Correlates of leaf optical properties in tropical forest sun and extreme-shade plants

Thirteens hade-adaptedr ain forest species were comparedw ith twelve sun-adaptedt ropical forest species for correlates to leaf optical properties (described previously in Amer. J. Bot. 73: 1100-1108). The two samples were similar in absorptance of quanta for photosynthesis, but the shade-adaptedt axa: 1) had significantlyl ower specificl eaf weights,i ndicatinga more metabolically efficient production of surface for quantum capture; 2) synthesized less chlorophyll per unit area; and 3) used less chlorophyll for capturing the same quanta for photosynthesis. The anatomical features that best correlate with this increased efficiency are palisade cell shape and chloroplast distribution. Palisade cells with more equal dimensions have more chloroplasts on their abaxial surfaces. This dense layer of chloroplasts maximizes the light capture efficiency limited by sieve effects. The more columnar palisade cells of sun-adapted taxa allow light to pass through the central vacuoles and spaces between cells, making chloroplasts less efficient in energy capture, but allowing light to reach chloroplasts in the spongy mesophyll. Pioneer species may be an exception to these two groups of species. Three pioneer taxa included in this study have columnar palisade cells that are extremely narrow and packed closely together. This layer allows little penetration of light, but exposure of the leaf undersurface may provide illumination of spongy mesophyll chloroplasts in these plants.

Vertical attenuation of light predicts the sequence to dawn song in a tropical forest avifuana

The concentration of avian song at first light (i.e., the dawn chorus) is widely appreciated but has an enigmatic functional significance. The most widely accepted explanation is that birds are active but light levels are not adequate for foraging. As a consequence, the time of first song should be predictable from the light level of individuals singing at dawn. To test this, I collected data from a tropical forest of Ecuador, involving 130 species. Light intensity at first song was a highly repeatable species' trait (r = 0.57). Foraging height was a good predictor of first song, with canopy birds singing at lower light levels than understory birds (r = -0.62). Although light level predicts the onset of singing in tropical and temperate bird communities, the structural complexity and trophic specializations in tropical forests may exert an important influence, which has been overlooked in research conducted in the temperate zone.

Forests as landscapes of social inequality: tropical forest cover and land distribution among shifting cultivators

Can social inequality be seen imprinted in a forest landscape? We studied the relationship between land holding, land use, and inequality in a peasant community in the Peruvian Amazon where farmers practice swidden-fallow cultivation. Longitudinal data on land holding, land use, and land cover were gathered through field-level surveys (n = 316) and household interviews (n = 51) in 1994/1995 and 2007. Forest cover change between 1965 and 2007 was documented through interpretation of air photos and satellite imagery. We introduce the concept of “land use inequality” to capture differences across households in the distribution of forest fallowing and orchard raising as key land uses that affect household welfare and the sustainability of swidden-fallow agriculture. We find that land holding, land use, and forest cover distribution are correlated and that the forest today reflects social inequality a decade prior. Although initially land-poor households may catch up in terms of land holdings, their use and land cover remain impoverished. Differential land use investment through time links social inequality and forest cover. Implications are discussed for the study of forests as landscapes of inequality, the relationship between social inequality and forest composition, and the forest-poverty nexus.

Conservation value, biodiversity value and methods of assessment in regenerating and human disturbed tropical forest

