Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.     

Posthurricane Seedling Structure in a Multi‐aged Tropical Dry Forest: Implications for Community Succession

Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Principles of Natural Regeneration of Tropical Dry Forests for Restoration

Tropical dry forests are the most threatened tropical terrestrial ecosystem. However, few studies have been conducted on the natural regeneration necessary to restore these forests. We reviewed the ecology of regeneration of tropical dry forests as a tool to restore disturbed lands. Dry forests are characterized by a relatively high number of tree species with small, dry, wind‐dispersed seeds. Over small scales, wind‐dispersed seeds are better able to colonize degraded areas than vertebrate‐dispersed plants. Small seeds and those with low water content are less susceptible to desiccation, which is a major barrier for establishment in open areas. Seeds are available in the soil in the early rainy season to maximize the time to grow. However, highly variable precipitation and frequent dry spells are important sources of mortality in seeds and seedlings. Collecting seeds at the end of the dry season and planting them when soil has sufficient moisture may increase seedling establishment and reduce the time they are exposed to seed predators. Germination and early establishment in the field are favored in shaded sites, which have milder environment and moister soil than open sites during low rainfall periods. Growth of established seedlings, however, is favored in open areas. Therefore, clipping plants around established seedlings may be a good management option to improve growth and survival. Although dry forests have species either resistant to fire or that benefit from it, frequent fires simplify community species composition. Resprouting ability is a noticeable mechanism of regeneration in dry forests and must be considered for restoration. The approach to dry‐forest restoration should be tailored to this ecosystem instead of merely following approaches developed for moister forests.     

Tree Regeneration in Church Forests of Ethiopia: Effects of Microsites and Management

Tree regeneration is severely hampered in the fragmented afromontane forests of northern Ethiopia. We explored how trees regenerate in remnant forests along the gradient from open field, forest edge to closed sites and canopy gaps inside the forest. We investigated the effects of seed sowing, litter removal, and weeding on the regeneration success along this gradient. Regeneration success was investigated for four indigenous tree species, and measured in terms of seedling establishment, growth, and survival. Species performed differently according to site conditions. Within the forest, local canopy openings facilitated seed germination ( Ekebergia ), seedling growth (all species except Olea ), or survival ( Ekebergia and Olea ), suggesting that all species benefited from local high light conditions in the forest. Outside the forest, germination (all species) and growth rates ( Juniperus and Olea ) were lower in the open field, most probably due to water stress in the dry season. Outer edge conditions favored growth for three of the four species. Natural seed germination was, however, zero at any site for Juniperus and Olea and low for Ekebergia and Prunus in the open field. Soil scarification influenced germination positively, while weeding did not have a positive effect. These results suggest that simple measures may improve seedling establishment, and that, for some species, forest edges are particularly useful for growth and survival after succesful establishment. Together with erecting fences, needed to protect seedlings against grazing, seed sowing, planting seedling, and soil scarification may contribute to maintain and restore church forests in the fragmented landscapes of northern Ethiopia.     

Survival and growth of dominant tree seedlings in seasonally tropical dry forests of Yucatan: site and fertilization effects

Aims Seasonally tropical dry forests of the Yucatan Peninsula are typically found in sites with nutrient-poor soils because of the recent geological origin of the region. The landscape is dominated by extensive karstic plates that shape environments where vegetation regeneration through seed germination may be limited by the availability of suitable microsites. In this study, we documented the survival and growth of seedlings from three dominant tree species (Bursera simaruba, Piscidia piscipula and Lysiloma latisiliquum) in seasonally tropical dry forests in Yucatan. Specifically, we evaluated the effect of nutrient addition (N and P, separately and in combination) on seedling survival and growth across three sites with differing levels of precipitation.Methods We conducted a nutrient addition experiment, whereby we established 12 plots of dimensions 10×10 m (100 m 2) at each site, from which three plots were randomly selected to receive one of four treatments: N addition, P addition, N and P addition and no nutrient addition (controls). Prior to treatment application, in each plot, we planted 10 seedlings of each species in October 2010 and subsequently conducted surveys of plant growth and survival every 20 days from November 2010 to April 2011.Important findings Overall, nutrient addition increased seedling survival and the magnitude of this effect was similar among sites. We did not observe an additive effect of the N + P treatment on survival. Similarly, we observed a positive effect of nutrient addition on seedling growth, but this effect was contingent upon site; regarding survival, the effects of N and P on seedling growth were not additive. These results suggest that seedling recruitment and growth in the three dominant species of trees in Yucatan are limited by nutrient availability but that the magnitude of this effect, particularly on seedling growth, is specific for species and site.     

