Savanna trees in Namibia—Factors controlling their distribution at the arid end of the spectrum

This study reviewed the distribution of ten common savanna trees in Namibia. Tree distributions were investigated in relation to bioclimatic, topographic and edaphic variables at a national scale. The factors of importance at these broad geographic scales appeared to be rainfall, substrate and, likely, the incidence of frost. Baikiaea plurijuga, Burkea africana, Guibourtia coleosperma and Pterocarpus angolensis seem to reach their bioclimatic limits in Namibia.At the local level, plant traits become important and contribute to explaining distribution patterns. High water and/or nitrogen use efficiency (Acacia erioloba, Colophospermum mopane), dual water obtaining strategies (Faidherbia albida), fire tolerance (e.g. Acacia species, Burkea africana and Pterocarpus angolensis) and drought tolerance (Boscia albitrunca) are some key attributes providing additional explanations for current distributions.Amongst the selected trees and at broad geographic scales, below-ground adaptations are governed by rainfall regime in combination with coarse-textured soils, whereby shallow-rooted trees prevail in the Kalahari sandveld. Deep-rooted species are found largely on non-sandy soils. Physiological performance of many trees appears to be directly linked to rainfall regime and trees may hence show varying performance throughout their distribution range. Insight into plant functional attributes of trees in Namibia is required to develop appropriate management strategies in the light of climate change. Modelling climate change impacts should consider the relative contribution of bioclimatic versus local environmental factors that explain the current distribution patterns of the selected trees.     

Rice yield in relation to electroultrafiltration extractable soil potassium

Summary Potassium in wetland rice soils from five different locations in the Philippines was analyzed using the electroultrafiltration (EUF) technique and by extraction withN NH₄ acetate (pH 7). The soils contained low exchangeable K and responded to K application. The K soil test values were calibrated against the rice response to K application under field conditions. EUF extractable soil K correlated highly significantly with the rice yield response to K fertilizer, whereas the NH₄ acetate extractable K (exchangeable K) did not. Under limiting K supply in soils, rice yield depends more on the EUF-K than on the exchangeable K. Maximum grain yields were obtained when the EUF-K values after harvest and before wetland preparation were above 30 ppm K.     

Ecological aspects of soil seed-banks in relation to bauxite mining. II. Twelve year old rehabilitated mines

Abstract Germinable seed stores were estimated for 12–13 year old rehabilitation sites in the jarrah forest over two seasons (autumn and spring). Collected soils were subjected to combinations of smoking and heating treatments before the germination procedure was commenced. The mean topsoil seed reserve to a depth of 10 cm was 1938 seeds m^?2. Of this total reserve, more than 80% was found in the upper 5 cm of soil. The topsoil seed reserve of these rehabilitation areas was almost seven times that of the adjacent native jarrah forest (292 seeds m^?2; Ward et al. 1997). There was a seasonal effect with autumn-collected soils having significantly more germinable seeds (2723 seeds m^?2) than soils collected in spring (1153 seeds m^?2). More than half (53%) of the topsoil seed reserve in rehabilitated areas was composed of annual weed species, dominated by Aira caryophyllea, Centaurium erythraea and the native Levenhookia pusilla. Of the total of 70 species identified, 13 species showed significantly higher germination in smoked trays while 11 species exhibited significantly higher germination in heated trays. Species responding to the smoking treatment tended to be annuals while those responding to the heating treatment were typically legume shrub species. Examination of the topsoil seed reserve and the vegetation present in these areas showed that while the species composition was similar between the seed store and the vegetation, there was a large difference in densities, with species occurring at much higher densities in the topsoil than in the vegetation. However, the rankings of species were significantly correlated between the topsoil seed reserve and the vegetation present at the site. The implications of these results to prescribed burning of these rehabilitated areas is discussed.     

Does soil determine the boundaries of monodominant rain forest with adjacent mixed rain forest and maquis on ultramafic soils in New Caledonia?

