Bamboo,fire and flood: consequences of disturbance for the vegetative growth of a clumping,clonal plant

The clumping growth form of tropical bamboos suggests a consolidator strategy in the face of intense competition rather than an ability to exploit disturbance. We investigated the annual growth and culm demographic responses to disturbance by fire and flood of vegetatively mature clumps of a riparian stand of Bambusa arnhemica. Linear-mixed models were employed to control for the non-independence of culm fates within clumps and clump growth among years. As the stand was monodominant and the species is gregariously monocarpic, responses can be interpreted independently of interspecific competition and trade-offs with sexual reproduction. Disturbance depressed clump growth but the culm demographic responses to fire and flood were quite different. Few culms were destroyed during the fire but damage depressed their subsequent survival, leading to declines in clump basal area. Fire also triggered the release of ramet buds, but the additional recruits were small and short-lived. Prolonged and early flooding in one of the study years depressed culm recruitment in clumps low on the lagoon bank but there was possible compensatory recruitment in the following year. There was no convincing evidence that sparse clumps recruited culms better than dense clumps, though culm recruitment was negatively correlated with culm survival. Fire and prolonged flooding are inevitable elements of the environment occupied by B. arnhemica, but we interpret the species’ response as persisting in the face of disturbance rather than exploiting the opportunities created by it.     

A multi-scale biogeographical analysis of Bambusa arnhemica, a bamboo from monsoonal northern Australia

Aims To identify the edaphic, environmental and historical factors influencing the patchy distribution of the semelparous bamboo Bambusa arnhemica F. Muell. at global, catchment and streambank scales. Location The entire range of B. arnhemica, a highly fire‐prone savanna matrix with generally infertile soils in the north‐west of the Northern Territory of Australia above the 1200 mm mean annual rainfall isohyet. Methods Distribution surveys were conducted by air, boat and on the ground. Plot data were collected throughout the range of the species. Results Bambusa arnhemica occurred predominantly in gallery forests on flood‐prone but nevertheless well‐drained and deep alluvial soils on sloping stream banks. It ranged widely along lentic watercourses from ephemeral headwater streams to the banks of major rivers and levees on the coastal floodplain. The species did not occur in savannas; savannas adjacent to B. arnhemica gallery forests were also flood‐prone and on deep alluvial soils, but were upslope on level ground. Bambusa arnhemica's infrequent non‐riparian occurrences were on a wide variety of substrates but generally on soils of moderate fertility and in coastal and/or rocky areas where at least partial topographic protection from fire is likely. Within and between catchments, the distribution of B. arnhemica was idiosyncratic, occurrence being almost always continuous downstream from highly variable ‘starting’ points to the poorly drained coastal floodplain. Main conclusions At local scales, B. arnhemica appears constrained by poor drainage and high fire‐frequencies. Enhanced soil fertility may increase its capacity to cope with fire. At the catchment and global scales, we propose that the distribution of B. arnhemica is the product of infrequent and as yet incomplete dispersal across and away from watercourses by seed that lacks specialized dispersal mechanisms, combined with passive dispersal along streams. From this we infer that B. arnhemica is neither a very recent, nor very ancient colonist from Asia. Bambusa arnhemica's circumscribed global distribution has no parallel amongst co‐occurring rain forest plants and may be the product of poor dispersal capacity and a substantial rock and floodplain barrier to the east. Limited dispersal capacity may be inextricably linked to local domination of space and the subsequent creation of regeneration space by parental death.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

Development of microsatellite markers for <Emphasis Type="Italic">Bambusa arnhemica</Emphasis> (Poaceae: Bambuseae), a bamboo endemic to northern Australia

Bambusa arnhemica is a bamboo species endemic to northern Australia. We isolated and characterized nine microsatellite loci from this species. The number of alleles ranged from 2 to 16 with an average of 6.8, and expected heterozygosities from 0.40 to 0.84 with an average of 0.69. The markers described here will be useful to investigate clump structure, evolution of the bamboo flowering wave, patterns of gene flow, and the biogeographic history of B. arnhemica in Australia.     

