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.

Early recruitment dynamics in tropical restoration.

Unassisted secondary succession in abandoned tropical pastures often results in species-poor forests of pioneer trees that persist for decades. We characterize recruitment rates of woody vegetation in planting treatments during the first 60 months of experimental restoration on thin, eroded soils at Los Tuxtlas, southern Mexico. We test the hypothesis that recruitment of later-successional trees is greater in fenced plots planted with native trees than in fenced controls that simulate natural succession, and further that recruitment of such species would be greater in plots planted with animal-dispersed trees than in those planted with wind-dispersed trees. Results indicated much greater recruitment of later-successional animal-dispersed trees in planted plots as compared with controls. Three censuses per year recorded 960 recruited individuals of 44 species of trees and shrubs from 20-60 months after cattle exclusion. Ninety-six percent of recruits were not of planted species. Repeated-measures analyses of variance indicated that recruited communities included more species of pioneers than of later-successional trees and shrubs, with more individuals and species dispersed by animals than by wind. Recruitment of pioneers did not differ between control and planted plots. Later-successional recruits dispersed by animals accumulated > 10 times faster in planted than control plots, with apparent acceleration after planted Cecropia obtusifolia and Ficus yoponensis first produced fleshy fruits 48 months after cattle exclusion. Sparse later-successional wind-dispersed recruits did not differ by treatment. Our preliminary results over the first five years after cattle exclusion indicate that planted stands clearly accelerate succession through accumulation of later-successional trees and shrubs dispersed by animals.

Challenges during the execution, results, and monitoring phases of ecological restoration: Learning from a country-wide assessment.

Outcomes from restoration projects are often difficult for policymakers and stakeholders to assess, but this information is fundamental for scaling up ecological restoration actions. We evaluated technical aspects of the interventions, results (ecological and socio-economic) and monitoring practices in 75 restoration projects in Mexico using a digital survey composed of 137 questions. We found that restoration projects in terrestrial ecosystems generally relied on actions included in minimal (97%) and maximal (86%) intervention, while in wetlands, the preferred restoration strategies were intermediate (75%) and minimal intervention (63%). Only a third of the projects (38%) relied on collective learning as a source of knowledge to generate techniques (traditional management). In most of the projects (73%), multiple criteria (>2) were considered when selecting plant species for plantings; the most frequently used criterion was that plant species were found within the restoration area, native or naturalized (i.e., a circa situm criterion; 88%). In 48% of the projects, the biological material required for restoration (e.g., seeds and seedlings) were gathered or propagated by project implementers rather than purchased commercially. Only a few projects (between 33 and 34%) reached a high level of biodiversity recovery (>75%). Most of the projects (between 69 to71%) recovered less than 50% of the ecological services. Most of the projects (82%) led to improved individual relationships. The analysis revealed a need to implement strategies that are cost-effective, the application of traditional ecological knowledge and the inclusion of indigenous people and local communities in restoration programs at all stages-from planning to implementation, through monitoring. We also identified the need to expand research to develop effective tools to assess ecosystems' regeneration potential and develop theoretical frameworks to move beyond short-term markers to set and achieve medium- and long-term goals. Cautious and comprehensive planning of national strategies must consider the abovementioned identified gaps.

Seedling recruitment under isolated trees in a tea plantation provides a template for forest restoration in eastern Africa.

Natural regeneration is less expensive than tree planting, but determining what species will arrive and establish to serve as templates for tropical forest restoration remains poorly investigated in eastern Africa. This study summarises seedling recruitment under 29 isolated legacy trees (14 trees comprised of three exotic species and 15 trees comprised of seven native species) in tea plantations in the East Usambara Mountains, Tanzania. Among the findings were that pioneer recruits were very abundant whereas non-pioneers were disproportionately fewer. Importantly, 98% of all recruits were animal-dispersed. The size of legacy trees, driven mostly by the exotic Grevillea robusta, and to some extent, the native Milicia excelsa, explained abundance of recruits. The distribution of bird-dispersed recruits suggested that some bird species use all types of legacy trees equally in this fragmented landscape. In contrast, the distribution of bat-dispersed recruits provided strong evidence that seedling composition differed under native versus exotic legacy trees likely due to fruit bats showing more preference for native legacy trees. Native, as compared to exotic legacy trees, had almost two times more non-pioneer recruits, with Ficus and Milicia excelsa driving this trend. Implications of our findings regarding restoration in the tropics are numerous for the movement of native animal-dispersed tree species in fragmented and disturbed tropical forests surrounded by farmland. Isolated native trees that bear fleshy fruits can attract more frugivores, resulting not only in high recruitment under them, but depending on the dispersal mode of the legacy trees, also different suites of recruited species. When selecting tree species for plantings, to maximize visitation by different dispersal agents and to enhance seedling recruit diversity, bat-dispersed Milicia excelsa and Ficus species are recommended.

Early seed fall and seedling emergence: precursors to tropical restoration.

