Application of Assisted Natural Regeneration to Restore Degraded Tropical Forestlands

Assisted natural regeneration (ANR) is a simple, low‐cost forest restoration method that can effectively convert deforested lands of degraded vegetation to more productive forests. The method aims to accelerate, rather than replace, natural successional processes by removing or reducing barriers to natural forest regeneration such as soil degradation, competition with weedy species, and recurring disturbances (e.g., fire, grazing, and wood harvesting). Compared to conventional reforestation methods involving planting of tree seedlings, ANR offers significant cost advantages because it reduces or eliminates the costs associated with propagating, raising, and planting seedlings. It is most effectively utilized at the landscape level in restoring the protective functions of forests such as watershed protection and soil conservation. ANR techniques are flexible and allow for the integration of various values such as timber production, biodiversity recovery, and cultivation of crops, fruit trees, and non‐timber forest products in the restored forest. This paper describes the steps of applying ANR and conditions under which it will be most effective. It also discusses ANR’s comparative advantages as well as some of its constraints.     

The Emerging Era of Novel Tropical Forests

In 1966 Eugene P. Odum delivered a speech before the Ecological Society of America that transformed the way ecologists looked at succession. His comparison of mature and successional systems lead ecologists to place secondary forests in an inferior position relative to mature ones to the point that today, prominent tropical biologists argue for and against the conservation value of secondary forests. Nevertheless, we live in the era of secondary forests that is rapidly giving way to a new era of novel tropical forests. Research in Puerto Rico documents the emergence of novel forests, which are different in terms of species composition, dominance, and relative importance of species from forests that were present before the island was deforested. These novel forests emerged without assistance. They are a natural response to the new environmental conditions created by human activity. Natural processes have remixed or reassembled native and introduced plant and animal species into novel communities adapted to anthropogenic environmental conditions. Novel forests are expected to protect soils, cycle nutrients, support wildlife, store carbon, maintain watershed functions, and mitigate species extinctions. The dawn of the age of tropical novel forests is upon us and must not be ignored.     

Hidden Costs of Passive Restoration

The first few years of tropical forest restoration can be expensive, especially when applied to expansive areas. In light of this, passive restoration has been recommended as a considerably cheaper or even free alternative. There are, however, both direct and indirect costs associated with passive restoration. First, the longer recovery time that is typically required in passive restoration can be perceived as project failure, especially when compared with nearby active restoration efforts. In the worst‐case scenario, this can lead to the premature termination of a project by a landowner who would like to see more rapid or visible results. Second, passive restoration may be viewed as land abandonment, and in developing nations where land tenure is not always strictly enforced this may invite unanticipated uses, such as ranchers who may unintentionally or intentionally allow livestock to take advantage of the “unused” forage grass, thus setting back recovery efforts. Lastly, passive restoration does have direct financial costs, including material costs for establishing fences and repairing them, and labor costs for site vigilance. These upfront investments may need to be made repeatedly in passive restoration efforts, and for a longer time period than for an active restoration project. Both the direct and indirect costs should be considered prior to choosing passive restoration as a strategy in a particular restoration project.     

Effects of Season and Vegetation Type on Community Organization of Dung Beetles in a Tropical Dry Forest1

Dung beetles are important components of most terrestrial ecosystems. In tropical rain forests, dung beetle communities can be very rich in number of species and individuals, and they are known to be useful bioindicators of habitat disturbance. In contrast, very little is known about the organization of dung beetle communities in tropical dry forests. The aim of this study was to describe in detail the dung beetle community of a Mexican tropical dry forest and to assess the relative importance of rainfall seasonality and forest structure in affecting the temporal and spatial dynamics of this community. Dung beetles were captured with pitfall traps at the beginning of the rainy season, the middle of the rainy season, and the middle of the dry season, in two distinct forest types: deciduous forest (DF) and semideciduous forest (SDF) at the Estación de Biología Chamela. Both rainfall seasonality and forest structure affected the community organization of dung beetles. During both rainy periods, 14 species were captured, but only three during the dry season. Dung beetles captured during the dry season were only found in the SDF. When comparing the beginning and the middle of the rainy season, differences in abundance and guild structure were also observed between both periods and between forest types, but these differences were much less pronounced.     