Aim To determine the soil characteristics of Nothofagus‐dominated rain forests in an ultramafic region (i.e. soils having high concentrations of metals including Mg, Fe and Ni), and whether soil characteristics may explain the location of monodominant rain forest in relation to adjacent mixed rain forest and maquis (shrub‐dominated vegetation). Location New Caledonia. Methods Soil characteristics were compared among six Nothofagus‐dominated rain forests from a range of altitudes and topographic positions. At four of these sites, comparisons were made with soils of adjacent mixed rain forest and maquis. Results Soil characteristics varied among the monodominant Nothofagus forests, largely due to differences between ultramafic soils and soils influenced by non‐ultramafic intrusions. The soils of all vegetation types had low concentrations of nutrients, particularly P, K and Ca (both total and extractable/exchangeable), and high total concentrations of Ni, Fe, Cr and Mn. There were significant differences between the rain forests and adjacent maquis in soil concentrations of several elements (N, P, Ca, Mg and Mn), more so in surface soils than at depth, but much of this pattern may be caused by effects of vegetation on the soil, rather than of soil on the vegetation. However, there were no significant differences in soil concentrations of any mineral elements between Nothofagus forest and adjacent mixed rain forest. Main conclusions We found no evidence for soil mediation of boundaries of Nothofagus rain forest with mixed rain forest, and little evidence for the boundaries of either forest type with maquis. We suggest that the local abrupt boundaries of these monodominant Nothofagus forests are directly related to temporal factors, such as time since the last wildfire and frequency of wildfire, and that disturbance is therefore a major causal factor in the occurrence of these forests.     

The effect of lime on phosphorus adsorption and barley growth in three acid soils

For three acid soils from Santa Catarina, Brazil, lime application and time of incubation with lime had little effect on the adsorption of added phosphorus. In two soils with high contents of exchangeable aluminium, solution P and isotopically exchangeable P were decreased by incubating with lime for 1 month: phosphorus was probably adsorbing on freshly precipitated aluminium hydrous oxides. In one soil with less exchangeable aluminium, P in solution was increased by liming. After 23 months lime increased solution and exchangeable P possibly due to crystallization of aluminium hydrous oxides reducing the number of sites for P adsorption. All these changes were however small. In a pot experiment, lime and phosphorus markedly increased barley shoot and root dry matter and P uptake. Although liming reduced P availability measured by solution P, isotopically exchangeable P and resin extractable P, it increased phosphorus uptake by reducing aluminium toxicity and promoting better root growth. The soil aluminium saturation was reduced by liming, but the concentration of aluminium in roots changed only slightly. The roots accumulated aluminium without apparently being damaged.     

Effect of landscape position on the acidification of loess‐derived soils under Pinus radiata

Abstract Interactions between landscape position and the acidifying effect of trees planted into loess‐derived grassland soils were studied in the Ventana region of Argentina. Forests of Pinus radiata planted at the end of 1940, were selected in two different positions from the landscape, plains and slopes. Samples of the soil surface mineral horizon were taken from landscape positions at four distances from the trees and compared with grassland soils. The values of the main soil chemical properties changed significantly with distance from trees, with a decrease in pH, base saturation, exchangeable Ca²⁺, Mg²⁺ and K⁺, and increase in Na⁺, Al³⁺ and particularly H⁺ closer to the trees. This pattern confirms the prominent role of vegetation in bringing about changes in soil properties. Regression models showed high levels of explanation (r² > 0.85) indicating that a high percentage of the spatial variability of soil chemical properties is systematic and predictable with distance from the trees. The pH in KCl proved an excellent tool for predicting the cationic composition of soils. Organic carbon and total nitrogen were significantly higher in the plains positions than in the slopes under the trees, whereas there was no difference under grassland. The slopes of the regression lines indicated that acidification is more intense in soils on the plains. Vegetation was the main factor influencing acidification of the studied soils. The landscape position regulates the bio‐hydrological factor and thus the speed of acidification process.     

Vegetation effects on phosphorus fractions in set-aside soils

As increasing amounts of arable land are being set aside, it is of importance to study the effect of vegetation on soil fertility. The fractionation of soil P under grassland, beech and spruce vegetation was investigated in sites previously fertilized with P by extracting sequentially with Resin, NaHCO₃, NaOH, HCl and finally NaOH after ultrasonic pretreatment. Under beech a large part of extractable P was found in inorganic fractions which are considered to be available for plants (Resin P₁ and Bicarbonate P₁). Under grass, a large part of the extractable P was found in potentially labile organic forms (Bicarbonate P_o and Fulvic acid P_o). After 25 years of permanent grass vegetation, the extractability of soil P was comparable to that from an adjacent arable plot. On spruce covered soils most of the added fertilizer P was rendered unextractable 20–30 years after application. However the available data does not allow a clear interpretation of this phenomena, as effects of soil parent material as well as vegetation may be taken into consideration. No decrease in P-extractability was found between beech and grass covered soils which had been fertilized for more than 200 years, when compared to less rich soils from the same area. On the basis of the current data it may be concluded that the vegetation affects the distribution of soil phosphorus fractions, and thus soil fertility. In the soils under investigation, grassland and beech vegetation conserved the phosphate availability to a high extent.     