Although the value of primary forests for biodiversity conservation is well known, the potential biodiversity and conservation value of regenerating forests remains controversial. Many factors likely contribute to this, including: 1. the variable ages of regenerating forests being studied (often dominated by relatively young regenerating forests); 2. the potential for confounding on-going human disturbance (such as logging and hunting); 3. the relatively low number of multi-taxa studies; 4. the lack of studies that directly compare different historic disturbances within the same location; 5. contrasting patterns from different survey methodologies and the paucity of knowledge on the impacts across different vertical levels of rainforest biodiversity (often due to a lack of suitable methodologies available to assess them). We also know relatively little as to how biodiversity is affected by major current impacts, such as unmarked rainforest roads, which contribute to this degradation of habitat and fragmentation. This thesis explores the potential biodiversity value of regenerating rainforests under the best of scenarios and seeks to understand more about the impact of current human disturbance to biodiversity; data comes from case studies from the Manu and Sumaco Biosphere Reserves in the Western Amazon. Specifically, I compare overall biodiversity and conservation value of a best case regenerating rainforest site with a selection of well-studied primary forest sites and with predicted species lists for the region; including a focus on species of key conservation concern. I then investigate the biodiversity of the same study site in reference to different types of historic anthropogenic disturbance. Following this I investigate the impacts to biodiversity from an unmarked rainforest road. In order to understand more about the differential effects of habitat disturbance on arboreal diversity I directly assess how patterns of butterfly biodiversity vary between three vertical strata. Although assessments within the canopy have been made for birds, invertebrates and bats, very few studies have successfully targeted arboreal mammals. I therefore investigate the potential of camera traps for inventorying arboreal mammal species in comparison with traditional methodologies. Finally, in order to investigate the possibility that different survey methodologies might identify different biodiversity patterns in habitat disturbance assessments, I investigate whether two different but commonly used survey methodologies used to assess amphibians, indicate the same or different responses of amphibian biodiversity to historic habitat change by people. The regenerating rainforest study site contained high levels of species richness; both in terms of alpha diversity found in nearby primary forest areas (87% ±3.5) and in terms of predicted primary forest diversity from the region (83% ±6.7). This included 89% (39 out of 44) of the species of high conservation concern predicted for the Manu region. Faunal species richness in once completely cleared regenerating forest was on average 13% (±9.8) lower than historically selectively logged forest. The presence of the small unmarked road significantly altered levels of faunal biodiversity for three taxa, up to and potentially beyond 350m into the forest interior. Most notably, the impact on biodiversity extended to at least 32% of the whole reserve area. The assessment of butterflies across strata showed that different vertical zones within the same rainforest responded differently in areas with different historic human disturbance. A comparison between forest regenerating after selective logging and forest regenerating after complete clearance, showed that there was a 17% greater reduction in canopy species richness in the historically cleared forest compared with the terrestrial community. Comparing arboreal camera traps with traditional ground-based techniques suggests that camera traps are an effective tool for inventorying secretive arboreal rainforest mammal communities and detect a higher number of cryptic species. Finally, the two survey methodologies used to assess amphibian communities identified contrasting biodiversity patterns in a human modified rainforest; one indicated biodiversity differences between forests with different human disturbance histories, whereas the other suggested no differences between forest disturbance types. Overall, in this thesis I find that the conservation and biodiversity value of regenerating and human disturbed tropical forest can potentially contribute to rainforest biodiversity conservation, particularly in the best of circumstances. I also highlight the importance of utilising appropriate study methodologies that to investigate these three-dimensional habitats, and contribute to the development of methodologies to do so. However, care should be taken when using different survey methodologies, which can provide contrasting biodiversity patterns in response to human disturbance.

Longevity of horseweed seed bank depending on the depth of burial.

2016

Directed seed dispersal by bellbirds in a tropical cloud forest

A fundamental goal of plant population ecology is to understand the consequences for plant fitness of seed dispersal by animals. Theories of seed dispersal and tropical forest regeneration suggest that the advantages of seed dispersal for most plants are escape from seed predation near the parent tree and colonization of vacant sites, the locations of which are unpredictable in space and time. Some plants may gain in fitness as a fortuitous consequence of disperser behavior if certain species of dispersers nonrandomly place seeds in sites predictably favorable for seedling establishment. Such patterns of directed dispersal by vertebrates long have been suggested but never demonstrated for tropical forest trees. Here we report the pattern of seed distribution and 1-year seedling survival generated by five species of birds for a neotropical, shade-tolerant tree. Four of the species dispersed seeds to sites near the parent trees with microhabitat characteristics similar to those at random locations, whereas the fifth species, a bellbird, predictably dispersed seeds under song perches in canopy gaps. The pattern of seedling recruitment was bimodal, with a peak near parent trees and a second peak, corresponding to bellbird song perches, far (>40 m) from parent trees. Seedling survival was higher for seeds dispersed by bellbirds than by the other species, because of a reduction in seedling mortality by fungal pathogens in gaps. Thus, bellbirds play a significant role in seed dispersal by providing directed dispersal to favorable sites and therefore may influence plant recruitment patterns and species diversity in Neotropical forests.

Forest restoration in an indigenous land considering a forest remnant influence (Avai, Sao Paulo State, Brazil)

The expansion of agricultural and pasture areas over native forest areas has been broadly documented and represents the main cause of deforestation that has occurred for the last decades. Such reality is not different in indigenous lands, and has been considered as an important obstacle for individuals who directly depend upon the appropriate management of natural resources to maintain their traditions. We investigated the seed rain, seed bank and natural regeneration of native woody species within a degraded pasture land in different distances from an adjacent seasonal semideciduous forest fragment to define methodological procedures based on ecological processes that might subsidize forest restoration in an indigenous land. Most seeds and seedlings picked from the seed rain and seed bank belonged to anemochoric and autochoric dispersing shrubby and herbaceous species originated in the pasture land. The woody species regeneration, on the other hand, reached higher levels, in terms of abundance and richness, as the forest fragment became closer. Zoochoric dispersal was predominant among such species and was mainly detected closer to the forest fragment. Several woody species picked in the forest fragment were also found in the pasture land, thus reinforcing their important role in supplying propagules and easing the successional process. (c) 2007 Elsevier B.V. All rights reserved.

