Rapid recovery of tropical forest diversity and structure after shifting cultivation in the Philippines uplands

Shifting cultivation is a widespread land‐use in the tropics that is considered a major threat to rainforest diversity and structure. In the Philippines, a country with rich biodiversity and high rates of species endemism, shifting cultivation, locally termed as kaingin, is a major land‐use and has been for centuries. Despite the potential impact of shifting cultivation on forests and its importance to many people, it is not clear how biodiversity and forest structure recover after kaingin abandonment in the country, and how well these post‐kaingin secondary forests can complement the old‐growth forests. We investigated parameters of forest diversity and structure along a fallow age gradient in secondary forests regenerating after kaingin abandonment in Leyte Island, the Philippines (elevation range: 445–650 m asl). We first measured the tree diversity and forest structure indices in regenerating secondary forests and old‐growth forest. We then measured the recovery of tree diversity and forest structure parameters in relation to the old‐growth forest. Finally, using linear mixed effect models (LMM), we assessed the effect of different environmental variables on the recovery of forest diversity and structure. We found significantly higher species density in the oldest fallow sites, while Shannon’s index, species evenness, stem number, basal area, and leaf area index were higher in the old‐growth forest. A homogeneous species composition was found across the sites of older fallow age. Multivariate analysis revealed patch size as a strong predictor of tree diversity and forest structure recovery after shifting cultivation. Our study suggests that, secondary forests regenerating after shifting cultivation abandonment can recover rapidly. Although recovery of forest structure was not as rapid as the tree diversity, our older fallow sites contained a similar number of species as the old‐growth forest. Many of these species are also endemic to the Philippines. Novel and emerging ecosystems like tropical secondary forests are of high conservation importance and can act as a refuge for dwindling tropical forest biodiversity.     

Assembly patterns of tree seedling communities in a human‐dominated Tropical landscape

Understanding how different components of species diversity of regenerating areas respond to both time of abandonment and landscape metrics may offer crucial information for both theoretical and practical purposes. Using 15 regeneration areas (from 4 to 30 years) and nine areas of mature forest, we assessed how tree seedling assemblage responds to time of abandonment and forest cover in terms of species diversity and taxonomic composition. We found that species diversity of seedlings responded positively to time of abandonment, but was not influence by forest cover. Diversity of rare and common species (⁰D and ¹D, respectively) but not dominant species (²D) increased with time of abandonment, reaching reference values very quickly (ca. 20 years). However, species composition seemed to respond to both ageing of forest stand and landscape structure represented by forest cover. Our findings reinforce that in human‐dominated landscapes, local processes related to recovery with time should be more important for plant community assembly than landscape structure, leading to a number of possibilities for multiple successional pathways. Abstract in Portuguese is available with online material.     

Recovery of plant species richness and composition after slash-and-burn agriculture in a tropical rainforest in Madagascar

Slash-and-burn agriculture is an important driver of deforestation and ecosystem degradation, with large effects on biodiversity and carbon sequestration. This study was conducted in a forest in Madagascar, which consists of fragments of slash-and-burn patches, within a matrix of secondary and primary forest. By recording species richness, abundance, and composition of trees, shrubs, and herbs in fallows of various age and slash-and-burn history, and in the secondary and primary forest, we show how slash-and-burn intensity (number of cycles, duration of abandonment), years since last abandonment, and environmental factors (distance to primary forest and topography) affect the natural succession and recovery of the forest ecosystem. We used ordination analyses to examine how the species composition varied between the different successions stages, and to examine tree recruitment. Our results show shrub dominance the first years after abandonment. Thereafter, a subsequent increase in species richness and abundance of tree seedlings and saplings suggests a succession towards the diversity and composition of the secondary and primary forest, although a big gap between the oldest fallows and the secondary forest shows that this will take much more than 30 years. A high number and frequency of slash-and-burn cycles decreased tree seedling and sapling richness and abundance, suggesting that reducing slash-and-burn intensity will increase the speed of tree recruitment and fallow recovery. Trees can be planted into fallows to speed up vegetation and soil recovery, such that fallows can be usable within needed time and thus the extension of cultivated areas reduced. We recommend further testing of six potential species for restoration based on their early colonization of the fallows and their survival through vegetation succession.     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

