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One-third of all Neotropical forests are secondary forests that regrow naturally after agricultural use through secondary succession. We need to understand better how and why succession varies across environmental gradients and broad geographic scales. Here, we analyze functional recovery using community data on seven plant characteristics (traits) of 1,016 forest plots from 30 chronosequence sites across the Neotropics. By analyzing communities in terms of their traits, we enhance understanding of the mechanisms of succession, assess ecosystem recovery, and use these insights to propose successful forest restoration strategies. Wet and dry forests diverged markedly for several traits that increase growth rate in wet forests but come at the expense of reduced drought tolerance, delay, or avoidance, which is important in seasonally dry forests. Dry and wet forests showed different successional pathways for several traits. In dry forests, species turnover is driven by drought tolerance traits that are important early in succession and in wet forests by shade tolerance traits that are important later in succession. In both forests, deciduous and compound-leaved trees decreased with forest age, probably because microclimatic conditions became less hot and dry. Our results suggest that climatic water availability drives functional recovery by influencing the start and trajectory of succession, resulting in a convergence of community trait values with forest age when vegetation cover builds up. Within plots, the range in functional trait values increased with age. Based on the observed successional trait changes, we indicate the consequences for carbon and nutrient cycling and propose an ecologically sound strategy to improve forest restoration success., Competing Interests: The authors declare no competing interest.

Tropical forests are converted at an alarming rate for agricultural use and pastureland, but also regrow naturally through secondary succession. For successful forest restoration, it is essential to understand the mechanisms of secondary succession. These mechanisms may vary across forest types, but analyses across broad spatial scales are lacking. Here, we analyse forest recovery using 1,403 plots that differ in age since agricultural abandonment from 50 sites across the Neotropics. We analyse changes in community composition using species-specific stem wood density (WD), which is a key trait for plant growth, survival and forest carbon storage. In wet forest, succession proceeds from low towards high community WD (acquisitive towards conservative trait values), in line with standard successional theory. However, in dry forest, succession proceeds from high towards low community WD (conservative towards acquisitive trait values), probably because high WD reflects drought tolerance in harsh early successional environments. Dry season intensity drives WD recovery by influencing the start and trajectory of succession, resulting in convergence of the community WD over time as vegetation cover builds up. These ecological insights can be used to improve species selection for reforestation. Reforestation species selected to establish a first protective canopy layer should, among other criteria, ideally have a similar WD to the early successional communities that dominate under the prevailing macroclimatic conditions.

Old-growth tropical forests harbor an immense diversity of tree species but are rapidly being cleared, while secondary forests that regrow on abandoned agricultural lands increase in extent. We assess how tree species richness and composition recover during secondary succession across gradients in environmental conditions and anthropogenic disturbance in an unprecedented multisite analysis for the Neotropics. Secondary forests recover remarkably fast in species richness but slowly in species composition. Secondary forests take a median time of five decades to recover the species richness of old-growth forest (80% recovery after 20 years) based on rarefaction analysis. Full recovery of species composition takes centuries (only 34% recovery after 20 years). A dual strategy that maintains both old-growth forests and species-rich secondary forests is therefore crucial for biodiversity conservation in human-modified tropical landscapes.

The nutrient demands of regrowing tropical forests are partly satisfied by nitrogen-fixing legume trees, but our understanding of the abundance of those species is biased towards wet tropical regions. Here we show how the abundance of Leguminosae is affected by both recovery from disturbance and large-scale rainfall gradients through a synthesis of forest inventory plots from a network of 42 Neotropical forest chronosequences. During the first three decades of natural forest regeneration, legume basal area is twice as high in dry compared with wet secondary forests. The tremendous ecological success of legumes in recently disturbed, water-limited forests is likely to be related to both their reduced leaflet size and ability to fix N 2 , which together enhance legume drought tolerance and water-use efficiency. Earth system models should incorporate these large-scale successional and climatic patterns of legume dominance to provide more accurate estimates of the maximum potential for natural nitrogen fixation across tropical forests.

