Natural regeneration as a tool for large‐scale forest restoration in the tropics: prospects and challenges

A major global effort to enable cost‐effective natural regeneration is needed to achieve ambitious forest and landscape restoration goals. Natural forest regeneration can potentially play a major role in large‐scale landscape restoration in tropical regions. Here, we focus on the conditions that favor natural regeneration within tropical forest landscapes. We illustrate cases where large‐scale natural regeneration followed forest clearing and non‐forest land use, and describe the social and ecological factors that drove these local forest transitions. The self‐organizing processes that create naturally regenerating forests and natural regeneration in planted forests promote local genetic adaptation, foster native species with known traditional uses, create spatial and temporal heterogeneity, and sustain local biodiversity and biotic interactions. These features confer greater ecosystem resilience in the face of future shocks and disturbances. We discuss economic, social, and legal issues that challenge natural regeneration in tropical landscapes. We conclude by suggesting ways to enable natural regeneration to become an effective tool for implementing large‐scale forest and landscape restoration. Major research and policy priorities include: identifying and modeling the ecological and economic conditions where natural regeneration is a viable and favorable land‐use option, developing monitoring protocols for natural regeneration that can be carried out by local communities, and developing enabling incentives, governance structures, and regulatory conditions that promote the stewardship of naturally regenerating forests. Aligning restoration goals and practices with natural regeneration can achieve the best possible outcome for achieving multiple social and environmental benefits at minimal cost.     

The Role of Microsite Conditions in Restoring Trembling Aspen (Populus tremuloides Michx) from Seed

Encouraging natural regeneration of Populus tremuloides Michx (trembling aspen) from seed is a largely unexplored means for reintroducing the species into reclamation areas. We evaluated the effects of microsite (surface contour and substrate type) on aspen seedling establishment and growth on a reclaimed coal mine. The 4.6 ha study site was divided into six 48 m‐wide strips that had 15 or 40 cm capping material salvaged from a nearby forest floor added to the mine surface. We surveyed 126 m long transects located in the center of each strip for microsite conditions, and the presence and height of aspen seedlings. We found that aspen seedlings generally preferred mineral‐organic substrates and concave microsites. To facilitate the regeneration of aspen by seed, we suggest that land managers increase small‐scale roughness and microtopographic diversity on reclaimed sites .     

Incorporating natural regeneration in forest landscape restoration in tropical regions: synthesis and key research gaps

Extensive tropical forest loss and degradation have stimulated increasing awareness at the international policy level of the need to undertake large‐scale forest landscape restoration (FLR). Natural regeneration offers a cost‐effective way to achieve large‐scale FLR, but is often overlooked in favor of tree plantations. The studies presented in this special issue show how natural regeneration can become an important part of FLR and highlight the ecological, environmental, and social factors that must be considered to effectively do so. They also identify major knowledge gaps and outline a research agenda to support the use of natural regeneration in FLR. Six central questions emerge from these studies: (1) What are the ecological, economic, and livelihood outcomes of active and passive restoration interventions?; (2) What are the tradeoffs and synergies among ecological, economic, and livelihood outcomes of natural regeneration, restoration and productive land uses, and how do they evolve in the face of market and climate shocks?; (3) What diagnostic tools are needed to identify and map target areas for natural regeneration?; (4) How should spatial prioritization frameworks incorporate natural regeneration into FLR?; (5) What legal frameworks and governance structures are best suited to encourage natural regeneration and how do they change across regions and landscapes?; (6) What financial mechanisms can foster low‐cost natural regeneration? Natural regeneration is not a panacea to solve tensions and conflicts over land use, but it can be advantageous under some circumstances. Identifying under what conditions this is the case is an important avenue for future research.     

Old field colonization by native trees and shrubs following land use change: Could this be Victoria’s largest example of landscape recovery?

Summary Natural regeneration of farmland areas following landuse change has the potential to reinstate native vegetation and landscape processes across larger scales than intentional works. However, few examples of large‐scale natural regeneration have been reported from southern Australia. In this study we use historical air photos to document the rate of establishment of natural regeneration in central Victoria following a change from agricultural to rural residential land use. In 2009, regrowth patches occupied 8185 ha, or 12.3% of the cleared landscape in the study region, mostly on relatively low fertility soils. Most of this area (6216 ha) supported Cassinia shrubland, with eucalypts encroaching as patches get older. On average, native vegetation has regenerated over nearly 1800 ha every decade since the mid‐1960s. If this trend continues, regrowth will occupy 20% of infertile soils on private land by 2025. This region now appears to support one of the largest examples of old field succession recorded from south‐eastern Australia. Regrowth patches are likely to provide many conservation benefits, although little information exists on habitat values provided by regrowth shrublands. Since regeneration is on private land, perceptions of whether regrowth is ‘good’ or ‘bad’ will vary according to landholder goals, as will future management of regrowth patches. Consequently, considerable ecological and social research is required to understand the ecosystem services and disservices which regrowth provides to both landholders and biota.     

