fire regimes in the Sierra Madre Occidental

Surface fire can modify spatial patterns and self-thinning in pine-oak ecosystems. Spatial pattern analyses were used to compare pattern development and interspecific spatial interactions in trees and seedlings in five Madrean pine-oak stands with different recent fire histories. Interspecific and intraspecific patterns were compared in small (< 15 cm dbh) and large (< 15 cm dbh) diameter classes of the pines (Pinus durangensis, P. teocote, and P. leiophylla) and oaks (Quercus sideroxylla, Q. crassifolia, and Q. laeta) that collectively dominated the five stands. Numbers of juvenile trees in 2.5 × 2.5 m subplots were correlated with cumulative distances to adult trees. Small pine and oak trees were intraspecifically clustered at all scales, irrespective of fire regime. Large pines were strongly clustered only in stands with longer fire-free intervals, and patterns of large versus small pine trees were regular or random in frequent fire stands. These patterns were consistent with fire-induced mortality of maturing trees under frequent fire. Large and small pines were segregated from small oaks at short and long distances in one stand with a 32-year fire-free interval, implying that two or more dynamic factors had produced regular patterns at different scales. Such regular spatial patterns at short distances were not seen in other stands. Therefore, there was little evidence for direct competition between oaks and pines. The results reported here are consistent with studies from other pine-oak ecosystems showing that different fire regime and site factors interact to influence stand development processes and relative dominance of pines and oaks. In some stands, the continued absence of fire could foster increasing tree densities and an intensification of local neighborhood effects, producing segregation of pine and oak species at longer distances. This revised version was published online in August 2006 with corrections to the Cover Date.     

On ecology of post-fire soil fungi: assessing impact of disturbance using species-abundance models as measure of community organization

Mediterranean ecosystems are among those most significantly modified by fires. Such fires lead to evident disturbance of the above- and below-ground ecosystem components, at various spatial and temporal scales, including soil microfungi. The ecological parameters used to measure the effects of disturbance on soil fungal communities include species-abundance distribution patterns, which reflect changes in the relationships between species numbers and their relative abundance, and serve as a critical measure of community organization. Species-abundance distribution patterns were used to assess the disturbance impact of experimental fires on soil fungal communities in Mediterranean maquis (southern Italy) in the short- to mid-term. The trend in the distribution patterns of heat-stimulated and xerotolerant soil fungi over time, their varying responses to low- and high-intensity fire, the efficiency of the soil germplasm bank, and the pivotal role of Neosartorya spp. in post-fire community structure in Mediterranean burned soils may all be used as tools to accurately assess the effects of fire on soil mycobiota.     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Where do seedlings go? A spatio‐temporal analysis of seedling mortality in a semi‐arid gypsophyte

Studies of seedling population dynamics often focus on survival because it provides an integrated measure of seedling performance. However, this approach involves a substantial loss of information because survival is the net result of a wide range of mechanisms. The present study overcomes these shortcomings by investigating spatial and temporal patterns in the causes of plant mortality in a population of Helianthemum squamatum seedlings. We use new point pattern analyses based on K functions combined with a new null model (“independent labeling”). A total of 871 seedlings of H.squamatum were mapped and regularly monitored over an 18‐month period. More than 60% of seedlings died during this period. Causes of mortality were spatially structured, and these structures shifted through time. Small differences in either the time of emergence or the environment surrounding H. squamatum seedlings had profound influences on their fate. Seedlings emerging late in the season under the canopy of adult plants died from drought more often than expected, whereas those emerging earlier in the same microsite survived more than expected. The identity of neighbors also affected the spatio‐temporal dynamics of mortality causes. Our results show that seedling‐adult interactions cannot be easily predicted from simple models, and that the time of seedling emergence, its age and the identity of its neighbors determine the sign and the spatial scale of these interactions. The new methods introduced in this article open an avenue for the detailed analyses of the spatio‐temporal dynamics of plant mortality and can help to disentangle the complexity of biotic interactions along environmental severity gradients.     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

The role of seed traits as segregation factors of hybrids in wild populations of Cistus (Cistaceae)