基于分层的典型中亚热带天然阔叶林分局部生物多样性

选择福建万木林保护区内两块典型中亚热带天然阔叶林样地为对象,按分层和不分层两种情形,研究了小尺度(20m)范围内各组成树种的单物种-面积关系(individual species-area relationship,ISAR),以探讨局部生物多样性格局及其维持机制。结果表明,两个群落的乔木层都可以划分出受光层和非受光层两个亚层。以受光层林木为圆心进行ISAR分析,发现在小尺度范围大多数受光层林木为局部生物多样性的中性种,只有个别树种在个别尺度上有显著的促进或排斥作用。研究途径避免了对非随机作用在维持局部生物多样性中的相对重要性的高估。不分层时,乔木层内同样以中性树种为主。研究结果支持小尺度范围内调查的典型中亚热带天然阔叶林的物种多样性由中性作用主导的结论。     

Secondary forest fragments offer important carbon and biodiversity cobenefits

Tropical forests store large amounts of carbon and high biodiversity, but are being degraded at alarming rates. The emerging global Forest and Landscape Restoration (FLR) agenda seeks to limit global climate change by removing carbon dioxide from the atmosphere through the growth of trees. In doing so, it may also protect biodiversity as a free cobenefit, which is vital given the massive shortfall in funding for biodiversity conservation. We investigated whether natural forest regeneration on abandoned pastureland offers such cobenefits, focusing for the first time on the recovery of taxonomic diversity (TD), phylogenetic diversity (PD) and functional diversity (FD) of trees, including the recovery of threatened and endemic species richness, within isolated secondary forest (SF) fragments. We focused on the globally threatened Brazilian Atlantic Forest, where commitments have been made to restore 1 million hectares under FLR. Three decades after land abandonment, regenerating forests had recovered ~20% (72 Mg/ha) of the above‐ground carbon stocks of a primary forest (PF), with cattle pasture containing just 3% of stocks relative to PFs. Over this period, SF recovered ~76% of TD, 84% of PD and 96% of FD found within PFs. In addition, SFs had on average recovered 65% of threatened and ~30% of endemic species richness of primary Atlantic forest. Finally, we find positive relationships between carbon stock and tree diversity recovery. Our results emphasize that SF fragments offer cobenefits under FLR and other carbon‐based payments for ecosystem service schemes (e.g. carbon enhancements under REDD+). They also indicate that even isolated patches of SF could help to mitigate climate change and the biodiversity extinction crisis by recovering species of high conservation concern and improving landscape connectivity.     

The role of remnant trees in carbon sequestration,vegetation structure and tree diversity of early succession regrowing fallows in eastern Sierra Leone

Remnant tree presence affects forest recovery after slash‐and‐burn agriculture. However, little is known about its effect on above‐ground carbon stocks, especially in Africa. We focused our study on Sierra Leone, part of the Upper Guinean forests, an important centre of endemism threatened by encroachment and forest degradation. We studied 99 (20‐m‐radius) plots aged 2–10 years with and without remnant trees and compared their above‐ground carbon stocks, vegetation structure (stem density, basal area) and tree diversity. Above‐ground carbon stocks, stem density, basal area, species richness and tree diversity increased significantly with fallow age. Remnant tree presence affected significantly tree diversity, species dominance and above‐ground carbon stocks, but not vegetation structure (stem density, basal area). Number of remnant trees and number of species of remnant trees were also important explanatory variables. Although other factors should be considered in future studies, such as the size and dispersal modes of remnant trees, our results highlight that more strategic inclusion of remnant trees is likely to favour carbon stock and forest recovery in old fallows. To our knowledge, this is the first study on early succession regrowing fallows in West Africa.     

Persistence of genetic variation in recolonized Tsuga canadensis (Eastern hemlock) populations following historic forest clearance

The recovery of genetic variation in newly recolonized populations is an important concern in forest conservation genetics. We examined the potential recovery of genetic diversity and changes to genetic structure in populations of the wind‐pollinated species Tsuga canadensis that naturally regenerated following the extensive 19th century regional forest clearance for agriculture in west‐central Massachusetts. We genotyped 264 individuals across six microsatellite loci and compared levels and patterns of genetic variation between primary forests (forests that were logged but never cleared) and secondary forests (stands that were recolonized following agricultural abandonment). We found no significant reductions in genetic diversity in secondary forests (A_R = 5.450; H_S = 0.718) compared to primary forests (A_R = 5.742; H_S = 0.730). Moreover, the population genetic differentiation was also not significantly reduced in secondary compared to primary forests, with no significant genetic structure observed among all populations. These results suggest rapid genetic recovery of T. canadensis populations in recolonized forests compared with other late‐successional temperate tree species.     

