Emerging forests on abandoned land: Puerto Rico’s new forests

The species composition of forests change continuously as the earth’s biota evolves and adjusts to environmental change. Humans are accelerating the rate of species turnover by moving species around the planet and dramatically changing environmental conditions. Our focus is on new forests in Puerto Rico that emerge naturally on abandoned lands previously converted to agriculture and degraded. These forest stands have combinations of species that are new to the island’s landscapes. New forests exhibit high species dominance during forest establishment, which includes dominance by alien tree species. These alien tree species establish and maintain forest cover, which may facilitate regeneration of native tree species. Landscape analysis and literature review revealed that these emerging stands are highly fragmented (60% were <1 ha in 1991), function as refugia for native organisms, and at 60–80 years old have similar species richness and structural features as native stands of similar age. However, the island’s new forests exhibit important differences from mature native forests on unconverted forestlands. New forests have fewer endemic species and fewer large trees (≥55 cm dbh) than mature native forests; they have higher soil bulk density and lower soil carbon and litter stocks; and they accumulate aboveground biomass, basal area, and soil carbon more slowly than native forests of similar age. We suggest that new forests will become increasingly prevalent in the biosphere in response to novel environmental conditions introduced to the planet by humans.     

热带次生林经营研究文献分析

热带次生林面积巨大且在热带森林资源中的作用越来越重要,因而拥有巨大的经营潜力和生态功能。次生林演替过程中的主要研究成果如下：土壤贮存的种子对次生林的再生非常重要,光投射的强度增加和温度提高将刺激种子萌芽,而残余植被群落强烈影响次生林种子的散布,食种子动物影响次生林的成林;次生林群落植物种类数量一般随演替的进程而增加;随着演替的进程,次生林内阳生树种趋于衰亡,耐荫树种不断增加;次生林的树种数量能接近原生林水平的时间过程更随着林分种类、过去土地的利用强度和利用种类,以及环境条件的不同而异;在一定环境中,人工林树种通过人工林树种改变光、温度和土壤表层的湿度,能够促进发芽和种子生长,改善林地和林下植被的状况,从而加速次生林演替。大多数食草动物喜欢在次生林中取食,因为次生林中先锋树种很少或没有机械和化学保护,且林内许多树种产生大量可食用的果实;环境和火对次生林演替有一定影响;在早期的演替过程中,更多的生物量分配到资源需求旺盛的组织（如叶和细根）,在演替晚期阶段,更多的生物量则分配到结构组织中（如木质部和根端）。次生林在15至20年内能迅速积累生物量,然后积累逐渐减慢;次生林被砍伐或火烧后,森林土壤的结构被破坏,导致土壤有机质和氮减少。     

Structure and species composition of novel forests dominated by an introduced species in northcentral Puerto Rico

The African tulip tree, Spathodea campanulata Beauv., is an introduced species forming novel forest types in Puerto Rico. These forests develop naturally after deforestation, agricultural use and land abandonment, and there are many questions as to their ecological characteristics. We sampled structure and species composition of large, small, and juvenile trees (≥10, ≥2.5 to <10, and <2.5 cm diameter at breast height, respectively) in nine secondary forests dominated by S. campanulata on alluvial, karst, and volcanic substrates in northcentral Puerto Rico. No differences were found in S. campanulata forest structure between substrates. Of a total of 79 species found, 17 were introduced. Forests on karst and alluvium had the highest and lowest global species richness, respectively. Species richness increased from large to small to juvenile trees in most sites, but more so on karst. The percentage of introduced species was inversely related to species richness of tree size classes on all substrate types. The dominance of S. campanulata in the large tree size class was highest and lowest in alluvial and volcanic sites, respectively, and decreased from large to small to juvenile trees on all substrate types. Species richness of S. campanulata forests is lower than that of native forests on equivalent substrates. Although land use history affects composition, the juvenile tree species established in S. campanulata forests seems to correspond to the geological substrate were sites are found. This study shows S. campanulata restores forest structure and native tree species on abandoned agriculture and grazing lands in Puerto Rico. Management of these novel forests should consider them as resources were natural processes have the potential of making them more diverse.     

