Original humus forms in a semi-deciduous tropical forest in Guadeloupe

Humus profiles underneath the canopy of dominant tree species in two secondary semi-evergreen forest sites in Grande-Terre (Guadeloupe) were analysed with a micromorphological method. In the vertisol of a tree plantation, the humus formed was rather similar under all tree species being an eumull and essentially due to the activity of the endoanecic earthworm Polypheretima elongata. In a natural secondary forest located on a steep slope and associated with a rendzina soil (without endoanecic earthworms), the humus forms were described at lower, mid- and upper slope. In this forest, two particular humus forms were observed. At the middle slope, underneath the canopy of Pisonia subcordata L. that produces nitrogen-rich litter, a calcareous amphimull, characterised by an OH horizon made of millipede faecal pellets, was formed. In the upper slope, underneath the canopy of Bursera simaruba (L.) Sarg. that produces a litter rich in resins and aromatic compounds that are poorly consumed by soil animals, a dysmull with a thick root mat (OFRh horizon) developed. Other humus forms were intermediate. The formation of these humus forms is discussed.     

Comparison of the mass and energy exchange of a pasture and a mature transitional tropical forest of the southern Amazon Basin during a seasonal transition

This research utilized tower‐based eddy covariance to quantify the trends in net ecosystem mass (CO₂ and H₂O vapor) and energy exchange of important land‐cover types of NW Mato Grosso during the March–December 2002 seasonal transition. Measurements were made in a mature transitional (ecotonal) tropical forest near Sinop, Mato Grosso, and a cattle pasture near Cotriguaçú, Mato Grosso, located 500 km WNW of Sinop. Pasture net ecosystem CO₂ exchange (NEE) was considerably more variable than the forest NEE over the seasonal transition, and the pasture had significantly higher rates of maximum gross primary production in every season except the dry–wet season transition (September–October). The pasture also had significantly higher rates of whole‐ecosystem dark respiration than the forest during the wetter times of the year. Average (±95% CI) rates of total daily NEE during the March–December 2002 measurement period were 26±15 mmol m^?2 day^?1 for the forest (positive values indicate net CO₂ loss by the ecosystem) and ?38±26 mmol m^?2 day^?1 for the pasture. While both ecosystems partitioned more net radiation (R_n) into latent heat flux (L_e), the forest had significantly higher rates of L_e and lower rates of sensible heat flux (H) than the pasture; a trend that became more extreme during the onset of the dry season. Large differences in pasture and forest mass and energy exchange occurred even though seasonal variations in micrometeorology (air temperature, humidity, and radiation) were relatively similar for both ecosystems. While the short measurement period and lack of spatial replication limit the ability to generalize these results to pasture and forest regions of the Amazon Basin, these results suggest important differences in the magnitude and seasonal variation of NEE and energy partitioning for pasture and transitional tropical forest.     

Leaf flushing phenology and herbivory in a tropical dry deciduous forest,southern India

Patterns of leaf-flushing phenology of trees in relation to insect herbivore damage were studied at two sites in a seasonal tropical dry forest in Mudumalai, southern India, from April 1988 to August 1990. At both sites the trees began to flush leaves during the dry season, reaching a peak leaf-flushing phase before the onset of rains. Herbivorous insects emerged with the rains and attained a peak biomass during the wet months. Trees that flushed leaves later in the season suffered significantly higher damage by insects compared to those that flushed early or in synchrony during the peak flushing phase. Species whose leaves were endowed with physical defenses such as waxes suffered less damage than those not possessing such defenses. There was a positive association between the abundance of a species and leaf damage levels. These observations indicate that herbivory may have played a major role in moulding leaf flushing phenology in trees of the seasonal tropics.     