Regrowth of tropical secondary forests following complete or nearly complete removal of forest vegetation actively stores carbon in aboveground biomass, partially counterbalancing carbon emissions from deforestation, forest degradation, burning of fossil fuels, and other anthropogenic sources. We estimate the age and spatial extent of lowland second-growth forests in the Latin American tropics and model their potential aboveground carbon accumulation over four decades. Our model shows that, in 2008, second-growth forests (1 to 60 years old) covered 2.4 million km(2) of land (28.1% of the total study area). Over 40 years, these lands can potentially accumulate a total aboveground carbon stock of 8.48 Pg C (petagrams of carbon) in aboveground biomass via low-cost natural regeneration or assisted regeneration, corresponding to a total CO2 sequestration of 31.09 Pg CO2. This total is equivalent to carbon emissions from fossil fuel use and industrial processes in all of Latin America and the Caribbean from 1993 to 2014. Ten countries account for 95% of this carbon storage potential, led by Brazil, Colombia, Mexico, and Venezuela. We model future land-use scenarios to guide national carbon mitigation policies. Permitting natural regeneration on 40% of lowland pastures potentially stores an additional 2.0 Pg C over 40 years. Our study provides information and maps to guide national-level forest-based carbon mitigation plans on the basis of estimated rates of natural regeneration and pasture abandonment. Coupled with avoided deforestation and sustainable forest management, natural regeneration of second-growth forests provides a low-cost mechanism that yields a high carbon sequestration potential with multiple benefits for biodiversity and ecosystem services.

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha(-1)), corresponding to a net carbon uptake of 3.05 Mg C ha(-1) yr(-1), 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha(-1)) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000-1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.

Habitat-specialist species may be restricted to a narrower range of microhabitats than habitat-generalist species. We addressed this hypothesis by comparing microhabitats of two pairs of congeners that differ in habitat specificity and co-occur in one distinct habitat type, Florida rosemary scrub. We characterized microhabitats of rosemary scrub specialists, Polygonella basiramia and Lechea cernua, their habitat-generalist congeners, Polygonella robusta and Lechea deckertii, and random points in the rosemary scrub habitat. Plants of both habitat specialists occurred in microhabitats with significantly more bare sand than plants of habitat-generalist species and random points. Plants of all four species occurred in microhabitats that were farther from dominant shrubs, Ceratiola and Quercus spp., than random points. Seedlings of both habitat specialists grew larger in bare sand microhabitats, whereas ground lichens and litter did not affect seedling growth of the habitat generalists. As the time since fire increases, bare sand cover decreases, Ceratiola density increases, Quercus density remains constant, and shrubs become taller. Physical characteristics, such as soil temperature, soil carbon, and soil moisture, differ slightly with respect to microhabitat. Our results suggest that P. basiramia and L. cernua are specialized on bare sand microhabitats that characterize their preferred habitat, rosemary scrub. Microhabitat specialization may limit the distribution of these rare species.

We tested the hypothesis that the tropical shrub Clidemia hirta appears more shade tolerant and is more abundant in its introduced than native range because of genetic differences in resource acquisition, allocation, and phenotypic plasticity between native and introduced genotypes. We examined growth, biomass allocation, and photosynthetic parameters of C. hirta grown in a greenhouse from seed collected from four populations in part of its native range (Costa Rica) and four populations in part of its introduced range (Hawaiian Islands). Six-month-old seedlings were placed in high (10.3-13.9 mol m(-2) day(-1)) or low (1.4-4.5 mol m(-2) day(-1)) light treatments and grown for an additional 6 months. Our study provided little evidence that Hawaiian genotypes of C. hirta differed genetically from Costa Rican genotypes in ways that would contribute to differences in habitat distribution or abundance. Some of the genetic differences that were apparent, such as greater allocation to stems and leaf area relative to whole plant biomass in Costa Rican genotypes and greater allocation to roots in Hawaiian genotypes, were contrary to predictions that genotypes from the introduced range would allocate more biomass to growth and less to storage than those from the native range. Hawaiian and Costa Rican genotypes displayed no significant differences in relative growth rates, maximal photosynthetic rates, or specific leaf areas in either light treatment. In the high light environment, however, Hawaiian genotypes allocated more biomass to reproductive parts than Costa Rican genotypes. Phenotypic plasticity for only 1 of 12 morphological and photosynthetic variables was greater for Hawaiian than Costa Rican genotypes. We conclude that genetic shifts in resource use, resource allocation, or plasticity do not contribute to differences in habitat distribution and abundance between the native and introduced ranges of C. hirta.