Radiocaesium in an organic soil and the effect of treatment with the fungicide ‘Captan’

Soil fungi accumulate radiocaesium from contaminated soil and it has been hypothesised that this may alter the plant availability and movement of the radionuclide in soil. The effect of twice-monthly addition of an aqueous suspension of the fungicide ‘Captan’ on the changes in a peaty podzol soil at 2 sites, contaminated 2 or 3 years earlier by the injection of ¹³⁴Cs, has been quantified. The sites had different soil acidity and vegetation cover. The less acid soil (pH_water 5.0) had been improved by the addition of lime and fertilizer and was reseeded with grass and clover. The more acid soil (pH_water 3.8) was under hill grasses, herbs and heather. On both sites the addition of fungicide did not alter the amount or concentration of radiocaesium in plant material sampled monthly or the depth distribution of radiocaesium in the soil profile. The concentration of the fungal constituent, ergosterol, in the soil, measured monthly, was unaffected by the fungicide treatment but evidence was obtained from a pot experiment to show that ergosterol decomposes slowly in cold, wet soils. On the more acid soil, two weeks after the last application of fungicide, there was a decline in active fungi as measured by fluorescein diacetate staining. Chloroform fumigation of the more acid soil resulted in a small increase in the amount of ¹³⁴Cs exchangeable with 1 M ammonium acetate. Radiocaesium in seven different fungi grown in pure culture was found to be almost entirely extractable (> 95%) with 1 M ammonium acetate. Another, Amanita rubescens, showed some retention and 88% was extractable. These findings do not preclude the fungal biomass as an important soil component controlling plant availability of radiocaesium from acid, organic soils by maintaining radiocaesium in a biological cycle, but make it unlikely that any fixation by fungi in a chemical sense is involved.     

Phenology of nine common woody species in semi-arid,deciduous Kalahari Sand vegetation

The phenological patterns of 9 woody species that are common components of Kalahari Sand communities were observed over a period of 32 months. The observations were made on plants adjacent to the soil moisture study sites which have been described earlier (Childes & Walker 1987). A pre-rain flush was noted in the 2 shrub/small tree species Ochna pulchra and Terminalia sericea. Leaf fall in all species was related to a decrease in soil moisture and minimum temperature. The rooting habits and other adaptations of the species resulted in a sequential leaf fall in the communities. Open woodland and scrub species such as Pterocarpus angolensis showed a generally synchronised pre-rain flowering, while the closed woodland species e.g. Baikiaea plurijuga had protracted asynchronous flowering periods. Fruit ripening generally occurred in the dry season when conditions were ideal for fruit and seed dispersal.     

The Importance of Nickel Phytoavailable Chemical Species Characterization in Soil for Phytoremediation Applicability

In this work Ni speciation in natural and spiked soils (with similar total concentration) was studied. Spiked soils were prepared by addition of NiSO₄.6H₂O to obtain concentration similar to the one in natural soils. In soils mobile species were determined with a simplified sequential extraction as follows: H₂O for water-soluble metal, KNO₃ for exchangeable species, DTPA for complexed/adsorbed species. Results show that in spiked soils the exchangeable and adsorbed Ni concentrations are considerably higher than in natural soils. A study of plant uptake was carried out in order to evaluate the relation between mobile species and phyto-availability. Alfalfa (Medicago sativa L.), even though it is not a hyperaccumulator, was selected for its tolerance to high metal concentrations in soil. Preliminary results show a very high correlation between Ni mobile species and Ni uptake by alfalfa. Significant differences were found between spiked and natural soils. In the latter, high levels of total Ni did not correspond to relevant uptake as in the case of spiked soil. Results stress the importance of evaluating preliminarly heavy metal speciation in soil before planning phytoremediation procedures.     

