Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.     

哀牢山中山湿性常绿阔叶林木径向生长季节动态及其对气候因子的响应

研究采用树木生长环在哀牢山中山湿性常绿阔叶林持续9年(2009—2017年)监测了2个常绿树种(厚皮香,Ternstroemia gymnanthera;南亚枇杷,Eriobotrya bengalensis)和2个落叶树种(西桦,Betula alnoides;珍珠花,Lyonia ovalifolia)的树干月生长量,采用逻辑斯蒂生长模型(Logistic model)模拟树木径向生长量和物候参数,并分析了年、季尺度上径向生长与主要气候因子的关系。结果表明:1)4个树种年平均生长量为6.3 mm,落叶树种年平均生长量(10.6 mm/a)显著高于常绿树种(3.0 mm/a);2)雨季(5—10月)是哀牢山中山湿性常绿阔叶林树木生长的主要时期,4个树种雨季平均生长量为5.9 mm,占全年总生长量的93%,其中落叶树种雨季生长量占全年的96%,而常绿树种雨季生长量占全年的86%;3)常绿树种生长季长度为169天,长于落叶树种(137天),而落叶树种最大生长速率(0.14 mm/d)显著高于常绿树种(0.03 mm/d),最大径向生长速率能很好地预测树种年生长量;4)低温、雾日和光合有效辐射是影响哀牢山亚热带常绿阔叶林4个研究树种径向生长的重要环境因子,其中温度对常绿树种径向生长具有显著影响,而雨日、雾日与空气湿度等水分因子对落叶树种径向生长更为重要。常绿树种年生长量对旱季气候因子的响应相比落叶树种更为敏感,树木旱季生长量除了受低温限制外,也受到水分供给的影响。气候变化可能改变不同物候类型树种在哀牢山中山湿性常绿阔叶林中的生长状态与分布格局。     

Cambial growth dynamics and climatic control of different tree life forms in tropical mountain forest in Ethiopia

Munessa Forest is a mountain forest in south-eastern Ethiopia experiencing seasonal rainfall variation. We investigated seasonal cambial activity and dormancy from increment rates of four different tree species belonging to varying life forms, namely, evergreen native conifer (Podocarpus falcatus), evergreen introduced conifer (Pinus patula), evergreen broadleaved tree (Prunus africana) and deciduous broadleaved tree (Celtis africana). Measurements of stem radius fluctuations were registered with the help of high-resolution electronic dendrometers. Daily amplitudes of stem diameter variations and daily and monthly net growth rates were determined and related to climatic variables measured at local climate stations. Thin sections of wood collected with a microcorer every 3–6 weeks allowed a visual control of newly formed wood cells during consecutive time intervals. Lack of water availability during the long dry season induced cambial dormancy of 5–7 months depending on life forms. After the onset of the short rainy season, stem swelling started quite synchronously with a variation of only single days in all studied species. Evergreen tree species were able to initiate wood formation during the short rainy season, whereas growth in the deciduous broadleaved species started in the long rainy season. The acquired data provide a basis for delineating the species-specific growth boundaries and the duration of the cambial growing season.     

What tree rings can tell us about the competition between trees and lianas? A case study based on growth,anatomy, density,and carbon accumulation