Forest Succession and Distance from Forest Edge in an Afro‐Tropical Grassland1

Forest succession on degraded tropical lands often is slowed by impoverished seed banks and low rates of seed dispersal. Within degraded landscapes, remnant forests are potential seed sources that could enhance nearby forest succession. The spatial extent that forest can influence succession, however, remains largely unstudied. In abandoned agricultural lands in Kibale National Park, Uganda, recurrent fires have helped perpetuate the dominance of tall (2–3 m) grasses. We examined the effects of distance from forest and grassland vegetation structure on succession in a grassland having several years of fire exclusion. At 10 and 25 m from forest edge, we quantified vegetation patterns, seed predation, and survival of planted tree seedlings. Natural vegetation was similar at both distances, as was seed (eight species) and seedling (six species) survival; however, distance may be important at spatial or temporal scales not examined in this study. Our results offer insight into forest succession on degraded tropical grasslands following fire exclusion. Naturally recruited trees and tree seedlings were scarce, and seed survival was low (20% after 7 mo). While seedling survival was high (95% after 6 to 8 mo), seedling shoot growth was very slow (x?= 0.5 cm/100 d), suggesting that survivorship eventually may decline. Recurrent fires often impede forest succession in degraded tropical grasslands; however, even with fire exclusion, our study suggests that forest succession can be very slow, even in close proximity to forest.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Testing for Janzen–Connell Effects in a West African Montane Forest

The Janzen–Connell hypothesis proposes that density dependent seed and seedling mortality, combined with increasing seed and seedling survival away from the parent tree, together promote regular spacing of species and thus α diversity. This hypothesis has rarely been tested in tropical Africa, and rarely in montane forests anywhere. We tested this hypothesis using a combination of field experiments and observations in the most floristically diverse dry submontane forest in Nigeria. We investigated distance effects on seedling herbivory, seedling survival and seedling height growth. We found a significant decrease in herbivory with distance from conspecific adult trees for all three species of experimentally planted seedlings (Entandrophragma angolense, Deinbollia pinnata and Sterculia setigera), and also for naturally occurring seedlings of Pouteria altissima but not of Newtonia buchananii or Isolona pleurocarpa. The relative density of large seedlings/saplings of P. altissima, N. buchananii and I. pleurocarpa increased significantly at greater distance from conspecific adult trees; however, we found no significant distance effect on survival or height growth over 3 mo for all three experimentally planted species. Taken together, our results are some of the first to show that Janzen–Connell effects occur on the African continent and in particular montane tropical forest and suggest that such effects may be pantropical.     

Distance‐dependent effects of soil‐derived biota on seedling survival of the tropical tree legume Ormosia semicastrata

Question: How do seed germination and subsequent seedling survival of O. semicastrata (Hance forma litchiifolia How) vary with respect to distance from parent trees and conspecific density in different types of tropical forest? Are there effects of soil biota on O. semicastrata that systematically depend on distance from parent trees and conspecific density? Do soil pathogens differently affect survival of O. semicastrata in different types of tropical forest? Location: Tropical lowland rain forest and tropical montane rain forest in Jianfengling National Nature Reserve, Hainan Island, China. Methods: Individual adult O. semicastrata trees were selected in lowland rain forest and montane rain forest. Soil was collected at a distance of 0‐5 m or 15‐20 m from the parent tree. Soil samples from each distance were combined into a bulk sample. Half of the soil sample was sterilized by autoclaving. Surface‐sterilized seeds were then added to the soil material in shade‐houses at both forests. Results: Germination of O. semicastrata seeds at low‐ or high‐seed density was barely affected by the sterilization procedure. In both forests, seedlings grown in non‐sterilized soil collected close to parent trees had significantly higher mortality compared to those in sterilized soil. In contrast, seedling survival with soil collected far from parent trees was not affected by the soil sterilization procedure. Conclusions: Host‐specific pathogens concentrated in the soil around parent trees may regulate community structure of tropical trees at the stage of seedling development.     