Restoration of dry tropical forests in Central America: A review of pattern and process

Much information on restoration and management exists for wet tropical forests of Central America but comparatively little work has been done in the dry forests of this region. Such information is critical for reforestation efforts that are now occurring throughout Central America. This paper describes processes of degradation due to land use and provides a conceptual framework for the restoration of dry tropical forest. Most of this forest type was initially harvested for timber and then cleared for cattle in the last century (1930-1970). Only 1.7% remains largely restricted to infertile soils and remote areas on the Pacific coastal side of Panama, Costa Rica, Nicaragua and Mexico. These cleared areas are again in a state of transition due to a combination of decreasing land productivity, and land speculation for tourism development. Some farms have been sold to new landowners who are interested in reforesting to increase biodiversity and forest cover. Attempts have therefore been made to reforest by protecting the land from fire and cattle, by supplementing natural regrowth with enrichment planting, or through use of tree plantations. Experimental studies have demonstrated the ability of these lands to grow back to forests because of native species ability to sprout after cutting, and the capacity of remnant trees in field and riparian zones to provide seeds and to moderate edge environment for seed germination and seedling establishment. However, research also shows that on sites with long histories of land clearance, species diversity will remain low with functional groups missing unless some active management occurs. Under-planting with late-successional native tree species can add structure and diversity; enrichment planting with large-fruited shade-intolerant species can initiate new islands of more diverse regeneration beneath their canopies; and plantings of fast-growing, nitrogen-fixing trees that provide light canopy shade can moderate the environment below, promoting regeneration establishment of late-successional species. Plantations are the only option for lands that have lost almost all remnants of native forest, and where soils and vegetation have changed to new states of structure and function. Conversion of pastures to tree plantations that can facilitate natural regeneration beneath them is appropriate when pastures are prone to fire and/or lack immediate seed sources nearby. After the grasses have been shaded out, natural recruitment can slowly occur over a 10-15 years period. Under-planting of shade-tolerant late-successional species can supplement species composition and structure.     

Past land-use and ecological resilience in a lowland Brazilian Atlantic Forest: implications for passive restoration

The ecological resilience of abandoned pastures has important implications for the potential of passive restoration in the Brazilian Atlantic Forest, especially in the face of the ambitious restoration targets for this biodiversity hotspot. In the present study, we aimed to investigate the influence of past land-use on natural regeneration in two secondary forests that established on abandoned pastures with different land use histories (Abandoned + Fire—15 years since the last fire event and Abandoned—36 years ago) in a lowland Brazilian Atlantic Forest. Both secondary forests exhibit low resilience for the three analyzed attributes: vegetation structure, richness, and species composition. Secondary forests affected by fire have the lowest values for species richness, aboveground biomass and leaf area index compared to old-growth forests. Secondary forests were dominated by Moquiniastrum polymorphum—Asteraceae, which was absent from the old-growth forests. Fire resistance, capacity to establish in nutrient-poor soils and efficient seed dispersal in human-modified landscapes are important attributes that may explain the dominance of this species. We demonstrate that even in sites considered appropriate for passive restoration a priori (e.g. located in a protected area, 36 years since abandonment and close to old-growth forests), secondary forests exhibited low aboveground biomass and communities were dominated by few species. Therefore, our results highlight the effects of past land-use on natural regeneration and suggest the necessity to use interventions such as enrichment plantings, nucleation techniques and assisted natural regeneration in order to accelerate forest restoration.     

Allometry, biomass, and chemical content of Novel African Tulip Tree (Spathodea campanulata) Forests in Puerto Rico

The African tulip tree, Spathodea campanulata, the most common tree in Puerto Rico, forms novel forest types with mixtures of native and other introduced tree species. Novel forests increase in area in response to human activity and there is no information about their biomass accumulation and nutrient cycling. We established allometric relationships and chemically analyzed plant parts of African tulip trees to determine the concentration and standing stock of chemical elements (C, N, P, K, Ca, Mg, S, Mn, Al, Fe, Na), and ash. Trees ranged in diameter at breast height from 8 to 85 cm and in height from 8.8 to 28 m. The concentrations of N, P, K, and Ca in leaves of the African tulip tree were similar to those of the native pioneer Cecropia schreberiana and higher than those of mature forest tree species in Puerto Rico. The over bark wood volume of African tulip trees in nine forest stands where it was dominant ranged from 163 to 849 m³/ha. Aboveground biomass ranged from 60 to 296 Mg/ha, and N and P stocks ranged from 190 to 988 and 32 to 137 kg/ha, respectively. Novel forests on abandoned agricultural lands can store more biomass and elements than native and plantation forest stands of similar age.     