Species and structural diversity affect growth of oak,but not pine,in uneven-aged mature forests

The effects of mixing tree species on tree growth and stand production have been abundantly studied, mostly looking at tree species diversity effects while controlling for stand density and structure. Regarding the shift towards managing forests as complex adaptive systems, we also need insight into the effects of structural diversity. Strict forest reserves, left for spontaneous development, offer unique opportunities for studying the effects of diversity in tree species and stand structure. We used data from repeated inventories in ten forest reserves in the Netherlands and northern Belgium to study the growth of pine and oak. We investigated whether the diversity of a tree's local neighbourhood (i.e., species and structural diversity) is important in explaining its basal area growth. For the subcanopy oak trees, we found a negative effect of the tree species richness of the local neighbours, which – in the studied forests – was closely related to the share of shade-casting tree species in the neighbourhood. The growth of the taller oak trees was positively affected by the height diversity of the neighbour trees. Pine tree growth showed no relation with neighbourhood diversity. Tree growth decreased with neighbourhood density for both species (although no significant relationship was found for the small pines). We found no overall diversity-growth relationship in the studied uneven-aged mature forests; the relationship depended on tree species identity and the aspect of diversity considered (species vs. structural diversity).     

Soil seed bank composition along a forest chronosequence in seasonally moist tropical forest,Panama

Abstract. We used a forest chronosequence at the Barro Colorado Nature Monument (BCNM) to examine changes in the abundance and species composition of seeds in the soil during forest succession. At each of eight sites varying from 20 yr to 100 yr since abandonment, and at two old-growth (> 500 yr) forest stands, we established two 160-m transects and sampled the surface 0–3 cm of soil in cores collected at each 5 m interval. Seed densities were estimated from the number of seedlings germinated from the soil over a six-week period. Contrary to expectation, neither the density of the soil seed bank, nor species richness or diversity were directly related to age since abandonment, but the density of the soil seed bank was correlated with the abundance of seed-bank-forming species in the standing vegetation. In marked contrast to published studies, herbaceous taxa were rare even in the youngest stands, and the common tree species, which accounted for most seeds in the soil, were present in all stands. The pioneer tree Miconia argentea (Melastomataceae) was the single most common species in the seed bank, accounting for 62% of seeds and present in 92% of soil samples. Rapid recovery of the vegetation of young regrowth stands on BCNM, when compared to sites elsewhere may be partly due to allochthonous seed rain from nearby mature forest stands and the lack of seed inputs of weeds and grasses from agricultural and pasture lands which may inhibit forest succession.     

Comparing forest structure and biodiversity on private and public land: secondary tropical dry forests in Costa Rica

Secondary forests constitute a substantial proportion of tropical forestlands. These forests occur on both public and private lands and different underlying environmental variables and management regimes may affect post‐abandonment successional processes and resultant forest structure and biodiversity. We examined whether differences in ownership led to differences in forest structure, tree diversity, and tree species composition across a gradient of soil fertility and forest age. We collected soil samples and surveyed all trees in 82 public and 66 private 0.1‐ha forest plots arrayed across forest age and soil gradients in Guanacaste, Costa Rica. We found that soil fertility appeared to drive the spatial structure of public vs. private ownership; public conservation lands appeared to be non‐randomly located on areas of lower soil fertility. On private lands, areas of crops/pasture appeared to be non‐randomly located on higher soil fertility areas while forests occupied areas of lower soil fertility. We found that forest structure and tree species diversity did not differ significantly between public and private ownership. However, public and private forests differed in tree species composition: 11 percent were more prevalent in public forest and 7 percent were more prevalent in private forest. Swietenia macrophylla, Cedrela odorata, and Astronium graveolens were more prevalent in public forests likely because public forests provide stronger protection for these highly prized timber species. Guazuma ulmifolia was the most abundant tree in private forests likely because this species is widely consumed and dispersed by cattle. Furthermore, some compositional differences appear to result from soil fertility differences due to non‐random placement of public and private land holdings with respect to soil fertility. Land ownership creates a distinctive species composition signature that is likely the result of differences in soil fertility and management between the ownership types. Both biophysical and social variables should be considered to advance understanding of tropical secondary forest structure and biodiversity.     