Hybridization has been postulated as a main speciation mechanism in Cistus, a hard-seeded group of plants widely distributed in fire-prone Mediterranean shrublands. However, ecological evidence about how the segregation of hybrids from parents might occur is nonexistent. The presence of two hybrid populations in Morocco and Spain from Cistus albidus and Cistus heterophyllus led us to investigate their seed trait differences (seed size, mass and coat thickness) with the parentals and their seed germinability. A sowing experiment to compare the seedling emergence ability and seedling fitness of hybrids and their parentals was also carried out. Hybrid seeds germinated in a similar proportion to those of the parents. The hybrid seeds were significantly lighter, smaller and had thinner seed coats in both populations. However, these seed trait differences did not affect the seedling fitness of hybrids. These results were interpreted as a successful shift in the regeneration niche of the hybrids due to a greater proportion of soft-coated seeds, able to emerge from the soil seed bank in absence of fire. Smaller seeds also would favour further and deeper seed dispersal to escape from a fire. Monitoring of the seed yield of the Spanish population showed a competitive ability of hybrids.     

Spatial patterns of soil pathogens in declining Mediterranean forests: implications for tree species regeneration

Soil-borne pathogens are a key component of the belowground community because of the significance of their ecological and socio-economic impacts. However, very little is known about the complexity of their distribution patterns in natural systems. Here, we explored the patterns, causes and ecological consequences of spatial variability in pathogen abundance in Mediterranean forests affected by oak decline. We used spatially explicit neighborhood models to predict the abundance of soil-borne pathogen species (Phytophthora cinnamomi, Pythium spiculum and Pythium spp.) as a function of local abiotic conditions (soil texture) and the characteristics of the tree and shrub neighborhoods (species composition, size and health status). The implications of pathogen abundance for tree seedling performance were explored by conducting a sowing experiment in the same locations in which pathogen abundance was quantified. Pathogen abundance in the forest soil was not randomly distributed, but exhibited spatially predictable patterns influenced by both abiotic and, particularly, biotic factors (tree and shrub species). Pathogen abundance reduced seedling emergence and survival, but not in all sites or tree species. Our findings suggest that heterogeneous spatial patterns of pathogen abundance at fine spatial scale can be important for the dynamics and restoration of declining Mediterranean forests.     

Germination behaviour of 14 Mediterranean species in relation to fire factors: smoke and heat

Fire is an ecological factor that has been present in the ecosystems of the Mediterranean region for thousands of years. Our study was undertaken to acquire knowledge of the effect of fire on the germination of Mediterranean species. We used high temperatures (up to 60°C) and smoke to determine the effect of these factors on the germination of species from the Mediterranean region. The species selected are characteristic of the central Mediterranean basin and are representative of both woody and herbaceous species: Rhamnus alaternus L., Cistus albidus L., Cistus monspeliensis L., Fumana ericoides (Cav.) Gand., Rosmarinus officinalis L., Melica ciliata L., Avena sterilis L., Bituminaria bituminosa (L.) C.H. Stirt., Anthyllis vulneraria L., Coronilla glauca L., Argyrolobium zanonii (Turra) P.W. Balland, Emerus major Mill., Genista scorpius (L.) D.C. and Spartium junceum L. The seeds were collected in Mediterranean shrubland (8) and woodland (6) ecosystems, around Montpellier, France (24°45′N and 3°50′E). Ten treatments were tested: a control, three smoke treatments and six heat treatments. The average germination level (germination percentage) and the average T₅₀ rates (time taken to reach 50% of germination) were calculated. The smoke and heat act in a different way on each of the species. The smoke enhanced the germination of two species, whilst moderate heat increased germination in all of the species excepting R. officinalis, F. ericoides, A. sterilis, A. vulneraria, and G. scorpius. Germination was fastest in M. ciliata and S. junceum and slowest in A. sterilis, E. major and C. albidus. The cues did not significantly affect the rate of germination. Fire modified the germination response of 12 of the 14 species studied.     

