Water status and development of tropical trees during seasonal drought

Summary Bud break, shoot growth and flowering of trees involve cell expansion, known to be inhibited by moderate water deficits. In apparent contradiction to physiological theory, many trees flower or exchange leaves during the 6 month-long, severe dry season in the tropical dry forest of Guanacaste, Costa Rica. To explore this paradox, changes in tree water status during the dry season were monitored in numerous trees. Water potential of stem tissues (_stem) was obtained by a modification of the pressure chamber technique, in which xylem tension was released by cutting defoliated branch samples at both ends. During the early dry season twigs bearing old, senescent leaves generally had a low leaf water potential (_leaf), while _stem varied with water availability. At dry sites, _stem was very low in hardwood trees (<–4 MPa), but near saturation (>–0.2 MPa) in lightwood trees storing water with osmotic potentials between –0.8 and –2.1 MPa. At moist sites trees bearing old leaves rehydrated during drought; their _stem increased from low values (<–3 MPa) to near saturation, resulting in differences of 3–4 MPa between _stem and _leaf. Indirect evidence indicates that rehydration resulted from osmotic adjustment of stem tissues and improved water availability due to extension of roots into moist subsoil layers. In confirmation of physiological theory, elimination of xylem tension by leaf shedding and establishment of a high solute content and high _stem were prerequisites for flowering and bud break during drought.     

Leaf productivity along a precipitation gradient in lowland Panama: patterns from leaf to ecosystem

Moisture availability has the potential to affect tropical forest productivity at scales ranging from leaf to ecosystem. We compared data for leaf photosynthetic, chemical and structural traits of canopy trees, litterfall production and seasonal availability of soil water at four sites across a precipitation gradient (1,800–3,500 mm year^–1) in lowland Panamanian forest to determine how productivity at leaf and ecosystem scales may be related. We found stronger seasonality in soil water potential at drier sites. Values were close to zero at all sites during the wet season and varied between a minimum of –2.5 MPa and –0.3 MPa at the driest and wettest sites, respectively, during the dry season. Leaf photosynthesis and nitrogen concentration decreased with increasing precipitation, whereas leaf thickness increased with increasing precipitation. Leaf toughness and fiber/N ratios increased with increasing precipitation indicating reduced nutritional content and palatability with precipitation. Seasonality of litter production and quality decreased with increasing precipitation, but the amount of litterfall produced was not substantially different among sites. It appears that in Neotropical forest, moisture availability is associated with leaf photosynthetic and defensive traits that influence litterfall timing and quality. Therefore, variation in leaf physiological traits has the potential to influence decomposition and nutrient cycling through effects on litter quality.     

Nitrogen transformations and nitrous oxide flux in a tropical deciduous forest in México

Summary Emissions of nitrous oxide and soil nitrogen pools and transformations were measured over an annual cycle in two forests and one pasture in tropical deciduous forest near Chamela, México. Nitrous oxide flux was moderately high (0.5–2.5 ng cm^–2 h^–1) during the wet season and low (<0.3 ng cm^–2 h^–1) during the dry season. Annual emissions of nitrogen as nitrous oxide were calculated to be 0.5–0.7 kg ha^–1 y^–1, with no substantial difference between the forests and pasture. Wetting of dry soil caused a large but short-lived pulse of N₂O flux that accounted for <2% of annual flux. Variation in soil water through the season was the primary controlling factor for pool sizes of ammonium and nitrate, nitrogen transformations, and N₂O flux.     

Aspects of tissue water relations and seasonal changes of leaf water potential components of evergreen and deciduous species coexisting in tropical dry forests

Summary This study compared the tissue water relations and seasonal changes in leaf water potential components of an evergreen tree,Morisonia americana, and two evergreen shrubs,Capparis verrucosa andC. aristiquetae, with two deciduous trees,Humboltiella arborea andLonchocarpus dipteroneurus, and the deciduous vineMansoa verrucifera. All these species coexist in a tropical dry forest in Venezuela. Leaves of the evergreen species are sclerophyllous, while those of the deciduous species are mesophytic. Leaf area to leaf weight ratios of fully mature leaves were about 75 and 170 cm² g^–1 in evergreen and deciduous species, respectively. Seasonal fluctuations of leaf water content per unit of dry weight, water potential, and turgor pressure were smaller in evergreen than in deciduous species. The analysis of tissue water relations using pressurevolume curves showed that evergreen species could develop a higher leaf turgor and lose turgor at lower leaf water potentials than deciduous species. This was related to a lower osmotic potential at full turgor in evergreen (-3.0 MPa)_than in deciduous (-2.0 MPa) species, rather than to the elastic properties of leaf tissue. The volumetric modulus of elasticity was 14 MPa in evergreen compared with 7–10 MPa in deciduous species. Thus, leaf characteristics are important in determining the drought resistance of evergreen species of this tropical dry forest.     