We explore processes of seed immigration and seedling recruitment before an experimental rainforest restoration matures enough to affect either. Twenty-four 30 × 30-m plots were fenced in 12 ha of pasture in 2006. Seeds were collected in ninety-six 1-m(-2) seed traps; recruits were censused in ~12,000 m(2) in which establishment was allowed. We tested effects of distance from forest, living trees, and stumps of trees cut during site preparation on seed rain in 2007 and effects of these and soil depth on recruits through June 2008. Seed fall and recruitment were not correlated with distance to forest 90-400 m away, nor to living shade trees outside the 160 × 485-m experimental grid. Recruitment differed for animal- and wind-dispersed species in a topographically complex landscape. Recruitment of wind-dispersed species was random with respect to soil depth or distance to recent stumps. Recruitment of animal-dispersed species was multimodal; partial correlations with number of stumps within 30 m of plots were significant with soil depth held constant (P < 0.025), as were correlations of recruitment with soil depth with number of stumps held constant (P < 0.01). Animal-dispersed recruits were often not conspecifics of adults that had been cut, indicating a legacy of attraction by fruiting trees of animals bearing seeds from distant sources. Ecological implications are that recruitment in pastures released from grazing reflects a mix of widely scattered wind-dispersed pioneers and, where animal-dispersed trees exist, multi-modal and decidedly non-random recruitment of pioneer and later successional animal-dispersed trees from seed banks.

Effect of initial soil properties on six-year growth of 15 tree species in tropical restoration plantings.

In restoration plantings in degraded pastures, initial soil nutrient status may lead to differential growth of tropical tree species with diverse life history attributes and capacity for N2 fixation. In 2006, we planted 1,440 seedlings of 15 native tree species in 16 fenced plots (30 × 30 m) in a 60-year-old pasture in Los Tuxtlas, Veracruz, Mexico, in two planting combinations. In the first year, we evaluated bulk density, pH, the concentration of organic carbon (C), total nitrogen (N), ammonia (NO3-), nitrate (NH4+), and total phosphorus (P) in the upper soil profile (0-20 cm in depth) of all plots. The first two axes of two principal component analyses explained more than 60% of the variation in soil variables: The axes were related to increasing bulk density, NO3-, NH4+, total N concentration, and pH. Average relative growth rates in diameter at the stem base of the juvenile trees after 6 years were higher for pioneer (45.7%) and N2-fixing species (47.6%) than for nonpioneer (34.7%) and nonfixing species (36.2%). Most N2-fixing species and those with the slowest growth rates did not respond to soil attributes. Tree species benefited from higher pH levels and existing litter biomass. The pioneers Ficus yoponensis, Cecropia obtusifolia, and Heliocarpus appendiculatus, and the N2-fixing nonpioneers Cojoba arborea, Inga sinacae, and Platymiscium dimorphandrum were promising for forest restoration on our site, given their high growth rates.

Coordinating ecological restoration options analysis and risk assessment to improve environmental outcomes.

Ecological risk assessment as currently practiced has hindered consideration of ecosystem services endpoints and restoration goals in the environmental management process. Practitioners have created barriers between procedures to clean up contaminated areas and efforts to restore ecosystem functions. In this article, we examine linkages between contaminant risk assessment approaches and restoration efforts with the aim of identifying ways to improve environmental outcomes. We advocate that project managers and other stakeholders use an ecological planning framework, with restoration options included upfront in the risk assessment. We also considered the opportunities to incorporate ecosystem services as potential assessment endpoints in the Problem Formulation stages of a risk assessment. Indeed, diverse perspectives of stakeholders are central to understand the relevance of social, cultural, economic, and regional ecology as influences on future use options for the landscape being restored. The measurement endpoints used to characterize the existing ecological conditions for selected ecosystem services can also be used to evaluate restoration success. A regional, landscape, or seascape focus is needed throughout the risk assessment process, so that restoration efforts play a more prominent role in enhancing ecosystem services. In short, we suggest that practitioners begin with the question of "how can the ecological risk assessment inform the decision on how best to restore the ecosystem?"

Effect of Ecological Restoration on Body Condition of a Predator.

Ecological restoration attempts to recover the structure and function of ecosystems that have been degraded by human activities. A crucial test of ecosystem recovery would be to determine whether individuals in restored environments are as healthy as those in conserved environments. However, the impact of restoration on physiology of terrestrial animals has never been tested. Here, we evaluated the effect of two restoration methods on body condition measured as body size, body mass, lipid and muscle content of the spider Nephila clavipes in a tropical dry forest that has suffered chronic disturbance due to cattle grazing. We used experimental plots that had been excluded from disturbance by cattle grazing during eight years. Plots were either planted with native trees (i. e. maximal intervention), or only excluded from disturbance (i. e. minimal intervention), and were compared with control conserved (remnants of original forest) and disturbed plots (where cattle is allowed to graze). We predicted (1) better body condition in spiders of conserved and restored sites, compared to disturbed sites, and (2) better body condition in plots with maximal intervention than in plots with minimal intervention. The first prediction was not supported in males or females, and the second prediction was only supported in females: body dry mass was higher in planted than in conserved plots for spiders of both sexes and also higher that in disturbed plots for males, suggesting that plantings are providing more resources. We discuss how different life histories and environmental pressures, such as food availability, parasitism, and competition for resources can explain our contrasting findings in male and female spiders. By studying animal physiology in restoration experiments it is possible to understand the mechanistic basis of ecological and evolutionary processes that determine success of ecological restoration.