Stand Structure and Regeneration of the Subalpine Hagenia abyssinica Forests of Mt. Kenya

The uppermost forest belt on Mt. Kenya (Kenya, East Africa), ranges from 2900 to 3400 m a.s.l. and is dominated by the evergreen “Kosso” tree, Hagenia abyssinica (Bruce) J.F. Cmel. (Rosaceae). The ecology of this tree, with emphasis on regeneration, was investigated. Twenty-five phytosociological relevés, representing several types of Hagenia forests were produced, and attributed to various associations of the alliances Hagenio abyssinicae — Hypericion revoluti Bussmann 1994 and Hagenio abyssinicae — Juniperion procerae Bussmann 1994 (Bussmann and Beck 1995a). Biometric data of young and adult Hagenia trees were collected. Kosso trees of the individual relevés were either of almost equal size, and presumably age, or could be grouped into only two size categories. This uniformity of the Hagenia populations suggests simultaneous regeneration after major disturbance. Charcoal horizons were found in soil profiles at various sites in the Hagenia zone of Mt. Kenya, indicating former burning of these forests. Therefore, fire is thought to be the disturbing factor, which triggers the regeneration of the Hagenia forests. Germination of Hagenia seeds was investigated under various ecological conditions at Mt. Kenya and in greenhouses. Crucial factors for successful germination were high temperatures and bare soil. Light intensity or potentially allelopathic decomposition compounds, leaching from the abundantly shed leaves, had no significant effect on the germination rates. Fire apparently promotes germination by clearing and heating the prospective seed bed. In areas suffering from a high activity of herbivores, the regeneration capability of Hagenia is greatly decreased or abolished. A fire-requiring regeneration cycle of the Hagenia forests of Mt. Kenya is concluded from the phytosociological and ecological data.     

Flower and Fruit Availability along a Forest Restoration Gradient

Frugivores and pollinators are two functional groups of animals that help ensure gene flow of plants among sites in landscapes under restoration and to accelerate restoration processes. Resource availability is postulated to be a key factor to structure animal communities using restoration sites, but it remains poorly studied. We expected that diverse forests with many plant growth forms that have less‐seasonal phenological patterns will provide more resources for animals than forests with fewer plant growth forms and strongly seasonal phenological patterns. We studied forests where original plantings included high tree species diversity. We studied resource provision (richness and abundance of flowers and fruits) of all plant growth forms, in three restoration sites of different ages compared to a reference forest, investigating whether plant phenology changes with restoration process. We recorded phenological data for reproductive plant individuals (351 species) with monthly sampling over 2 years, and found that flower and fruit production have been recovered after one decade of restoration, indicating resource provision for fauna. Our data suggest that a wide range of plant growth forms provides resource complementarities to those of planted tree species. Different flower phenologies between trees and non‐trees seem to be more evident in a forest with high non‐tree species diversity. We recommend examples of ideal species for planting, both at the time of initial planting and post‐planting during enrichment. These management actions can minimize shortage and periods of resource scarcity for frugivorous and nectarivorous fauna, increasing probability of restoring ecological processes and sustainability in restoration sites.     

Research Priorities for Neotropical Dry Forests1

Our understanding of the human and biophysical dimensions of tropical dry forest change and its cumulative effects is still in the early stages of academic discovery. The papers in this special section on Neotropical dry forests cover a wide range of sites and problems ranging from the use of multispectral and hyperspectral remote sensing platforms to the impact of hurricanes on tropical dry forest regeneration. Here, we present to the scientific community the results of a workshop on which research priorities for tropical dry forests were discussed. This discussion focuses on the need to develop linkages between remote sensing, ecological, and social science research. The incorporation of social sciences into ecological research could contribute dramatically to our understandings of tropical dry forests by providing important contextual information to ecologists, and by helping to develop an important science–policy–public nexus on which environmental management can succeed.     