Phenology of Acacia albida trees in Zambezi riverine woodlands

The phenology of Acacia albida trees was studied over eight years in Mana Pools National Park, Zimbabwe. Leaf flush occurred towards the end of the rains. It was followed by flowering, with ripe fruit-fall in the late dry season. Leaf-fall occurred early in the following wet season. After seasons of low rainfall, trees were severely defoliated by caterpillars soon after leaf flush and this prompted a second flush of leaves. In these years there were also two periods of flowering and fruit production. During dry years, the proportion of trees which failed to set fruits after flowering increased, fruit production by those trees which did bear fruit was reduced and fruit-fall started earlier in the year.     

Abundance and Leaf Damage Caused by Gall‐Inducing Insects in a Mexican Tropical Dry Forest1

We evaluated the temporal and spatial patterns of abundance and the amount of damage caused by gall‐inducing insects (GII) in deciduous and riparian habitats in a seasonal tropical dry forest in Mexico. Plants occurring in these habitats differ in their phenology and moisture availability. Deciduous habitats are seasonal and xeric, while riparian habitats are aseasonal and mesic. We found 37 GII species and each one was associated with a specific plant species. In total, 19 species (51.3%) were present in deciduous habitats, 13 species (35.2%) in riparian habitats, and only 5 species (13.5%) occurred in both. Abundance and leaf damage by GII were greater in deciduous than in riparian habitats during the wet season. For each GII species that occurred in both habitats, host plant species supported greater abundance and leaf damage by GII in deciduous habitats during the wet season. These results indicate a greater association of GII species with host plants in deciduous than in riparian habitats during the wet season. In riparian habitats, 11 plant species (61.1%) had greater density of GII in the dry than in the wet season. Similarly, leaf damage by GII was significantly greater in the dry than in the wet season in riparian habitats for 12 plant species (66.7%). Dry forest plants of riparian habitats presented two peaks of leaf‐flushing: GII colonized leaves produced in the first peak at the beginning of the wet season, and accumulated or recolonized leaves in the second peak at the beginning of the dry season. The levels of leaf damage by GII detected in this study in the rainy season were considerably higher than those obtained for folivorous insects in other neotropical forests, suggesting that this GII guild might have an important impact on their host plant species in this tropical community.     

Monsoonal influences on evapotranspiration of savanna vegetation of northern Australia

Data from savannas of northern Australia are presented for net radiation, latent and sensible heat, ecosystem surface conductance (G_s) and stand water use for sites covering a latitudinal range of 5° or 700 km. Measurements were made at three locations of increasing distance from the northern coastline and represent high- (1,750 mm), medium- (890 mm) and low- (520 mm) rainfall sites. This rainfall gradient arises from the weakened monsoonal influence with distance inland. Data were coupled to seasonal estimates of leaf area index (LAI) for the tree and understorey strata. All parameters were measured at the seasonal extremes of late wet and dry seasons. During the wet season, daily rates of evapotranspiration were 3.1-3.6 mm day^-1 and were similar for all sites along the rainfall gradient and did not reflect site differences in annual rainfall. During the dry season, site differences were very apparent with evapotranspiration 2-18 times lower than wet season rates, the seasonal differences increasing with distance from coast and reduced annual rainfall. Due to low overstorey LAI, more than 80% of water vapour flux was attributed to the understorey. Seasonal differences in evapotranspiration were mostly due to reductions in understorey leaf area during the dry season. Water use of individual trees did not differ between the wet and dry seasons at any of the sites and stand water use was a simple function of tree density. G_s declined markedly during the dry season at all sites, and we conclude that the savanna water (and carbon) balance is largely determined by G_s and its response to atmospheric and soil water content and by seasonal adjustments to canopy leaf area.     

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.     