Disturbance History of a Seasonal Tropical Forest in Western Thailand: A Spatial Dendroecological Analysis

Disturbances play an important role in forest dynamics across the globe. Researchers have mainly focused on the temporal context of disturbances, but have largely ignored the spatial patterns of tree recruitment they create. Geostatistical tools enable the analysis of spatial patterns and variability in tropical forest disturbance histories. Here, we examine the potential of combining dendroecological analysis and spatial statistics to reconstruct the disturbance history of a seasonal dry evergreen tropical forest plot at the Huai Kha Khaeng Wildlife Sanctuary (HKK), western Thailand. We used tree‐ring‐derived age estimates for 70 individuals of the shade‐intolerant pioneer species Melia azederach (Meliaceae) and tree locations across a 316‐ha study plot to identify the timing and spatial extent of past disturbances. Although the age distribution for Melia suggested that regeneration had been continuous over the past 60 yr, spatial analyses (mark correlation function and kriging) demonstrated the presence of three spatially discrete age cohorts. Two of these cohorts suggested a severe disturbance ~20 yr before present. A third cohort appears to have established ~50 years ago. Using historical records, we conclude that fire disturbance is the most likely disturbance factor affecting HKK. Nevertheless, we do not rule out other disturbance factors. The combined application of tree‐ring analysis and spatial statistics as applied in this study could be readily applied to reconstruct disturbance histories in other tropical regions where tree species with annual growth rings are present.     

地中海沿岸沙丘微生境对幼苗出现时空格局的影响

实验样地设在地中海沿岸沙丘,选择了3个不同的微生境代表：(1)稳定沙丘上的开阔地片段,(2)稳定沙丘上的灌丛下区域,(3)不稳定沙丘的路径区域;从2001年11月至2002年4月的整个生长季节,每一个微生境出现的幼苗在4个日期被监测,并在每一个取样日把每一株幼苗鉴定、计数后,用剪刀把地上部分移去;研究调查了3种微生境幼苗出现的时空分布格局,并分析了雨量与幼苗出现数量的关系。结果发现：在地中海沿岸沙丘生态系统,幼苗出现在时间上具有明显的分布特征,大多数幼苗出现在第一观测期,整个生长季幼苗都不断出现,但幼苗出现的数目却逐渐下降;各功能群的幼苗占幼苗总数的比例分别为：1年生阔叶草47．4％,多年生阔叶草2．5％,豆科植物17,0％,菊科植物14．5％,1年生禾草为11．7％,多年生禾草为1．9％,十字花科植物3．7％,伞形科植物1,4％。在空间上,总幼苗密度、物种丰富度和物种多样性等显示出重要的微生境差异,开阔地区域具有最大的幼苗密度、物种丰富度和物种多样性;3个微生境的幼苗出现不是同步的,微生境影响种子萌发的时间分布格局,灌丛下种子萌发具有滞后现象;大多数功能群的幼苗密度分布基本上具有显著的微生境差异,主要物种的幼苗分布也具有显著的微生境差异。雨量和萌发的幼苗数量间未发现显著的关系。     

Fire Risk and Vegetation Structural Dynamics in Mediterranean Shrubland

Phytomass structural characteristics are highly related to vegetation flammability. In fire-prone species like Mediterranean gorse, which accumulate standing dead fuel, susceptibility to fire is a function of fuel load, vegetation composition and fuel cover, and these characteristics change with time. Thus, for effective fuel control management, knowledge of the vegetation structural dynamics related to fire risk is crucial for preventing future fires. This study analyses structural dynamics in the above-ground phytomass of Ulex parviflorus shrublands in relation to different stages of flammability, i.e., the amount of time elapsed since the last fire. For this, 152 plants were cut from shrublands at different stages of development (young, mature and senescent), and various dimensional measurements were taken on each. The phytomass was separated into living or dead fuel fractions as well as into twigs or branches depending on the stem diameter. Basal diameter is the variable that best predicted Ulex parviflorus total phytomass as well as that of the different fractions. Both dimensional and phytomass variables increased with plant development. In the young shrublands Ulex parviflorus constitutes 54% of total phytomass, and Ulex parviflorus's dead twigs fraction accounts for 5% of total phytomass. In the mature and senescent shrublands, this species represents 80% of total shrubland phytomass, and dead twigs reach values greater than 40%. Our results show that structural changes in the fuel over short periods of time (young and mature) reveal critical periods in shrub development. Identification of these stages is a necessary tool for planning fuel control programmes.