Tropical Dry Forests of Venezuela: Characterization and Current Conservation Status1

Tropical dry forests are located predominantly in the northern portion of Venezuela, above 6°N. Although their potential extent covers ca 400,000 km² (44% of the land), they currently occupy about 10 percent of this area. The diversity and complexity of Venezuelan dry forests increases from north to south along a gradient of decreasing severity of the dry season. A typical dry forest in Venezuela presents ca 110–170 species of plants from ca 40 to 50 families within an area of approximately 10 ha. Species composition and forest structure, however, are dependent on local landscape conditions (e.g., soil type, topography), and nearby forest types can be very different. Our analysis of five dry forest variants showed a maximum family similarity of 67 percent, although most values fell in the 50–60 percent interval. They are currently considered as one of Venezuela's most threatened ecosystems, but only 5 percent of extant dry forests are included in protected areas; this represents 0.5 percent of their potential extent. It is fundamental to promote the creation of at least 3 or 4 more large protected areas (ca 5000 ha), with different climatic and orographic characteristics, in combination with the recovery of threatened species, the restoration of degraded systems, and the implementation of sustainable development projects. Their apparent high resilience suggests that with the proper management we can restore and maintain the integrity of Venezuelan dry forests.     

Deforestation Trends of Tropical Dry Forests in Central Brazil

Tropical dry forests are the most threatened forest type in the world yet a paucity of research about them stymies development of appropriate conservation actions. The Paranã River Basin has the most significant dry forest formations in the Cerrado biome of central Brazil and is threatened by intense land conversion to pastures and agriculture. We examined changes in Paranã River Basin deforestation rates and fragmentation across three time intervals that covered 31 yr using Landsat imagery. Our results indicated a 66.3 percent decrease in forest extent between 1977 and 2008, with an annual rate of forest cover change of 3.5 percent. Landscape metrics further indicated severe forest loss and fragmentation, resulting in an increase in the number of fragments and reduction in patch sizes. Forest fragments in flatlands have virtually disappeared and the only significant forest remnants are mostly found over limestone outcrops in the eastern part of the basin. If current patterns persist, we project that these forests will likely disappear within 25 yr. These patterns may be reversed with creation of protected areas and involvement of local people to preserve small fragments that can be managed for restoration.     

Dictyostelid Cellular Slime Molds in Canopy Soils of Tropical Forests1

The occurrence and distribution of dictyostelid cellular slime molds (CSM) in the mantle of dead organic matter (literally a “canopy soil”) at the bases of large epiphytes were studied in the Luquillo Experimental Forest of north-eastern Puerto Rico. CSM were isolated from 18 of 50 samples collected from this microhabitat, and four different species were recovered. Dictyostelium purpureum was the single most abundant species and represented almost half (48%) of all clones isolated during the study. Total densities (clones/g) averaged only 38 in the five forest types examined, but densities > 75 were recorded for two forest types. Relative abundance of CSM in canopy soils of the five forest types followed the same general pattern displayed by these organisms in forest floor litter, but a particular species was not necessarily common to both microhabitats in a given forest type.     

Dry Season Den Use by Pygmy Spotted Skunk (Spilogale pygmaea) in a Tropical Deciduous Forest of Mexico

Dry season den use by the pygmy spotted skunk Spilogale pygmaea , an endemic and threatened species of western Mexico, was evaluated at the micro and macrohabitat level, in the tropical deciduous forest of Chamela, Jalisco, Mexico. During the dry seasons of 1997, 1998, and 2000, 79 skunk den sites (29 for females and 50 males) were located using Radiotelemetry. Dens were underground excavations with one to five access points, located mostly on open ground or under live or dead trees. Skunks exhibited a continued reuse of dens but evidence of simultaneous occupation by two or more skunks was not found. Microhabitat den-site analysis showed that skunks showed flexibility and individual variation in usage of the microhabitat surrounding den sites inside the forest. Macrohabitat den-site analysis showed that skunks were not selecting either tropical deciduous or tropical semideciduous forest; however, no dens were located outside the forest in transformed habitats, strengthening the hypothesis of the association of this species with the tropical deciduous forests on the Pacific coast of Mexico. The current deforestation rates of the tropical deciduous forest in the coast of Jalisco could be reducing the potential den sites for pygmy spotted skunks and exposing them to adverse environmental conditions and predation.     

