Short-term changes in tree populations in a tropical evergreen forest at Varagalaiar, Western Ghats, India

Within a 30-ha permanent plot of a tropical evergreen forest in the Anamalais, Indian Western Ghats, all the 13415 trees 30cmgbh (belonging to 153 species) were identified and tagged during 1997–1998. This communication reports the results of tree population changes based on two annual censuses (1999 and 2000) of the plot after the initial census, and compares the results with other tropical forest sites. One species, Ficus beddomei, disappeared from the plot and there was no new addition of species. Of the 96 species that showed changes in their population density, for 12 species recruitment and mortality matched. Tree recruitment (5 treesha^–1year^–1) exceeded mortality (4 treesha^–1year^–1) during the two-year period. Four modes of tree deaths were recorded in our site. The per cent mortality of trees differed between tree size classes during each recensus. The plot tree density increased progressively in the two-year period, indicating that shifts in species composition and density in natural forests, without major catastrophic disturbance, occur slowly.     

Dynamics,productivity and species richness of tropical rainforests along elevational and edaphic gradients on Mount Kinabalu,Borneo

We studied the dynamics of nine tropical rainforests on Mount Kinabalu, Borneo, at four elevations (700, 1,700, 2,700 and 3,100 m) on various edaphic conditions for four 2-year periods over 8 years (1995–2003), and examined the relationships with above-ground productivity. Mean growth rate of stem diameter, basal area turnover rate and estimated recruitment rate (using growth rate and size distribution) correlated with productivity among the nine forests in all periods. These rates based on growth rates of surviving stems appeared to be good measures of stand turnover. However, observed recruitment rate and mortality (and turnover rate as mean of these rates) based on direct observation of recruits and deaths did not correlate with productivity in some periods. These rates may not be useful as measures of stand turnover given small sample size and short census interval because they were highly influenced by stochastic fluctuation. A severe drought associated with the 1997–1998 El Niño event inflated mortality and depressed mean growth rate, recruitment rate and basal area turnover rate, but had little effect on the correlations between these rates (except mortality) and productivity. Across broad elevational and edaphic gradients on Mount Kinabalu, forest turnover, productivity and species richness correlated with each other, but the causal interpretation is difficult given the different histories and species pools among forests at different elevations.     

Fine root biomass under light gap openings in an Amazon rain forest

Summary Belowground processes in light gap openings are poorly understood, particularly in tropical forests. Fine roots in three zones of light gap openings and adjacent intact forest were regularly measured in buried bags and surface litter envelopes for 2 years. Fine root biomass does not vary significantly within gaps for either buried bags or for surface litter envelopes. When entire gaps are compared without regard for within gap zones, root growth into both surface litter and buried bags is significantly different between gaps, with highest rates of fine root biomass accumulation in the smallest gap. These results suggest that the aboveground within-gap zones do not result in a congruent pattern of below-ground zonation. Gap size, decomposition of the fallen tree, and pre-gap fine root growth rates should be considered to determine fine root growth patterns following the formation of light gap openings.     

Recruitment,mortality and growth of mangrove (<Emphasis Type="Italic">Rhizophora</Emphasis> sp.) seedlings in Ulugan Bay,Palawan, Philippines

Propagule dispersal, establishment and recruitment to the sapling stage are critical steps in the life cycle of mangroves. Specific (i.e., per capita) rates of recruitment and mortality, and the growth rates of Rhizophora seedlings in three mangrove stands in Ulugan Bay (Palawan, Philippines) were estimated between March 1999 and February 2001. Recruitment and mortality were variable in space and time, with mortality exceeding rates of recruitment at all sites. The specific rates of seedling recruitment and mortality were higher in Buenavista (0.66 year^–1 and –1.67 year^–1) than in Umalagan (0.05 year^–1 and –0.33 year^–1) and Oyster Bay (0.13 year^–1 and –0.24 year^–1). The annual rate of production of internodes by the main stem was similar at the three sites (5.4–5.5 internodes year^–1), but the annual rate of elongation of the main stem was higher in Buenavista (10.6 cm year^–1) than in Oyster Bay (7.6 cm year^–1) and Umalagan (5.6 cm year^–1).     

An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.     