In tropical forest, landscape fragmentation and the consequent degradation of disturbed forests increase the incidence of light and dry hot winds, causing a disturbance on natural regeneration. Under these conditions, lianas (woody vines) development is stimulated instead of other species, which are more suited to mature forest and under less influence of the edge effect. For this, lianas colonization is an important variable for assessing the disturbance level of a forest. In this context, it becomes important to understand the nature of the competitive relationships between hyper-abundant lianas and ring growth of the host trees. Here, we selected trees with occupation or absence of lianas from two tropical species – Pinus caribaea var. hondurensis (Caribbean pine) and Tectona grandis (teak) – localized in a semideciduous forest fragment in southeastern Brazil, aiming to compare growth, climatic response, anatomy (vessels and intra-annual density fluctuations), wood density and carbon, by tree-ring analysis. The results showed that the lianas caused a change in tree-ring anatomy of host trees in last 10 years, mainly. We observed that trees occupied by lianas had a decrease the radial growth and carbon in the two species, an increase of the vessels size in teak and a decrease of the IADF frequency in Caribbean pine. In teak, the climate-tree relationship indicated that trees with lianas had lower response to rainfall and higher response to temperature in the summer (rainy and hottest period); in Caribbean pine, we observed that trees with lianas had a 2-month delay in the radial growth response to rainfall in the dry season. In the teak group, we observed that host trees had higher wood density values than liana-free tree in the outer rings, and the opposite was showed for pine. These findings show that tree-ring growth of host trees are a strong bioindicator of forest disturbance caused by aggressive colonization of lianas. We believe that these methods are applicable to future studies relating to the effects of habitat fragmentation and forest degradation on biodiversity and ecosystem services, particularly in the context of global climate change.     

Species-Specific Growth Responses to Climate Variations in Understory Trees of a Central African Rain Forest

Basic knowledge of the relationships between tree growth and environmental variables is crucial for understanding forest dynamics and predicting vegetation responses to climate variations. Trees growing in tropical areas with a clear seasonality in rainfall often form annual growth rings. In the understory, however, tree growth is supposed to be mainly affected by interference for access to light and other resources. In the semi-deciduous Mayombe forest of the Democratic Republic of Congo, the evergreen species Aidia ochroleuca, Corynanthe paniculata and Xylopia wilwerthii dominate the understory. We studied their wood to determine whether they form annual growth rings in response to changing climate conditions. Distinct growth rings were proved to be annual and triggered by a common external factor for the three species. Species-specific site chronologies were thus constructed from the cross-dated individual growth-ring series. Correlation analysis with climatic variables revealed that annual radial stem growth is positively related to precipitation during the rainy season but at different months. The growth was found to associate with precipitation during the early rainy season for Aidia but at the end of the rainy season for Corynanthe and Xylopia. Our results suggest that a dendrochronological approach allows the understanding of climate–growth relationships in tropical forests, not only for canopy trees but also for evergreen understory species and thus arguably for the whole tree community. Global climate change influences climatic seasonality in tropical forest areas, which is likely to result in differential responses across species with a possible effect on forest composition over time.     

Evaluation of X-ray densitometry to identify tree-ring boundaries of two deciduous species from semi-arid forests in Brazil

The presence of visible annual rings in semi-arid tropical trees may allow the application of dendrochronological methods. However, variation in water availability may cause the formation of narrow, irregular ill- or non-defined annual rings hindering the correct dating of tree-ring series. We aimed to evaluate X-ray densitometry as a method to identify tree rings of two deciduous tree species from the Caatinga forest, a semi-arid region in the northeast of Brazil, and compare with two other methods commonly used in dendrochronology, the sliding-stage micrometer and image analysis. Xylem was observed macro- and microscopically and wood anatomical features were assessed in Aspidosperma pyrifolium and Poincianella pyramidalis trees. In both species, tree-ring boundaries were identified considering intra-annual density patterns and wood anatomical features. No significant differences in tree-ring widths were found among methods. X-ray densitometry measurements showed a positive correlation with the measurements obtained with image analysis and sliding-stage micrometer in A. pyrifolium and P. pyramidalis, revealing the high reliability of the methods used. However, inter-correlation of tree-ring width series showed differences in the accuracy of crossdating across measuring methods. The maximum, mean and minimum density values were species-dependent, with mean wood density of A. pyrifolium lower than P. pyramidalis. Our results highlight X-ray densitometry as an important and complementary tool to identify tree-rings boundaries in semi-arid tree species, especially in A. pyrifolium. Along with other measuring methods, it may provide higher accuracy in dendrochronological studies in semi-arid or subtropical environments.     