Successional Change and Resilience of a Very Dry Tropical Deciduous Forest Following Shifting Agriculture

We analyzed successional patterns in a very dry tropical deciduous forest by using 15 plots differing in age after abandonment and contrasted them to secondary successions elsewhere in the tropics. We used multivariate ordination and nonlinear models to examine changes in composition and structure and to estimate forest recovery rates and resilience. A shrub phase characterized early succession (0–3 yr); afterwards, the tree Mimosa acantholoba became dominant. Below its canopy, sprouts and seed-regenerated individuals of mature forest species slowly accumulated. Canopy height, plant density, and crown cover stabilized in less than 15 yr, whereas species richness, diversity, and basal area continued to increase. The pioneer species group has very low diversity and the long-lived pioneer phase typical of humid forests is absent; species composition may therefore recover soon as suggested by convergence toward mature forest species composition. The time trend of plant density also differed from humid forests for it lacked its characteristic density decline, presumably because of differences in regeneration mechanisms between very dry and other less water-stressed forest types. As opposed to the prevailing hypothesis, resilience was not higher than in moister forests, and thus factors other than structure relative simplicity must be accounted for when assessing resilience.     

Recovery of amphibian,reptile, bird and mammal diversity during secondary forest succession in the tropics

In tropical regions, many studies have focused on how vegetation and ecosystem processes recover following the abandonment of anthropogenic activities, but less attention has been given to the recovery patterns of vertebrates. Here we conduct a meta‐analysis (n = 147 studies) of amphibian, reptile, bird and mammal recovery during tropical secondary forest succession (i.e. natural regeneration). For each taxonomic group, we compared changes in species richness and compositional similarity during natural secondary succession to reference forests (mature or old growth forest). In addition, we evaluated the response of forest specialists and the change in bird and mammal functional groups during natural secondary succession in the tropical moist forest biome. Overall, species richness of all groups reached levels of the reference forests during natural secondary succession, but this was not the case for species compositional similarity. The delay in recovery of forest specialists may be the reason for the delay in recovery of species compositional similarity. Overall, vertebrate recovery increased with successional stage, but other potential predictors of diversity recovery, such as, the geographical setting (amphibian and reptile species compositional similarity recovered more rapidly on islands), rainfall (mammal species richness and compositional similarity recovered faster in regions of low rainfall), and the landscape context (amphibian, reptile and mammal species compositional similarity recovered faster in regions with more forest patches) influenced vertebrate recovery. These results demonstrate the important role of secondary forests in providing habitat for many vertebrates, but the slow recovery of species compositional similarity, forest specialists and some functional groups (e.g. insectivorous birds) highlighted the challenge of secondary forest persistence, and strongly argues for the continued protection of old growth/mature forest as habitat for forest specialists and as sources for secondary forest sites.     

Vegetation recovery and plant facilitation in a human-disturbed lava field in a megacity: searching tools for ecosystem restoration

Unplanned urban development threatens natural ecosystems. Assessing ecosystem recovery after anthropogenic disturbances and identifying plant species that may facilitate vegetation regeneration are critical for the conservation of biodiversity and ecosystem services in urban areas. At the periphery of Mexico City, illegal human settlements produced different levels of disturbance on natural plant communities developed on a lava field near the Ajusco mountain range. We assessed natural regeneration of plant communities 20 years after the abandonment of the settlements, in sites that received low (manual harvesting of non-timber forest products), medium (removal of aboveground vegetation), and high (removal of substrate and whole vegetation) disturbance levels. We also tested the potential facilitative role played by dominant tree and shrub species. Plant diversity and vegetation biomass decreased as disturbance level increased. Sites with high disturbance level showed poor regeneration and the lowest species similarity compared to the least disturbed sites. Six dominant species (i.e., those with the highest abundance, frequency, and/or basal area) were common to all sites. Among them, three species (the tree Buddleja cordata, and two shrubs, Ageratina glabrata and Sedum oxypetalum) were identified as potential facilitators of community regeneration, because plant density and species richness were significantly higher under their canopies than at open sites. We propose that analyzing community structural traits of the successional vegetation (such as species diversity and biomass) and identifying potential facilitator species are useful steps in assessing the recovery ability of plant communities to anthropogenic disturbances, and in designing restoration strategies.     

