Woody overstorey and herbaceous understorey biomass in Acacia harpophylla (brigalow) woodlands

Extensive areas of Acacia harpophylla (brigalow) woodlands have been cleared for pasture production in Queensland. The woody regrowth of A. harpophylla influences pasture production and composition following initial development of these woodlands. Biomass component regressions were developed using tree basal area as the predictor variable and used to estimate component yields of regrowing A. harpophylla communities. Leaf biomass of A. harpophylla reached a maximum of 12 t ha^-1 with a leaf area index of 2.5 when regrowing plants were 2.5 m high and about 10 years old. Pasture production, pasture basal area and the proportion of sown pasture species were lower at higher tree basal area of A. harpophylla. The greatest decrease was seen between 0 and 2 m²ha^-2tree basal area. Annual grasses and broadleaf herbaceous species represented a greater proportion of pasture biomass at high tree basal area. The poor control achieved by herbicide sprays on A. harpophylla plants that are 2–2.5 m tall would be partly due to poor coverage of the herbicide spray, as leaf area index is at a maximum at this stage. A density of 1000 plants ha^-1is a threshold value above which regrowth control would be necessary at some stage to maintain acceptable pasture production and composition. The non-linear regression relationship between pasture production and tree basal area was similar to that observed in Eucalyptus spp. However, in the equation that relates pasture yield (Y) to tree basal area (X), Y = A + B*e^(-kX)the value of k for A. harpophylla communities was generally higher than observed in Eucalyptus spp. communities with the same potential pasture production in the absence of trees. Within even-aged stands, the use of tree leaf biomass to predict pasture production showed no advantage over the use of tree basal area as the predictor. However, there are advantages in using tree leaf biomass as a predictor when comparing communities with different size- or age-class structures.     

Eucalyptus recruitment in degraded woodlands: no benefit from elevated soil fertility

Grasses and forbs compete heavily with young tree seedlings for available resources, greatly reducing tree seedling establishment success. Soil nutrient enrichment associated with agricultural intensification can increase the growth of both herbaceous and woody lifeforms growing in isolation, but may change the balance of competitive advantage when growing together. The effects of nitrogen and phosphorus enrichment on pasture biomass and competition with two Australian grassy woodland trees (Eucalyptus albens and Eucalyptus microcarpa) was investigated in a field plot trial. Soil nutrients increased pasture biomass, but had no measurable effect on tree growth in our experiment. Competition from pasture species, even at low levels, led to high tree seedling mortality and greatly reduced tree seedling growth compared with pasture-free plots. However, when pasture-free plots were excluded from the analysis, tree seedling leaf area was not strongly correlated with herbaceous biomass. Tree seedling establishment was severely restricted even at the lowest levels of pasture biomass. We conclude that increased soil fertility resulted in a competitive advantage to the pasture, and does not improve tree seedling establishment when grown either with or without exotic herbaceous pasture (grassy understorey) species.     

Effects of Quercus emoryi on herbaceous vegetation in a semi-arid savanna

Ten trees (5–70 m² canopy area) were selected to determine effects of tree size (crown area) on herbaceous species composition and biomass in a Quercus emoryi savanna in southeastern Arizona. Consistent with most studies in temperate savannas, herbaceous biomass was reduced beneath the canopy relative to grassland areas. However, tree size appeared to exert no influence over herbaceous biomass. In contrast to most temperate savannas, Q. emoryi trees did not affect distribution of herbaceous species.     

Does designation of protected areas ensure conservation of tree diversity in the Sudanian dry forest of Burkina Faso?