Leaf area index determination in an alder forest: a comparison of three methods

Leaf area index (LAI) of a stand of adult black alder trees(Alnus glutinosa L., Gaertn.) was determined by means of threeindependent methods. (1) The seasonal course of LAI was directlyobtained by counting leaves in situ and adding up their areas,estimated from harvested subsamples of leaves. (2) The seasonalvariation of LAI in the stand was estimated using the Li-CorLAI-2000 PCA in parallel and with this instrument a VegetationArea Index (VAI, projected area of all phyto-elements) was actuallymeasured. (3) Maximum LAI was calculated from leaf litter collectionstaking into account specific leaf area within different layersof the alder crown. Direct LAI estimates (1) and calculationsfrom leaf litter (3) revealed the same figure of maximum LAI(4.8). This LAI was reached in August. The LAI-2000 PCA capturedthe seasonal variation and underestimated, by 11% on average,the LAI obtained directly. Compared with results gained withother broad-leaved tree species the LAI-2000 PCA values foralder were reliable. It is suggested that this is due to thehorizontal homogeneous structure of the main leaf layer. Thisis in the periphery of the crown, where 90% of the light interceptionoccurs. Taking the het-erogeneity into account a satisfactorycompatibility of the three methods applied to the alder standwas achieved. Key words: Alnus glutinosa, leaf area index, in situ counting, LAI-2000 PCA, litter collections     

Biogeochemical properties of a tropical swamp forest ecosystem in southern Thailand

The distributions of organic matter in the tropical swamps in southern Thailand are reported. The concentrations of particulate and dissolved organic carbon (POC and DOC) in the Bang Nara River, which drains swamp forests and nearby paddy fields, were 2.9 ± 2.0 and 6.2 ± 1.3 mg C l⁻¹, respectively. Although the variation was large, DOC concentration in the Bang Nara River seemed to be higher than POC in November 1992 (DOC/POC ratio, 2.8 ± 2.2). River waters from the upland areas were characterized by low POC and DOC concentrations as compared with Bang Nara River water. The δ¹³C values of POC and river sediments were useful to distinguish between organic matter originating in upland and swamp areas. It is suggested that the distributions of organic matter and its isotopic composition reflect the difference in drainage characteristics between lowland swamp and upland areas. Isotopic analyses of plant leaves and soils revealed that the swamp forest ecosystems were characterized by low δ¹³C and low δ¹⁵N values, which suggested low efficiency of water use by plants and large contributions of atmospheric deposition of nitrogen, respectively. Although CO₂ recycling in the forest might be an important factor determining the δ¹³C values of understory plants, the main process in carbon metabolism of tropical swamp forests would be CO₂ exchange between the atmosphere and forest canopy. Received: May 1, 2001 / Accepted: September 28, 2001     

Leaf litter decomposition in a tropical peat swamp forest in Peninsular Malaysia

It has long been assumed that the peat underlying tropical peat swamp forests accumulates because the extreme conditions (water logged, nutrient poor, anaerobic and acidic—pH 2.9–3.5) impede microbial activity. Litterbag studies in a tropical Malaysian peat swamp (North Selangor peat swamp forest) showed that although the sclerophyllous, toxic leaves of endemic peat forest plants (Macaranga pruinosa, Campnosperma coriaceum, Pandanus atrocarpus, Stenochlaena palustris) were barely decomposed by bacteria and fungi (decay rates of only 0.0006–0.0016 k day⁻¹), leaves of M. tanarius, a secondary forest species were almost completely decomposed (decay rates of 0.0047–0.005 k day⁻¹) after 1 year. Thus it is intrinsic properties of the leaves (that are adaptations to deter herbivory in the nutrient poor environment) that impede microbial breakdown. The water of the peat swamp was very high in dissolved organic carbon (70–84 mg l⁻¹ DOC). Laboratory studies revealed initial rapid leaching of DOC from leaves (up to 1,720 mg l⁻¹ from 4 g of leaves in 7 days), but the DOC levels then fell rapidly. The leaching of DOC resulted in weight loss but the physical structure of the leaves remained intact. It is suggested that the DOC is used as a substrate for microbial growth hence lowering the concentration of DOC in the water and transferring energy from the leaves to other trophic levels. This would explain how nutrient poor tropical peatswamps support diverse, abundant flora and fauna despite low nutrient levels and lack of rapid litter cycling such as occurs in other types of tropical rainforests.     