Contrasting patterns of diameter and biomass increment across tree functional groups in Amazonian forests

Species' functional traits may help determine rates of carbon gain, with physiological and morphological trade-offs relating to shade tolerance affecting photosynthetic capacity and carbon allocation strategies. However, few studies have examined these trade-offs from the perspective of whole-plant biomass gain of adult trees. We compared tree-level annual diameter increments and annual above-ground biomass (AGB) increments in eight long-term plots in hyper-diverse northwest Amazonia to wood density (rho; a proxy for shade tolerance), whilst also controlling for resource supply (light and soil fertility). rho and annual diameter increment were negatively related, confirming expected differences in allocation associated with shade tolerance, such that light-demanding species allocate a greater proportion of carbon to diameter gain at the expense of woody tissue density. However, contrary to expectations, we found a positive relationship between rho and annual AGB increment in more fertile sites, although AGB gain did not differ significantly with rho class on low-fertility sites. Whole-plant carbon gain may be greater in shade-tolerant species due to higher total leaf area, despite lower leaf-level carbon assimilation rates. Alternatively, rates of carbon loss may be higher in more light-demanding species: higher rates of litterfall, respiration or allocation to roots, are all plausible mechanisms. However, the relationships between rho and AGB and diameter increments were weak; resource availability always exerted a stronger influence on tree growth rates.     

Can functional classification of tropical trees predict population dynamics after disturbance?

Question: How far can we simplify the fioristic complexity of a tropical rainforest into functional groups in order to predict tree population dynamics after logging‐induced disturbance? Location: Paracou experimental site, French Guiana. Methods: We used data from over 15 years in control and disturbed plots from a silvicultural trial started in 1984. We selected 53 common tree species assigned to five functional groups based on potential size and light requirement. For each species, we quantified: the fate, i.e. variation in population size, and dynamic processes, i.e. mortality, recruitment and growth, driving this fate. We investigated the links between dynamic processes, fate and functional groups. Results: Disturbance stimulated growth and recruitment for most species, but had a heterogeneous impact on mortality. Species fate in disturbed plots depended on recruitment and was more favourable than in control plots. The functional classification was more predictive for most separate dynamic processes than for species fate: after disturbance, significant differences were found between all functional groups for growth. Pioneer+heliophilous species showed significantly higher recruitment rates. Mortality of shade‐tolerant species slightly increased and of mid‐tolerant and heliophilous species decreased. Conclusions: A combination of three species classifications separately built from the growth, recruitment and mortality processes is more informative than a global classification combining the processes. Identifying the pioneer+heliophilous species on the basis of their growth rate is crucial to predict species fate after disturbance. We showed that potential growth rate could be used as a reliable indicator to identify this group.     

Wood growth patterns of <Emphasis Type="Italic">Macrolobium acaciifolium</Emphasis> (Benth.) Benth. (Fabaceae) in Amazonian black-water and white-water floodplain forests

Macrolobium acaciifolium (Benth.) Benth. (Fabaceae) is a dominant legume tree species occurring at low elevations of nutrient-poor black-water (igapó) and nutrient-rich white-water floodplain forests (várzea) of Amazonia. As a consequence of the annual long-term flooding this species forms distinct annual tree rings allowing dendrochronological analyses. From both floodplain types in Central Amazonia we sampled cores from 20 large canopy trees growing at identical elevations with a flood-height up to 7 m. We determined tree age, wood density (WD) and mean radial increment (MRI) and synchronized ring-width patterns of single trees to construct tree-ring chronologies for every study site. Maximum tree age found in the igapó was more than 500 years, contrary to the várzea with ages not older than 200 years. MRI and WD were significantly lower in the igapó (MRI=1.52±0.38 mm year^?1, WD=0.39±0.05 g cm^?3) than in the várzea (MRI=2.66±0.67 mm year^?1, WD=0.45±0.03 g cm^?3). In both floodplain forests we developed tree-ring chronologies comprising the period 1857–2003 (n=7 trees) in the várzea and 1606–2003 (n=13 trees) in the igapó. The ring-width in both floodplain forests was significantly correlated with the length of the terrestrial phase (vegetation period) derived from the daily recorded water level in the port of Manaus since 1903. In both chronologies we found increased wood growth during El Niño events causing negative precipitation anomalies and a lower water discharge in Amazonian rivers, which leads to an extension of the terrestrial phase. The climate signal of La Niña was not evident in the dendroclimatic proxies.     