Improving Planting Stocks for the Brazilian Atlantic Forest Restoration through Community-Based Seed Harvesting Strategies

High-diversity reforestation can help jumpstart tropical forest restoration, but obtaining viable seedlings is a major constraint: if nurseries do not offer them, it is hard to plant all the species one would like. From 2007 to 2009, we investigated five different seed acquisition strategies employed by a well-established tree nursery in southeastern Brazil, namely (1) in-house seed harvesters; (2) hiring a professional harvester; (3) amateur seed harvesters; or (4) a seed production cooperative, as well as (5) participating in a seed exchange program. In addition, we evaluated two strategies not dependent on seeds: harvesting seedlings from native tree species found regenerating under Eucalyptus plantations, and in a native forest remnant. A total of 344 native tree and shrub species were collected as seeds or seedlings, including 2,465 seed lots. Among these, a subset of 120 species was obtained through seed harvesting in each year. Overall, combining several strategies for obtaining planting stocks was an effective way to increase species richness, representation of some functional groups (dispersal syndromes, planting group, and shade tolerance), and genetic diversity of seedlings produced in forest tree nurseries. Such outcomes are greatly desirable to support high-diversity reforestation as part of tropical forest restoration. In addition, community-based seed harvesting strategies fostered greater socioeconomic integration of traditional communities in restoration projects and programs, which is an important bottleneck for the advance of ecological restoration, especially in developing countries. Finally, we discuss some of the limitations of the various strategies for obtaining planting stocks and the way forward for their improvement.

The importance of landscape structure for seed dispersal in rain forest fragments

Questions What are the main features of the seed rain in a fragmented Atlantic forest landscape? Can seed rain species attributes (life form, dispersal mode, successional status) relate to the spatial arrangement (size and number of fragments, edge density and presence of corridor) of forest fragments in the landscape? How does the rain forest landscape structure affect the seed rain? Location Atlantic rainforest, Sao Paulo State, Southeastern Brazil. Methods Seed rain samples were collected monthly throughout 1yr, counted, identified and classified according to species dispersal mode, successional status and life form. Seed rain composition was compared with woody species near the seed traps. Relationships between seed rain composition and landscape spatial arrangement (fragment area, presence of corridor, number of fragments in the surroundings, proximity of fragments, and edge density) were tested using canonical correspondence analysis (CCA). Results We collected 20142 seeds belonging to 115 taxa, most of them early successional and anemochorous trees. In general, the seed rain had a species composition distinct from that of the nearby forest tree community. Small isolated fragments contained more seeds, mainly of anemochorous, epiphytic and early-successional species; large fragments showed higher association with zoochorous and late-successional species compared to small fragments. The CCA significantly distinguished the species dispersal mode according to fragment size and isolation, anemochorous species being associated to small and isolated fragments, and zoochorous species to larger areas and fragment aggregation. Nevertheless, a gradient driven by proximity (PROX) and edge density (ED) segregated lianas (in the positive extremity), early successional and epiphyte species (in the negative end); large fragments were positively associated to PROX and ED. Conclusions The results highlight the importance of the size and spatial arrangement of forest patches to promote habitat connectivity and improve the flux of animal-dispersed seeds. Landscape structure controls seed fluxes and affects plant dispersal capacity, potentially influencing the composition and structure of forest fragments. The seed rain composition may be used to assess the effects of landscape spatial structure on plant assemblages, and provide relevant information for biodiversity conservation.

Frugivores, seed dispersal and tree regeneration along a human disturbance gradient in East African tropical rainforests