Understory diversity and composition after planting of teak and mahogany in Yogyakarta,Indonesia

ABSTRACT

Forest rehabilitation is when a desired tree species is planted in degraded forests or lands. Rehabilitation by planting a single tree species is a common way to restore exploited forests to maintain ecological processes. We compared woody and herbaceous understory vegetation between forests rehabilitated by mahogany (N = 12) or teak (N = 12) planted from 1941 until 2003 in Yogyakarta, Indonesia. Understory vegetation of these areas was compared with that of three native forests. Species richness, species diversity, density of plants and proportion of native plants did not differ between the rehabilitated areas and the native forest. Recently rehabilitated areas were different from the native forests while 41–74 yr after rehabilitation, characteristics of understory vegetation approached those of native forest. We described species composition using ordination, and found it to differ between areas rehabilitated with teak and with mahogany and, particularly, between the rehabilitated areas and the native forests. Time since rehabilitation and tree species planted were important for the species composition of understory vegetation. We conclude that the selection of species for rehabilitation and letting rehabilitated areas mature are important for understory development and species diversity.     

Effects of multiple factors on plant diversity of forest fragments in intensive farmland of Northern Portugal

The joint and independent effects of dominant tree species, forest patch spatial attributes, and forest structure and management as drivers of plant species richness and composition in small forest patches scattered within an intensive agricultural landscape were addressed.In a landscape with scattered urban and intensive dairy agricultural areas in north-west Portugal, within which small forest patches (dominated by pines, eucalypts, or both) represent semi-natural habitat islands, 50 small forest patches, with areas ranging between 0.3 and 3 ha, were selected and surveyed for vascular plant diversity, within dairy farming landscape mosaics dominated by annual forage crops. Explanatory variables were composed of three datasets derived either from GIS mapping or field observations: forest type (dominant tree species), forest patch spatial attributes (patch area and shape index), and measures of forest management and structure (diameter at breast height, tree density per hectare, and percentage cover of vegetation strata). Variations in these forest patch attributes were assessed across forest types, and related to measures of plant diversity (total, native, alien, woody, and herbaceous species richness). Redundancy analysis with variance partitioning was applied to evaluate the joint and independent effects of the three sets of variables on species assemblages. The recent shift in canopy dominance from pine to eucalypt observed in the region appears to be related to a (nonsignificant) tendency for the increase of patch area and to the decrease of patch complexity, as well as several changes in forest structure and management, expressed as a trend to denser tree canopies and lower cover of understory plants. Dominant tree species and attributes related to forest structure and management were the most important factors determining plant diversity. The joint effect of the dominant tree species and forest structure and management resulted in lower levels of plant species richness in eucalypt plantations. These were also more prone to invasion by alien species, probably due to decreased biotic resistance from unsaturated native plant assemblages. Our results draw attention to the importance of dominant tree species and management practices for the maintenance of plant diversity levels (species richness and composition) in dairy landscape mosaics, highlighting the importance of the remnants of semi-natural forests as refuges for plant diversity in the landscape context. Nonetheless, forest plant diversity could further be fostered by promoting naturalness of pine stands and the regeneration of native oak woodlands in some forest areas. This would also diversify the range of ecosystem services that could be provided by forest areas in these peri-urban farmlands.     