Tree dieback and regeneration in secondary Acacia zanzibarica woodlands on an abandoned cattle ranch in coastal Tanzania

Questions: Bush encroachment is a major problem when African savanna ecosystems are used for cattle ranching. How do secondary woodlands develop after ranching is abandoned? What are the patterns and rates of tree mortality and regeneration? Location: Mkwaja Ranch (now part of Saadani National Park) in coastal Tanzania. Methods: Ninety‐seven circular plots (4‐m radius) were set in secondary Acacia zanzibarica woodland along a gradient of tree density. Variables relating to tree and grass layers and soil characteristics of plots were recorded. Seedlings were counted twice in the wet seasons, and resprouts once. Tree flowering and pod production were assessed during the fruiting season, while survival of trees initially present was recorded after 12 and 32 months. Interrelationships among variables were investigated using multiple linear regression, binary logistic regression and mixed effects models. Results: After 32 months, over one‐third of trees in plots had died. Most died after fire, especially on heavy soils, and mortality was significantly related to the tree live biomass ratio and soil conditions. Seed production was very low, especially in denser stands. Numbers of seedlings correlated with soil and grass variables but not with seed rain. Half of trees killed above‐ground produced coppice shoots from the base; in contrast, root suckering was independent of topkill. By the end of the study, no seedlings survived and no resprouts emerged above the grass layer. Conclusions: A. zanzibarica woodlands at Mkwaja Ranch were able to develop because of ranching, and can only persist under intensive grazing. The woodlands do not represent a successional stage towards forest and will probably revert to predominantly grassland vegetation within 10–20 years unless grazing pressure from wild ungulates increases considerably and/or fire regimes change.     

Early life-history dynamics of Caribbean coral species on artificial substratum: the importance of competition, growth and variation in life-history strategy

The development of a coral community was monitored for 6 years (1998–2004) on 46 m² of artificial settlement substrate in Curaçao, Netherlands Antilles. Growth and survival of recruits (n=1385) belonging to 16 different species were quantified in relation to characteristics of the benthic community developing around them. The early life history dynamics (i.e. growth rate, growth strategy and survival) of corals differed among species although these differences were small for species occupying similar habitats (i.e. underside versus topside of substratum). In contrast to recruit survival, juvenile growth rates were highly variable and unrelated to benthic community structure, at least at the scale of this study. Competing benthic organisms affected coral recruitment success through space preemption (mainly by macroalgae) or recruit overgrowth (mainly by sponges). The results highlight the small spatial scale (mm–cm) at which the processes responsible for recruitment success or failure occur and emphasize the need to include such small-scale observations in studies of coral early life-phase dynamics.     

Growth dynamics of Dracaena cinnabari under controlled conditions as the most effective way to protect endangered species

Dracaena cinnabari Balf. fil. is an endangered endemic species growing on the Yemeni island of Soqotra. Dracaena woodlands are considered as one of the oldest forest communities on Earth. Uncontrolled grazing unfortunately caused a lack of naturally occurring regeneration. Our two-year research was focused on the growth dynamics of Dracaena seedlings from two separate populations. One hundred of germinated seeds from two different altitudes from the island were sown and planted under the same conditions. Average increment and difference between the growth dynamics of plants from the two localities were investigated. The observed data on this plant species revealed very interesting, hitherto unknown results. (1) The seedlings germinated within a time period from four to ten weeks. Germination rate was 90% on the Firmihin highland plateau and 78% on the Scand Mountain. (2) Average plant length from both localities was almost the same (24.9 cm) at the end of measurement. Differences in values between the two populations proved as non-significant. (3) A significant difference was found in the number of leaves and in the sum of lengths of all leaves on one plant. While the seedlings from Firmihin featured a wide spreading above-ground part with a large number of leaves, the plants from Scand invested more energy into faster leaves elongation rate. (4) Growth dynamics reflected seasonal changes. Increments were slower or ceased during the period of vegetative rest from autumn to spring. (5) Average mortality rate was 13%. Most of the plants died during the period of vegetative rest. Further study on germination and regeneration under artificial conditions seems like the only way to prevent species extinction.     

Effect of vesicular-arbuscular mycorrhizae on seedling growth of four tree species from the tropical deciduous forest in Mexico

The influence of vesicular-arbuscular mycorrhizae on the growth of seedlings of Caesalpinia eriostachys, Cordia alliodora, Ipomoea wolcottiana and Pithecellobium mangense was investigated in a greenhouse experiment conducted at the Biological Station of Chamela on the Pacific coast of Mexico. Dry biomass production, relative growth rate, root/shoot ratio and mycorrhizal dependency were quantified for 75-day-old seedlings. With the exception of the pioneer species I. wolcottiana, mycorrhizal infection resulted in increases in biomass production, relative growth rate and leaf area. The root/shoot ratios attained for the species, however, did not show a consistent trend with infection. Nevertheless, all species had root/shoot ratios below 1 with infection and only one, Cordia alliodora, had a ratio greater than 1 without infection. The two late successional species from the mature part of the forest, Caesalpinia eriostachys and P. mangense, showed a larger mycorrhizal dependency than the two associated with disturbed environments.