The frailty of tropical restoration plantings

Conserving and restoring biodiversity are compelling challenges in the face of deforestation and fragmentation of tropical forests. Establishing restoration plantings that act as stepping‐stone corridors for animals and develop into forest islets is one way we can reconnect forest fragments split by active agricultural landscapes. However, this strategy's success is contingent on dispersal agents attracted from the forests that vary greatly in their dispersal services, diet, and mobility. Dispersal agents capable of traversing the agricultural matrix that also provide high‐quality and high‐quantity seed dispersal are often a small subset of the present fauna. They also tend to be large‐bodied birds with broad diets (e.g. toucans). This subset of dispersal agents (here termed “effective restoration agents”) plays a key role in driving succession in restoration plantings. Their absence or low numbers can compromise the strategy of using plantings to enhance connectivity in landscapes fragmented by crops, orchards, or extensive pastures. In this event, additional intervention may be required to attract other dispersal agents that would otherwise not disperse seeds at or play a significant role in restoration plantings.     

Genetic diversity of reintroduced tree populations in restoration plantations of the Brazilian Atlantic Forest

Long‐term ecological success of large‐scale restoration programs planned for the next decades will rely on genetic diversity (GD) of reintroduced or colonizing species, a limiting factor in highly fragmented landscapes. In small and isolated natural remnants or restoration areas, substantial reduction in population's size or connectivity may lead to local extinctions due to the accumulation of deleterious recessive alleles and ongoing reduction of fecundity, plant vigor, recruitment success, and adaptive potential. Despite the paramount role of GD for species persistence, its levels in restoration programs are poorly known. We assessed the GD of four model tree species (different succession stages, dispersal, and pollination syndromes) from the Brazilian Atlantic Forest, comparing two high‐diversity restoration plantations, one forest fragment and one conserved remnant. Contrary to the expectation that the plantation strategies adopted in the restoration programs could result in genetic composition homogenization, we found that restoration areas established heterogeneous genetic groups with similar levels of neutral GD and inbreeding to those observed in natural forest remnants. This pattern was consistent across the four functionally different tree species, despite some species idiosyncrasies. For instance, we observed lower allelic richness in early successional species in restoration sites, suggesting that some species may be more prone to reintroduction with lower GD. Thus, we advocate the use of high GD levels in restoration to support biodiversity conservation in human‐modified landscapes, thus reinforcing the role of ecological restoration for recovering the diversity of genes—the basic constituent of biodiversity.     

Small seed bank in grasslands and tree plantations in former grassland sites in the South Brazilian highlands

The soil seed bank can be an important source for vegetation regeneration, and data on the similarity between aboveground vegetation and the seed bank can provide information about successional pathways after disturbances or land-use change. We conducted this study in natural grasslands in the subtropical highland region in southern Brazil. We evaluated the effect of silviculture on richness, density, and composition of the seed bank at former grassland sites converted to pine plantations 25 years ago. We worked at six grassland sites and three pine plantation sites and used the seedling emergence method. Seed bank density and richness in grasslands were lower than those reported in similar environments in other regions. Species richness and density varied considerably within each vegetation type; therefore, richness and density were not statistically significant, while composition varied among vegetation types. In terms of species, the pine plantation seed bank was a small subset of the grassland seed bank. Seeds of typical grassland species were missing in the pine plantation, but also had only low abundances in the grassland, and similarity of seed bank and vegetation were low (less than 20%). The low seed density found in this study, including in grasslands areas, indicates that regeneration of species from the soil seed bank likely is of a limited role for the maintenance of plant populations after disturbances in this system. Our data further suggest that natural regeneration after tree planting in grasslands is reduced due to seed limitation.     

Carbon stocks in above‐ground biomass of monoculture plantations,mixed species plantations and environmental restoration plantings in north‐east Australia