Tree community structure, dynamics, and diversity partitioning in a Bornean tropical forested landscape

Human-modified forested landscapes are prevalent in the tropics, and the role of complex mosaics of diverse vegetation types in biodiversity conservation remains poorly understood. Demographic traits and the spatial pattern of biodiversity are essential information when considering proper forest management and land use strategies. We compared the tree community structure (stem density, basal area, tree diversity, abundance of rare, endemic, and upper-layer trees, and species composition) and the forest dynamics (mortality, recruitment rate, and increments of basal area, and above- and below-ground biomass) of 39–46 plots among five dominant forest types: young and old fallows, rubber plantations, and fragmented and old-growth forests in Sarawak, Malaysia. We also explored how tree diversity was distributed across different spatial scales using additive partitioning of diversity. Swidden cultivation and rubber plantations showed decreased stem density, basal area, tree diversity, abundance of rare, endemic, and upper-layer trees, and increments of above- and below-ground biomass, which affected tree mortality, dominant trees, and species composition. Little distinction in species composition was observed among young and old fallows and rubber plantations, indicating a relatively quick recovery of the tree community in the early stages. The highest diversity was found among forest types, indicating that the whole forested landscape comprises a suitable scale for tree biodiversity conservation in the region. Our results suggest that although fragmented and old-growth forests have an irreplaceable role and a high priority in conserving biodiversity and sustaining the function of the forest ecosystem, secondary forests may also have a reinforcing role in maintaining tree diversity in the region, especially under the current circumstances in which a large portion of the landscape is human-modified and faces an increasing threat from the expansion of oil palm plantations.     

Tree diversity,composition, forest structure and aboveground biomass dynamics after single and repeated fire in a Bornean rain forest

Forest fires remain a devastating phenomenon in the tropics that not only affect forest structure and biodiversity, but also contribute significantly to atmospheric CO2. Fire used to be extremely rare in tropical forests, leaving ample time for forests to regenerate to pre-fire conditions. In recent decades, however, tropical forest fires occur more frequently and at larger spatial scales than they used to. We studied forest structure, tree species diversity, tree species composition, and aboveground biomass during the first 7 years since fire in unburned, once burned and twice burned forest of eastern Borneo to determine the rate of recovery of these forests. We paid special attention to changes in the tree species composition during burned forest regeneration because we expect the long-term recovery of aboveground biomass and ecosystem functions in burned forests to largely depend on the successful regeneration of the pre-fire, heavy-wood, species composition. We found that forest structure (canopy openness, leaf area index, herb cover, and stem density) is strongly affected by fire but shows quick recovery. However, species composition shows no or limited recovery and aboveground biomass, which is greatly reduced by fire, continues to be low or decline up to 7 years after fire. Consequently, large amounts of the C released to the atmosphere by fire will not be recaptured by the burned forest ecosystem in the near future. We also observed that repeated fire, with an inter-fire interval of 15 years, does not necessarily lead to a huge deterioration in the regeneration potential of tropical forest. We conclude that burned forests are valuable and should be conserved and that long-term monitoring programs in secondary forests are necessary to determine their recovery rates, especially in relation to aboveground biomass accumulation.     

Recovery of Tawa-dominated forest fragments in the Rotorua Basin, New Zealand, after cessation of livestock grazing

Summary   Tawa ( Beilschmiedia tawa )-dominated forest fragments on farms within the Rotorua Basin were surveyed to quantify the likely recovery processes following exclusion of domestic livestock grazing, using a space-for-time substitution approach. Vegetation structure, plant diversity and soil fertility were measured at 24 sites within 15 forest fragments on six farms, covering a range in time since exclusion from grazing of 1–53 years. The forest fragments were compared with a large area of ungrazed forest in the nearby Lake Okataina Scenic Reserve. As time since exclusion from grazing increased, indigenous plant species diversity increased (up to 30–35 years); ground fern and epiphyte abundance increased (up to 30–35 years); tree seedling and sapling numbers, and litter cover also increased (up to 10–15 years); and overall tree numbers increased, while average tree diameter at breast height and overall tree basal area did not differ significantly. The soil fertility status was highly variable, obscuring clear patterns, although Olsen P status decreased with time since grazing exclusion. Once grazing of forest fragments ceases, significant changes in their diversity, structure and soil characteristics can be expected, which indicate recovery of these plant communities towards the conditions observed in ungrazed forest.     