The importance of conservation status of the forest (protected versus unprotected) at two sites with differing human population density (high versus low) on the tree diversity of a Sudanian dry forest in Burkina Faso was studied. All woody species were recorded in 127 circular plots (area = 456.16 m²), and density, dominance, frequency, importance value indices and a variety of diversity measures were calculated to assess the species composition, structure and heterogeneity. A total of 69 species, representing 26 families and 52 genera, were found. Combretaceae, Leguminosae subfamily Caesalpinioideae and Rubiaceae were the dominant families. Neither human pressure nor forest conservation status significantly influenced the tree species richness. Stem density and basal area were significantly higher at the site with high population density than otherwise. Fisher’s diversity index revealed the unprotected forest at the site with low population density as the most diverse. We identified species with high conservation importance that should be enriched to maintain a viable population size. In conclusion, the current designation of protected areas seems inefficient at ensuring the conservation of tree diversity in the forest reserve. Thus, participatory conservation programme should be initiated.     

Plant diversity,structure and regeneration potential in tropical forests of Western Ghats,India

Diversity, stand structure and regeneration potential are the key elements of any forest ecosystem. For the present study, seven sites were selected with the aims of assessing plant diversity, structure and regeneration potential in tropical forests across Kanyakumari Wildlife Sanctuary (KWLS), Western Ghats, India. The sites were classified based on the similarity: tropical dry deciduous sites (TDDs I and II), tropical semi-evergreen sites (TSEs I and II) and tropical evergreen sites (TEFs I-III). The phytosociological survey was done by laying a total of 70 plots (10 plots in each study site). Standard methods were followed for the assessment of diversity, structure and regeneration patterns. A total of 267 species (205 genera, 70 families) were recorded. The tree species richness ranged 24 (TDD II) – 76 (TEF III). Of the vegetation spectrum, trees, vines and understorey accounted 56.5, 15.3 and 28.2% respectively to the total flora documented. A total of 66 species were endemic. The total tree density and tree basal area (seedlings, saplings, juveniles and adults) were 18,790 individuals (mean 2684) and 137.6 m² (mean 19.7 m²) in 70 plots respectively. The mean tree adult density and basal area ranged 370 (TDD I) – 900 (TEF I) individuals/ha and 24.2 (TDD I) – 75.3 (TEF III) m²/ha respectively. The overall species richness was highest in TDD I, but TEF III had the highest tree species richness. The diameter class-wise distribution showed the characteristic ‘reverse J-shaped’ curve. Most tree species were ‘newly recruited’. The dominant species had ‘fair’ to ‘good’ regeneration potential. However, 12 tree species showed ‘no’ regeneration. The overall regeneration pattern of trees was ‘good’, but ‘no’ or ‘poor’ regeneration patterns in some tree species, especially endemics is a point of concern. Since a majority of tree species were ‘new recruits’, species composition may likely change in the future. The results obtained would help in understanding diversity patterns, structural attributes and regeneration potential in tropical forests of protected areas for better forest management and conservation.     

Savanna tree influence on understory vegetation and soil nutrients in northwestern Kenya

Abstract. Contrary to observations and models in which trees and herbaceous plants are viewed as competitors, we found that trees in an African savanna have positive impacts on herbaceous biomass production and composition, and on soil nutrient status. In the Turkana District of northwestern Kenya, we investigated vegetation and soil gradients along equi-angular transects radiating from the boles of individual Acacia tortilis trees. Total herbaceous biomass averaged 260 ± 17(se) g/m² at the bole and declined to 95 ± 8 g/m² in the tree interspaces. Soil organic carbon and total nitrogen concentrations were greatest (0.72 % and 0.083 %, respectively) in shallow soils near the bole and declined rapidly toward the interspaces and with increasing depth. Transects were also established between tree pairs to assess effects of differential canopy proximities. Grass production averaged 220 ± 21 g / m² below overlapping canopies, 150 ± 15 g / m² under individual canopies, and 95 ± 8 g / m² in interstitial areas. Detrended correspondence analysis revealed that shifts in species composition were correlated with distance from tree bole out to the edge of the canopy. Species response, in terms of relative cover, to increasing distance from the bole, seemed to fall into five general classes: 1) greatest at the bole, 2) increasing with distance from the bole, 3) greatest in the mid canopy zone, 4) least at the bole and 5) no response. Trees did not influence herbaceous compositionbeyondtree canopies. It is assumed that shade cast by the tree canopy with subsequent reductions of understory water stress and temperature and increased nutrient concentrations may be the most important factors affecting understory soil and vegetation.     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