Variations and Trade‐offs in Functional Traits of Tree Seedlings during Secondary Succession in a Tropical Lowland Rain Forest

The seedling stage is generally the most important bottleneck for the successful regeneration of trees in forests. The traits of seedlings, particularly biomass allocation and root traits, are more easily quantified than the traits of adults. In this study, we tested the hypothesis that seedling traits vary and trade‐off tracking the changing environment during secondary succession. We measured the major morphological traits of 27 dominant species and the major environmental factors in a chronosequence (30‐yr‐old fallow, 60‐yr‐old fallow, and old growth forest) after shifting cultivation in a tropical lowland rain forest on Hainan Island, China. The 30‐yr‐old fallow had higher light and nutrient availability, and the older forests had higher soil water content. Redundancy analysis based on species abundance and environmental factors revealed groups of seedlings that dominate in different stages of succession. Seedlings in different stages of succession had different strategies of biomass allocation for harvesting resources that varied in availability. Species characteristic of younger forest had higher allocation to roots and higher specific leaf area, while species characteristic of older forest had higher allocation to leaves. Our study suggests that the variations and trade‐offs in the major functional traits of tree seedlings among successional classes may reflect changes in environmental conditions during succession.     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

Changes of community composition at multiple trophic levels due to hunting in Nigerian tropical forests

Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make predictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. This was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non‐hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera ctions, triggered by the loss of seed dispersers. The results predict a rarity of primate‐dispersed trees in future tropical forest canopies; a forest less diverse in timber and non‐timber resources.     

中国东北森林生长演替模拟模型及其在全球变化研究中的应用

 NEWCOP模型是一个新的适于模拟东北森林的种类组成动态的林窗类计算机模拟模型,它通过模拟在每一个林分斑块上的每株树木的更新、生长和死亡的全过程来反映森林群落的中长期生长和演替动态。由于 NEWCOP模型是一个由气候变量驱动的生态系统模型,故可用于评价气候变化对东北森林生长和演替的影响。在东北大兴安岭、小兴安岭和长白山地区对NEWCOP模型进行了验证和校准。沿环境梯度对NEWCOP模型的数字模拟实验表明：它能准确地再现顶极森林中树种组成及其在东北地区的垂直分布规律和水平分布规律;能准确地再现大兴安岭、小兴安岭和长白山的主要类型森林的生长和演替规律;在一定的场合NEWCOP还可反映林分的径级结构;NEWCOP模型还具有对现有森林的跟踪模拟能力。应用NEWCOP模型评估了东北森林生态系统对可能气候变化的敏感性。在GFDL 2×CO2和GISS 2×CO2气候变化情景下,东北森林的种类组成将发生很大变化,落叶阔叶树将取代目前长白山、小兴安岭的红松(Pinus koraiensis)和大兴安岭的兴安落叶松(Larix gmelinii)成为东北森林主要树种,而针叶树将在地带性森林中占很小的比重,阔叶树中蒙古栎(Quercus mongolica)将是最重要的树种,它将成为小兴安岭和大兴安岭最主要树种;东北地区适于森林生长的区域将大幅度减少,这些变化主要发生在气候变化过渡期。东北森林对不同的气候变化情景有不同响应。但是,总的趋势是未来东北森林中落叶阔叶树的比重将大幅度增加。这些结论对在全球气候变化背景下,我国东北合理地选择造林树种和制定现有森林的保护经营策略具有一定参考价值。     