Soil phosphorus heterogeneity promotes tree species diversity and phylogenetic clustering in a tropical seasonal rainforest

The niche theory predicts that environmental heterogeneity and species diversity are positively correlated in tropical forests, whereas the neutral theory suggests that stochastic processes are more important in determining species diversity. This study sought to investigate the effects of soil nutrient (nitrogen and phosphorus) heterogeneity on tree species diversity in the Xishuangbanna tropical seasonal rainforest in southwestern China. Thirty‐nine plots of 400 m² (20 × 20 m) were randomly located in the Xishuangbanna tropical seasonal rainforest. Within each plot, soil nutrient (nitrogen and phosphorus) availability and heterogeneity, tree species diversity, and community phylogenetic structure were measured. Soil phosphorus heterogeneity and tree species diversity in each plot were positively correlated, while phosphorus availability and tree species diversity were not. The trees in plots with low soil phosphorus heterogeneity were phylogenetically overdispersed, while the phylogenetic structure of trees within the plots became clustered as heterogeneity increased. Neither nitrogen availability nor its heterogeneity was correlated to tree species diversity or the phylogenetic structure of trees within the plots. The interspecific competition in the forest plots with low soil phosphorus heterogeneity could lead to an overdispersed community. However, as heterogeneity increase, more closely related species may be able to coexist together and lead to a clustered community. Our results indicate that soil phosphorus heterogeneity significantly affects tree diversity in the Xishuangbanna tropical seasonal rainforest, suggesting that deterministic processes are dominant in this tropical forest assembly.     

Bryophyte Species Diversity in Secondary Forests Dominated by the Introduced Species Spathodea campanulata Beauv. in Puerto Rico

The introduced tree species Spathodea campanulata (Bignoniaceae) forms novel forests in Puerto Rico, these having emerged after the abandonment of fields in the mid‐20th century and resulting in forests with a new species composition. We assessed bryophyte species richness in these novel forests and sought correlations with geological substrate, past land use, forest edge and patch area, forest structure, elevation, microhabitat diversity, tree species richness, and microclimatic conditions. Transects were established (edge and forest interior) in nine moist forest patches dominated by Spathodea in north‐central Puerto Rico. These Spathodea forest patches ranged from 0.6 to 9 ha. ANOVA, Chi‐square, correlation, and cluster analyses were used in data analyses. We found 57 bryophyte species. There was a significant difference in bryophyte richness among patches. Those on karst exhibited highest bryophyte richness due to microhabitat diversity, past land use, and shorter hydroperiods. Alluvial sites scored lowest in bryophyte species richness, and forest structure was important for bryophyte communities on these sites. Significant differences in temperature, relative humidity, and light intensity were observed between edge and forest interior. These appeared important for establishing bryophyte species cover but not richness and composition. Microhabitat diversity, patch area, and forest age were more related to bryophyte species richness than elevation, exposed edge, and tree species richness, regardless of geologic substrate. Collectively, Spathodea patches were similar to mature forests on the Island with respect to bryophyte species richness and composition. Novel Spathodea forests have conservation value due to their habitat suitability for bryophyte communities.     

Vegetation Structure,Composition, and Species Richness Across a 56‐year Chronosequence of Dry Tropical Forest on Providencia Island,Colombia1

We compared vegetation structure and species richness across a 56‐yr chronosequence of six replicated age classes of dry tropical forest on the island of Providencia, Colombia, in the Southwest Caribbean. Stand age classes were determined using sequential, orthorectified panchromatic aerial photos acquired between 1944 and 1996 and Landsat 7 ETM + satellite imagery from 2000. Along the chronosequence we established 59 plots of 2 × 50 m (0.01 ha) to document changes in species richness, basal area, tree height, stem density, and sprouting ability. All woody trees and shrubs >2.5 cm diameter at breast height (DBH) were censused and measured. Although woody species density reached a peak in stands from 32 to 56 yr old, rarefaction analysis showed that species richness increased linearly with stand age and was highest in stands 56 yr old or greater. Nonparametric, abundance‐based estimators of species richness also showed positive and linear associations with age. Basal area and mean tree height were positively associated with age since abandonment, while sprouting ability showed a negative relationship. Our results indicate rapid recovery of woody species richness and structural characteristics along this tropical dry forest chronosequence.