High butterfly beta diversity between Brazilian cerrado and cerrado–caatinga transition zones

Tropical dry forests are among the most diverse and threatened habitats in the world, yet they are rarely protected and remain poorly studied. In Brazil, dry forests are naturally fragmented and embedded within various biomes, thus making it important to assess biotic homogeneity among geographically separated forest fragments. We sampled 7732 individuals belonging to 48 species to quantify the diversity of fruit-feeding butterfly communities at four Brazilian dry forest sites, and found differences in community structure between northern and central sites. Species richness per plot was the same in both areas, but abundance per plot was higher in northern sites. Species composition differed between sites mostly due to species of Satyrinae. Additive partitioning showed that beta diversity corresponded to 70.1% of all diversity. Rather than species loss, beta diversity primarily represented species turnover that was potentially driven by differences in the surrounding habitats. Butterfly community composition and abundance were influenced by vegetation where abundance increased with tree density and basal area, and decreased with the average tree height. Butterfly species richness and abundance were higher in the wet season than in the dry season, and all species sampled in the dry season were present in the wet season. Differences in community structure across relatively short geographic distances in the same type of habitat highlight the importance of conserving tropical dry forest fragments to ensure the maintenance of butterfly diversity and, presumably, other insects.     

Use of Tropical Dry Forests and Agricultural Areas by Neotropical Bird Communities

Agricultural activities represent a global threat to biodiversity, yet little is known about the relative importance of different agricultural land uses in relation to their wildlife communities. We explored bird community structure, diversity, and composition in a landscape dominated by primary tropical dry forest, and in three agricultural systems (i.e., tree orchards, herbaceous cropfields, cattle pastures) to evaluate the way in which birds use different habitats within the landscape. Tropical dry forests had the highest species richness and community evenness, although the bird community in tree orchards was also relatively species rich and even. Cattle pastures had more bird species than cropfields but both habitats were comparatively species poor with low evenness. Our results are related to habitat structure and the spatial location of sites within the landscape matrix. Based on our observations, we recommend including native tree species within agricultural systems and surrounding areas to provide additional resources for birds. Finally, we suggest promoting natural recruitment of native trees and shrubs within cattle pastures to provide suitable habitat for species that use tropical dry forest plants. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Savannas after afforestation: Assessment of herbaceous community responses to wildfire versus native tree planting

Afforestation and fire exclusion are pervasive threats to tropical savannas. In Brazil, laws limiting prescribed burning hinder the study of fire in the restoration of Cerrado plant communities. We took advantage of a 2017 wildfire to evaluate the potential for tree cutting and fire to promote the passive restoration of savanna herbaceous plant communities after destruction by exotic tree plantations. We sampled a burned pine plantation (Burned Plantation); a former plantation that was harvested and burned (Harvested & Burned); an unburned former plantation that was harvested, planted with native trees, and treated with herbicide to control invasive grasses (Native Tree Planting); and two old-growth savannas which served as reference communities. Our results confirm that herbaceous plant communities on post-afforestation sites are very different from old-growth savannas. Among post-afforestation sites, Harvested & Burned herbaceous communities were modestly more similar in composition to old-growth savannas, had slightly higher richness of savanna plants (3.8 species per 50-m²), and supported the greatest cover of native herbaceous plants (56%). These positive trends in herbaceous community recovery would be missed in assessments of tree cover: whereas canopy cover in the Harvested & Burned site was 6% (less than typical of savannas of the Cerrado), the Burned Plantation and Native Tree Planting supported 34% and 19% cover, respectively. By focusing on savanna herbaceous plants, these results highlight that tree cutting and fire, not simply tree planting and fire exclusion, should receive greater attention in efforts to restore savannas of the Cerrado.     