Summary The emergence of carbon markets has provided a potential source of funding for reforestation projects. However, there is concern amongst ecologists that these markets will promote the establishment of monoculture plantations rather than more diverse restoration plantings, on the assumption that fast‐growing monocultures are likely to store more carbon than restoration plantings. We examined the validity of this assumption for three predominantly rainforest plantation types established in the moist tropical uplands of north‐east Australia: monoculture plantations of native rainforest conifers (n = 5, mean age 13 years); mixed species plantations of rainforest cabinet timber species, rainforest conifers and eucalypts (n = 5, mean age 13 years); and, environmental restoration plantings comprised mostly of a diverse range of rainforest trees (n = 10, mean age 14 years). We found that restoration plantings stored significantly more carbon in above‐ground biomass than monoculture plantations of native conifers (on average, 106 t vs 62 t carbon per ha); and tended to store more carbon than mixed species timber plantations which were intermediate in value (86 t carbon per ha). Carbon stocks were higher in restoration plantings than in monoculture and mixed species plantations for three reasons. First, and most importantly, restoration plantings were more densely stocked than monoculture and mixed species plantations. Second, there were more large diameter trees in restoration plantings than monoculture plantations. Third, the trees used in restoration plantings had a higher average wood density than the conifers used in monoculture plantations. While, on average, wood density was higher in mixed species plantations than restoration plantings, the much higher stocking rate of restoration plantings meant they stored more carbon than mixed species plantations. We conclude that restoration plantings in the moist tropics of north‐east Australia can accumulate relatively high amounts of carbon within two decades of establishment. Comparison with reference rainforest sites suggests that restoration plantings could maintain their high stocking rates (and therefore high biomass) as they develop in future decades. However, because restoration plantings are currently much more expensive to establish than monoculture plantations, restoration plantings are unlikely to be favoured by carbon markets. Novel reforestation techniques and designs are required if restoration plantings are to both provide habitat for rainforest biota and store carbon in biomass at a cost comparable to monoculture plantations.     

Rescue tree monocultures! A phylogenetic ecology approach to guide the choice of seedlings for enrichment planting in tropical monoculture plantations

Monoculture tree plantations have been preferred by farmers and companies because they maximize the production of specific products and are easier to manage, while multispecies plantations can provide multiple ecosystem services but have still been adopted at very limited scales. One potential midway to enhance the ecological contribution of monoculture plantations is to introduce native tree species in the plantation understory. We conducted phylogenetically explicit enrichment planting experiments beneath the monocultures of two tree species native to the Brazilian Atlantic Forest to assess if the evolutionary distance of each seedling species to the overstory tree explained variations in seedling performance. We planted in the understory of two replicate plantations of species Esenbeckia leiocarpa Engl. (Rutaceae; 38 and 25 years old) and Zeyheria tuberculosa (Vell) (Bignoniaceae; both replicates were 30 years old) six native tree species, ranging in their evolutionary distances to these overstory species. We assessed individual seedling survival and height at 1, 7, and 19 months after planting. We found a weak yet significant, positive correlation between seedling performance (growth and survival) and the phylogenetic distance of the planted seedling to the overstory species, which was driven by the poor performance of conspecifics. Seedlings of the same order and class as the overstory trees showed a significantly lower risk of death compared to conspecifics and to the furthest relative, Cryptocarya aschersoniana. These findings highlight that very close relatives to the overstories should be avoided when planting beneath monoculture plantations, but that knowledge on species‐specific requirements is still the most important for a successful planting.     

Seeding the future – the issues of supply and demand in restoration in Australia

It has been almost 15 years since concerns about the limited capacity of remnant native vegetation to supply the volumes of seed required to meet increasing restoration demands were first raised. Since that time little progress has been made towards addressing this constraint with the ongoing decline of native vegetation communities, especially since 2000, further challenging seed supply. We provide examples of the size of this demand for seed, as well as major issues associated with seed sourcing. We also discuss how invoking the concept of market forces to drive seed supply and demand is inappropriate and highlight the need for an industry body to oversee seed collection and utilisation standards. We further propose key actions that are required to secure the seed supply chain within the next 20 years to meet existing and future restoration targets. We argue that concerted, coordinated action at Commonwealth, State and regional levels are required to underpin effective future restoration outcomes.     

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.     

Recovery of floristic diversity and basal area in natural forest regeneration and planted plots in a Costa Rican wet forest

When compared to planted reforestation, natural unassisted regeneration is often reported to result in slow recovery of biomass and biodiversity, especially early in succession. In some cases, naturally regenerating forests are not comparable to the community structure of primary forests after many decades. However, direct comparison of the outcomes of tropical forest restoration and natural regeneration is hindered by differences in metrics of forest recovery, inconsistency in land use histories, and dissimilarities in experimental design. We present the results of a replicated reforestation experiment comparing natural regeneration and polyculture tree planting at multiple diversity levels (3, 6, 9, or 12 native tree species), with uniform land use history and initial edaphic conditions. We compare the recovery of basal area and floristic diversity in these treatments after 5 yr of succession. Total basal area was higher in planted plots than in naturally regenerating plots, but it but did not vary among the different planted diversity levels. The basal area of woody recruits did not differ among treatments. The diversity of woody recruits increased substantially over time but did not vary among planting treatments. Species composition trajectories showed directional turnover over time, with no consistent differences among treatments. The convergence of restoration trajectories and similarity of floristic community diversity and composition across all treatments, after only 5 yr, provides evidence of the viability of natural regeneration for rapid restoration of forest biodiversity.     

