Constraints to seedling success of savanna and forest trees across the savanna-forest boundary

Tropical savannas and closed forests are characterized by distinct tree communities, with most species occurring almost exclusively in only one of the two environments. The ecology of these two groups of species will largely determine the structure and dynamics of the savanna-forest boundary, but little is known about the ecological and physiological differences that might control their distributions. We performed field and nursery experiments to compare seedling establishment success, predawn leaf water potential, biomass allocation, and root carbohydrate concentration of congeneric species, each composed of one savanna species and one forest species. Seedling establishment of savanna and forest species responded differently to vegetation cover, with forest species having lowest establishment success in the open savanna and savanna species having lowest success in forest. Subsequent survival followed similar patterns, resulting in even greater differences in cumulative success. The low survival of forest species in the savanna appears related to drought stress, as seedlings of forest species had lower predawn leaf water potential than savanna species. Seedlings of savanna species had greater root: shoot ratios and root total nonstructural carbohydrate (TNC) concentration, particularly among evergreen genera. Among evergreen genera, root TNC per shoot mass, which may largely determine resprout capacity, was seven times higher in savanna species than forest species. Although water availability and microclimate may reduce the success of forest species, these factors appear unable to completely exclude forest seedling establishment in savanna. Fire, on the other hand, appears to be a much more absolute constraint to success of forest species in savanna.     

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

西双版纳热带季节雨林林窗内幼树叶生长与虫食动态

 研究了西双版纳热带季节雨林6种乔木幼树在林窗中的叶生长与叶虫食动态。6种幼树叶生长主要在雨季(5～10月),在旱季雾凉期(11～2月)叶停止生长。阳性树种旱季干热期(3～4月)开始叶生长,顶极树种至5月雨季初才开始叶生长。表明雾凉期低温抑制两类树种叶生长,干热期水分不足抑制顶极树种叶生长。叶生长同步性(每两个月的叶生长量变异系数C．V．)顺序为：望天树(Shorea chinensis,1.42)>金钩花(Pseuduvaria indochinensis,1.41)>八宝树(Duabanga grandiflora, 1.02)>云南石梓(Gmelina arborea,0.98)>团花树(Anthocephalus chinensis,0.84)和铁刀木(Cassia siamea,0.84)。旱季造成一些种类幼树出现落叶高峰,严重叶虫食也导致云南石梓在7～8月出现落叶高峰。叶虫食主要出现在雨季,团花树和云南石梓在干热期叶生长能减少昆虫取食,但此时铁刀木叶同步生长却不能降低食叶昆虫危害。6种幼树叶生长量年进程与叶虫食量年进程间存在正相关,其中望天树、金钩花、八宝树和团花树分别达到显著(p<0.05)或极显著(p<0.01)水平。望天树和金钩花在雨季初叶同步生长能减轻叶虫食。食叶昆虫偏爱取食幼叶,6种幼树平均幼叶的虫食量占总虫食量的72.9％,幼叶虫食速率平均为成熟叶的4.3倍。     

Assessing onset and length of greening period in six vegetation types in Oaxaca,Mexico, using NDVI-precipitation relationships

Variations in the normalized vegetation index (NDVI) for the state of Oaxaca, in southern Mexico, were analyzed in terms of precipitation anomalies for the period 1997-2003. Using 10-day averages in NDVI data, obtained from AVHRR satellite information, the response of six types of vegetation to intra-annual and inter-annual fluctuations in precipitation were examined. The onset and temporal evolution of the greening period were studied in terms of precipitation variations through spectral analysis (coherence and phase). The results indicate that extremely dry periods, such as those observed in 1997 and 2001, resulted in low values of NDVI for much of Oaxaca, while good precipitation periods produced a rapid response (20-30 days of delay) from a stressed to a non-stressed condition in most vegetation types. One of these rapid changes occurred during the transition from dry to wet conditions during the summer of 1998. As in many parts of the tropics and subtropics, the NDVI reflects low frequency variations in precipitation on several spatial scales. Even after long dry periods (2001-2002), the various regional vegetation types are capable of recovering when a good rainy season takes place, indicating that vegetation types such as the evergreen forests in the high parts of Oaxaca respond better to rainfall characteristics (timing, amount) than to temperature changes, as is the case in most mid-latitudes. This finding may be relevant to prepare climate change scenarios for forests, where increases in surface temperature and precipitation anomalies are expected.     