Species diversity and community structure in sal (<Emphasis Type="Italic">Shorea robusta</Emphasis>) forests of two different rainfall regimes in West Bengal,India

Knowledge on the structure and composition of the plant communities has enormous significance in conservation and management of forests. The present study aimed to assess the community attributes, viz., structure, composition and diversity in the moist and dry sal (Shorea robusta) forests in the West Bengal province of India and compare them with the other sal forests of India. The phytosociological data from these forests were quantitatively analysed to work out the species richness, diversity, evenness, dominance, importance value, stand density and the basal area. The analysis showed that plant richness and diversity in moist sal forests of northern West Bengal are higher than the dry sal forests of south-west Bengal; a total of 134 tree (cbh ≥30 cm), 113 shrub and 230 herb species were recorded in the moist sal forest compared to 35 tree, 41 shrub and 96 herb species in dry sal forest. Papilionaceae was observed to be the dominant family. Dry sal forests had higher tree dominance (0.81) and stand density (1,006 stems ha⁻¹) but lower basal area (19.62 m²ha⁻¹) while moist sal forest had lower tree dominance (0.18) and stand density (438 stems ha⁻¹) but higher basal area (56.52 m²ha⁻¹). Tree species richness and stem density across girth classes in both the types decreased from the smallest to largest trees, while the occurrence rate of species increased with increase in girth class. A t-test showed significant differences in species richness, basal area and the stand density at 95% confidence level (p = <0.05) in the two forest types. The CCA indicated very low overall match (canonical correlation value = 0.40) between the two sets of variables from moist and dry sal types. The differences in these forests could be attributed to the distinct variations in climatic conditions- mainly the rainfall, disturbance regimes and the management practices.     

Soil organic C and nutrient contents under trees with different functional characteristics in seasonally dry tropical silvopastures

Aims

The selection of tree characteristics is critical for the outcome of the tree effects on soil fertility in silvopastoral pastures. This study aims to quantify the effects of trees on soil nutrient and C stocks, as well as assessing differences on the effects between legume (Albizia saman; Enterolobium cyclocarpum) and non-legume tree species (Tabebuia rosea; Guazuma ulmifolia).

Methods

In Central Nicaragua, soil was sampled (0–10 cm deep) in paired plots, under both a canopy and in open grassland, in 12 sites per tree species and analysed for organic C, total N stocks, available P and extractable K⁺, Ca²⁺ and Mg²⁺. To assess the effects of herbaceous composition and cattle to soil proprieties, we recorded the cover of plant groups and assessed the mass of dung in each plot.

Results

Soil organic C and N, available P and extractable K⁺ and Ca²⁺ were higher under the tree canopy than under paired open grassland. The basal area of trees was positively related with the canopy effect on soil variables, thus suggesting that the age or sizes of the trees are relevant factors associated with the content of soil C and nutrients. No specific effects related to the legume species group were detected.

Conclusions

Our results indicate that in fertile seasonally dry subtropical pastures, scattered trees have an overall effect on soil fertility, and that the magnitude of the effect depends more on the tree characteristics (i.e. basal area, crown area) than on whether the species is a legume or not.     

Relationship between plant species richness and biomass in an arid sub-alpine grassland of the central Himalayas,Nepal

