Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity?

Results from the humid tropics of Australia demonstrate that diverse plantations can achieve greater productivity than monocultures. We found that increases in both the observed species number and the effective species richness were significantly related to increased levels of productivity as measured by stand basal area or mean individual tree basal area. Four of five plantation species were more productive in mixtures with other species than in monocultures, offering on average, a 55% increase in mean tree basal area. A general linear model suggests that species richness had a significant effect on mean individual tree basal area when environmental variables were included in the model. As monoculture plantations are currently the preferred reforestation method throughout the tropics these results suggest that significant productivity and ecological gains could be made if multi-species plantations are more broadly pursued.

Growth and physiological response of six Australian rainforest tree species to a light gradient

An understanding of growth and photosynthetic potential of subtropical rainforest species to variations in light environment can be useful for determining the sequence of species introductions in rainforest restoration projects and mixed species plantations. We examined the growth and physiology of six Australian subtropical rainforest tree species in a greenhouse consisting of three artificial light environments (10%, 30%, and 60% full sunlight). Morphological responses followed the typical sun-shade dichotomy, with early and late secondary species (Elaeocarpus grandis, Flindersia brayleyana, Flindersia schottiana, and Gmelina leichhardtii) displaying higher relative growth rate (RGR) compared to mature stage species (Cryptocarya erythroxyion and Heritiera trifoliolatum). Growth and photosynthetic performance of most species reached a maximum in 30-60% full sunlight. Physiological responses provided limited evidence of a distinct dichotomy between early and late successional species. E. grandis and F brayleyana, provided a clear representation of early successional species, with marked increase in Am in high light and an ability to down regulate photosynthetic machinery in low light conditions. The remaining species (F. schottiana, G. leichhardtii, and H. trifoliolatum) were better represented as failing along a shade-tolerant continuum, with limited ability to adjust physiologically to an increase or decrease in light, maintaining similar A(max) across all light environments. Results show that most species belong to a shade-tolerant constituency, with an ability to grow and persist across a wide range of light environments. The species offer a wide range of potential planting scenarios and silvicultural options, with ample potential to achieve rapid canopy closure and rainforest restoration goals.

Modelling predicts positive and negative interactions between three Australian tropical tree species in monoculture and binary mixture

Computer modelling promises to be an important tool for analysing and predicting interactions between trees within mixed species forest plantations. This study explored the use of an individual-based mechanistic model as a predictive tool for designing mixed species plantations of Australian tropical trees. The `spatially explicit individually based-forest simulator' (SeXI-FS) modelling system was used to describe the spatial interaction of individual tree crowns within a binary mixed-species experiment. The three-dimensional model was developed and verified with field data from three forest tree species grown in tropical Australia. The model predicted the interactions within monocultures and binary mixtures of Flindersia brayleyana, Eucalyptus pellita and Elaeocarpus grandis, accounting for an average of 42% of the growth variation exhibited by species in different treatments. The model requires only structural dimensions and shade tolerance as species parameters. By modelling interactions in existing tree mixtures, the model predicted both increases and reductions in the growth of mixtures (up to +/-50% of stem volume at 7 years) compared to monocultures. This modelling approach may be useful for designing mixed tree plantations.

Screening selected myrtaceous tree species for production of essential oils in northern Queensland

Trials to identify alternative cropping options to Melaleuca alternifolia for northern Queensland essential oil growers were established at Dimbulah and Innot Hot Springs in 2001. Seed sources of Asteromyrtus symphyocarpa (1,8-cineole form), Eucalyptus staigeriana (citral), Melaleuca cajuputi subsp. cajuputi (trans-nerolidol), M. ericifolia (d-linalool), M. quinquenervia (trans-nerolidol and viridiflorol forms) and M. viridiflora (methyl cinnamate) with potential to produce commercial foliar oils were evaluated. Information was gathered on their adaptability, growth and oil yields over 49 months and 52 months (two harvests) from planting at Dimbulah and Innot Hot Springs, respectively. Of the species and chemotypes evaluated, M. quinquenervia showed potential for commercial production of trans-nerolidol, a compound used in perfumery. It had a very high survival rate (96%) and yields could be expected to improve dramatically from the average 100 kg/ha per harvest achieved in these trials with further research into selection of seed source, control of insect damage and breeding for genetic improvement. M. cajuputi subsp. cajuputi gave a similar performance to M. quinquenervia. The rarity of the trans-nerolidol form of this species and remoteness of its natural occurrence are impediments to further planting and research. E. staigeriana, with second harvest yields of ~600 kg/ha, performed exceptionally well on both sites but potential for development is limited by the ready availability of competitively priced E. staigeriana oil produced in South America. Survival of M. ericifolia ranged from 62% to 82% at 32 months (second harvest) at Innot Hot Springs and was deemed a failure at Dimbulah with poor growth and low survival, raising a major question about the suitability of this species for cultivation in the seasonally dry tropics. Planting of this species on a wider scale in northern Queensland cannot be recommended until more is known about factors affecting its survival. A. symphyocarpa and M. viridiflora were too slow-growing to warrant further consideration as potential oil-producing species at this time.