覆网保护和埋藏对坚果树种种子直播的影响

作者以都江堰林区6种代表性坚果树种为对象,研究了覆网保护和埋藏等措施对直播种子存活和幼苗建成的影响,同时调查了鼠类在春秋两季的数量变化。结果表明:秋季鼠类对种子的捕食压力大,使得种子在无保护的情况下很少存活到出苗;而秋季覆网保护(第二年春季解除覆网)和完全保护(覆网一直保留到第二年10月调查时为止)均增加了直播种子的存活和出苗率;将种子埋藏在土壤1–4cm也显著增加直播种子的存活率和出苗率,这与鼠类密度(12%)在秋季显著高于春季(2%)有直接关系。此外,坚果树种的种子存活和出苗与其种子特征有一定联系。因此,作者认为如果根据鼠类数量的季节波动规律而采取适当育林措施(如保护和埋藏等)能够增加森林种子直播的效率。     

Beyond hectares: four principles to guide reforestation in the context of tropical forest and landscape restoration

New climate change agreements emerging from the 21st Conference of the Parties and ambitious international commitments to implement forest and landscape restoration (FLR) are generating unprecedented political awareness and financial mobilization to restore forests at large scales on deforested or degraded land. Restoration interventions aim to increase functionality and resilience of landscapes, conserve biodiversity, store carbon, and mitigate effects of global climate change. We propose four principles to guide tree planting schemes focused on carbon storage and commercial forestry in the tropics in the context of FLR. These principles support activities and land uses that increase tree cover in human‐modified landscapes, while also achieving positive socioecological outcomes at local scales, in an appropriate contextualization: (1) restoration interventions should enhance and diversify local livelihoods; (2) afforestation should not replace native tropical grasslands or savanna ecosystems; (3) reforestation approaches should promote landscape heterogeneity and biological diversity; and (4) residual carbon stocks should be quantitatively and qualitatively distinguished from newly established carbon stocks. The emerging global restoration movement and its growing international support provide strong momentum for increasing tree and forest cover in mosaic landscapes. The proposed principles help to establish a platform for FLR implementation and monitoring based on a broad set of socioenvironmental benefits including, but not solely restricted, to carbon mitigation and wood production.     

Impacts of cyclone Larry on the vegetation structure of timber plantations,restoration plantings and rainforest on the Atherton Tableland,Australia

Abstract We examined the impact of severe cyclone ‘Larry’ on the vegetation structure of monoculture and mixed species timber plantations, restoration plantings and reference sites in upland rainforests on the Atherton Tableland, north Queensland, Australia. Sites were initially assessed in 2000 and resurveyed in 2006, 6–8 months after the cyclone traversed the region. In both surveys, timber plantations had a relatively open canopy, grassy understorey and few shrubs or small‐sized trees; whereas restoration plantings had a relatively closed canopy, an understorey of bare ground, leaf litter and rainforest seedlings, a high density of small‐diameter trees and a moderate representation of special life forms characteristic of rainforest. Cyclone damage varied with tree size, site type, proximity to the cyclone and stem density. First, the proportion of trees that were severely damaged by the cyclone (major branches broken, stem snapped or pushed over) increased with the diameter of trees across all site types. Second, damage to larger‐sized trees (>10 cm d.b.h., >20 cm d.b.h.) was proportionally highest in monoculture plantations, intermediate in mixed species plantations and rainforest, and lowest in restoration plantings. Third, within site types, damage levels decreased with distance from the cyclone track and with stem density. There was no evidence that topographical position influenced damage levels, at least for timber plantations. We tentatively attribute the high levels of damage experienced by timber plantations to their relatively open structure and the large size of stems in plantations. Restoration plantings generally escaped severe damage by the cyclone, but their continued development towards rainforest conditions may require a coordinated monitoring and maintenance programme to address the potential threat of weed invasion.     