Seasonal patterns of gas exchange,water relations and growth of it Roupala montana,an evergreen savanna species

Roupala montana is an evergreen species widespread in the seasonal savannas of the central plains of Brazil. I examined the degree of coupling of photosynthetic gas-exchange characteristics, water relations and growth responses of R. montana with regard to seasonal changes in soil water availability. Despite a rainless period of over three months soil water potential at 60 cm depth reached values of only about -1.0 MPa, while pre-dawn leaf water potential (^l) reached about -0.4 MPa by the end of the three-month drought. Thus, R. montana had access to deep soil water in the dry period, but pre-dawn ^l did not reach the high wet season values of -0.2 MPa. Most of the shoot growth was concluded in the onset of the rainy season. Although some individual branches might have shown some extension thereafter, most of them remained inactive during the rest of the rainy season and the subsequent dry season. New leaf production was also restricted to the first part of the wet period. R. montana remained evergreen in the dry season, but there was a 27% decrease in the number of leaves and herbivory removed about 16% of the leaf area still present in the plant. CO₂-exchange rates of these leaves reached only ca. 55% of the maximum rainy season values of 14 µmol m^-2 s^-1. Thus, the estimated potential daily carbon gain was about 34% of the maximum by the end of the dry period. These values will be even lower, if we considered the decrease in photosynthetic rates that occurred around midday. These reductions in photosynthetic rates as a result of partial stomatal closure were measured both in the wet and dry season and they were related to increases in the evaporative demand of the atmosphere. In conclusion, the combined effect of herbivory, leaf loss and reductions in photosynthetic rates limited plant productivity in the dry season.     

Seasonality of weather and tree phenology in a tropical evergreen mountain rain forest

Flowering and fruiting as phenological events of 12 tree species in an evergreen tropical mountain rain forest in southern Ecuador were examined over a period of 3–4 years. Leaf shedding of two species was observed for 12 months. Parallel to the phenological recordings, meteorological parameters were monitored in detail and related to the flowering and fruiting activity of the trees. In spite of the perhumid climate of that area, a high degree of intra- and inter-specific synchronisation of phenological traits was apparent. With the exception of one species that flowered more or less continuously, two groups of trees could be observed, one of which flowered during the less humid months (September to October) while the second group started to initiate flowers towards the end of that phase and flowered during the heavy rains (April to July). As reflected by correlation coefficients, the all-time series of meteorological parameters showed a distinct seasonality of 8–12 months, apparently following the quasi-periodic oscillation of precipitation and related cloudiness. As revealed by power spectrum analysis and Markov persistence, rainfall and minimum temperature appear to be the only parameters with a periodicity free of long-term variations. The phenological events of most of the plant species showed a similar periodicity of 8–12 months, which followed the annual oscillation of relatively less and more humid periods and thus was in phase or in counter-phase with the oscillations of the meteorological parameters. Periods of unusual cold or dryness, presumably resulting from underlying longer-term trends or oscillations (such as ENSO), affected the homogeneity of quasi-12-month flowering events, fruit maturation and also the production of germinable seeds. Some species show underlying quasi-2-year-oscillations, for example that synchronise with the development of air temperature; others reveal an underlying decrease or increase in flowering activity over the observation period, influenced for instance by solar irradiance. As Ecuador suffers the highest rate of deforestation in South America, there is an urgent need for indigenous plant material for reforestation. A detailed knowledge of the biology of reproduction in relation to governing external factors (mainly climate) is thus required.     

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.     