Ecological outcomes of agroforests and restoration 15 years after planting

Large‐scale forest restoration relies on approaches that are cost‐effective and economically attractive to farmers, and in this context agroforestry systems may be a valuable option. Here, we compared ecological outcomes among (1) 12–15‐year‐old coffee agroforests established with several native shade trees, (2) 12–15‐year‐old high‐diversity restoration plantations, and (3) reference old‐growth forests, within a landscape restoration project in the Pontal do Paranapanema region, in the Atlantic Forest of southeastern Brazil. We compared the aboveground biomass, canopy cover, and abundance, richness, and composition of trees, and the regenerating saplings in the three forest types. In addition, we investigated the landscape drivers of natural regeneration in the restoration plantations and coffee agroforests. Reference forests had a higher abundance of trees and regenerating saplings, but had similar levels of species richness compared to coffee agroforests. High‐diversity agroforests and restoration plantations did not differ in tree abundance. However, compared to restoration plantations, agroforests showed higher abundance and species richness of regenerating saplings, a higher proportion of animal‐dispersed species, and higher canopy cover. The abundance of regenerating saplings declined with increasing density of coffee plants, thus indicating a potential trade‐off between productivity and ecological benefits. High‐diversity coffee agroforests provide a cost‐effective and ecologically viable alternative to high‐diversity native tree plantations for large‐scale forest restoration within agricultural landscapes managed by local communities, and should be included as part of the portfolio of reforestation options used to promote the global agenda on forest and landscape restoration.     

Microsite factors influencing Eucalyptus regeneration in temperate woodlands

Natural regeneration of Eucalyptus is a major problem for the restoration and preservation of temperate woodlands. South‐eastern Australia experienced a period of above average rainfall from mid‐2010 to early 2011, creating favourable conditions for localised Eucalyptus regeneration. We conducted a case–control study comparing sites where natural regeneration from seed was present and absent. Vegetation cover, soil condition and hydrophobicity, as well as shelter objects and landscape position, were recorded to identify what influences natural regeneration from seed. Using conditional logistic regression, we found the odds of regeneration occurring increased with uphill slope angle and herbaceous cover. Our results suggest that, during periods of high rainfall, stable microsites that capture seed and resources are most likely to support Eucalyptus regeneration.     

Using historical woodland creation to construct a long‐term,large‐scale natural experiment: the WrEN project

Natural experiments have been proposed as a way of complementing manipulative experiments to improve ecological understanding and guide management. There is a pressing need for evidence from such studies to inform a shift to landscape‐scale conservation, including the design of ecological networks. Although this shift has been widely embraced by conservation communities worldwide, the empirical evidence is limited and equivocal, and may be limiting effective conservation. We present principles for well‐designed natural experiments to inform landscape‐scale conservation and outline how they are being applied in the WrEN project, which is studying the effects of 160 years of woodland creation on biodiversity in UK landscapes. We describe the study areas and outline the systematic process used to select suitable historical woodland creation sites based on key site‐ and landscape‐scale variables – including size, age, and proximity to other woodland. We present the results of an analysis to explore variation in these variables across sites to test their suitability as a basis for a natural experiment. Our results confirm that this landscape satisfies the principles we have identified and provides an ideal study system for a long‐term, large‐scale natural experiment to explore how woodland biodiversity is affected by different site and landscape attributes. The WrEN sites are now being surveyed for a wide selection of species that are likely to respond differently to site‐ and landscape‐scale attributes and at different spatial and temporal scales. The results from WrEN will help develop detailed recommendations to guide landscape‐scale conservation, including the design of ecological networks. We also believe that the approach presented demonstrates the wider utility of well‐designed natural experiments to improve our understanding of ecological systems and inform policy and practice.     