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A max , stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.

Six speices of shrubs and one large herb with contrasting life history patterns were used as bioassays of nutrient availability in a Costa Rican lowland rain forest soil. Growth responses of the herb (Phytolacca rivinoides, Phytolaccaceae) confirmed soil measurements indicating high availability of N and potentially limiting levels of P, K, Mg and Ca. Growth responses of the shrub species (Miconia spp., Melastomataceae and Piper spp., Piperaceae) to a complete nutrient fertilizer were generally less than that of Phytolacca. Lack of a strong shrub response to +P fertilization is probably due to mycorrhizal associations and slower growth rates of woody species. In general, increased growth did not occur at the expense of phenolic production in the leaves. The results emphasize that assessment of specific nutrient limitations to plant growth vary depending on species selected for the bioassay, even among species from the same community.

In a series of field experiments using Costa Rican rain forest plants, we examined the effect of accessibility on fruit removal rates. We compared the effects of fruit placement in terminal and axillary infructescences on diurnal and nocturnal removal rates, visitation rates, and incidence of fruit damage. We used three different species of berries (Phytolacca rivinoides, Psychotria brachiata, and Psychotria pitteri) and worked in three different habitats (fallow fields, treefall gaps, and forest understory) and in two different seasons (July-September, a season of fruit abundance and December-January, a season of fruit scarcity.)We found that in oldfields especially, diurnal removal rates by birds were significantly greater from axillary than from terminal infructescences. Nocturnal removal from axillary infructescences-presumably by rodents-is also occassionally significant. From these data, from observations on climbing ability and fruit use in captive rodents, and from reports in the literature, we suggest that rodents are significant sources of fruit and seed loss in tropical shrubs. We hypothesize that placement of the infructescence on the plant affects fruit removal by both seed-dispersing birds and by the less agile, often seed-destroying rodents. The balance between the two rates is an important component of a plant's dispersal success.Diurnal fruit removal rates were higher during the season of fruit scarcity than during the season of fruit abundance and higher in old fields than in forest gaps or understory. Fruit damage rates-probably due to orthopterans-were slightly greater in gaps and understory than in old fields.

I suggest that between-community variations in diversity patterns during succession in plant communities are due to the effects of selection on life history strategies under different disturbance regimes. Natural disturbances to plant communities are simultaneously a source of mortality for some individuals and a source of establishment sites for others. The plant community consists of a mosaic of disturbance patches (gaps) of different environmental conditions. The composition of the mosaic is described by the size-frequency distribution of the gaps and is dependent on the rates and scales of disturbance. The life-history strategies of plant species dependent on some form of disturbance for establishment of propagules should reflect this size-frequency distribution of disturbance patches. An extension of island biogeographic theory to encompass relative habitat area predicts that a community should be most rich in species adapted to growth and establishment in the spatially most common patch types. Changes in species diversity during succession following large scale disturbance reflect the prevalent life history patterns under historically common disturbance regimes. Communities in which the greatest patch area is in large-scale clearings (e.g. following fire) are most diverse in species establishing seedlings in xeric, high light conditions. Species diversity decreases during succession. Communities in which such large patches are rare are characterized by a large number of species that reach the canopy through small gaps and realtively few which regenerate in the large clearings. Diversity increases during succession following a large scale disturbance.Evidence from communities characterized by different disturbance regimes is summarized from the literature. This hypothesis provides an evolutionary mechanism with which to examine the changes in plant community structure during succession. Diversity peaks occurring at "intermediate levels" of disturbance as discussed by Connell and Huston are interpreted in this context.

Physician distribution continues as a major national issue despite the projected oversupply of physicians by 1990. Kirksville College of Osteopathic Medicine (KCOM) in Kirksville, Missouri, has a high percentage of its graduates going into rural primary care. In this study of physicians who graduated from KCOM from 1930 to 1974, the authors sought to identify the factors influencing physicians to select rural primary care. The size of the physician's hometown, KCOM curricular experiences, and faculty role models were the most important factors influencing a physician to select rural primary care. While these findings are similar to other studies, this is the first to examine osteopathic medicine.