Early Regeneration of Tropical Dry Forest from Abandoned Pastures: Contrasting Chronosequence and Dynamic Approaches

Old field secondary succession of tropical dry forests (TDFs) is poorly understood, particularly regarding the dynamics of seedlings, saplings, and sprouts (regenerative communities). We used chronosequence and dynamic approaches to: (1) document successional trajectories of regenerative communities during the first dozen years of regeneration in abandoned pastures at Chamela, Mexico; (2) test the usefulness of chronosequences to predict the dynamics of regenerative communities along time; and (3) assess the influence of surrounding forest matrix, stand density, and understory light availability (in the rainy season) as driving factors of such dynamics. More than 1000 plants and 95 species of shrubs and trees 10–100 cm tall were monitored between 2004 and 2007 in nine abandoned pastures (0–12 yr since abandonment) and two old‐growth forest (OGF) sites; gain and loss rates of plants, species, and plant cover were obtained. Chronosequence predicted a rapid and asymptotic increase of plant density, species density, and plant cover toward the OGF values. Such prediction did not match with dynamic data that showed negative or neutral net community rates of change, independently of fallow age. Recruitment and species gain rates increased with the amount surrounding forest matrix. No other effect of the explored factors was detected. Strong rainfall shortenings could be responsible for the high loss and low gain rates of plants and species recorded in most sites. We highlight the critical role of supra‐annual rainfall variability on the dynamics of TDF regenerative communities and the poor predictive value of chronosequences in forest systems subjected to strong environmental temporal variation. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

The Conservation of Saproxylic Insects in Tropical Forests: A Research Agenda

Invariably, insects are overlooked when tropical forest management issues are discussed, because there are so many species, they are taxonomically intractable and so poorly known. Often people take the view that if you look after the vegetation and vertebrates, the insects will look after themselves. This may be true for some functional groups, but for saproxylic insects, this seems unlikely. Their study deserves high priority, since they are dependent on the very resource – wood – whose removal from the ecosystem is the usual object of forest management. Given the current international effort to develop 'criteria and indicators' to monitor sustainable forest management for biodiversity values, there is a window of opportunity for sound ecological research on saproxylic insects to influence the formulation of forest policy such that their needs can be taken into account. There is already a large body of knowledge on temperate and boreal region saproxylic insects, and on the effects that logging has on them, but knowledge of the tropical forest situation lags far behind. This paper proposes a research agenda to enable the needs of saproxylic insects to be taken into account in natural forest management in the tropics. Basic questions, such as whether logging has so far had an impact on tropical saproxylic insects, and whether there are workable sampling techniques to investigate this, still remain to be addressed and deserve high priority. The links between the responses of saproxylic insects and more 'charismatic' study species need to be investigated. We also need to know whether there is a correlation between the intensity of logging and the response of saproxylic insects, and, critically, whether we would be justified in measuring some surrogate aspects of forest structure (as potential habitat for saproxylic insects) rather than the saproxylic insects themselves, and modelling this to determine likely impacts of different management regimes. We consider such an ambitious research agenda as justified given the scale of impact that forest use and management is likely to have on tropical forest insects in the future.     