Productivity of plantation hardwood tree species in north-eastern Australia: A report from the Forest Adaptation and Sequestration Alliance

The report summarises data from a large number of trials of species with potential for use by the plantation forest industry in north-eastern Australia and provides information aimed at improving the understanding of growth rates, pest and disease risks and carbon sequestration. Data is summarised and presented at a regional level as opposed to individual trial or plot level. As well, nutritional impediments to tree growth and impacts on forest health are also reported. This report is intended to contribute to policy deliberations about developing forestry opportunities that can that can be integrated into the landscape, with particular consideration given to lower rainfall regions. There are several examples in north-eastern Australia where production forests have developed sub-optimally; this has often been due to poor selection of tree species as little information has been available. This report helps address this deficiency.

Is prosopis a curse or a blessing? : An ecological economic analysis of an invasive alien tree species in Sudan.

The issue of the usefulness of different prosopis species versus their status as weeds is a matter of hot debate around the world. The tree Prosopis juliflora had until 2000 been proclaimed weedy in its native range in South America and elsewhere in the dry tropics. P. juliflora or mesquite has a 90-year history in Sudan. During the early 1990s a popular opinion in central Sudan and the Sudanese Government had begun to consider prosopis a noxious weed and a problematic tree species due to its aggressive ability to invade farmlands and pastures, especially in and around irrigated agricultural lands. As a consequence prosopis was officially declared an invasive alien species also in Sudan, and in 1995 a presidential decree for its eradication was issued. Using a total economic valuation (TEV) approach, this study analysed the impacts of prosopis on the local livelihoods in two contrasting irrigated agricultural schemes. Primarily a problem-based approach was used in which the derivation of non-market values was captured using ecological economic tools. In the New Halfa Irrigation Scheme in Kassala State, four separate household surveys were conducted due to diversity between the respective population groups. The main aim was here to study the magnitude of environmental economic benefits and costs derived from the invasion of prosopis in a large agricultural irrigation scheme on clay soil. Another study site, the Gandato Irrigation Scheme in River Nile State represented impacts from prosopis that an irrigation scheme was confronted with on sandy soil in the arid and semi-arid ecozones along the main River Nile. The two cases showed distinctly different effects of prosopis but both indicated the benefits to exceed the costs. The valuation on clay soil in New Halfa identified a benefit/cost ratio of 2.1, while this indicator equalled 46 on the sandy soils of Gandato. The valuation results were site-specific and based on local market prices. The most important beneficial impacts of prosopis on local livelihoods were derived from free-grazing forage for livestock, environmental conservation of the native vegetation, wood and non-wood forest products, as well as shelterbelt effects. The main social costs from prosopis were derived from weeding and clearing it from farm lands and from canalsides, from thorn injuries to humans and livestock, as well as from repair expenses vehicle tyre punctures. Of the population groups, the tenants faced most of the detrimental impacts, while the landless population groups (originating from western and eastern Sudan) as well as the nomads were highly dependent on this tree resource. For the Gandato site the monetized benefit-cost ratio of 46 still excluded several additional beneficial impacts of prosopis in the area that were difficult to quantify and monetize credibly. In River Nile State the beneficial impact could thus be seen as completely outweighing the costs of prosopis. The results can contributed to the formulation of national and local forest and agricultural policies related to prosopis in Sudan and also be used in other countries faced with similar impacts caused by this tree.