Soil CO2 efflux in a tropical forest in the central Amazon

This study investigated the spatial and temporal variation in soil carbon dioxide (CO₂) efflux and its relationship with soil temperature, soil moisture and rainfall in a forest near Manaus, Amazonas, Brazil. The mean rate of efflux was 6.45±0.25 SE μmol CO₂ m^?2s^?1 at 25.6±0.22 SE°C (5 cm depth) ranging from 4.35 to 9.76 μmol CO₂ m^?2s^?1; diel changes in efflux were correlated with soil temperature (r²=0.60). However, the efflux response to the diel cycle in temperature was not always a clear exponential function. During period of low soil water content, temperature in deeper layers had a better relationship with CO₂ efflux than with the temperature nearer the soil surface. Soil water content may limit CO₂ production during the drying‐down period that appeared to be an important factor controlling the efflux rate (r²=0.39). On the other hand, during the rewetting period microbial activity may be the main controlling factor, which may quickly induce very high rates of efflux. The CO₂ flux chamber was adapted to mimic the effects of rainfall on soil CO₂ efflux and the results showed that efflux rates reduced 30% immediately after a rainfall event. Measurements of the CO₂ concentration gradient in the soil profile showed a buildup in the concentration of CO₂ after rain on the top soil. This higher CO₂ concentration developed shortly after rainfall when the soil pores in the upper layers were filled with water, which created a barrier for gas exchange between the soil and the atmosphere.     

Nitrogen addition reduces soil respiration in a mature tropical forest in southern China

Response of soil respiration (CO₂ emission) to simulated nitrogen (N) deposition in a mature tropical forest in southern China was studied from October 2005 to September 2006. The objective was to test the hypothesis that N addition would reduce soil respiration in N saturated tropical forests. Static chamber and gas chromatography techniques were used to quantify the soil respiration, following four‐levels of N treatments (Control, no N addition; Low‐N, 5 g N m^?2 yr^?1; Medium‐N, 10 g N m^?2 yr^?1; and High‐N, 15 g N m^?2 yr^?1 experimental inputs), which had been applied for 26 months before and continued throughout the respiration measurement period. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates found in the warm and wet growing season (April–September) and the lowest rates in the dry dormant season (December–February). Soil respiration rates showed a significant positive exponential relationship with soil temperature, whereas soil moisture only affect soil respiration at dry conditions in the dormant season. Annual accumulative soil respiration was 601±30 g CO₂‐C m^?2 yr^?1 in the Controls. Annual mean soil respiration rate in the Control, Low‐N and Medium‐N treatments (69±3, 72±3 and 63±1 mg CO₂‐C m^?2 h^?1, respectively) did not differ significantly, whereas it was 14% lower in the High‐N treatment (58±3 mg CO₂‐C m^?2 h^?1) compared with the Control treatment, also the temperature sensitivity of respiration, Q₁₀ was reduced from 2.6 in the Control with 2.2 in the High‐N treatment. The decrease in soil respiration occurred in the warm and wet growing season and were correlated with a decrease in soil microbial activities and in fine root biomass in the N‐treated plots. Our results suggest that response of soil respiration to atmospheric N deposition in tropical forests is a decline, but it may vary depending on the rate of N deposition.     

15N natural abundance in forest and pasture soils of the Brazilian Amazon Basin

The natural abundance of ¹⁵N was examined in soil profiles from forests and pastures of the Brazilian Amazon Basin to compare tropical forests on a variety of soil types and to investigate changes in the sources of nitrogen to soils following deforestation for cattle ranching. Six sites in the state of Rondônia, two sites in Pará and one in Amazonas were studied. All sites except one were chronosequences and contained native forest and one or more pastures ranging from 2 to 27 years old. Forest soil ¹⁵N values to a depth of 1 m ranged from 8 to 23 and were higher than values typically found in temperate forests. A general pattern of increasing ¹⁵N values with depth near the soil surface was broadly similar to patterns in other forests but a decrease in ¹⁵N values in many forest profiles between 20 and 40 cm suggests that illuviation of ¹⁵N-depleted nitrate may influence total soil ¹⁵N values in deeper soil where total N concentrations are low. In four chronosequences in Rondônia, the ¹⁵N values of surface soil from pastures were lower than in the original forest and ¹⁵N values were increasingly depleted in older pastures. Inputs of atmospheric N by dinitrogen fixation could be an important N source in these pastures. Other pastures in Amazonas and Pará and Rondônia showed no consistent change from forest values. The extent of fractionation that leads to ¹⁵N enrichment in soils was broadly similar over a wide range of soil textures and indicated that similar processes control N fractionation and loss under tropical forest over a broad geographic region. Forest ¹⁵N profiles were consistent with conceptual models that explain enrichment of soil ¹⁵N values by selective loss of ¹⁴N during nitrification and denitrification.     