Diversity of naturally-regenerated native woody species in forest plantations in the Ethiopian highlands

Woody species diversity and ground layervegetation cover were studied in plantations ofPinus patula, Cupressus lusitanica,Grevillea robusta, and Juniperusprocera, and in surrounding natural forests inWondo Genet, Ethiopia. In the understory of theplantations, a total of 53 naturallyregenerated tree and shrub species belonging to31 families were recorded. Important indigenoustimber species were also represented. Treesaccounted for 72% of all naturally-regeneratedwoody plant species. On the average, 78% ofthe woody species found in plantations werealso found in the natural forest. There was nosignificant variation in understory woodyspecies richness among plantations. C.lusitanica plantations had a significantlyhigher abundance of woody species than those ofP. patula, but the difference wasnot significant in relation to J. procera,G. robusta and the adjacent natural forest.The herbaceous ground cover percentage in G. robusta and P. patula stands wasconsiderably higher than in C. lusitanicaand J. procera stands. Relative toplantations of other species, plantations ofthe native tree species J. procera hadhigher understory species richness, mediumwoody species abundance and low ground cover.Although there was a marked variation inoverstory plantation density, its effect onunderstory species richness and abundance wasnot significant. The density ofnaturally-regenerated woody species inplantations was over three times the usualplanting density in Ethiopia, indicating a highpotential of forest plantations for restoringthe natural forest ecosystems on degraded landsat a comparatively low cost.     

Floristic and structural recovery of a laurel forest community after clear-cutting: A 60 years chronosequence on La Palma (Canary Islands)

We analyzed a post-clearcut chronosequence (0.5 to 60 years after harvesting) in the laurel forest of La Palma island (Canarian Archipelago) to determine the recovery of the stands with respect to species composition, richness, life strategies and structural parameters of the canopy. Multivariate analysis showed that exotic species, as well as annual ruderal species were confined to early-successional stages, while native perennials, typical of laurel forests, dominated the late-successional stages. Total species richness decreased significantly with time after clear-cutting. The relative fast recovery of understory native species may be due to low forest floor disturbance during harvesting. Shade-intolerant pioneer, pioneer-remnant and shade-tolerant late-successional species were the main life strategies of native tree species. Most structural parameters showed a continuous and monotonic increase (basal area, biomass) or decrease (density, percentage of photosynthetic biomass) during succession. Once clear-cutting, here performed with an interval of 8 years, is abandoned, the recovery of the laurel forest seems possible due to careful logging that protects the soil and a rapid asexual regeneration of native tree species, revealing this to be a sustainable management practice.     

Spatio-temporal variation of environmental signals inducing seed germination in temperate conifer plantations and natural hardwood forests in northern Japan

To document the spatial and temporal variation of environmental signals inducing seed germination in temperate forests, we measured temporal patterns of environmental signals and seed germination of six pioneer tree species in unthinned and thinned stands of conifer forests (Cryptomeria japonica plantations) and in the understory and gaps of hardwood forests in Japan. We also conducted germination experiment in laboratory for the six pioneer species to test the effects of red:far-red (R:FR) light ratio and temperature fluctuations on the seed germination. In conifer forests, the photosynthetic photon flux density (PPFD), the R:FR ratio, and the amplitude of temperature fluctuations in thinned stands were 2, 1.5, and 3 times higher, respectively, than those of unthinned stands. The PPFD and R:FR ratios just above forest floor also increased after the removal of thick litter accumulation. As a result, higher seed germination was observed in thinned compared to unthinned stands for three photoblastic species, whereas little differences were observed for three non-photoblastic species. These findings suggest that thinning, which frequently reduces litter accumulation, can substantially affect the regeneration of pioneer species and the resultant species diversity in conifer plantations. None of the measured environmental signals changed seasonally in unthinned stands of conifer forests, but they all changed remarkably in the understory of the hardwood forests. In this system, all signals were high and nearly identical to those in the gaps in early spring prior to canopy closure. Thus, the percent germination of the three photoblastic species was enhanced by high R:FR ratios and/or large temperature fluctuations even beneath the canopy and was nearly equal to that in the thinned conifer stands where the environmental conditions were nearly identical to those in the gaps. However, all of the environmental signals decreased with the expansion of canopy leaves and reached minimums at canopy closure. Even in the thinned stands and the gaps, the PPFD and magnitude of temperature fluctuations decreased over time due to shading by growing herbs and/or emerging canopy leaves. In these temporally changing environments, the germination of all photoblastic species ceased simultaneously. This study clearly demonstrated that the environmental signals inducing seed germination of photoblastic pioneer species spatially and temporally change in temperate forests, particularly in deciduous hardwood forests. Furthermore, these signals, PPFD, R:FR ratio, and the amplitude of temperature fluctuations, appear to play a very important role in tree regeneration and subsequent species diversity.     