Samenausbreitung und Regeneration von Bäumen sind wichtig für den langfristigen Bestand von Baum- und Frugivorengemeinschaften in tropischen Regenwäldern. Zunehmende Rohdung und Degradation gefährden den Ablauf dieser mutualistischen Prozesse in diesem Ökosystem. Um den Einfluss von kleinräumiger menschlicher Störung auf die Frugivorengemeinschaft und die zentralen Ökosystemprozesse Samenausbreitung und Regeneration zu erforschen, habe ich 1) die Frugivorengemeinschaft und die Samenausbreitungsrate von Celtis durandii (Ulmaceae) und 2) den Zusammenhang zwischen Baumarten mit fleischigen Früchten, Frugivoren und der Etablierung von Keimlingen dieser Baumarten in unterschiedlich stark gestörten Flächen dreier ostafrikanischer tropischer Regenwälder untersucht. Insgesamt konnte ich 40 frugivore Vogel- und Primatenarten in den drei untersuchten Waldgebieten nachweisen. Auf gering gestörten Flächen wurden mehr Frugivore als auf stark gestörten Flächen aufgenommen. Auch die Beobachtungen an C. durandii ergaben mehr frugivore Besucher in Bäumen auf gering gestörten als auf stark gestörten Flächen. Dies führte zu einer marginal signifikant höheren Samenausbreitungsrate auf den gering gestörten Flächen. Diese Ergebnisse waren auf regionaler Ebene in allen drei untersuchten Wäldern konsistent. Dies zeigt, dass kleinräumige Störung einen umfassenderen negativen Einfluss auf Frugivore und ihre Funktion als Samenausbreiter hat als zuvor angenommen. Bei der Vegetationserfassung nahm ich 131 verschiedene Baumarten mit fleischigen Früchten in den drei Regenwäldern auf. Kleinräumige menschliche Störung erhöhte den Artenreichtum dieser Baumarten marginal signifikant, hatte jedoch keinen direkten Einfluss auf die Frugivorendichte und den Artenreichtum von Keimlingen dieser Baumarten. Der Artenreichtum von Baumarten mit fleischigen Früchten zeigte einen marginal signifikant positiven Einfluss auf die Frugivorendichte, allerdings nicht auf die Keimlinge. Allerdings führte die Dichte der Frugivoren zu signifikant erhöhtem Artenreichtum der Keimlinge. Folglich scheint kleinräumige Störung die Keimlingsetablierung indirekt durch erhöhten Baumartenreichtum und erhöhte Frugivorendichte zu beeinflussen. Die Frugivorendichte hatte einen größeren Einfluss auf die Waldregeneration als kleinräumige Störung und Baumartenreichtum. Demnach scheint kleinräumige menschliche Störung sowohl positive als auch negative Effekte auf Samenausbreitung und Regeneration zu haben. Somit sind weitere Studien notwendig, die den Einfluss von kleinräumiger menschlicher Störung auf Mutualismen tropischer Regenwälder aufklären.

Forest structure and live aboveground biomass variation along an elevational gradient of tropical Atlantic moist forest (Brazil)

Live aboveground biomass (AGB) is an important source of uncertainty in the carbon balance from the tropical regions in part due scarcity of reliable estimates of live AGB and its variation across landscapes and forest types. Studies of forest structure and biomass stocks of Neotropical forests are biased toward Amazonian and Central American sites. In particular, standardized estimates of aboveground biomass stocks for the Brazilian Atlantic forest are rarely available. Notwithstanding the role of environmental variables that control the distribution and abundance of biomass in tropical lowland forests has been the subject of considerable research, the effect of short, steep elevational gradients on tropical forest structure and carbon dynamics is not well known. In order to evaluate forest structure and live AGB variation along an elevational gradient (0-1100 m a.s.l.) of coastal Atlantic Forest in SE Brazil, we carried out a standard census of woody stems >= 4.8 cm dbh in 13 1-ha permanent plots established on four different sites in 2006-2007. Live AGB ranged from 166.3 Mg ha(-1) (bootstrapped 95% CI: 1444,187.0) to 283.2 Mg ha(-1) (bootstrapped 95% CI: 253.0,325.2) and increased with elevation. We found that local-scale topographic variation associated with elevation influences the distribution of trees >50 cm dbh and total live AGB. Across all elevations, we found more stems (64-75%) with limited crown illumination but the largest proportion of the live AGB (68-85%) was stored in stems with highly illuminated or fully exposed crowns. Topography, disturbance and associated changes in light and nutrient supply probably control biomass distribution along this short but representative elevational gradient. Our findings also showed that intact Atlantic forest sites stored substantial amounts of carbon aboveground. The live tree AGB of the stands was found to be lower than Central Amazonian forests, but within the range of Neotropical forests, in particular when compared to Central American forests. Our comparative data suggests that differences in live tree AGB among Neotropical forests are probably related to the heterogeneous distribution of large and medium-sized diameter trees within forests and how the live biomass is partitioned among those size classes, in accordance with general trends found by previous studies. In addition, the elevational variation in live AGB stocks suggests a large spatial variability over coastal Atlantic forests in Brazil, clearly indicating that it is important to consider regional differences in biomass stocks for evaluating the role of this threatened tropical biome in the global carbon cycle. (C) 2010 Elsevier B.V. All rights reserved.