大兴安岭北坡火烧迹地自然与人工干预下的植被恢复模式初探

针对大兴安岭北坡林区森林植被在不同火烧强度、火烧时间的火烧迹地上的恢复状况进行了调查研究。结果表明,轻度火烧区的植被自然更新恢复良好;中度火烧区的森林植被依靠人工促进更新要比自然更新更早达到预期目标;重度火烧区的森林植被如果完全依靠自然更新,恢复到预期目标会非常缓慢,而通过人工更新则可跨越几个演替阶段,较快接近本地的顶极群落。     

Drivers of native species regeneration in the process of restoring natural forests from mono‐specific,even‐aged tree plantations: a quantitative review

A substantial proportion of the existing tree plantations has been established following clearing of native forests. This form of conversion has become widely unaccepted, and there are increasing demands to reverse it through ecological restoration. Yet, there is a lack of integrated knowledge on how best to restore. Here, we reviewed 68 studies to identify the main factors determining establishment success of regeneration of native woody species when restoring natural forests from plantation forests using active and passive approaches, beneath existing canopies, and following their removal. According to the evidence collected, herbivory, within‐gap position, soil properties, and ground cover type and structure had limited influence on regeneration, showing significant effects in less than 26% of cases in which their influence was tested. In contrast, spatial landscape configuration, overstorey structure, ground vegetation structure, overstorey composition, and climate and geomorphology had significant effects in 67, 47, 47, 52, and 63% of cases, respectively. Regeneration diversity and abundance increased with proximity to natural vegetation remnants and seed sources. Lower canopy and understorey stocking levels positively influenced regeneration, as did interventions to reduce them. Canopy cover reduction proved especially effective in warmer regions, in stands of broadleaved species, younger ages (<30 years), higher densities (>1,000 trees/ha), and taller canopies (>20 m). Restoration of native forests can be optimized by adopting interventions that prove most effective, and prioritizing more responsive stand types. However, the specific stand attributes and environmental factors described should be further studied to understand the mechanisms underlying their influence on regeneration.     

Colonization of woody seedlings in the understory of actively and passively restored tropical moist forests

The status of woody seedling colonization gives clues about the self‐sustainability of restored forests, a tenet of restoration success. Little is known about woody seedling colonization in restored afrotropical forests. We evaluated effects of restoration methods (active vs. passive), sampling year, restoration age, and distance from old‐growth forests on seedling colonization in restored afrotropical moist forests. Seedlings were measured in 2011 and 2014 in 71 clusters of 284 permanent sampling plots (12.6 m² each) in actively (initially 3–16 years old) and 21 clusters of 63 plots in passively restored forests (initially 16 years old) in Kibale National Park, western Uganda. Seedlings were also measured in nearby old‐growth forests in three clusters of five plots in 2014. We determined species diversity, richness, abundance per plot, and species composition as measures of seedling colonization in restored and old‐growth forests. We found that diversity, richness, and abundance of seedlings were significantly higher in passively than actively restored forests. Diversity and richness but not abundance significantly increased between sampling years and with restoration age. Distance from old‐growth forests did not significantly affect diversity, richness, and abundance. Species composition of actively and passively restored forests was different from that of old‐growth forests after 19 years since restoration started. Our results show that passive restoration should be the preferred method for recovering afrotropical forests, and highlight the effect of continued management on biodiversity of restored forests.     

Performance of 11 tree species under different management treatments in restoration plantings in a tropical dry forest

Ecological restoration in tropical dry forests urgently needs to incorporate experimental evidence to increase effectiveness. The main barriers for tree establishment are adverse microenvironmental conditions and competition with exotic grasses. Therefore, management should address such barriers in order to enhance tree performance. We evaluated the effect of plastic mulching, grass removal, and no management on survival after 2 months and stem volume and canopy size after 2 years and integrated response index (IRI) in plantings of 11 native tree species with different growth rates in pastures near the tropical dry forest of Chamela, Mexico. Results revealed that: (1) initial seedling mortality was minimal in all treatments (8%) and lowest under no management (2%); (2) plastic mulching, but not grass removal, leads to increased size for most species, irrespective of their growth rank; (3) a trade‐off between initial plant survival and size after 2 years occurred due to plastic mulching; and (4) most species showed similar values of the IRI because of high survival, stem volume, or canopy cover. Grass removal decreased early survival of all species and increased stem volume only for one slow‐growing species. The use of plastic mulching increased stem volume for slow‐growing species, whereas fast‐growing species developed larger canopies with that treatment. Effects of grass removal and mulching seem to be very species‐specific and not dependent in growth rank of species, although overall mulching seems to provide better conditions for seedling performance than grass removal alone.     