Patterns of forest use and endemism in resident bird communities of north-central Michoacán, Mexico

We compared breeding avian communities among 11 habitat types in north-central Michoacán, Mexico, to determine patterns of forest use by endemic and nonendemic resident species. Point counts of birds and vegetation measurements were conducted at 124 sampling localities from May through July, in 1994 and 1995. Six native forest types sampled were pine, pine–oak, oak–pine, oak, fir, and cloud forests; three habitat types were plantations of Eucalyptus, pine, and mixed species; and the remaining two habitats were shrublands and pastures. Pastures had lower bird-species richness and abundance than pine, oak–pine, and mixed-species plantations. Pine forests had greater bird abundance and species richness than oak forests and shrublands. Species richness and abundance of endemics were greatest in fir forests, followed by cloud forests. Bird abundance and richness significantly increased with greater tree-layer complexity, although sites with intermediate tree complexity also supported high abundances. When detrended correspondence-analysis scores were plotted for each site, bird species composition did not differ substantially among the four native oak-and-pine forest types, but cloud and fir forests, Eucalyptus plantations, and mixed-species plantations formed relatively distinct groups. Plantations supported a mixture of species found in native forests, shrublands, and pastures. Pastures and shrublands shared many species in common, varied greatly among sites in bird-species composition, and contained more species specific to these habitats than did forest types.     

Effects of logging, liana tangles and pasture on seed fate of dry forest tree species in Central Brazil

We assessed seed fate of six tree species in a seasonally deciduous forest of Central Brazil. Seed germination, predation, removal and death caused by pathogens or dessication were followed in an undisturbed forest site, a logged forest site, and an active pasture. In both forest types we sowed seeds under patches of liana tangles (Low Forest habitat) and under patches of mature forest (High Forest habitat). Seeds were monitored weekly for up to 8 months. Overall, small-seeded species (Astronium fraxinifolium and Tabebuia impetiginosa) had high germination, although germination was lower in the pasture due to insect predation. The hard-coated seeds of Erythrina sp. had higher germination rates in the pasture. The large-seeded Cavanillesia arborea and Swartzia multijuga had high seed removal (ca. 90%) in the pasture and under High Forest, but only 23–34% under Low Forest in the logged site. Lower removal of large seeds in Low Forest suggests that this habitat conceals the seeds and constrains the movement of large mammals. Both species were favored in Low Forest, suggesting that caution should be used when managing lianas. When protected from predation, seeds with a soft coat and high water content, such as the seeds of Eugenia dysenterica and S. multijuga, died from desiccation in the pasture. Restoration efforts must take into consideration high mortality caused by desiccation or insect predation in abandoned pastures of dry forests.     

Diversity and succession of adventive and indigenous vascular understorey plants in Pinus radiata plantation forests in New Zealand

The vegetation of Pinus radiata plantation forests in New Zealand was studied to examine how the indigenous flora has responded to this novel habitat. A chronosequence of stands about 5, 16 and 27 years was assessed in each of four different biogeographic regions to test the effects of several stand and site factors on the succession of vascular understorey plant communities. A total of 202 indigenous and 70 adventive vascular plant species were found across all study areas, with considerable geographic variation among forests in species composition, species richness (range 48–135 species), and the percentage of indigenous species (50–86%). Both richness and cover of adventive species decreased significantly over time, whereas richness and cover of indigenous species was highest in the oldest stands, and overall species richness was lowest at mid-rotation. The guild composition changed from dominance of grasses and forbs in young stands to dominance of ferns and understorey trees in mature stands. These temporal changes were accompanied by a decrease in light-demanding pioneer species and an increase in shade tolerant, later seral species adapted to a forest environment. Measurements of the degree of canopy closure in stands with low or high stocking and modelling of temporal changes of canopy closure indicated that these understorey plant dynamics are influenced by changes in light availability as stands age. Despite the successional changes within forests, geographic variation more strongly influenced understorey communities because stands within a forest area were grouped together in DCA and TWINSPAN analyses, along rainfall and temperature gradients. Although the canopy species of such intensively managed plantation forests is an alien element in the New Zealand flora, the sheltered forest environment of older stands allows the establishment of a mostly indigenous forest understorey community with considerable similarities to indigenous forests located nearby.     