Resprouting Ability of Dry Forest Tree Species after Disturbance Does Not Relate to Propagation Possibility by Stem and Root Cuttings

Tropical dry forest tree species are recognized for their high resprouting ability after disturbance. We tested whether species that commonly produce root and stem suckers can be propagated by large stem and root cuttings, a useful method for landscape restoration programs. We performed four experiments: (1) In a greenhouse, we tested the propagation of six species using large stem cuttings collected from early successional sites. We used the following treatments: (i) dry season collection and planting; (ii) dry season collection, storage in humid soil, and wet season planting; (iii) wet season collection and planting; and (iv) wet season collection and planting after treatment with commercial NAA auxin. (2) Stem cuttings of Myracrodruon urundeuva were planted in a pasture during the rainy season after either NAA, IBA, or no auxin treatment. (3) As a control experiment, we also planted cuttings of Spondias mombin, a species known for successfully regenerating from cuttings. (4) Root cuttings of six species were collected in recently plowed pastures and planted in the greenhouse with and without treatment with NAA auxin. No root cuttings rooted. Only M. urundeuva and Astronium fraxinifolium stem cuttings rooted. Maximum success was obtained for stem cuttings collected and planted in the dry season (23%). Only 13% of M. urundeuva had sprouted by the 15th month of the field experiment. As a result, large cuttings are not recommended for propagation of the studied species. Future studies should include development of suitable methods of root harvesting and prospection of traditional knowledge for species selection.     

神农架地区常绿落叶阔叶混交林树种更新研究

神农架山区山体的中部,选取代表性的常绿落叶阎叶混交林样地,总面积为２．４ｈｍ＾２。在样地中对出现的林窗斑块以及相邻的对照的非林窗宙样方进行群落学调查。记录乔木树种的种类、胸径和高度以及它的幼苗和幼树的高度、数量。按照树种在林宙内外重要值的位序差值将群落出现的乔木层树种划分为４类生态种组,其中,对林窗强烈正更新反应的树种有６种,强烈负更新反应的树种９种,中等更新反应的９种,不明显更新反应的１０种。林窗内外乔木树种生态种组的组成明显不同。不同生态种组树种的幼苗在林宙与非林窗斑块中的更新表现出显著的差别。对神农架地区常绿落叶阎叶混交林树种更新的反应和过程的研究结果支持了林窗—分享假说的观点。     

猴耳环天然更新特性

为研究药用植物猴耳环（Archidendron clypearia）的天然更新特性,2015年7月和2016年7月对广东省广州市、惠州市和深圳市主要森林公园、自然保护区和生态公益林内天然分布的42株母树进行了天然更新状况调查,并采用熵值法对更新效果进行评价。结果表明：（1）母树的上下和左右冠幅十分接近,但97.6%的母树存在偏冠现象,尤以向坡下偏冠更为明显,上坡方向树冠宽度约为下坡的1/2;偏冠是影响种子散布范围的主要原因之一;（2）与母树的距离与不同方向对种子雨量影响显著,但两者对苗木的数量和径高生长没有显著影响;（3）林冠下≤ 1 a的苗木占总数的87.8%,且林下无5年及以上的幼树,预示猴耳环依靠天然更新维持种群增长困难较大;（4）用信息熵理论导出的更新指数来评价与母树不同距离的更新效果显示,与母树距离越远更新效果越差;距母树0~3 m范围内更新效果最好;（5）猴耳环具有诸多"极小种群野生植物"的特征,亟需对其资源分布现状及生物生态学特性开展进一步的深入调查和研究。     

Love of Nurse Plants is Not Enough for Restoring Oak Forests in a Seasonally Dry Tropical Environment

The highest concentration of oak species in the world occurs in Mexico, but human activities have strongly degraded these oak forests. Mexican oaks have high economic, social, and cultural value, and restoring these forests is of paramount importance for the people of Mexico. Here, we propose a method for restoring oak forests using native shrubs that colonize degraded areas as nurse plants for oak seedlings. To test the viability of this proposal, seedling transplant experiments were performed in a degraded area near a protected oak forest relict. Two pioneer shrubs were identified as potential nurse species: Mimosa luisana and Senecio sp. The target oak species was Quercus castanea . Oak seedlings were located beneath the canopies of both shrubs and in the surrounding area without shrub cover. Water is a limiting resource for oak establishment in seasonally dry environments; therefore, we included irrigation systems in our experimental design to determine whether the combination of nurse plants plus watering led to higher rate of survival than the presence of nurse species alone. Seedling survival without watering was less than 20% both beneath nurse species and in the surrounding habitat. When water was supplied, survival rate beneath nurse species increased up to 58% while survival in the surrounding habitat did not differ from that observed in treatments without watering. Our results indicate that survival rate of oak seedlings is increased by the presence of nurse plants only when water is supplied. This suggests that restoration of oak forests in these degraded areas requires both nurse plants and watering.     