Seasonal patterns of canopy development and carbon gain in nineteen warm desert shrub species

Summary Canopy development and photosynthetic rate were measured at monthly intervals over a period of one year in 19 shrub and subshrub species of the Mojave and upper Sonoran Deserts. Thirteen of these species realized a substantial fraction of their total net carbon assimilation via twig photosynthesis. The twig contribution to whole plant yearly carbon gain reached a maximum of 83% in species such as Thamnosma montana, Salizaria mexicana, and Baccharis brachyphylla. This large contribution by twigs was due to both low levels of leaf production and the greater longevity of twig tissues. In some other species, however, leaf and twig organs had similar lifespans. During the year of this study (which had an unusually warm, mild winter), no species showed a pattern of winter deciduousness. The reduction in total photosynthetic area between maximal spring canopy development and mid August summer dormancy ranged from 32 to 94%. Some herbaceous perennial species died back to the ground, but none of the woody shrubs were totally without green canopy area at any time of the year. No species studied were capable of high rates of photosynthesis at low plant water potentials in July and August, but, in those species which maintained a substantial canopy area through the drought period, previously stressed tissues showed substantial recovery after fall rains. Photosynthetic rate was significantly correlated with both plant water potential and tissue nitrogen content over the entire year, but only weakly so. This is due in part to the winter months when plant water potentials and tissue nitrogen contents were high, but photosynthetic rates were often low.     

Habitat-related variation in tree leaf form in four tropical forest types on Pulau Ubin,Singapore

Abstract. Sun leaves from 37 species of tree and shrub were collected in four forest types on Pulau Ubin, Singapore. Evidence of habitat-related variation in the form of leaves is presented. The species were from the habitats: mangrove, beach forest, adinandra belukar (secondary forest on degraded soils) and secondary forest on undegraded soils. The mangrove forest species sampled had thicker leaves with a lower specific leaf area than the beach forest species. Leaves of species from the adinandra belukar were thicker and smaller, with more dry weight per unit area, than those of pioneers from undegraded sites. This is interpreted as oligotrophic xeromorphy in adinandra belukar.     

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity (k_L) of upper branches was positively correlated with maximum rates of net CO₂ assimilation per unit leaf area (A_area) and stomatal conductance (g_s) across 20 species of canopy trees. Maximum k_L showed stronger correlation with A_area than initial k_L suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k_L was negatively correlated with A_area/g_s and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics (k_theoretical) was positively related to A_area and k_L, consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k_L, A_area, net CO₂ assimilation per unit leaf mass (A_mass), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.     

基于多角度植被指数的马尾松林LAI反演方法

CHRIS/PROBA是目前具有最高空间分辨率（17 m×17 m）的星载多角度高光谱数据,该款数据在反演植被垂直结构参数,如树高、叶面积指数（leaf area index,LAI）等方面具有重要的应用前景。基于四尺度几何光学模型得到马尾松（Pinus massoniana Lamb.）冠层的归一化差分植被指数（normalized difference vegetation index,NDVI）各向异性分布规律,利用CHRIS红光特征波段和近红外特征波段构建一种新型多角度植被指数（normalized hotspot-dark-spot difference vegetation index,NHDVI）,并将其应用于CHRIS数据对马尾松林的LAI遥感估算上。结果显示：（1）相比归一化差分植被指数（NDVI）与土壤调节植被指数（soil adjusted vegetation index,SAVI）而言,NHDVI能很好地融合光谱信息与角度信息,与地面实测LAI的决定系数达到0.7278;（2）利用NHDVI-LAI统计回归模型方法来反演LAI值,将得到的LAI值与地面实测值进行相关性分析,结果拟合优度达到0.8272,均方根误差RMSE为0.1232。与传统植被指数相比,包含角度信息的多角度植被指数对LAI的反演在精度上有较大提升,同时比基于辐射传输模型的反演方法更简易、实用。     