Understorey vegetation in young naturally regenerated and planted birch (Betula spp.) stands on abandoned agricultural land

The abandonment of agricultural lands in Northern and Eastern Europe increases the area covered by first generation forests, which are either formed as an outcome of secondary succession or established as plantations. However, questions remain as to how these new stands develop and what kind of species they favour, which in turn has impacts on their ecological and economical value. Our aim was to compare understorey vascular plant and bryophyte vegetation characteristics between naturally regenerated and planted birch stands on abandoned agricultural sites in Estonia, focusing on the aspects of species richness and forest understorey recovery. Species richness and diversity of vascular plants were similar in both stand types but the number of forest vascular plant species was significantly higher in naturally regenerated stands. The bryophyte layer of naturally regenerated stands had a higher species richness, diversity, and number of forest bryophyte species. The higher number of forest vascular plant and bryophyte species in naturally regenerated stands can be explained by the longer undisturbed succession period. The recovery of the forest understorey was unaffected by former agricultural land use (crop field or grassland). The influence of soil properties on the recovery of the forest understorey was not detected, but the number of vascular plant species that grow in forests as well as in grasslands was negatively correlated with distance from forest. Overall, understorey vegetation of natural and planted birch stands did not reveal substantial differences. However, in the case of vigorous natural birch regeneration in the vicinity of forest land, unassisted reforestation should be favoured.     

Direct seeding of late-successional trees to restore tropical montane forest

Natural regeneration of large-seeded, late-successional trees in fragmented tropical landscapes can be strongly limited by a lack of seed dispersal resulting in the need for more intensive restoration approaches, such as enrichment planting, to include these species in future forests. Direct seeding may be an alternative low-cost approach to planting nursery-raised tree seedlings, but there is minimal information on its efficacy or when in the successional process this technique will be most successful. We tested directly seeding five native tree species into habitats representing passive and active restoration approaches: (1) recently abandoned pasture; (2) naturally establishing, young secondary forests; and (3) young, mixed-species (fast-growing N-fixers and commercially valuable species) tree plantations established to facilitate montane forest recovery in southern Costa Rica. We monitored germination, survival, growth, and above- and below-ground biomass over a 2-year period. Germination in pastures, secondary forests, and tree plantations was similar (∼43%). Seedling survival after one and two years was significantly higher under tree plantations (91% year 1, 75% year 2) compared to secondary forests (76, 44%) or pastures (74, 41%). Moreover, seedlings had greater total biomass and lower root:shoot ratios in the plantations, suggesting higher nutrient availability in that treatment. Costs for direct seeding were 10- to 30-fold less per 100 seedlings after 2-year compared to nursery-raised seedlings planted at the same sites; however, there are important trade-offs to the two restoration approaches. Planting nursery-raised seedling is a more effective but higher cost approach for rapidly establishing canopy cover and restoring large areas whereas direct seeding is a more efficient way to enrich an existing system. We particularly recommend using direct seeding as a complimentary measure to the more intensive restoration approach of planting fast-growing and N-fixing trees.     