Lowland tapirs facilitate seed dispersal in degraded Amazonian forests

The forests of southeastern Amazonia are highly threatened by disturbances such as fragmentation, understory fires, and extreme climatic events. Large‐bodied frugivores such as the lowland tapir (Tapirus terrestris) have the potential to offset this process, supporting natural forest regeneration by dispersing a variety of seeds over long distances to disturbed forests. However, we know little about their effectiveness as seed dispersers in degraded forest landscapes. Here, we investigate the seed dispersal function of lowland tapirs in Amazonian forests subject to a range of human (fire and fragmentation) and natural (extreme droughts and windstorms) disturbances, using a combination of field observations, camera traps, and light detection and ranging (LiDAR) data. Tapirs travel and defecate more often in degraded forests, dispersing much more seeds in these areas [9,822 seeds per ha/year (CI_95% = 9,106; 11,838)] than in undisturbed forests [2,950 seeds per ha/year (CI_95% = 2,961; 3,771)]. By effectively dispersing seeds across disturbed forests, tapirs may contribute to natural forest regeneration—the cheapest and usually the most feasible way to achieve large‐scale restoration of tropical forests. Through the dispersal of large‐seeded species that eventually become large trees, such frugivores also contribute indirectly to maintaining forest carbon stocks. These functions may be critical in helping tropical countries to achieve their goals to maintain and restore biodiversity and its ecosystem services. Ultimately, preserving these animals along with their habitats may help in the process of natural recovery of degraded forests throughout the tropics. Abstract in Portuguese is available with online material.     

Vertebrate‐mediated seed rain and artificial perches contribute to overcome seed dispersal limitation in a Mediterranean old field

Natural regeneration of vegetation is a frequent outcome of land abandonment, although the rate and diversity of such regeneration may be severely restricted by seed dispersal limitation, among other factors. In spite of this, studies aiming to quantify seed rain and test methods to enhance it, such as artificial perches, are still underrepresented in the Mediterranean. In our study, we quantified seed rain density and richness and tested the effects of artificial perches on such rain over a distance gradient on seven Mediterranean island old fields. In each of the seven sites, we positioned three sampling stations, each consisting of 1 seed trap under an artificial perch and 1 as a control on the ground, distributed at 30, 60, and 90 m from natural vegetation remnant. All traps received seeds, suggesting no overall dispersal limitation. Of the 11 seed species found, 10 were fleshy‐fruited and dispersed by vertebrates. Seed traps under perches received significantly higher seed rain of fleshy‐fruited species dispersed by birds, while ground traps received significantly more seeds of the species also dispersed by mammals, especially Rubus ulmifolius. The distance from the seed source was nonsignificant in all cases. Our study demonstrates the key role of vertebrate‐mediated seed dispersal services to overcome dispersal limitation in old fields, as well as the effective contribution of even small artificial perches in contrasting such limitation. The lack of differences over the distance gradient reveal that the upper spatial limit of dispersal limitation was not achieved.     

The role of natural regeneration to ecosystem services provision and habitat availability: a case study in the Brazilian Atlantic Forest

Natural regeneration provides multiple benefits to nature and human societies, and can play a major role in global and national restoration targets. However, these benefits are context specific and impacted by both biophysical and socioeconomic heterogeneity across landscapes. Here, we investigate the benefits of natural regeneration for climate change mitigation, sediment retention and biodiversity conservation in a spatially explicit way at very high resolution for a region within the global biodiversity hotspot of the Atlantic Forest. We classified current land‐use cover in the region and simulated a natural regeneration scenario in abandoned pasturelands, areas where potential conflicts with agricultural production would be minimized and where some early stage regeneration is already occurring. We then modeled changes in biophysical functions for climate change mitigation and sediment retention, and performed an economic valuation of both ecosystem services. We also modeled how land‐use changes affect habitat availability for species. We found that natural regeneration can provide significant ecological and social benefits. Economic values of climate change mitigation and sediment retention alone could completely compensate for the opportunity costs of agricultural production over 20 yr. Habitat availability is improved for three species with different dispersal abilities, although by different magnitudes. Improving the understanding of how costs and benefits of natural regeneration are distributed can be useful to design incentive structures that bring farmers’ decision making more in line with societal benefits. This alignment is crucial for natural regeneration to fulfill its potential as a large‐scale solution for pressing local and global environmental challenges.     