Epiphytic bromeliad communities in secondary and mature forest in a tropical premontane area

We analyzed the differences in species richness, community composition, population structure and within-tree location of epiphytic bromeliads in contiguous secondary and mature forests in a premontane area in Costa Rica. Diversity in the mature forest was highest, and the communities differed in their composition as well as in the recruitment rates of the dominant species. Guzmania monostachia and Catopsis nutans dominated the secondary forests, whereas Tillandsia fasciculata and T. tricolor were more abundant in the mature forest. The secondary forest species showed high rates of seedling recruitment while the opposite was found for the mature forest species. Species presence and abundance among and within habitats did not correlate with their physiological (i.e. CAM vs. C₃ photosynthesis) or morphological attributes. The spatial distribution patterns were similar among habitats; bromeliads tended to aggregate on a few relatively large phorophytes. The species shared a similar vertical stratification within habitats, except for the two dominant species in the early and mid-successional stages, although its ecological implication is not clear. With some exceptions, conspecifics of different ages were located on similar substrate types (i.e. stems, primary, secondary, or tertiary branches) within the tree-crowns, which suggests limited within-tree dispersion. Differences in species composition and rates of seedling recruitment among secondary and mature forest may arise from ecophysiological differences among species; however, the combined effect of seed availability and dispersal differences may have a larger influence. Thus, epiphyte community assembly can only be understood when the differences in habitat conditions, the availability of propagules, their dispersal characteristics and requirements for seedling establishment are known.     

Leaf age and the timing of leaf abscission in two tropical dry forest trees

We used experimental defoliations to examine the effect of leaf age on the timing of leaf shedding in two tropical dry forest trees. Trees of the deciduous Bombacopsis quinata (bombacaceae, a.k.a. Pachira quinata) and the brevi-deciduous Astronium graveolens (anacardiaceae) were manually defoliated for three times during the rainy season. All trees started to produce a new crown of leaves 2 weeks after defoliation, and continued expanding leaves throughout the rainy season. At the transition to the dry season, the experimental groups consisted of trees with known differences in maximum leaf age. Defoliations resulted in declines in stem growth but did not affect the mineral content or water relations of the leaves subsequently produced. There was no effect of leaf age on the timing of leaf abscission in B. quinata. In A. graveolens, the initiation of leaf shedding followed in rank order, the maximum leaf age of the four treatments, but there was substantial coherence among treatments in the major period of leaf abscission such that trees completed leaf shedding at the same time. In the two species, leaf water potential (Ψ_L) and stomatal conducantce (g _S) declined with the onset of the dry season, reaching minimum values of –0.9 MPa in P. quinata and <–2.0 MPa in A. graveolens. Within each species, leaves of different age exhibited similar Ψ_L and g _S at the onset of drought, and then decreased at a similar rate as the dry season progressed. Overall, our study suggests that the environmental factors were more important than leaf age in controlling the timing of leaf shedding.     

Leaf and photosynthetic characteristics of pioneer and forest species in tropical montane habitats

Gas exchange and chlorophyll a fluorescence measurements of expanding and adult leaves of four plant species were compared under field conditions. The pioneer species (PS) tended to have thinner leaves with lower nitrogen content and higher stomatal density compared to forest species (FS). Expanding leaves featured lower photosynthetic pigment contents and gas exchange capacity than adult leaves consistent with an immature photosynthetic apparatus. At the time of maximum irradiance, sun-exposed leaves of both PS and FS showed alteration of initial, variable, and maximum fluorescence as well as their ratios indicating photoinhibition. However, leaves recovered to some extent at predawn, suggesting the activation of photoprotective mechanisms. Sun-exposed leaves had comparable responses to high irradiance.     