Growth response of Halimium halimifolium at four sites with different soil water availability regimes in two contrasted hydrological cycles

The relationship between water availability and plant growth response in Halimium halimifolium (L.) Willk throughout two years with contrasted precipitation (300 and 850 mm) was examined by measuring vegetative growth and midday leaf water potential in four sandy soil sites with different water availability in Doñana National Park, Spain. H. halimifolium, Cistaceae is mostly restricted to sandy substrates close to coastal Mediterranean areas. At Doñana National Park this species is the main component of stable sand vegetation. Vegetative growth started in March, ending in July. The maximum shoot elongation rate (110 mm/year) and number of branches (8.3 branches/year) occurred in plants growing in the most hygrophytic site (MN) in both the wet and dry years. Plants at this site exhibited higher midday water potentials throughout the year. In contrast, the minimum shoot elongation rate (40 mm/year) and midday water potential (–4.0 MPa) occurred in Monte Intermedio plants (MI) in the dry year, although the water table was shallower than in Monte Blanco (MB). In the wet year the minimum shoot and branch elongation were recorded in MB. The number of leaves per branch for a single main shoot was higher (55 leaves/branch) in the driest area (MB), but these leaves had the smallest area. ANOVA showed significant differences in growth response and midday water potential between the four sites. A stepwise multiple linear regression showed that midday water potential, water table depth and monthly average temperature were the variables most closely associated with shoot elongation rate. We conclude that under severe dryness, the expected patterns of greater growth in sites with better water supply may differ from compared with the expected growth pattern in normal conditions.     

Effects of High Nutrient Supply on the Growth of Seven Bamboo Species

Over the last decade, bamboo has emerged as an interesting plant for the treatment of various polluted waters using plant-based wastewater treatment systems. In these systems, nitrogen and phosphorous concentrations in wastewater can exceed plant requirements and potentially limit plant growth. The effects of two nutrient rates on the growth of seven bamboo species were assessed in a one-year experiment: Dendrocalamus strictus, Thyrsostachys siamensis, Bambusa tuldoides, Gigantochloa wrayi, Bambusa oldhamii, Bambusa multiplex and Bambusa vulgaris. Nutrient rates were applied with a 20:20:20 NPK fertilizer as 2.6 and 13.2 t.ha.yr^?1NPK to three-year-old bamboo planted in 70 L containers. Morphological characters, photosynthetic responses, and NPK content in bamboo tissues were investigated. Under high-nutrient supply rate, the main trend observed was an increase of culm production but the culms’ diameters were reduced. For the seven species, the aboveground biomass yield tended to increase with high-nutrient rate. Increasing in nutrient rates also improved the photosynthetic activity which is consistent with the increase of nitrogen and phosphorus contents measured in plant tissues. All the bamboo species tested appears suitable for wastewater treatment purposes, but the species Bambusa oldhamii and Gigantochloa wrayi showed the higher biomass yield and nutrient removal.     

Seasonality of insect herbivory on the leaves of Neoboutonia macrocalyx in the Kibale National Park, Uganda

The seasonality of herbivory on the leaves of Neoboutonia macrocalyx Pax. in Kibale Forest National Park, Uganda, was studied. A total of 2929 fallen leaves was collected during 15 months under randomly-selected trees in three different habitats; natural forest and two selectively cut forest sites. The percentage of leaf area eaten and leaf size were estimated. Leaf herbivory was highly seasonal and correlated with rainfall in the previous 2 months, but less than 100 mm monthly rainfall had no effect. There was no correlation between leaf size and rainfall. Although Kibale Forest has two wet seasons, insect feeding on leaves had only one peak during the major rainy season from September to December. Three to four months after peak herbivory, leaves had very low rates of insect damage. Habitat had only a small effect on the amount of insect feeding. The sampling time accounted for 71% of variation in leaf herbivory. New leaves were formed continuously year-round. The constant leaf production by Neoboutonia trees may be an adaptation to escape generalist herbivorous insects which might be synchronized with the major wet season when the leaf flush of the most other deciduous species occurs. Thus, the availability of fresh leaves is not acting as a regulating factor in seasonality of Neoboutonia herbivory.     

Clonal regeneration of an arrow bamboo, <Emphasis Type="Italic">Fargesia qinlingensis</Emphasis>, following giant panda herbivory