Role of Soils in Determining Sites for Lowland Heathland Reconstruction in England

No guidelines are currently available that advise which soil properties of arable land can be used to suggest suitable locations for heathland reconstruction. This paper reviews studies comparing soil properties of heathland or semi-natural grassland with those of adjacent arable fields, investigations in the autecology of the dominant heathland plant, Calluna vulgaris (common heather), and long-term experiments of fertilizer inputs on arable soils. Three properties must be assessed before the suitability of a field can be determined: extractable phosphorus, exchangeable calcium, and pH. A number of other nutrients may also be important, but evidence is currently insufficient to substantiate this. Natural changes in levels of extractable phosphorus, exchangeable calcium, and pH appear to be very slow, so nutrient stripping and acidification will be necessary where recommended levels are exceeded to successfully restore heathland vegetation.     

Floristic characterisation of alkali soils in northwestern India

Summary Floristic composition studied in a 40 hectares representative site of alkali soils in the Indo-gangetic plains of Northwestern India revealed a remarkably restricted spectrum of natural vegetation. Soil analysis of specific niches, occupied by aggregations of different species, was employed to identify alkali halophytes. Response functions of eight prominent species to increasing levels of soil alkalinity, studied in pot culture, showed that most of them were adapted to alkali soils. Based on the computed importance value indices and observed biomass production maxima, it has been inferred thatSporobolus marginatus, Sporobulus coromandelianus, Diplachne fusca andChloris barbata, qualify as plant indicators of high-alkali soil conditions.     

Edaphic Distribution of Some Species of the Fern Genus Adiantum in Western Amazonia1

The distribution patterns of Adiantum species were inventoried at twelve noninundated rain forest (tierra firme) sites in Amazonian Peru and Ecuador. Six species belonging to the genus were found, and four of these were abundant at some of the sites. At six of the sites none of the species was particularly abundant. Both the occurrence and the abundance of the species varied among sites with regard to soil texture and concentration of exchangeable bases and within sites with regard to topography. Adiantum tomentosum) Klotzsch was restricted to loamy soils with intermediate to low content of extractable bases and was locally most abundant in the well drained parts of slopes and ridge tops. Adiantum terminatum Kunze ex Miq. and A. humile Kunze occurred at sites with intermediate to high content of extractable bases, but their relative abundances differed among and within sites. Adiantum pulverulentum L. was restricted to soils with relatively high base concentration. Ecological differences among the species obviously explained many of the differences in their distribution patterns at the local and regional scales, and may be important for the understanding of such patterns at biogeographical scales as well.     

Climatic signals in tree rings of <Emphasis Type="Italic">Burkea africana</Emphasis> and <Emphasis Type="Italic">Pterocarpus angolensis</Emphasis> from semiarid forests in Namibia

Tree-ring studies contribute worldwide to the understanding of climate and its relation to tree growth. Long tree-ring chronologies serve as climate proxies for the reconstruction of past, pre-instrument climate and its recent change. In tropical regions, the availability of exactly dated tree-ring chronologies is limited. The dendroclimatic potential of two dominant species from dry forests in northern Namibia was examined in the study presented in this paper. Both species (Burkea africana Hook and Pterocarpus angolensis DC) were sampled at two sites (ca. 900 km apart), and the response to several climatic variables, including ENSO indices, is studied. All specimens showed distinct growth rings and cross-dating between radii was successful for all trees. Species-specific mean curves were built for both sites. The mean curves of different species of the same site synchronised significantly, allowing the construction of a site-specific chronology. Synchronisation between sites was not possible, but spectral analysis of the chronologies implied that both show similar long-term (6.7 year) oscillation patterns. B. africana is more sensitive to rainfall variation than P. angolensis at both sites. Growth response to rainfall was positive, but a time-lag in the reaction occurred between the sites, corresponding to the time-lag of the beginning of the rainy season. Air temperature showed a negative correlation with stem increment at both sites. The response at the westernmost site to two ENSO indices indicates a tree growth decrease during El Niño years, which are generally dry in southern Africa.     