Phenology of temperate trees in tropical climates

Several North American broad-leaved tree species range from the northern United States at 47°N to moist tropical montane forests in Mexico and Central America at 15–20°N. Along this gradient the average minimum temperatures of the coldest month (T _Jan), which characterize annual variation in temperature, increase from –10 to 12°C and tree phenology changes from deciduous to leaf-exchanging or evergreen in the southern range with a year-long growing season. Between 30 and 45°N, the time of bud break is highly correlated with T _Jan and bud break can be reliably predicted for the week in which mean minimum temperature rises to 7°C. Temperature-dependent deciduous phenology—and hence the validity of temperature-driven phenology models—terminates in southern North America near 30°N, where T _Jan>7°C enables growth of tropical trees and cultivation of frost-sensitive citrus fruits. In tropical climates most temperate broad-leaved species exchange old for new leaves within a few weeks in January-February, i.e., their phenology becomes similar to that of tropical leaf-exchanging species. Leaf buds of the southern ecotypes of these temperate species are therefore not winter-dormant and have no chilling requirement. As in many tropical trees, bud break of Celtis, Quercus and Fagus growing in warm climates is induced in early spring by increasing daylength. In tropical climates vegetative phenology is determined mainly by leaf longevity, seasonal variation in water stress and day length. As water stress during the dry season varies widely with soil water storage, climate-driven models cannot predict tree phenology in the tropics and tropical tree phenology does not constitute a useful indicator of global warming.     

Forest structure and carbon dynamics in Amazonian tropical rain forests

Living trees constitute one of the major stocks of carbon in tropical forests. A better understanding of variations in the dynamics and structure of tropical forests is necessary for predicting the potential for these ecosystems to lose or store carbon, and for understanding how they recover from disturbance. Amazonian tropical forests occur over a vast area that encompasses differences in topography, climate, and geologic substrate. We observed large differences in forest structure, biomass, and tree growth rates in permanent plots situated in the eastern (near Santarém, Pará), central (near Manaus, Amazonas) and southwestern (near Rio Branco, Acre) Amazon, which differed in dry season length, as well as other factors. Forests at the two sites experiencing longer dry seasons, near Rio Branco and Santarém, had lower stem frequencies (460 and 466 ha^–1 respectively), less biodiversity (Shannon–Wiener diversity index), and smaller aboveground C stocks (140.6 and 122.1 Mg C ha^–1) than the Manaus site (626 trees ha^–1, 180.1 Mg C ha^–1), which had less seasonal variation in rainfall. The forests experiencing longer dry seasons also stored a greater proportion of the total biomass in trees with >50 cm diameter (41–45 vs 30% in Manaus). Rates of annual addition of C to living trees calculated from monthly dendrometer band measurements were 1.9 (Manaus), 2.8 (Santarém), and 2.6 (Rio Branco) Mg C ha^–1 year^–1. At all sites, trees in the 10–30 cm diameter class accounted for the highest proportion of annual growth (38, 55 and 56% in Manaus, Rio Branco and Santarém, respectively). Growth showed marked seasonality, with largest stem diameter increment in the wet season and smallest in the dry season, though this may be confounded by seasonal variation in wood water content. Year-to-year variations in C allocated to stem growth ranged from nearly zero in Rio Branco, to 0.8 Mg C ha^–1 year^–1 in Manaus (40% of annual mean) and 0.9 Mg C ha^–1 year^–1 (33% of annual mean) in Santarém, though this variability showed no significant relation with precipitation among years. Initial estimates of the C balance of live wood including recruitment and mortality as well as growth suggests that live wood biomass is at near steady-state in Manaus, but accumulating at about 1.5 Mg C ha^–1 at the other two sites. The causes of C imbalance in living wood pools in Santarém and Rio Branco sites are unknown, but may be related to previous disturbance at these sites. Based on size distribution and growth rate differences in the three sites, we predict that trees in the Manaus forest have greater mean age (~240 years) than those of the other two forests (~140 years).     