Characteristics of giant panda herbivory sites and clonal regeneration of an arrow bamboo Fargesia qinlingensis following giant panda grazing were studied in the Qinling Mountains of China. Three types of plots were located in a pandas’ summer habitat in 2002: herbivory (naturally grazed by giant pandas), clipped (simulated panda herbivory), and control. Average area of herbivory sites was 2.92 m² and average distance from herbivory sites to the closest tree (dbh > 10 cm) was 1.0 m. Pandas avoided thin bamboo culms with basal diameters <5 mm. Average height of stumps of culms grazed by panda was 0.67 m and average density of grazed culms was 9.0 culms m⁻². Annual culm mortality rate was significantly greater in herbivory and clipped plots than in control plots while annual recruitment rate was not significantly different among the three plot types in 2003. Neither recruitment rate nor mortality rate were significantly different among the three plot types in 2004. Annual recruitment rate was significantly greater than annual mortality rate only in control plots in both 2003 and 2004, suggesting static ramet dynamics in disturbed plots (herbivory and clipped). Density of new shoots was not significantly different, but basal diameter of new shoots was significantly less in herbivory plots compared to control plots in 2002. Differences of annual mortality rate and growth of new shoots found between control plots and herbivory plots suggest that clonal regeneration of F. qinlingensis culms was negatively affected by giant panda grazing. Therefore, no evidence of a clonal integration compensatory response to panda herbivory was found in F. qinlingensis.     

Potential of the mite-pathogenic fungus Neozygites floridana (Entomophthorales: Neozygitaceae) for control of the cassava green mite Mononychellus tanajoa (Acari: Tetranychidae)

The cassava green mite, Mononychellus tanajoa (Bondar), is an exotic pest in Africa and is the target of a classical biological control programme. Field data from the Neotropics, where it is indigenous, are presented for the first time, charting the variation in abundance of M. tanajoa over several seasons. This was highly variable, with a characteristic trough mid-year and a peak at the turn of the year. This pattern corresponded positively with rainfall levels, appearing to fit a phenology also characteristic of African studies, where rainfall at the start of the wet season promotes a leaf flush and so growth in M. tanajoa populations. Analyses implied some impact of leaf-inhabiting predatory mites (predominantly Neoseiulus idaeus Denmark & Muma) and a considerable impact of the fungal pathogen Neozygites floridana Fisher on M. tanajoa populations. This pathogen was not observed in the host population for several (generally dry) periods implying survival outside the host, perhaps as resting spores. This is a particularly desirable characteristic of a biological control agent. It is therefore proposed that N. floridana might be of particular use in drier cassava-growing areas where rainfall at the outset of the wet season is not sufficiently intense to cause heavy M. tanajoa mortality but may be sufficient to stimulate epizootics of the fungal pathogen, protecting the flush of new cassava growth.     

Combined effects of light and water availability on photosynthesis and growth of Arisaema heterophyllum in the forest understory and an open site

Photosynthetic characteristics, leaf longevity and biomass accumulation of a threatened herb species, Arisaema heterophyllum, were studied in the understory of a riparian forest and at a neighboring deforested open site for 3 years in order to understand the combined effects of light and water availability. Light availability was 2- to 4-fold higher at the deforested than at the forest site during the growing season of the species, and precipitation varied considerably over the 3 years. Despite the difference in water availability among the years (dry in 1994 and 1996, and wet in 1995), the species showed a strong acclimation to the different light environments. Light-saturated assimilation rate on a leaf area basis, leaf mass area ratio (LMA), and relative growth rate (RGR) were all higher at deforested site. While a positive correlation between individual RGR and microsite light availability was found in the wet year, no correlation was found in the dry years, and mean RGR was significantly lower in the dry year at both sites. Leaf longevity, photosynthetic capacity on a leaf mass basis, dark respiration rate, and leaf conductance, varied considerably from year to year, especially in the plants at the open site, probably depending on water availability. In the dry years plants at the deforested sites showed a lower photosynthetic rate and leaf conductance under unwatered than under watered conditions. These results suggest that the water availability in a given year may strongly affect light acclimation and annual RGR of the herb species in natural habitats, even under mesic climate conditions. Received: 15 February 1997 / Accepted: 20 May 1997     

Effects of precipitation and soil water potential on drought deciduous phenology in the Kalahari