Recovering more than tree cover: herbivores and herbivory in a restored tropical dry forest.

Intense and chronic disturbance may arrest natural succession, reduce environmental quality and lead to ecological interaction losses. Where natural succession does not occur, ecological restoration aims to accelerate this process. While plant establishment and diversity is promoted by restoration, few studies have evaluated the effect of restoration activities on ecological processes and animal diversity. This study assessed herbivory and lepidopteran diversity associated with two pioneer tree species growing in 4-year-old experimental restoration plots in a tropical dry forest at Sierra de Huautla, in Morelos, Mexico. The study was carried out during the rainy season of 2010 (July-October) in eleven 50 x 50 m plots in three different habitats: cattle-excluded, cattle-excluded with restoration plantings, and cattle grazing plots. At the beginning of the rainy season, 10 juveniles of Heliocarpus pallidus (Malvaceae) and Ipomoea pauciflora (Convolvulaceae) were selected in each plot (N = 110 trees). Herbivory was measured in 10 leaves per plant at the end of the rainy season. To evaluate richness and abundance of lepidopteran larvae, all plants were surveyed monthly. Herbivory was similar among habitats and I. pauciflora showed a higher percentage of herbivory. A total of 868 lepidopteran larvae from 65 morphospecies were recorded. The family with the highest number of morphospecies (9 sp.) was Geometridae, while the most abundant family was Saturnidae, with 427 individuals. Lepidopteran richness and abundance were significantly higher in H. pallidus than in I. pauciflora. Lepidopteran richness was significantly higher in the cattle-excluded plots, while abundance was significantly higher in the non-excluded plots. After four years of cattle exclusion and the establishment of plantings, lepidopteran richness increased 20 -fold in the excluded plots compared to the disturbed areas, whereas herbivory levels were equally high in both restored and disturbed sites. Restoration with plantings and exclusion of cattle and plantings was shown to be a successful strategy for attracting lepidopterans and cattle exclusion was the main factor explaining lepidopteran diversity.

Roles of birds and bats in early tropical-forest restoration.

Restoration of tropical forest depended in large part on seed dispersal by fruit-eating animals that transported seeds into planted forest patches. We tested effectiveness of dispersal agents as revealed by established recruits of tree and shrub species that bore seeds dispersed by birds, bats, or both. We documented restoration of dispersal processes over the first 76 months of experimental restoration in southern Mexico. Mixed-model repeated-measures randomized-block ANOVAs of seedlings recruited into experimental controls and mixed-species plantings from late-secondary and mature forest indicated that bats and birds played different roles in the first years of a restoration process. Bats dispersed pioneer tree and shrub species to slowly regenerating grassy areas, while birds mediated recruitment of later-successional species into planted stands of trees and to a lesser extent into controls. Of species of pioneer trees and shrubs established in plots, seven were primarily dispersed by birds, three by bats and four by both birds and bats. Of later-successional species recruited past the seedling stage, 13 were of species primarily dispersed by birds, and six were of species dispersed by both birds and bats. No later-successional species primarily dispersed by bats established in control or planted plots. Establishment of recruited seedlings was ten-fold higher under cover of planted trees than in grassy controls. Even pre-reproductive trees drew fruit-eating birds and the seeds that they carried from nearby forest, and provided conditions for establishment of shade-tolerant tree species. Overall, after 76 months of cattle exclusion, 94% of the recruited shrubs and trees in experimental plots were of species that we did not plant.

Planting densities and bird and rodent absence affect size distributions of four dicots in synthetic tallgrass communities.

Variability in the size distributions of populations is usually studied in monocultures or in mixed plantings of two species. Variability of size distributions of populations in more complex communities has been neglected. The effects of seeding density (35 or 350 seeds/species/m2) and presence of small vertebrates on the variability of size distributions were studied for a total of 1,920 individuals of 4 species in replicated synthetic communities of 18 species in northern Illinois. End-of season height and above-ground biomass were measured for prairie perennials Dalea purpurea (purple prairie clover), Echinacea purpurea (purple coneflower), Desmanthus illinoensis (Illinois bundleflower) and Heliopsis helianthoides (early sunflower). Variability in biomass distribution of the four target species was twice as great at low than at high densities when small vertebrates were excluded. Our results suggest that inter- and intraspecific competition may affect all individuals more under high-density conditions, thereby reducing the variability in their biomass distributions within this community. This result, a consequence of plant-plant interaction, is obscured when small birds or mammals are present, presumably because either or both add variance that overwhelms the pattern.