The significance of episodic rains for reproductive phenology and productivity of trees in semiarid regions of northwestern Venezuela

The semiarid regions of northwestern Venezuela have extremely low and highly unpredictable precipitation, yet these conditions support species with contrasting phenology and leaf longevity. Episodic rains significantly increased leaf water potential (from –5 to –2.5 MPa) in several species and, in some cases, triggered flowering, leading us to hypothesize that the coexistence of species with contrasting phenology is due to differences in their ability to utilize small rainfall events. Irrigation treatments were used to simulate brief rainfall events, and the response of three species (Erythrina velutina [deciduous], Croton heliaster [semideciduous], and Capparis odoratissima [evergreen]) was monitored over a period of 14 months. To partition the effects of water reaching the canopy versus the soil, irrigation was supplied either in the form of mist to the canopy or by minisprinklers near the base of the trees. Nonirrigated trees were used as controls. Productivity (estimated as aboveground litter production) and water potential were enhanced by soil irrigation in two species. However, in the evergreen species canopy irrigation had a greater effect on water relations and productivity than soil irrigation, as indicated by higher predawn water potential, higher total annual flower (40 g m^–2 year^–1) and fruit (5 g m^–2 year^–1) production, and longer leaf longevity (410 days in control trees versus 520 days in canopy-irrigated trees). Canopy irrigation augmented flower and fruit production in all three species. Our findings suggest that reproductive phenology in these species is driven by episodic rains and that evergreen species may sustain productivity by their ability to make use of water deposited on leaf surfaces.     

Modification of Vegetative Phenology in a Tropical Semi‐deciduous Forest by Abnormal Drought and Rain1

The control of vegetative phenology in tropical trees is not well understood. In dry forest trees, leaf abscission may be enhanced by advanced leaf age, increasing water stress, or declining photoperiod. Normally, it is impossible to dissect the effects of each of these variables because most leaves are shed during the early dry season when day length is near its minimum and leaves are relatively old. The 1997 El‐Niño Southern Oscillation caused a ten‐week long, severe abnormal drought from June to August in the semi‐deciduous forests of Guanacaste, Costa Rica. We monitored the effect of this drought on phenology and water status of trees with young leaves and compared modifications of phenology in trees of different functional types with the pattern observed during the regular dry season. Although deciduous trees at dry sites were severely water stressed (Ψstem < ‐7MPa) and their mesic leaves remained wilted for more than two months, these and all other trees retained all leaves during the abnormal drought. Many trees exchanged leaves three to four months earlier than normal during the wet period after the abnormal drought and shed leaves again during the regular dry season. Irrigation and an exceptional 70 mm rainfall during the mid‐dry season 1998/1999 caused bud break and flushing in all leafless trees except dormant stem succulents. The complex interactions between leaf age and water stress, the principal determinants of leaf abscission, were found to vary widely among trees of different functional types.     

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.     

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae werecompared to test in part a hypothesis that the forest understorey plays anintegral role in drought response. At Danum, Sabah, the relatively commonspecies Dimorphocalyx muricatus is associated with ridgeswhilst another species, Mallotus wrayi, occurs widely bothon ridges and lower slopes. Sets of subplots within two 4 -hapermanent plots in this lowland dipterocarp rain forest, were positioned onridges and lower slopes. Soil water potentials were recorded in1995–1997,and leaf water potentials were measured on six occasions. Soil water potentialson the ridges (–0.047 MPa) were significantly lower than onthe lower slopes (–0.012 MPa), but during the driest periodin May 1997 they fell to similarly low levels on both sites (–0.53MPa). A weighted 40-day accumulated rainfall index was developedtomodel the soil water potentials. At dry times, D.muricatus(ridge) had significantly higher pre-dawn (–0.21 v.–0.57 MPa) and mid-day (–0.59 v.–1.77 MPa) leaf water potentials than M.wrayi (mean of ridge and lower slope). Leaf osmotic potentials ofM. wrayi on the ridges were lower (–1.63MPa) than on lower slopes (–1.09 MPa), withthose for D. muricatus being intermediate (–1.29MPa): both species adjusted osmotically between wet and dry times.D. muricatus trees were more deeply rooted thanM. wrayi trees (97 v. 70cm). M. wrayi trees had greaterlateral root cross-sectional areas than D. muricatus treesalthough a greater proportion of this sectional area for D.muricatus was further down the soil profile. D.muricatus appeared to maintain relatively high water potentialsduring dry periods because of its access to deeper water supplies and thus itlargely avoided drought effects, but M. wrayi seemed to bemore affected yet tolerant of drought and was more plastic in its response. Theinteraction between water availability and topography determines these species'distributions and provides insights into how rain forests can withstandoccasional strong droughts.     