Forest Regeneration from Pasture in the Dry Tropics of Panama: Effects of Cattle, Exotic Grass, and Forested Riparia

We currently have the opportunity to restore one of the most threatened tropical ecosystems on the Pacific coast of Panama as a consequence of land use change. Factors that influence succession must be understood in order to capitalize on natural regeneration mechanisms. In this study, we determined the effects of exotic grass removal, cattle removal, proximity to forested riparian zones, and topography (upslope vs. downslope) on the initial stages of forest regeneration from pasture in a dry tropical region. After 3 years, basal area, stem density, and species richness of plants (trees, shrubs, vines, and herbs) were significantly and positively affected by exclusion of cattle, presence of exotic grass (no herbicide application), and presence of adjacent forested riparian zones ( p < 0.01). Interactions between factors were not significant. Cattle foraged and stomped on vegetation, whereas herbicide application, although effectively removing grass, also killed tree and shrub sprouts, the major source of regrowth. Proximity to forested riparian zones had the greatest effect on species diversity. Shannon's index for diversity ( H ) equaled 3.23 in plots adjacent to forested riparian zones as compared to 2.78 in plots not associated with these areas. Our recommendations during the early stages of forest succession are to (1) exclude cattle, (2) make site-specific decisions about herbicide application based on the presence or absence of forested riparia and prevalence of coppicing, and (3) actively conserve and protect riparian zones, which function as a critical source of diverse propagules.     

Effects of Habitat, Cattle Grazing and Selective Logging on Seedling Survival and Growth in Dry Forests of Central Brazil

We studied the effect of forest gaps and cattle grazing on survival and growth of seedlings of seven native tree species, planted in dry forest fragments with different intensities of past logging in Central Brazil. Seedling survival after 12 mo ranged from 35 to 77 percent among species. Survival of most species was similar in gaps and understory in minimally disturbed forest remnants, but was lower in the gaps of more heavily logged forests. In contrast, growth was much higher in gaps than in understory in minimally disturbed forests, but for most species was similar in gaps and understory in more heavily logged forests. We did not detect an effect of cattle on seedling survival. Seedlings of the most commonly logged species show a high potential to survive and grow when planted in logged forests.     

The Role of Sprouts in the Restoration of Atlantic Rainforest in Southern Brazil

The relative importance of sexual reproduction (seed) and sprouting as sources for regeneration in Brazilian Atlantic Forest was evaluated in three different successional forest stages: young forest, immature forest, and late‐successional forest. Young plants (10–100 cm tall) of tree species were classified into the following categories: (1) seedlings that are nonsprouting—plants that originated through sexual reproduction as seeds; (2) stem base sprouting—plants that sprouted at the base of an existing plant; and (3) underground stem sprouting—plants that sprouted from subterranean stems of an existing plant. A total of 1,030 individuals of 48 species were collected. Underground stem sprouting is the rarest form of propagation, with stem base sprouting somewhat more common and possibly associated with recovery of damaged parts. The greatest contribution to regeneration was due to seeds: 92% of the individuals counted in 67% of the plant species. However, 13 species were “facultative” sprouters as seedlings and sprouters were observed in this group. The three forest ages differed in the proportion of regeneration strategies; in immature forest, sprouting was more common (15%) than in young (7%) and late‐successional (3%; p < 0.05) forest. In these three forest stages, germinating seeds are the major source of new plants; although sprouting as a reproductive strategy is rare, it is related to recovery after damage of an already existing plant and may be due to previous land use history (agriculture) and low soil fertility. Restoration using natural regeneration should consider these factors to understand seed arrival as seed is the main source of regeneration.