The hump-shaped relationship between plant species richness and biomass is commonly observed at fine scale for herbaceous vegetation in temperate climates. This relationship predicts that herbaceous species richness is highest at an intermediate level of biomass that corresponds to moderate competition or disturbance. However, this relationship has not previously been investigated in high arid sub-alpine mountain grasslands. We tested the humped-back prediction in the arid Trans-Himalayan mountain grassland with a seasonal grazing system. The study area is located in the bottom of a U-shaped valley, in the Manang district (3500 m a.s.l.). We sampled two hundred plots (1m × 1m) in two different types of pastures: common pasture and old field, which both have similar grazing practices. There was a significant unimodal relationship between species richness and biomass only in the common pasture, and when the two sites were analyzed together. The species turnover is estimated by DCA in standard deviation unit. The turnover was lower in the old field than in the common pasture. The unimodal relationship between plant species richness and biomass did not disappear after accounting for unknown environmental gradients expressed as DCA (detrended correspondence analysis) axes and spatial variables. The species richness is highest at 120 ± 40 g/m². The results indicate that a hump-shaped relationship is also found in arid Trans-Himalayan grasslands.     

Microsite determinants of variability in seedling and cutting establishment in tropical forest restoration plantations

Plantations are frequently established on abandoned pasture lands to speed forest recovery. This strategy requires matching a tree species mix with the prevailing microenvironmental conditions. In four degraded pastures of the Mexican Lacandon rainforest, we planted 2,400 trees of 6 species (Guazuma ulmifolia, Inga vera, Ochroma pyramidale, Trichospermum mexicanum, Bursera simaruba, and Spondias mombin) to (1) test survival, initial growth, and establishment costs; (2) evaluate whether vegetative cuttings outperform direct seeding or transplants of nursery‐raised seedlings; (3) determine tree response to herbaceous dominance and soil compaction; and (4) scrutinize the results' consistency across sites and sampling scales of tree–microenvironment interactions (individual tree vs. averaged plot responses). After 2 years, overall survival and growth rates were high for 2 of 3 nursery‐raised species. Contrary to expectations, all seedlings outperformed the cuttings while direct seeding resulted in a cost‐effective option of intermediate efficacy. The impact of soil resistance to root penetration on tree biomass accumulation was species dependent while bulk density was not relevant. Soil‐covering, herbaceous vegetation accelerated growth in 3 of 4 tested species during the dry season. At this initial stage of forest restoration in abandoned pastures, Guazuma and Trichospermum were the most restoration‐effective species. Costs can be reduced by using direct‐seeding Inga and avoiding weeding during the dry season. Finally, our results demonstrate how species selection trials can be misleading due to site variations in tree response and to sampling scales that fail to account for small‐scale environmental heterogeneity. We recommend ways to improve the design of restoration trials.     

Influence of tree cover on herbaceous layer development and carbon and water fluxes in a Portuguese cork-oak woodland

Facilitation and competition between different vegetation layers may have a large impact on small-scale vegetation development. We propose that this should not only influence overall herbaceous layer yield but also species distribution and understory longevity, and hence the ecosystems carbon uptake capacity especially during spring. We analyzed the effects of trees on microclimate and soil properties (water and nitrate content) as well as the development of an herbaceous community layer regarding species composition, aboveground biomass and net water and carbon fluxes in a cork-oak woodland in Portugal, between April and November 2011.The presence of trees caused a significant reduction in photosynthetic active radiation of 35 mol m⁻² d⁻¹ and in soil temperature of 5 °C from April to October. At the same time differences in species composition between experimental plots located in open areas and directly below trees could be observed: species composition and abundance of functional groups became increasingly different between locations from mid April onwards. During late spring drought adapted native forbs had significantly higher cover and biomass in the open area while cover and biomass of grasses and nitrogen fixing forbs was highest under the trees. Further, evapotranspiration and net carbon exchange decreased significantly stronger under the tree crowns compared to the open during late spring and the die back of herbaceous plants occurred earlier and faster under trees. This was most likely caused by interspecific competition for water between trees and herbaceous plants, despite the more favorable microclimate conditions under the trees during the onset of summer drought.     