Water uptake and transport in lianas and co-occurring trees of a seasonally dry tropical forest

Water uptake and transport were studied in eight liana species in a seasonally dry tropical forest on Barro Colorado Island, Panama. Stable hydrogen isotope composition (D) of xylem and soil water, soil volumetric water content (_v), and basal sap flow were measured during the 1997 and 1998 dry seasons. Sap flow of several neighboring trees was measured to assess differences between lianas and trees in magnitudes and patterns of daily sap flow. Little seasonal change in _v was observed at 90–120 cm depth in both years. Mean soil water D during the dry season was –19 at 0–30 cm, –34 at 30–60 cm, and –50 at 90–120 cm. Average values of xylem D among the liana species ranged from –28 to –44 during the middle of the dry season, suggesting that water uptake was restricted to intermediate soil layers (30–60 cm). By the end of the dry season, all species exhibited more negative xylem D values (–41 to –62), suggesting that they shifted to deeper water sources. Maximum sap flux density in co-occurring lianas and trees were comparable at similar stem diameter (DBH). Furthermore, lianas and trees conformed to the same linear relationship between daily sap flow and DBH. Our observations that lianas tap shallow sources of soil water at the beginning of the dry season and that sap flow is similar in lianas and trees of equivalent stem diameter do not support the common assumptions that lianas rely primarily on deep soil water and that they have higher rates of sap flow than co-occurring trees of similar stem size.     

Effect of liana cutting on water potential and growth of adult<Emphasis Type="Italic"> Senna multijuga</Emphasis> (Caesalpinioideae) trees in a Bolivian tropical forest

Lianas, or woody climbing plants, are a major constituent of seasonally dry tropical forests, and are thought to impact negatively their host trees. In this study we evaluated whether liana presence was associated with reduced leaf water potentials and growth in adult Senna multijuga trees during the dry season in a lowland Bolivian forest. We used leaf water potentials in trees as a first approach to assess trees’ water status, under the assumption that leaf water potentials become more negative when water losses (via transpiration) exceed gains (by uptake). We measured relative growth in girth at 1.5 m height (gbh) to quantify tree growth. At the beginning of the 1996 dry season (early June), we selected 20 S. multijuga trees 10–20 cm dbh, and measured their gbh. We also recorded pre-dawn and mid-day leaf water potentials in these trees. In ten experimental trees all lianas were then cut, while the remaining trees were used as controls. Pre-dawn and mid-day water potentials were re-measured 1 day after liana-cutting, and then every week in all trees for 1 month and then at 3 and 5 months, until the beginning of the next rainy season (November); gbh was measured again in July 1997 to estimate relative growth rate. Liana removal was associated with less negative pre-dawn (–0.3 vs –0.4 MPa) and mid-day (–0.5 vs –0.7 MPa) water potentials in trees during the dry season. This difference appeared as early as 1 day after cutting, and disappeared once the rainy season began. Liana-cut trees grew more (0.4 mm/mm year) than liana-uncut trees (0.2 mm/mm year). These findings suggest that lianas may interfere with water availability to these trees during the dry season, and may also hinder tree growth. Received: 16 November 1999 / Accepted: 23 March 2000     

Contribution of vines to the evapotranspiration of a secondary forest in eastern Amazonia