Allelopathic potential of the neotropical dry-forest tree Acacia pennatula Benth.: inhibition of seedling establishment exceeds facilitation under tree canopies

Secondary succession after land abandonment in tropical dry forests has been suggested to be favoured by the facilitation effects for seedling establishment exerted by pioneer trees isolated in these savannah-like landscapes. However, it has also been noticed that these pioneer species may sometimes have an encroaching effect and arrest succession for several decades. We investigated in this study whether allelopathy can play a role in limiting seedling establishment of co-occurring tree species under the canopy of Acacia pennatula by means of lab bioassays and field experiments in north-west Nicaragua. Leaf extracts of A. pennatula did not affect seed germination but reduced the general growth and especially the development of the root compartment in seedlings, shifting their biomass allocation model to a reduced root/shoot ratio. Survival of planted seedlings under the canopy of A. pennatula was about 20–30% lower than outside, and this reduction was particularly pronounced as the dry season progressed, despite the milder conditions (e.g. higher soil moisture) being experienced in the inner positions under the canopy. Altogether, our results suggest that, rather than facilitating, A. pennatula may inhibit the establishment of seedlings under its canopy probably by means of an allelopathic interference in the development of the root system with critical negative consequences for young seedlings in terms of overcoming the dry season. This article warns about overemphasizing the nucleation effect that remnant and isolated trees may have to facilitate secondary succession in these highly disturbed savannah-like tropical dry forests.     

Tree seedling dynamics in two abandoned tropical dry forests of differing successional status in Yucatán,Mexico: a field experiment with N and P fertilization

The effects of nitrogen (N) and phosphorus (P) fertilization on seedling survival, recruitment, species richness and diversity in two abandoned tropical dry forests (10-yr old, young forest, and c. 60-yr old, old forest) in Yucatán, Mexico, were studied over two years. The seedling dynamics in the control plots were found to be highly seasonal with highest recruitment and lowest death rates during the rainy season. A low percentage of seedlings were resprouts; this important mechanism for forest regeneration had a higher-than-expected survival when compared to seedlings regenerated from seeds. Nutrient addition had significant effects on seedling dynamics in both of the forest regeneration stages. In the young forest, N fertilization facilitated the increase of seedling density. In the old forest, the addition of P decreased seedling diversity, while it increased the recruitment of only a few species. In both forests, P fertilization increased the survival time of seedlings when interacting with light availability and bulk density on the topsoil layer. Results suggest that low nutrient availability combined with low light availability constrain forest succession in Yucatán, Mexico. This revised version was published online in August 2006 with corrections to the Cover Date.     

Non‐Native Grass Removal and Shade Increase Soil Moisture and Seedling Performance during Hawaiian Dry Forest Restoration

Invasive non‐native species can create especially problematic restoration barriers in subtropical and tropical dry forests. Native dry forests in Hawaii presently cover less than 10% of their original area. Many sites that historically supported dry forest are now completely dominated by non‐native species, particularly grasses. Within a grass‐dominated site in leeward Hawaii, we explored the mechanisms by which non‐native Pennisetum setaceum, African fountain grass, limits seedlings of native species. We planted 1,800 seedlings of five native trees, three native shrubs, and two native vines into a factorial field experiment to examine the effects of grass removal (bulldozed vs. clipped plus herbicide vs. control), shade (60% shade vs. full sun), and water (supplemental vs. ambient) on seedling survival, growth, and physiology. Both grass removal and shade independently increased survival and growth, as well as soil moisture. Seedling survival and relative growth rate were also significantly dependent on soil moisture. These results suggest that altering soil moisture may be one of the primary mechanisms by which grasses limit native seedlings. Grass removal increased foliar nitrogen content of seedlings, which resulted in an increase in leaf‐level photosynthesis and intrinsic water use efficiency. Thus in the absence of grasses, native species showed increased productivity and resource acquisition. We conclude that the combination of grass removal and shading may be an effective approach to the restoration of degraded tropical dry forests in Hawaii and other ecologically similar ecosystems.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.