Fruit Production and Seed Predation in Two Miombo Woodland Trees in Zambia

During a six year period (1990–1995) fruiting frequency among marked trees and seed production and predispersal predation by beetle larvae of lsoberlinia angolensis and Julbernardia globiflora of the Leguminosae family and Caesalpinoideae subfamily were studied at two dry miombo woodland sites in central Zambia. Fruit production varied from year to year and was not related to annual rainfall. On average I. angolensis trees fruited once every two years while J. globiflora trees fruited once every two and one-half years. Number of fruits per tree was correlated with tree size for both species. In a peak year fruit biomass represented two to four percent of total above ground biomass but for J. globiflora about 5.5 and 4.5 percent of nitrogen and potassium, respectively, were in fruits compared to 0.6 and 1.8 percent for I. angolensis. The lower fruiting frequency in J. globifora was probably related to the depletion of N reserves during a fruiting year which required a longer replenishment period than in I. angolensis. Predispersal seed predation was low for J. globiflora (10%) and very high in I. angolensis (65–85%) and for the latter species predation rate increased from August to November 1991 and significant variations were observed among years and pod types. Seed predation rate was lowest in a peak fruiting year and this observation provides support for the satiation hypothesis. The results of the study also suggest that I. angolensis and J. globiflora may have different regeneration strategies although reproductive allocation in both species was within the range reported for other iteroparous plants.     

Effects of slope aspect and vegetation types on selected soil properties in a dryland Hirmi watershed and adjacent agro‐ecosystem,northern highlands of Ethiopia

This study aimed to determine the spatial variation of topsoil properties as influenced by slope aspects and vegetation types. Three vegetation types and two slope aspects were considered. Disturbed and undisturbed samples were taken from the top soils. A two‐way analysis of variance was used to test significant mean differences (α = 0.05). Results showed that contents of bulk density, total porosity, organic matter and exchangeable magnesium were significantly different with changes in slope aspect (P < 0.05). Textural classes, organic matter, available phosphorus, carbon‐to‐nitrogen ratio, cation‐exchange capacity, per cent base saturation and exchangeable bases demonstrated significant variation among vegetation types (P < 0.05). The south‐facing aspect and the grass vegetation showed higher bulk density but lower organic matter and total porosity. As a remedy for the identified problems, there is a need to undertake plantation of indigenous trees.     

Diets of two sympatric colobines in Zaire: More evidence on seed-eating in forests on poor soils

Recent studies have shown that the so- called folivorous African colobines eat a significant amount of seeds. There is disagreement as to whether seed-eating is due to the poor quality of tree foliage, due in turn to poor soils, or to the fact that seeds are a normal part of colobine diets. To test these hypotheses, we studied feeding of red colobus monkeys, Colobus badius tholloni,and black-and-white colobus monkeys, Colobus angolensis angolensis,in a tropical rain forest of Central Zaire (Salonga National Park). We conducted studies on soil properties, vegetation composition, and the availability and chemical contents of food plants simultaneously. Soils were very acid, with a high percentage of sand, very low cation exchange capacity, and very low exchangeable bases. The forest was dominated by legumes (45.6% of trees), among which the Caesalpinioideae were the best represented (85%). C. badiusfed mostly on leaves (61%) and seeds (33%), legumes making up 65% of their diet. C. angolensisfed mostly on seeds (50%) and leaves (27%); 39% of their diet came from legumes. The two species tended to select items richer in crude protein or lipids or both. Total phenolics and condensed tannins were abundant in the foliage and seeds but were poor predictors of colobine choice of food. Intersite comparisons show that colobines in Zaire ate a higher proportion of seeds than all other related species so far studied in Africa and that the Salonga forest had among the poorest soils and harbored the highest percentage of Caesalpinioideae. Our results confirm that seed- eating is more common among colobines living in areas where soils are poor. They strongly suggest that this link is mediated through forest composition, especially the abundance of legumes, and that the development of seed- eating results both from the high availability of nutrient-rich seeds and from the poor quality of mature tree foliage.     

Calcicole–calcifuge plant strategies limit restoration potential in a regional semi‐arid flora