Atlantic Forest topsoil nutrients can be resistant to disturbance and forest clearing

Human impacts can affect the soil properties through erosion and leaching, the ecosystem functions and, consequently, the capacity of a forest to regenerate. Here, we determine the effects of forest disturbance and succession on selected soil chemical properties using two different approaches, before‐after‐control‐impact (BACI) and space‐for‐time (SFT) substitution, and the threatened Atlantic Forest biome as model. We assessed with BACI the long‐term (37‐year) effects of clear cutting on soil properties by comparing data from two topsoil surveys (1978–2017) divided into two treatments: a preserved old growth forest (control) and an adjacent forest that was experimentally cleared with full tree removal (clear‐cut). We examined with SFT the relationship between stand age and soil properties using soil data from three old growth and 13 s growth forests ranging from 7 to 33 years. We found no significant differences between treatments for any soil property or significant changes in phosphorus, potassium, and calcium + magnesium over time. In contrast, pH increased and aluminum decreased in both areas. No relation was found between forest age and most of soil properties, with the exception of potassium which returned to old growth forest levels after 20 years of natural succession, and pH. BACI indicated that deforestation of old growth forest caused no significant effects on soil chemical properties after 37 years of regeneration. SFT demonstrated that soil properties did not change significantly during forest regeneration on formerly disturbed lands. Our findings indicate that natural nutrient‐depleted lowland forests were overall resistant to deforestation followed by passive regeneration at landscape scale. Abstract in Portuguese is available with online material.     

植物自然更新研究进展

植物自然更新是一个复杂的生态学过程,它对种群的增殖、扩散、延续和群落稳定及演替具有重要的作用,是植被动态研究的热点。目前国内外在这方面的多数研究主要从更新过程中的某一或几个阶段入手,分析各生态因子对更新的影响,或从林窗、火烧、动物等干扰对自然更新影响的角度揭示植被自然更新的机制。本文从植物自然更新方式、干扰对更新的影响及群落的更新等方面进行了总结,并提出了今后该领域的研究建议。     

Natural regeneration and biodiversity: a global meta‐analysis and implications for spatial planning

Natural regeneration offers a cheaper alternative to active reforestation and has the potential to become the predominant way of restoring degraded tropical landscapes at large‐scale. We conducted a meta‐analysis for tropical regions and quantified the relationships between both ecological and socioeconomic factors and biodiversity responses in naturally regenerating areas. Biogeographic realms, past disturbance, and the human development index (HDI) were used as explanatory variables for biodiversity responses. In addition, we present a case study of large‐scale natural regeneration in the Brazilian Atlantic Forest and identify areas where different forms of restoration would be most suitable. Using our dataset for tropical regions, we showed that natural regeneration was predominantly reported within: the Neotropical realm; areas that were intensively disturbed; and countries with medium HDI. We also found that biodiversity in regenerating forests was more similar to the values found in old growth forests in: countries with either low, high, or very high HDI; less biodiverse realms; and areas of less intensive past disturbance. Our case study from Brazil showed that the level of forest gain resulting from environmental legislation, in particular the Brazilian Forest Code, has been reduced, but remains substantial. Complementary market incentives and financial mechanisms to promote large‐scale natural regeneration in human‐modified agricultural landscapes are also needed. Our analysis provides insights into the factors that promote or limit the recovery of biodiversity in naturally regenerating areas, and aids to identify areas with higher potential for natural regeneration.     

Demographic costs and benefits of natural regeneration during tropical forest restoration

For tropical forest restoration to result in long‐term biodiversity gains, native trees must establish self‐sustaining populations in degraded sites. While many have asked how seedling recruitment varies between restoration treatments, the long‐term fate of these recruits remains unknown. We address this research gap by tracking natural recruits of 27 species during the first 7 years of a tropical forest restoration experiment that included both planted and naturally regenerating plots. We used an individual‐based model to estimate the probability that a seedling achieves reproductive maturity after several years of growth and survival. We found an advantage for recruits in naturally regenerating plots, with up to 40% increased probability of reproduction in this treatment, relative to planted plots. The demographic advantage of natural regeneration was highest for mid‐successional species, with relatively minor differences between treatments for early‐successional species. Our research demonstrates the consequences of restoration decision making across the life cycle of tropical tree species.     

枯枝落叶层对森林天然更新的障碍

探讨枯枝落叶层对树种天然更新的障碍机制,对于实现天然次生林的科学管理以及采取适当营林措施促进目的树种天然更新具有重要的理论和实践意义.从枯枝落叶层的物理阻断影响 (物理因子)、枯枝落叶层的化感作用(化学因子)、动物侵害和微生物致病(生物因子)等方面,论述了枯枝落叶层对天然更新方面的障碍作用,揭示枯枝落叶层物理阻断对天然更新影响规律,以及主要林型的化感作用与天然更新关系的作用机制.建议今后应加强对不同林型枯枝落叶层的结构、枯枝落叶层下光量子流密度的变化规律以及红光与远红光比值的变化规律等方面研究.     