The contribution of vines to the evapotranspiration (ET) of a secondary forest in eastern Amazonia was estimated based on field measurements of vine and tree transpiration, and seasonal changes in soil water content to 12 meters depth. Transpiration of vines and trees was measured with sapflow gauges placed around stems or branches. Total ET of the secondary forest was estimated as the sum of rainfall and reductions in soil moisture measured using Time Domain Reflectometry sensors installed in the walls of soil shafts. Our results suggest that vines transpire more than trees with stems of similar diameter, and with similar leaf crown exposure to sunlight. Trees experienced a smaller reduction in transpiration from the wet to the dry season than did vines. During the dry season, vines represented 8% (0.4 mm d^–1) of total secondary forest ET (5.4 mm d^–1), but they represented only 5.5% (0.5 m² ha^–1) of total secondary forest basal area (9.6 m² ha^–1). Considering that transpiration corresponds to 66–90% of forest ET, vines may contribute 9–12% to the transpiration of the forest. Hence, vine cutting, which is a commonly recommended management practice to favor the growth of tropical timber trees, may result in a proportionally larger reduction in evapotranspiration than in forest basal area.     

Certain phenological characters of the shrub layer of Kumaun Himalayan forests

The phenology of 49 shrub species in five forest types occurring along an altitudinal gradient (350–2150 m) in Kumaun Himalaya has been studied. The evergreen leaf-exchanging taxa accounted for nearly half of the species, the remaining half was nearly equally divided between an evergreen continual leaf drop type and deciduous taxa. The percentage of species with lengthy leaf drop increased with elevation and finally leveled off. At each site the maximum leaf drop period coincided with the warm dry period. Percentage of species with multiple leaf flushing was low for all forests. The degree of extended leafing decreased with increasing elevation along which summer dryness also decreased. Earliest leaf initiation was observed for evergreen continual leaf drop species, followed by evergreen leaf-exchanging, and deciduous types.For each forest, two peaks of flowering activity occurred, one during the warm dry period and the other in the warm wet period. The percentage of species with multiple flowering increased with increasing elevation. Nearly half of the species bore fleshy fruits. The mature fruit retention period for different forests ranged from about 2–3 months.The proportion of deciduous species was similar in trees and shrubs; leaf drop was common during the summer season for trees, while it was common during the winter season for shrubs; the proportion of species with multiple leafings was greater and leaf initiation earlier in shrubs than trees; and generally shrubs showed two flowering peaks and trees only one.Nomenclature follows Osmaston (1926).Financial support from the Gaula Catchment Eco-development project and the Department of Science and Technology, Government of India, is gratefully acknowledged. We thank Dr. Y. P. S. Pangtey for his help in plant identification.     

Seasonal variations in water requirement and influence of intermittent watering on grazing Yankasa sheep

Eighty male Yankasa sheep were used in two different experiments. A set of forty sheep, divided into four groups of ten rams each, were used in each experiment. The animals weighed, on average, 20 kg and were 15 months old. The animals were selected from the sheep flock of the Ahmadu Bello University Farm. They were grazed for 6 h daily for a period of 50 days during which they were subjected to intermittent watering. Watering intervals were 24 h, 48 h, 72 h and 96 h for Treatments 1, 2, 3 and 4, respectively. The experiment was carried out during the wet season of June to August and repeated in the hot, dry season of March to April, using another set of 40 Yankasa male sheep to find seasonal variations. The animals were not allowed to drink on the field when grazing during the rainy season.Seasonal variations had significant (P < 0.05) effects on the water intake and weight gains of Yankasa sheep. The animals lost 18.8 g, 115.2 g and 172 g daily on 48 h, 72 h and 96 h watering intervals in the dry season, but, on daily watering, they gained 66 g. During the rainy season the animals gained 135.6 g, 124.4 g, 108 g and 92.2 g body weight daily under 24 h, 48 h, 72 h and 96 h watering intervals, respectively.Slight variations were noted in rectal temperatures between treatments with a range of 39.07°C–39.59°C for the dry season and 38.73°C–38.95°C for the wet season. Respiratory rate was significantly (P < 0.05) affected by watering intervals, especially during the dry season, with average values of 32.4, 25.8, 20.5 and 19.9 for 24 h, 48 h, 72 h and 96 h watering intervals. Jaw movements were reduced with long watering intervals, and also pulse rate, especially in the dry season.The results suggest that grazing Yankasa sheep do not have to be watered daily during the wet season but free drinking water is very essential, at least every 24 h (daily) during the dry season.     