Vertebrate herbivores are the main cause of seedling mortality in a logged African rainforest—implications for forest restoration

High seedling mortality can limit tropical forest regeneration after disturbance, such as selective logging. However, causes of seedling mortality are often poorly understood although this information is critical for designing appropriate restoration methods. We studied seedling mortality in a rainforest in Uganda where natural recovery has failed after selective logging carried out 40 years earlier and the sites are dominated by herbaceous vegetation. We established an experiment where seeds of 12 indigenous tree species were sown into different‐sized logging gaps in order to capture factors that cause early seedling mortality. We examined whether seedling mortality was influenced by vegetation cover (VC), vertebrate herbivores (VHs), gap size, or forest compartment and recorded causes of mortality. We found that seedling mortality was in general high but showed high tree species‐specific variation (ranging from 19 to 92% six months after emergence). VHs, particularly rodents, were the most important cause of mortality. Rotting caused mortality particularly in smaller gaps, but elephant trampling, drying, being washed away by rain and falling debris were only minor mortality causes. Dense VC was not a considerable barrier to seedling survival, but by providing shelter for rodents, it indirectly increased seedling herbivory. Our results show that high seedling mortality on logged sites can limit forest regeneration and rodents in particular can be a significant factor causing high seedling mortality.     

Natural forest regeneration and ecological restoration in human‐modified tropical landscapes

In human‐modified tropical landscapes (HMLs) the conservation of biodiversity, functions and services of forest ecosystems depends on persistence of old growth forest remnants, forest regeneration in abandoned agricultural fields, and restoration of degraded lands. Understanding the impacts of agricultural land uses (ALUs) on forest regeneration is critical for biodiversity conservation in HMLs. Here, we develop a conceptual framework that considers the availability of propagules and the environment prevailing after field abandonment as two major determinants of forest regeneration in HMLs. The framework proposes that regeneration potential decreases with size, duration and severity of agricultural disturbance, reducing propagule availability and creating ill‐suited environmental conditions for regeneration. We used studies from Southern Mexico to assess this framework. First, we identify regeneration bottlenecks that trees face during transit from seed to follow‐up life stages, using demographic analysis of dominant pioneer species in recently abandoned fields. Then, we explore effects of ALUs on forest regeneration at the field and landscape scales, addressing major legacies. Finally, we integrate agricultural disturbance with landscape composition to predict attributes of successful second growth forests in HMLs, and provide indicators useful to select tree native species for active restoration. An indicator of disturbance inflicted by ALUs, based on farmers’ information, predicted better regeneration potential than measurements of soil and microclimate conditions at time of abandonment. Cover of cattle pastures in the landscape was a stronger indicator of forest regenerating attributes than cover of old growth forest remnants. To conclude, we offer recommendations to promote forest regeneration and biodiversity conservation in HMLs.     

Restoring the Victorian Western (Basalt) Plains grassland. 1. Laboratory trials of viability and germination, and the implications for direct seeding

Summary   The Victorian Western (Basalt) Plains grassland is one of Australia's most threatened plant communities. Practitioners using seed for its restoration need to know whether seed can be sown fresh or whether it requires an after-ripening period. This study assessed the viability and germination of freshly harvested wild seed from 64 grassland species indigenous to the Basalt Plains of western Victoria. The seed was collected as part of a broader experiment that examined the potential of direct-sown complex seed mixes for the restoration of grassland communities. The germination of fresh seed at 25°C varied widely between species. Comparisons with tetrazolium viability tests for each species indicated varying levels of dormancy within the species pool. Germination separated into three broad responses at day 28. One-third of the species failed to germinate, one-third germinated at 1% to 50% and the remaining species germinated between 51% and 100%. Therefore, if the aim of a sowing was the rapid and synchronous establishment of most of the sown species, the use of fresh seed in restoration could be problematic. After 3 months of dry storage, eight species were re-tested for germination. Each of the selected species had shown high viability but low initial germination. Only two species significantly increased their total germination at 25°C. The annual species, Triptilodiscus pygmaeus , increased its total germination from 6% as fresh seed to 99% after dry storage. Testing the viability and germination capacity of freshly harvested seed from a large and diverse sample of native grassland species demonstrated that many of the species were unlikely to germinate rapidly or synchronously when sown in complex seed mixes soon after harvest. This finding has implications for the scheduling and management of restoration projects that rely on the use of such seed.