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.     

1987年大兴安岭特大火灾后北坡森林景观生态恢复评价

通过空间直观景观模型(LAND IS)来探讨大兴安岭北坡1987年特大森林火灾后,在目前这种恢复方案下以及完全依靠天然更新下森林景观的长期动态,通过对比研究来评价目前所采取的恢复措施是否能够有效地恢复森林资源。研究结果表明:1987年大火后所采取的恢复措施可以在很大程度上增加针叶树种在该区所占的比例,而相应地降低阔叶树种的比重。由于在目前的恢复措施下,不仅大面积地种植针叶树种,同时也采伐了大量的成、过熟林,使落叶松和樟子松的蓄积量在开始阶段不仅没有增加,反而有大幅度的下降。但最终,所更新的幼苗逐渐成材,落叶松和樟子松的蓄积量逐渐超过天然更新下的。在演替前期,白桦的蓄积量在这2种模拟方案下相差不大;而在演替后期,白桦在天然更新方案下的蓄积量要高于在目前恢复措施下的蓄积量。另外,物种在不同的区域的变化动态又有较大的差异。在重度火烧区,由于人为种植大量的针叶树种,使它们所占的面积比例明显高于完全依靠天然更新下其在重度火烧区所占的比例;而阔叶树种则相反,在完全依靠天然更新方案下所占的面积百分比明显高于在目前恢复方案下的比例。而在未火烧和轻中度火烧区,由于火后人为采伐大量的落叶松和樟子松,使这2个树种在该区所占的比例在开始100多年里要低于完全依靠天然更新下的比例,随后逐渐高于天然更新下所占的比例,但相差不大;阔叶树种则正好相反。另外,目前恢复方案不仅极大地改变了主要物种在该区所占的比例,而且还明显的影响了它们的空间分布格局。     

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.     

Genotyping validates photo‐identification by the head scale pattern in a large population of the European adder (Vipera berus)

Capture‐mark‐recapture procedures are a basic tool in population studies and require that individual animals are correctly identified throughout their lifetime. A method that has become more and more popular uses photographic records of natural markings, such as pigmentation pattern and scalation configuration. As with any other marking tool, the validity of the photographic identification technique should be evaluated thoroughly. Here, we report on a large‐scale double‐marking study in which European adders (Vipera berus) were identified by both microsatellite genetic markers and by the pattern of head scalation. Samples that were successfully genotyped for all nine loci yielded 624 unique genotypes, which matched on a one‐to‐one basis with the individual assignments based on the head scalation pattern. Thus, adders considered as different individuals by their genotypes were also identified as different individuals by their head scalation pattern, and vice versa. Overall, variation in the numbers, shape, and arrangement of the head scales enabled us to distinguish among 3200+ photographed individual snakes. Adders that were repeatedly sequenced genetically over intervals of 2–3 years showed no indication whatsoever for a change in the head scale pattern. Photographic records of 900+ adders that were recaptured over periods of up to 12 years showed a very detailed and precise match of the head scale characteristics. These natural marks are thus robust over time and do not change during an individual's lifetime. With very low frequency (0.3%), we detected small changes in scalation that were readily discernible and could be attributed to physical injury or infection. Our study provides a conclusive validation for the use of photo‐identification by head scale patterns in the European adder.     

Fishing‐down within populations harms seed dispersal mutualism

Large fish are often the most effective seed dispersers, but they are also the preferred target for fisheries. We recently started to comprehend the detrimental impacts of the extirpation of large frugivorous fish species on natural forest regeneration, but we lack a general understanding of how intraspecific size‐selective harvest affects fish–fruit mutualism. Our literature review demonstrated that large individuals within populations positively affect diverse aspects of seed dispersal, from consuming a higher diversity of seeds to enhancing germination. Furthermore, we filled a research gap by studying how individual size variations within two small frugivorous fish species (<16 cm) affect seed dispersal in flooded savannas. Even within small‐bodied species, large individuals swallow a higher number of intact seeds, but not necessarily a higher proportion. Overall, our results demonstrate the disproportional role of large‐bodied individuals as key seed dispersers in flooded habitats. Consequently, fishing‐down within both large‐ and small‐bodied species can negatively affect seed dispersal and natural regeneration in overfished wetlands.