Regulation of xylem sap flow in an evergreen, a semi-deciduous, and a deciduous Meliaceae species from the Amazon

The significance of phenological characteristics, stomatal conductance of the leaves, and stem water storage fluctuations for the regulation of xylem sap flow in an evergreen (Carapa guianensis Aubl.), in a semi-deciduous (Swietenia macrophylla King), and in a deciduous (Cedrela odorata L.) Meliaceae species was studied in a 7-year-old plantation near Manaus, Brazil. The study responds to the increasing demand for knowledge on the water relations of highly exploited timber trees of the Amazon. Xylem sap flow measurements indicated that the daily sap flow of Carapa (3.8 l day^-1 tree^-1 to 16.4 l day^-1 tree^-1) exceeded the daily sap flow of Swietenia (2.4 l day^-1 tree^-1 to 7.0 l day^-1 tree^-1) and Cedrela (1.6 l day^-1 tree^-1 to 11.6 l day^-1 tree^-1) during the entire year, although the highest flux densities were measured in Cedrela. The decrease in xylem sap flow observed in periods with low soil water potentials and high atmospheric vapor saturation deficits was more pronounced in the deciduous (Cedrela) and semi-deciduous species (Swietenia) than the evergreen species (Carapa). Carapa, which has the highest daily sap flow, had the highest biomass and sapwood portion. The high flux densities measured in Cedrela most likely result from the large earlywood vessels in this species. The seasonal variation of xylem sap flow of the three species was correlated with the stomatal conductance of the leaves measured by infiltration experiments. Stem water storage fluctuations in Carapa and Swietenia were predominantly due to transpiration; in Cedrela it was predominantly due to evaporative water loss on the stem surface during dry periods.     

Roadless wilderness area determines forest elephant movements in the Congo Basin

A dramatic expansion of road building is underway in the Congo Basin fuelled by private enterprise, international aid, and government aspirations. Among the great wilderness areas on earth, the Congo Basin is outstanding for its high biodiversity, particularly mobile megafauna including forest elephants (Loxodonta africana cyclotis). The abundance of many mammal species in the Basin increases with distance from roads due to hunting pressure, but the impacts of road proliferation on the movements of individuals are unknown. We investigated the ranging behaviour of forest elephants in relation to roads and roadless wilderness by fitting GPS telemetry collars onto a sample of 28 forest elephants living in six priority conservation areas. We show that the size of roadless wilderness is a strong determinant of home range size in this species. Though our study sites included the largest wilderness areas in central African forests, none of 4 home range metrics we calculated, including core area, tended toward an asymptote with increasing wilderness size, suggesting that uninhibited ranging in forest elephants no longer exists. Furthermore we show that roads outside protected areas which are not protected from hunting are a formidable barrier to movement while roads inside protected areas are not. Only 1 elephant from our sample crossed an unprotected road. During crossings her mean speed increased 14-fold compared to normal movements. Forest elephants are increasingly confined and constrained by roads across the Congo Basin which is reducing effective habitat availability and isolating populations, significantly threatening long term conservation efforts. If the current road development trajectory continues, forest wildernesses and the forest elephants they contain will collapse.     

Siluro-Devonian miospore biostratigraphy of the Urubu River area, western Amazon Basin, northern Brazil

Well-preserved miospores have been observed in the Urubu River area from the western part of the Amazon Basin, northern Brazil, in rocks belonging to the uppermost Pitinga and Manacapuru Formations. A previous palynological work based on chitinozoans has dated these strata as Late Ludlow, Early Pridoli and Early Lochkovian. Although samples are relatively poor in miospores, the present study reveals the most diversified Silurian-Earliest Lochkovian palynoflora ever reported from South America, an interval where miospore information is still very rare. Sixty-four species have been identified, of which two are new (Artemopyra urubuense and Retusotriletes amazonensis). One single specimen attributed to the genus Grandispora, identical to unpublished forms identified by one of us (P.S.) in Lochkovian sections of Saudi Arabia, has been found in a Lochkovian sample from the Urubu River. These constitute the oldest geological record of Grandispora. Among the abundant acritarchs, specimens of the genus Schizocystia have been observed for the first time in pre-Devonian rocks.Although the Late Silurian-Early Lochkovian miospore biostratigraphy is still in its infancy in South America, it seems that Amazon Basin assemblages do not show significant differences in composition in comparison to coeval palynofloras from other areas of northwestern Gondwana. This is surprising since the miospore localities under consideration are widely distributed over high to low paleolatitudes, i.e. from cool temperate to dry subtropical belts. Conversely, Siluro-Devonian Gondwanan assemblages differ considerably from the contemporary palynofloras of the Old Red Sandstone Continent and adjacent areas, thus contradicting recent phytogeographic interpretations based on megafloras.