Maintenance of forest species diversity and latitudinal gradient

The species diversity of trees maintained in tropical rain forests is much higher than in temperate, boreal, or seasonally dry tropical forests. Many hypotheses have been proposed for higher diversity in tropical rain forests, including: (i) higher specialization of resource use, (ii) different mode of disturbance, (iii) smaller opportunity for competition on oligotrophic soil, (iv) higher productivity, (v) more active specific herbivores and pathogens, (vi) evolutionary/ecological history. In this paper we report mathematical models for tree-by-tree replacement. First the analysis of random drift model shows that the effect of gap size to species diversity is not very strong. Second we study phenological segregation model, which has the following assumptions: Basic mechanism for many species to coexist in the community is assumed given by the storage effect of lottery model, as species differ in seasonality in peak fruit production and in the subsequent period of high regeneration ability. Gaps formed during unfavorable season accumulate and become available for regeneration in the beginning of the growing season. The resulting synchronization of regeneration opportunity jeopardizes the coexistence of many similar species in seasonal environments. Analysis of a mathematical model shows: (1) the existence of unfavorable season can greatly reduce the diversity of coexisting species. (2) Diversity in the equilibrium community can be high when niche width of each species is broad and resource use is strongly overlapped. (3) Equilibrium community may include several distinct groups of species differing in phenology of regeneration. Effect of unequal niche width and frequency dependent regeneration are also examined.     

Leaf structure of the cerrado (Brazilian savanna) woody plants

With the aim of recognizing the commonest leaf pattern found in the woody flora of the cerrado (the Brazilian savanna) we analyzed the leaf anatomy of 30 representative species. The leaves are mostly dorsiventral and hypostomatic and covered by trichomes and a thick layer of wax and cuticle; the vascular bundles are surrounded by a sheath of fibers. The mesophyll has a developed palisade tissue, dispersed sclerified cells and idioblasts bearing crystals and phenolic compounds. We compared the results with those reported for other species (60 species) from the same biome and for the families that the studied species belong. The present study suggests that the xeromorphism observed for the cerrado leaves is related to the evolutionary history of this biome, since its first floristic elements must have faced deficient water conditions as well as the consequent soil acidity and toxicity. Therefore we may infer that the leaf anatomical pattern here observed was already present in the first elements of the cerrado and was selected to guarantee the survival of those species in the new environment. Furthermore, the xeromorphic features present in those leaves continue nowadays to help the plants protecting themselves from the different biotic and abiotic factors they are subjected to.     

Nitrogen and phosphorus resorption in trees of a Mexican tropical dry forest

Resorption efficiency (RE) and proficiency, foliar nutrient concentrations, and relative soil nutrient availability were determined during 3 consecutive years in tree species growing under contrasting topographic positions (i.e., top vs. bottom and north vs. south aspect) in a tropical dry forest in Mexico. The sites differed in soil nutrient levels, soil water content, and potential radiation interception. Leaf mass per area (g m^–2) increased during the growing season in all species. Soil P availability and mean foliar P concentrations were generally higher at the bottom than at the top site during the 3 years of the study. Leaf N concentrations ranged from 45.4 to 31.4 mg g^–1. Leaf P varied from 2.3 to 1.8 mg g^–1. Mean N and P RE varied among species, occasionally between top and bottom sites, and were higher in the dry than in the wet years of study. Senesced-leaf nutrient concentrations (i.e., a measure of resorption proficiency) varied from 13.7 to 31.2 mg g^–1 (N) and 0.4 to 3.3 mg g^–1 (P) among the different species and were generally indicative of incomplete nutrient resorption. Phosphorus concentrations in senesced leaves were higher at the bottom than at the top site and decreased from the wettest to the the driest year. Soil N and P availability were significantly different in the north- and south-facing slopes, but neither nutrient concentrations of mature and senesced leaves nor RE differed between aspects. Our results suggest that water more than soil nutrient availability controls RE in the Chamela dry forest, while resorption proficiency may be interactively controlled by both nutrient and water availability.     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.