Composition and ecology of vascular epiphyte communities along an altitudinal gradient in central Veracruz,Mexico

Abstract. Vascular epiphytes were studied in forests at altitudes from 720 to 2370 m on the Atlantic slope of central Veracruz, Mexico. The biomass of all trees of each species > 10 cm diameter at breast height within plots between 625 and 1500 m² was estimated. The number of species per plot ranged between 22 and 53, and biomass between 9 and 249 g dry weight/m². The highest values, both of species and biomass, were found at an intermediate altitude (1430 m). Habitat diversity may contribute to epiphyte diversity in humid forests, but the importance of this effect could not be distinguished from the influence of climate. A remarkably high number of bromeliads and orchids grew in relatively dry forests at low altitudes. In wet upper montane forests, bromeliads were replaced by ferns, while orchids were numerous at all sites, except for a pine forest. The number of epiphytic species and their biomass on a tree of a given site were closely related to tree size. According to Canonical Correspondence Analysis, the factor determining the composition of the epiphytic vegetation of a tree was altitude and to some extent tree size, whereas tree species had practically no influence. The only trees which had an evidently negative effect on epiphytes were pines, which were particularly hostile to orchids and to a lesser degree to ferns, and Bursera simaruba, which generally had few epiphytes due to its smooth and defoliating bark.     

Early Plant Succession in Abandoned Pastures in Ecuador1

We compared early plant succession in four abandoned pastures of differing age since abandonment and a nearby secondary forest site in northwestern Ecuador. Two “Open” pastures had no tree canopy covering, and two “Guava” pastures had a well-developed canopy cover of Psidium guajava. No site had been seeded with pasture grasses. All pastures were compared in a chronological sequence; two were monitored for 18 months. Species richness was consistently higher in Guava sites than in Open sites and it continued to increase over time, whereas it remained static in Open sites. Species richness was highest in secondary forest. Recruitment of tree saplings in Guava sites was lower than in secondary forest; however, it was nearly absent in Open sites. The seed bank contained predominantly herbaceous species at all sites, and was highly dissimilar to aboveground vegetation. Dominance-diversity curves for Guava sites showed a more equitable distribution of species that increased over time. In contrast, dominance-diversity curves for Open sites were static and indicated dominance by a few aggressive species. Soil characteristics among sites were variable; however, a principal components analysis on soils isolated the older Open site from all others. The older Open site had the lowest species richness and was dominated by Baccharis trinervis, an aggressive shrub species. The site appears to be in a state of arrested succession and some form of restorative intervention may be necessary to initiate succession toward a forested condition. Succession in Guava sites appears headed toward secondary forest, whereas it does not in Open sites.     

Differences in Fine-Root Biomass of Trees and Understory Vegetation among Stand Types in Subtropical Forests

Variation of total fine-root biomass among types of tree stands has previously been attributed to the characteristics of the stand layers. The effects of the understory vegetation on total fine-root biomass are less well studied. We examined the variation of total fine-root biomass in subtropical tree stands at two sites of Datian and Huitong in China. The two sites have similar humid monsoon climate but different soil organic carbon. One examination compared two categories of basal areas (high vs. low basal area) in stands of single species. A second examination compared single-species and mixed stands with comparable basal areas. Low basal area did not correlate with low total fine-root biomass in the single-species stands. The increase in seedling density but decrease in stem density for the low basal area stands at Datian and the quite similar stand structures for the basal-area contrast at Huitong helped in the lack of association between basal area and total fine-root biomass at the two sites, respectively. The mixed stands also did not yield higher total fine-root biomasses. In addition to the lack of niche complementarity between tree species, the differences in stem and seedling densities and the belowground competition between the tree and non-tree species also contributed to the similarity of the total fine-root biomasses in the mixed and single-species stands. Across stand types, the more fertile site Datian yielded higher tree, non-tree and total fine-root biomasses than Huitong. However, the contribution of non-tree fine-root biomass to the total fine-root biomass was higher at Huitong (29.4%) than that at Datian (16.7%). This study suggests that the variation of total fine-root biomass across stand types not only was associated with the characteristics of trees, but also may be highly dependent on the understory layer.     