We utilized an ecosystem process model to investigate the influence of precipitation and soil water potential on vegetation phenology in the semi‐arid, drought‐deciduous ecosystems in the Kalahari region of South Africa. The timing of leaf flush was assumed to be the first day during which a rainfall event exceeded that day's estimate of potential evapotranspiration after a defined dry season. Leaf senescence was assumed to be a dynamic feedback between soil water potential and net plant carbon gain and was determined by dynamically modeling the effects of concomitant trends in soil water potential and net primary production on leaf area index (LAI). Model predictions of LAI were compared with satellite‐derived normalized difference vegetation indices (NDVI) for 3 years at two sites along the Kalahari transect. The mean absolute error for the prediction of modeled leaf flush date compared with leaf flush dates estimated from NDVI were 10.0 days for the Maun site and 39.3 days for the Tshane site. Correlations between model predicted 10‐day average LAI and 10‐day composite NDVI for both Maun and Tshane were high (ρ=0.67 and 0.74, respectively, P<0.001), suggesting that this method adequately predicts intra‐annual leaf area dynamics in these dry tropical ecosystems.     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Soil nutrient status after bamboo flowering and death in a seasonal tropical forest in western Thailand

We have examined the surface (0–10 cm) soil characteristics of sites after bamboo (Cephalostachyum pergracile) mass flowering and death (DB sites) in comparison with sites with living bamboo (Bambusa tulda) (LB sites) in a seasonal tropical forest in Thailand. One year after bamboo flowering the DB sites were acidic with lower concentrations of exchangeable Ca and Mg and soil nitrogen than the LB sites. Therefore, although leaf and root litter of the dead bamboo was deposited in the DB sites after bamboo flowering, soil nutrient status decreased.     

Do seasonal changes in light availability influence the inverse leafing phenology of the neotropical dry forest understory shrub Bonellia nervosa (Theophrastaceae)?

In tropical dry forests most plants are deciduous during the dry season and flush leaves with the onset of the rains. In Costa Rica, the only species displaying the opposite pattern is Bonellia nervosa. To determine if seasonal changes in light availability are associated with the leaf and reproductive phenology of this species, we monitored leaf production, survival, and life span, as well as flower and fruit production from April 2000 to October 2001 in Santa Rosa National Park. Leaf flushing and flower bud production took place shortly after the autumnal equinox when day length starts to decrease. Leaves began expansion at the end of the wet season, and plants reached 70 % of their maximum leaf area at the beginning of the dry season, maintaining their foliage throughout the entire dry period. Leaf shedding occurred gradually during the first three months of the wet season. Leaf flushing and shedding showed high synchrony, with leaf numbers being related to light availability. Maximum leaf production coincided with peaks in radiation during the middle of the dry season. Decreasing day length induces highly synchronous flower bud emergence in dry forest species, but this is the first study indicating induction of leaf flushing by declining day length.     

Effect of rainfall pattern on nitrogen mineralization and leaching in a green manure experiment in South Rwanda

The effects of green manures, sorghum residues and farmyard manure on N dynamics and crop yields were studied during three dry and wet seasons on a Typic Sombriudox in South Rwanda. In addition, a resin core study was conducted within a 4-year green manure field experiment to follow the seasonal pattern of N mineralization and leaching after application of residues from Tephrosia vogelii, Sorghum bicolor, a mixture of both materials, and farmyard manure.During the dry season, topsoil (0–20 cm) mineral N remained constant. At the beginning of the wet season, the rainfall pattern determined N availability. With low rainfall intensities a mineralization flush occurred, doubling topsoil mineral N concentrations within 5 days after wetting. In contrast, under heavy rains at the onset of the rainy season, topsoil mineral N decreased by 50–70% within the first two weeks.The application of organic fertilizers has a strong influence on N availability, but the effects can be negated by heavy rainfall. Incorporation of leaves from Tephrosia vogelii (2.7 t dm ha^-1) and farmyard manure (7 t dm ha^-1) doubled the mineralization flush after the first rains. During the rest of the wet season, N release by the green manure was small, whereas the farmyard manure was found to mobilize N after a period of N immobilization. Incorporation of sorghum residues had only a small effect, while mixing the straw with green and farmyard manure immobilized N temporarily.Nitrogen leaching, measured by exchange resins at a depth of 20 cm, was increased up to 50% by the incorporation of green and farmyard manure. This points to rapid N translocation of easily mineralizable N. The additional incorporation of sorghum residues reduced N leaching of both materials significantly. Since rainfall is often unpredictable, the synchronization of N released from crop residues with crop N demand may require additional management practices.