Floristic and structural patterns along a chronosequence of secondary forest succession in Argentinean subtropical montane forests

We studied forest structure and composition along a chronosequence of secondary forest succession in Northwest Argentina's montane forests (‘Yungas’) at 27°S, between 700 and 900 m. Early herbaceous stages, forested stages of 11, 25, 45, and 50 years after abandonment, and old-growth forests were surveyed. Secondary forests included stands that originated in abandoned herbaceous crops and in abandoned fruit orchards. Basal area and species composition differed between 50-year-old secondary forests and old-growth forests. In contrast, tree density and species diversity were similar in the 50-year-old and in the old-growth forests. The previous use (herbaceous crops or fruit orchards) was an important influence on secondary forest composition. Whereas stands originating in herbaceous fields were dominated by wind-dispersed native species such as Heliocarpus popayanensis, Tecoma stans, Parapiptadenia excelsa, and Tipuana tipu, fruit-orchard-originated stands were dominated by animal-dispersed species. Among the animal-dispersed species, the exotic tree Morus alba was the most abundant, and its abundance in secondary forests seems to slow the succession toward old-growth forest composition. Overall, after accounting for differences attributable to pre-abandonment conditions, secondary forest succession showed a trend toward compositional convergence, with the rate of succession apparently regulated by the demography of long-lived pioneer species.     

Woody understory plant diversity in pure and mixed native tree plantations at La Selva Biological Station,Costa Rica

Tree plantations can be an important tool for restoration of abandoned pasturelands in the tropics. Plantations can help speed up secondary forest succession by improving soil conditions, attracting seed-dispersal agents, and providing shade necessary for understory growth. In this study, abundance and richness of understory regeneration was measured in three native tree plantations 15–16 years of age at La Selva Biological Station in the Costa Rican Caribbean lowlands. Each plantation contained tree species in pure plots, a mixture of the species, and natural regeneration plots (no trees planted). The greatest abundance of regeneration was found in the understory of pure plots of Jacaranda copaia (Aubl.) D.Don., Vochysia guatemalensis Donn.Sm., Dipteryx panamensis Benth, Vochysia ferruginea Mart., and in two mixed stands, while the lowest was found in the natural regeneration treatments with about half the values as in the plantation stands. There was a significant negative correlation between percent canopy openness and abundance of regeneration in the understory. Two distinctive clusters separated the regeneration treatments from the mixed and pure plantations at a very low Bray–Curtis similarity value. The natural regeneration treatments are separated from mixed and pure plantations in the two-dimensional ordination. The lack of difference between the understory make-up of pure and mixed plantations in abundance, species richness, and seed-dispersal syndromes of understory species suggests that planting mixed stands is not necessarily superior to planting pure stands for promoting understory diversity of woody species. While regeneration of woody species can be faster under pure- or mixed-species plantations than in open pastures, the abundance, richness and species composition depends on each plantation species, or species assemblages in case of the mixtures.     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

Early seedling establishment of two tropical montane cloud forest tree species: The role of native and exotic grasses

Tropical montane cloud forest has been undergoing a drastic reduction because of its widespread conversion to pastures. Once these forests have been cleared exotic grasses are deliberately introduced for forage production. Exotic grass species commonly form monodominant stands and produce more biomass than native grass species, resulting in the inhibition of secondary succession and tree regeneration. The purpose of this study was to assess the effect of native vs. exotic grass species on the early establishment of two native tree seedlings (Mexican alder, Alnus acuminata and Jalapa oak, Quercus xalapensis) on an abandoned farm in central Veracruz, Mexico. Seedling survival and growth were monitored (over 46 weeks) in relation to grass cover and height, and available photosynthetic active radiation (PAR). More seedlings survived in the presence of the native grass Panicum glutinosum than those growing with the exotic grass Cynodon plectostachyus (92% vs. 48%). The causes of seedling mortality varied between species; Q. xalapensis was affected by herbivory by voles but mainly in the exotic grass-dominated stands, whereas A. acuminata seedlings died due to competition with the exotic grass. A. acuminata seedlings increased more in height in the exotic grass-dominated stands (102 ± 7.8 cm) compared to native grass-dominated stands (51 ± 4.7 cm). Grass layer height, cover and available PAR were correlated (Pearson; p < 0.05). In the exotic grass dominated plots, grass layer height was correlated with the relative height growth rates of Q. xalapensis (Pearson; p < 0.05). These results indicate that the exotic grass may be affecting tree regeneration directly (grass competition) and indirectly (higher herbivory). Passive restoration may occur once P. glutinosum dominated pastures are abandoned. However, when C. plectostachyus dominates, introduction of early and mid successional tree seedlings protected against vole damage is needed.