AimTo examine calcicole and calcifuge plant strategies, as well as nutrient‐acquisition strategies, as drivers of the distribution of species in response to edaphic factors, and the degree to which these strategies may act as filters to species establishment in ecological restoration on heavily altered or reconstructed substrates.LocationAn 82,000‐ha area within a major mining province in the Mid‐West region of Western Australia, harboring vegetation communities ranging from species‐poor halophytic scrub on saline flats to dense biodiverse shrubland on the skeletal soils of ancient Banded Ironstone Formations (BIF).MethodsUnivariate and multivariate analyses were employed to examine how variation in soil chemistry and landscape position (undulating plains, slopes, and BIF crests and ridges) influenced patterns of floristic diversity, calcifuge plant strategies, and nutrient‐acquisition strategies in 538 plant species from 830 relevés.ResultsLandscape position was the strongest driver of species richness and vegetation functional composition. Soils became increasingly acidic and P‐impoverished along an increasing elevational gradient. Vegetation from different landscape positions was not compositionally dissimilar, but vegetation of BIF crests and ridges was up to twice as biodiverse as vegetation from adjacent lower‐relief areas and harbored higher proportions of calcifuge species and species with mycorrhizal associations.Main conclusionsTopographic and edaphic complexity of BIF landforms in an otherwise relatively homogenous landscape has likely facilitated species accumulation over long time periods. They represent musea of regional floristic biodiversity, excluding only species that cannot establish or are inferior competitors in heavily weathered, acidic, skeletal, and nutrient‐impoverished soils. Plant strategies likely represent a major filter in establishing biodiverse, representative vegetation on postmining landforms in geologically ancient regions.     

The effect of phosphate rock dissolution on soil chemical properties and wheat seedling root elongation

Soils of the Appalachian region of the United States are acidic and deficient in P. North Carolina phosphate rock (PR), a highly substituted fluoroapatite, should be quite reactive in these soils, allowing it to serve both as a source of P and a potential ameliorant of soil acidity. An experiment was conducted to evaluate the influence of PR dissolution on soil chemical properties and wheat (Triticum aestivum cv. Hart) seedling root elongation. Ten treatments including nine rates of PR (0, 12.5, 25, 50, 100, 200, 400, 800, and 1600 mg P kg^-1) and a CaCO₃ (1000 mg kg^-1) control were mixed with two acidic soils, moistened to a level corresponding to 33 kPa moisture tension and incubated for 30 days. Pregerminated wheat seedlings were grown for three days in the PR treated soils and the CaCO₃ control. Root length was significantly (P<0.05) increased both by PR treatments and CaCO₃, indicating that PR dissolution was ameliorating soil acidity. The PR treatments increased soil pH, exchangeable Ca, and soil solution Ca while lowering exchangeable Al and 0.01 M CaCl₂ extractable soil Al. Root growth in PR treatments was best described by an exponential equation (P<0.01) containing 0.01 M CaCl₂ extractable Al. The PR dissolution did not reduce total soil solution Al, but did release Al complexing anions into soil solution, which along with increased pH, shifted Al speciation from toxic to nontoxic forms. These results suggest that North Carolina PR should contribute to amelioration of soil acidity in acidic, low CEC soils of the Appalachian region.     

Recovery Deficiency Following Tree Mortality in Mangroves of Two Caribbean Islands: Field Survey and Statistical Classification

Mangrove species are well adapted to the harsh ecological conditions of their environment throughout the tropics. However, in the islands of Guadeloupe and Martinique (Lesser Antilles), deficient forest recovery was evidenced in 43 mangrove sites (>1000 m²) affected by apparently natural tree mortality. Such sites were recorded from four chronological sets of aerial photographs between 1950 and 1995, and field-investigated in terms of environment and vegetation characteristics. Given the speculative relationship between the slow-regenerating vegetation and its present environment within non-steady state, disturbed sites, statistical analyses arbitrarily matching physicochemical and biological data were primarily avoided. On one hand, principal component analysis (PCA), combined with an agglomerative hierarchical classification, was performed on environmental, rank-ordered data; on the other hand, multidimensional scaling (MDS) was implemented on vegetation data. Discriminant analyses (DA) further characterized the environment/vegetation interrelationships for each site type. Ultimately, three main types of mortality sites have been distinguished among the study areas. One type clustered the sites showing the lowest salinity values and the highest surge vulnerability whose dominant mangrove species is Rhizophora. Another type presented highly saline sites having clayey soils with pure, stunted, Avicennia stands. The last type consisted of peculiar forest gaps on deep, compact, peat soils. During the rainy season, these sites turned into shallow ponds scattered with living, young Rhizophora and large, standing, dead Avicennia. The authors suggest that this classification may serve as a comprehensive framework to test subsequent hypotheses (hurricanes, droughts…) on the origins of natural massive tree mortality and the causes of recovery deficiency in mangroves of the Caribbean.