Photosynthetic responses of tree seedlings in grass and under shrubs in early-successional tropical old fields, Costa Rica

Only recently have studies addressed the effect of early-colonizing vegetation on tree seedling survival and growth during secondary succession in tropical old fields, and few studies have elucidated the physiological responses of tree seedlings to different vegetational communities. We compared growth and various photosynthetic parameters for seedlings of four rain-forest tree species, Cedrela tonduzii, Inga punctata, Ocotea whitei, and Tapirira mexicana, growing in areas of pasture grass and shrubs in early-successional abandoned pasture in Costa Rica; in addition, we made measurements for two species in forest gaps. We tested the general hypothesis that early-colonizing shrubs facilitate growth of forest tree seedlings. Specifically, we measured microclimate, growth, CO₂ assimilation, stomatal conductance, photosystem II quantum yield (KPSII), and xanthophyll pigment pools for all seedlings. Photosynthetic flux density (PFD) was higher under grass than shrubs or forest gaps, but was highly variable in each growth environment. For three of the four species, height growth was greatest in the grass compared to the shrubs and forest gaps; growth was similar below grass and shrubs for O. whitei. Photosynthetic capacity, apparent quantum yield, and stomatal conductance did not vary across habitats, but light compensation point and PFD at light saturation tended to be higher in the grass compared to forest and shrub growth environments. Water use efficiency differed across growth environments for three of the species. For plants in ambient PFD and dark-adapted plants, the efficiency of excitation energy transfer through PSII was lowest for plants in the grass compared to shrubs and forest gaps and also differed across species. Measurement of steady-state responses of KPSII to increasing PFD indicated a significant effect of growth environment at low PFD for all species and significant effects at high PFD only for I. punctata. All species exhibited a high degree of midday xanthophyll de-epoxidation in the different growth environments. Xanthophyll pigment pool size on an area basis was highest in the grass compared to shrubs and forest gaps for all four species. The results suggest that shrubs do not provide a facilitative effect for growth or photosynthesis for ~1.5-year-old seedlings of these four species. We conclude that site differences in success of tree seedlings during succession are a result of complex interactions of facilitation and competition and are not simply based on physiological responses to PFD.     

Large-Scale Wind Disturbances Promote Tree Diversity in a Central Amazon Forest

Canopy gaps created by wind-throw events, or blowdowns, create a complex mosaic of forest patches varying in disturbance intensity and recovery in the Central Amazon. Using field and remote sensing data, we investigated the short-term (four-year) effects of large (>2000 m²) blowdown gaps created during a single storm event in January 2005 near Manaus, Brazil, to study (i) how forest structure and composition vary with disturbance gradients and (ii) whether tree diversity is promoted by niche differentiation related to wind-throw events at the landscape scale. In the forest area affected by the blowdown, tree mortality ranged from 0 to 70%, and was highest on plateaus and slopes. Less impacted areas in the region affected by the blowdown had overlapping characteristics with a nearby unaffected forest in tree density (583±46 trees ha⁻¹) (mean±99% Confidence Interval) and basal area (26.7±2.4 m² ha⁻¹). Highly impacted areas had tree density and basal area as low as 120 trees ha⁻¹ and 14.9 m² ha⁻¹, respectively. In general, these structural measures correlated negatively with an index of tree mortality intensity derived from satellite imagery. Four years after the blowdown event, differences in size-distribution, fraction of resprouters, floristic composition and species diversity still correlated with disturbance measures such as tree mortality and gap size. Our results suggest that the gradients of wind disturbance intensity encompassed in large blowdown gaps (>2000 m²) promote tree diversity. Specialists for particular disturbance intensities existed along the entire gradient. The existence of species or genera taking an intermediate position between undisturbed and gap specialists led to a peak of rarefied richness and diversity at intermediate disturbance levels. A diverse set of species differing widely in requirements and recruitment strategies forms the initial post-disturbance cohort, thus lending a high resilience towards wind disturbances at the community level.