Carbon and water balance in Polylepis sericea,a tropical treeline species

Polylepis sericea trees grow well above the continuous forest line in the Venezuelan Andes. In these environments, extreme daily temperature ranges can occur at any time of the year and trees experience a 4 month dry period. The purpose of this work was to study carbon and water relations of this species in the field during wet and dry seasons in order to understand this species' success at such high altitudes. Leaf gas exchange (portable system in open mode) and leaf water potential (pressure chamber) were measured at 1–2 h intervals during several daily courses at 4000 m elevation in the Páramo de Piedras Blancas. CO₂ assimilation versus leaf temperature curves were also obtained for this species in the laboratory. Clear differences in the measured parameters were observed between seasons. For a wet season day, maximum CO₂ assimilation rate was 7.4 mol m^-2 s^-1 and leaf conductance was relatively constant (approximately 100 mmol m^-2 s^-1)In the dry season day, maximum CO₂ assimilation rate was 5.8 molm^-2 s^-1 and leaf conductance was close to 60 mmolm^-2 s^-1. Minimum leaf water potentials measured were -1.3 MPa for the wet and -2.2 MPa for the dry season. The CO₂ assimilation-leaf temperature relationship showed a 13.4°C leaf temperature optimum for photosynthesis with maximum and minimum compensation points of 29.5 and -2.8°C, respectively. Maximum night-time respiration was relatively high (2.7 (imol) m^-2 s^-1)Our results show thatP. sericea maintains a highly positive carbon balance through all daily courses, even though there is a slight water stress effect during the dry season; this suggests that its carbon assimilation machinery is well adapted to the low temperatures and seasonal water stress found in the high tropical mountains.     

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.     

Soil nutrient dynamics in response to irrigation of a Panamanian tropical moist forest

We measured concentrations of soil nutrients (0–15 and 30–35 cm depths) before and after the dry season in control and dry-season irrigated plots of mature tropical moist forest on Barro Colorado Island (BCI) in central Panama to determine how soil moisture affects availability of plant nutrients. Dry-season irrigation (January through April in 1986, 1987, and 1988) enhanced gravimetric soil water contents to wet-season levels (ca. 400 g kg^–1 but did not cause leaching beyond 0.8 m depth in the soil. Irrigation increased concentrations of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺), but it had little effect on concentrations of inorganic N (NH₄ ^+C, NO₃ ^– and S (SO₄ ^2–). These BCI soils had particularly low concentrations of extractable P especially at the end of the dry season in April, and concentrations increased in response to irrigation and the onset of the rainy season. We also measured the response of soil processes (nitrification and S mineralization) to irrigation and found that they responded positively to increased soil moisture in laboratory incubations, but irrigation had little effect on rates in the field. Other processes (plant uptake, soil organic matter dynamics) must compensate in the field and keep soil nutrient concentrations at relatively low levels.     

Vernal nitrogen and phosphorus retention by forest understory vegetation and soil microbes

Northern hardwood forests experience annual maximal loss of nutrients during spring. The vernal dam hypothesis predicts that spring ephemeral herbs in northern hardwood forests serve as sinks for nutrients during this season and reduce the loss of nutrients from the terrestrial ecosystem. Soil microbes of northern hardwood forests also sequester nutrients during spring. We compared the vernal nutrient acquisition ability of a soil microbial community and an understory plant community with species of mixed leaf phenology. We monitored nitrogen and phosphorus pool sizes in understory vegetation and soil microbes during spring from 1999 through 2001 in a northern hardwood forest in the Catskill Mountains, New York. Vegetation nutrient content increased during two spring seasons by an average of 3.07 g N m^–2 and 0.19 g P m^–2 and decreased during one spring by 0.81 g N m^–2 and 0.10 g P m^–2. Evergreen, wintergreen, and deciduous plant species were able to sequester nutrients during spring. Soil microbial nutrient content decreased during one spring by 1.29 g N m^–2 and remained constant during the other two springs. Streamwater nitrogen losses were not correlated with biotic nutrient uptake suggesting a temporal disconnect between the two processes. We conclude that understory vegetation is a larger potential sink for vernal nutrients than are soil microbes in this northern hardwood forest and understory and species representing multiple phenologies are capable of vernal nutrient uptake.     

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size

Summary We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.