Effect of rodents on seed fate of five hornbill-dispersed tree species in a tropical forest in north-east India

Hornbills are important dispersers of a wide range of tree species. Many of these species bear fruits with large, lipid-rich seeds that could attract terrestrial rodents. Rodents have multiple effects on seed fates, many of which remain poorly understood in the Palaeotropics. The role of terrestrial rodents was investigated by tracking seed fate of five horn bill-dispersed tree species in a tropical forest in north-cast India. Seeds were marked inside and outside of exclosures below 6-12 parent fruiting trees (undispersed seed rain) and six hornbill nest trees (a post-dispersal site). Rodent visitors and seed removal ere monitored using camera traps. Our findings suggest that several rodent species. especially two species of porcupine were major on-site seed predators. Scatter-hoarding was rare (1.4%). Seeds at hornbill nest trees had lower survival compared with parent fruiting trees, indicating that clumped dispersal by hornbills may not necessarily improve seed survival. Seed survival in the presence and absence of rodents varied with tree species. Some species (e.g. Polyalthia simiarum) showed no difference, others (e.g. Dysoxylum binectariferum) experienced up to a 64%. decrease in survival in the presence of rodents. The differing magnitude of seed predation by rodents can have significant consequences at the seed establishment stage.

Plant secondary compounds and soil microbial processes in carbon and nitrogen cycling in relation to tree species

The aim of this study was to explore soil microbial activities related to C and N cycling and the occurrence and concentrations of two important groups of plant secondary compounds, terpenes and phenolic compounds, under silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) as well as to study the effects of volatile monoterpenes and tannins on soil microbial activities. The study site, located in Kivalo, northern Finland, included ca. 70-year-old adjacent stands dominated by silver birch, Norway spruce and Scots pine. Originally the soil was very probably similar in all three stands. All forest floor layers (litter (L), fermentation layer (F) and humified layer (H)) under birch and spruce showed higher rates of CO2 production, greater net mineralisation of nitrogen and higher amounts of carbon and nitrogen in microbial biomass than did the forest floor layers under pine. Concentrations of mono-, sesqui-, di- and triterpenes were higher under both conifers than under birch, while the concentration of total water-soluble phenolic compounds as well as the concentration of condensed tannins tended to be higher or at least as high under spruce as under birch or pine. In general, differences between tree species in soil microbial activities and in concentrations of secondary compounds were smaller in the H layer than in the upper layers. The rate of CO2 production and the amount of carbon in the microbial biomass correlated highly positively with the concentration of total water-soluble phenolic compounds and positively with the concentration of condensed tannins. Exposure of soil to volatile monoterpenes and tannins extracted and fractionated from spruce and pine needles affected carbon and nitrogen transformations in soil, but the effects were dependent on the compound and its molecular structure. Monoterpenes decreased net mineralisation of nitrogen and probably had a toxic effect on part of the microbial population in soil, while another part of the microbes seemed to be able to use monoterpenes as a carbon source. With tannins, low-molecular-weight compounds (also compounds other than tannins) increased soil CO2 production and nitrogen immobilisation by soil microbes while the higher-molecular-weight condensed tannins had inhibitory effects. In conclusion, plant secondary compounds may have a great potential in regulation of C and N transformations in forest soils, but the real magnitude of their significance in soil processes is impossible to estimate.

Juvenile diversity of tree species in Theobroma cacao L. agroforestry and primary forest – Central Sulawesi, Indonesia

This thesis studies the tree species’ juvenile diversity in cacao (Theobroma cacao L.) based agroforestry and in primary forest in a natural conservation forest environment of Lore Lindu National Park, Sulawesi, Indonesia. Species’ adult composition in Lore Lindu National Park is relatively well studied, less is known about tree species’ diversity in seedling communities particularly in frequently disturbed cacao agroforestry field environment. Cacao production forms a potentially serious thread for maintaining the conservation areas pristine and forested in Sulawesi. The impacts of cacao production on natural environment are directly linked to the diversity and abundance of shade tree usage. The study aims at comparing differences between cacao agroforestry and natural forest in the surrounding area in their species composition in seedling and sapling size categories. The study was carried out in two parts. Biodiversity inventory of seedlings and saplings was combined with social survey with farmer interviews. Aim of the survey was to gain knowledge of the cacao fields, and farmers’ observations and choices regarding tree species associated with cacao. Data was collected in summer 2008. The assessment of the impact of environmental factors of solar radiation, weeding frequency, cacao tree planting density, distance to forest and distance to main park road, and type of habitat on seedling and sapling compositions was done with Non-metric Multidimensional Scaling (NMS). Outlier analysis was used to assess distorting variables for NMS, and Multi-Response Permutation Procedures (MRPP) analysis to differentiate the impact of categorical variables. Sampling success was estimated with rarefaction curves and jackknife estimate of species richness. In the inventory 135 species of trees and shrubs were found. Only some agroforestry related species were dominating. The most species rich were sapling communities in forest habitat. NMS was showing generally low linear correlation between variation of species composition and environmental variables. Solar radiation was having most significance as explaining variable. The most clearly separated in ordination were cacao and forest habitats. The results of seedling and sapling inventory were only partly coinciding with farmers’ knowledge of the tree species occurring on their fields. More research with frequent assessment of seedling cohorts is needed due to natural variability of cohorts and high mortality rate of seedlings.