Spatio-temporal patterns of throughfall and solute deposition in an open tropical rain forest

The brief interaction of precipitation with a forest canopy can create a high spatial variability of both throughfall and solute deposition. We hypothesized that (i) the variability in natural forest systems is high but depends on system-inherent stability, (ii) the spatial variability of solute deposition shows seasonal dynamics depending on the increase in rainfall frequency, and (iii) spatial patterns persist only in the short-term. The study area in the north-western Brazilian state of Rondonia is subject to a climate with a distinct wet and dry season. We collected rain and throughfall on an event basis during the early wet season (n = 14) and peak of the wet season (n = 14) and analyzed the samples for pH and concentrations of NH4+, Na+, K+, Ca2+ Mg2+,, Cl-, NO3-, SO42- and DOC. The coefficient 3 4 cient of variation for throughfall based on both sampling intervals was 29%, which is at the lower end of values reported from other tropical forest sites, but which is higher than in most temperate forests. Coefficients of variation of solute deposition ranged from 29% to 52%. This heterogeneity of solute deposition is neither particularly high nor particularly tow compared with a range of tropical and temperate forest ecosystems. We observed an increase in solute deposition variability with the progressing wet season, which was explained by a negative correlation between heterogeneity of solute deposition and antecedent dry period. The temporal stability of throughfall. patterns was Low during the early wet season, but gained in stability as the wet season progressed. We suggest that rapid plant growth at the beginning of the rainy season is responsible for the lower stability, whereas less vegetative activity during the later rainy season might favor the higher persistence of ""hot"" and ""cold"" spots of throughfall. quantities. The relatively high stability of throughfall patterns during later stages of the wet season may influence processes at the forest floor and in the soil. Solute deposition patterns showed less clear trends but all patterns displayed a short-term stability only. The weak stability of those patterns is apt to impede the formation of solute deposition -induced biochemical microhabitats in the soil. (C) 2008 Elsevier B.V. All rights reserved.

Repeated disturbances and canopy disturbance regime in a tropical semi-deciduous forest

The canopy disturbance regime and the influence of gap methods on the interpretation of forest structure and dynamics were evaluated in a tropical semi-deciduous forest in south-eastern Brazil. We encountered a gap density of 11.2 gaps ha(-1) and an average size which varied from 121 to 333 m(2) depending on the gap delimitation method considered (minimum gap size was 10 m(2)). Although average size was slightly higher, the median value obtained (78 m(2)) was comparable to other tropical forest sites and the gap size-class distribution found supported the pattern described for such forest sites. Among 297 gap makers, snapping and uprooting were the most common modes of disturbance. The number and basal area of gap makers were good predictors of gap size. Almost 25% of all gaps suffered from repeated disturbance events that brought about larger gap sizes. Such processes, along with delimitation methods, strongly influenced the estimation of turnover rate and therefore the interpretation of forest dynamics. These results demonstrated the importance of further studies on repeated disturbances, which is often neglected in forest studies.

Ecosystem dynamics at disturbed and undisturbed sites in North Queensland wet tropical rain forest. III. Nutrient returns to the forest floor through litterfall

The nutrient contents and accessions in litterfall over a period of 3 y are reported for undisturbed areas and at two sites disturbed by selective harvesting in tropical rain forest in North Queensland, Australia. Mean concentrations (mg g(-1) dry weight) of nutrients in litterfall ranged from 10 to 12 for nitrogen; 0.33 to 0.43 for phosphorus; 3.6 to 4.3 for potassium; 6.0 to 10.5 for calcium and 1.7 to 2.6 for magnesium. These concentrations are in the middle to lower part of the spectrum of values recorded for tropical forest. Accessions of nutrients in litterfall (kg ha(-1) y(-1)) ranged from 59 to 64 N; 1.9 to 2.4 P; 20 to 24K; 34 to 63 Ca; and 9 to 16 Mg. These rates, particularly for IN and P, are among the lowest recorded for tropical forests. There were no consistent between-site differences in total nutrient accessions in small litterfall. In terms of the contribution of litterfall to the accessions of nutrients to the forest floor, this suggests that the logged sites have recovered from the effects of selective harvesting within 25 y. Nutrient accessions at each site were distinctly seasonal, with maximum accessions occurring in the late dry season to late in the wet season. Leaf-fall accounted for the largest proportion of nutrient accessions over the study period, although at certain times accessions in both reproductive material and wood were significant. A cyclone which crossed the coast near the study sites resulted in large nutrient accessions over a short period but had little effect on the total annual accession. A comparison with previous studies of litterfall in Australian tropical rainforests indicates that nutrient return in litterfall is directly related to soil fertility.