Phenology of tree species of tropical moist forest of Uttara Kannada district, Karnataka, India

Phenological observations on tree species in tropical moist forest of Uttara Kannada district (13ℴ55′ to 15ℴ31′ N lat; 74ℴ9′ to 75ℴ10′ E long) during the years 1983–1985 revealed that there exists a strong seasonality for leaf flush, leaf drop and reproduction. Young leaves were produced in the pre-monsoon dry period with a peak in February, followed by the expansion of leaves which was completed in March. Abscission of leaves occurred in the post-monsoon winter period with a peak in December. There were two peaks for flowering (December and March), while fruit ripening had a single peak in May–June, preceding the monsoon rainfall. The duration of maturation of leaves was the shortest, while that of full ripening of fruits was the longest. Mature flowers of evergreen species lasted longer than those of deciduous species; in contrast the phenophase of ripe fruits of deciduous species was longer than that of evergreen species.

Scale-dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity. 2. Here, we conduct an analysis of relationships between tree species richness, biomass and productivity in 25 forest plots of area 8-50ha from across the world. The data were collected using standardized protocols, obviating the need to correct for methodological differences that plague many studies on this topic. 3. We found that at very small spatial grains (0.04ha) species richness was generally positively related to productivity and biomass within plots, with a doubling of species richness corresponding to an average 48% increase in productivity and 53% increase in biomass. At larger spatial grains (0.25ha, 1ha), results were mixed, with negative relationships becoming more common. The results were qualitatively similar but much weaker when we controlled for stem density: at the 0.04ha spatial grain, a doubling of species richness corresponded to a 5% increase in productivity and 7% increase in biomass. Productivity and biomass were themselves almost always positively related at all spatial grains. 4. Synthesis. This is the first cross-site study of the effect of tree species richness on forest biomass and productivity that systematically varies spatial grain within a controlled methodology. The scale-dependent results are consistent with theoretical models in which sampling effects and niche complementarity dominate at small scales, while environmental gradients drive patterns at large scales. Our study shows that the relationship of tree species richness with biomass and productivity changes qualitatively when moving from scales typical of forest surveys (0.04ha) to slightly larger scales (0.25 and 1ha). This needs to be recognized in forest conservation policy and management.

Temporal variability of forest communities: empirical estimates of population change in 4000 tree species

Long-term surveys of entire communities of species are needed to measure fluctuations in natural populations and elucidate the mechanisms driving population dynamics and community assembly. We analysed changes in abundance of over 4000 tree species in 12 forests across the world over periods of 6-28years. Abundance fluctuations in all forests are large and consistent with population dynamics models in which temporal environmental variance plays a central role. At some sites we identify clear environmental drivers, such as fire and drought, that could underlie these patterns, but at other sites there is a need for further research to identify drivers. In addition, cross-site comparisons showed that abundance fluctuations were smaller at species-rich sites, consistent with the idea that stable environmental conditions promote higher diversity. Much community ecology theory emphasises demographic variance and niche stabilisation; we encourage the development of theory in which temporal environmental variance plays a central role.

An estimate of the number of tropical tree species

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between similar to 40,000 and similar to 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of similar to 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of similar to 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

The Asian elephant is amongst the top three frugivores of two tree species with easily edible fruit

Large animal species are prone to local extirpation, but ecologists cannot yet predict how the loss of megaherbivores affects ecosystem processes such as seed dispersal. Few studies have compared the quantity and quality of seed dispersal by megaherbivores versus alternative frugivores in the wild, particularly for plant species with fruit easily consumed by many frugivorous species. In a disturbed tropical moist forest in India, we examine whether megaherbivores are a major frugivore of two tree species with easily edible, mammal-dispersed fruit. We quantify the relative fruit removal rates of Artocarpus chaplasha and Careya arborea, by the Asian elephant (Elephas maximus) and alternative dispersers. Through focal watches and camera trapping, we found the elephant to be amongst the top three frugivores for each tree species. Furthermore, seed transects under A. chaplasha show that arboreal frugivores discard seeds only a short distance from the parental tree, underscoring the elephant's role as a long-distance disperser. Our data provide unprecedented support for an old notion: megaherbivores may be key dispersers for a broad set of mammal-dispersed fruiting species, and not just fruit inaccessible to smaller frugivores. As such, the elephant may be particularly important for the functional ecology of the disturbed forests it still inhabits across tropical Asia.