Recent growth increase in endemic Juglans boliviana from the tropical Andes

The spatial coverage of tree-ring chronologies in tropical South America is low compared to the extratropics, particularly in remote regions. Tree-ring dating from such tropical sites is limited by the generally weak temperature seasonality, complex coloration, and indistinct anatomical morphology in some tree species. As a result, there is a need to complement traditional methods of dendrochronology with innovative and independent approaches. Here, we supplement traditional tree-ring methods via the use of radiocarbon analyses to detect partial missing rings and/or false rings, and wood anatomical techniques to precisely delineate tree-ring boundaries. In so doing we present and confirm the annual periodicity of the first tree-ring width (TRW) chronology spanning from 1814 to 2017 for Juglans boliviana (‘nogal’), a tree species growing in a mid-elevation tropical moist forest in northern Bolivia. We collected 25 core samples and 4 cross-sections from living and recently harvested canopy-dominant trees, respectively. The sampled trees were growing in the Madidi National Park and had a mean age of 115 years old, with certain trees growing for over 200 years. Comparison of (residual and standard) TRW chronologies to monthly climate variables shows significant negative relationships to prior year May-August maximum temperatures (r = −0.54, p < 0.05) and positive relationships to dry season May-October precipitation (r = 0.60, p < 0.05) before the current year growing season. Additionally, the radial growth of Juglans boliviana shows a significant positive trend since 1979. Our findings describe a new and promising tree species for dendrochronology due to its longevity and highlight interdisciplinary techniques that can be used to expand the current tree-ring network in Bolivia and the greater South American tropics.     

Effects of seasonal rainfall on radial growth in two tropical tree species

 Seasonal drought may limit primary productivity in most of the tropics, but the determinants of tree growth are not well known. A 10-year study of the deciduous trees Cochlospermum vitifolium (Willd.) Spreng. (Cochlospermaceae) and Cnidoscolus spinosus Lundell (Euphorbiaceae) in southwestern México showed radial growth to be highly correlated (both r>0.85) only with precipitation during an interval of <2 months in the mid-wet season. Growth was not affected by total annual precipitation or by an early starting or late ending of the wet season, or by heavy rainfall in the dry season. Annual mean girth increments ranged from 0.03 to 3.31 cm and −0.1 to 2.01 cm, respectively. The best model for growth (r ²>0.85) was a linear combination of mid-summer precipitation (positive coefficient) and total precipitation over the previous 2 years (negative coefficient). Comparison with other species showed heterogeneous responses of wood production to climate variation, and suggests that the range of functional types of dry forest trees is still unknown. Received: 9 September 1996 / Accepted: 4 March 1997     

Do the phenology and functional stem attributes of woody species allow for the identification of functional groups in the semiarid region of Brazil?

The phenology of tree species in environments that are subject to strong climatic seasonality is mainly determined by water availability, which may vary as a function of wood density. The relationship among phenology, water potential, wood density and the capacity of water storage in the stem were determined for woody species of caatinga vegetation (dry forest) in the semiarid region of NE Brazil. Leaf flush and fall, flowering and fruiting events were recorded over a 31-month period, and the water potential was measured over a two-year period. These data were related to precipitation, water availability in the soil and photoperiod. Seven deciduous species exhibited low wood density (DLWD,?<0.5?g?cm^?3), high capacity of water storage in the stem (until 250?% of the dry weight) and high water potential during the year, as opposed to 15 deciduous species that showed high wood density (DHWD,?≥0.5?g?cm^?3). Leaf flush, flowering and the fruiting of DHWD species were related to precipitation, whereas these phenological events occurred at the end of the dry season and/or the beginning of the rainy season for DLWD species and were related to the photoperiod. The two evergreen species showed variations of water potential that were intermediate between those of DHWD and DLWD deciduous species, leaf flush during the dry season and flowering at the end of dry season. These results suggest the existence of three functional groups: evergreen species, DHWD deciduous species and DLWD deciduous species.     

An assessment of growth ring identification in subtropical forests from northwestern Argentina

Most subtropical forests in South America are located in regions with a marked seasonality in precipitation, which may induce the formation of annual bands in woody species. Due to the lack of precise information on tree-ring visibility, we evaluated the wood characteristics of 37 tree species in the subtropical Yungas and Chaco forests from northwestern (NW) Argentina. Anatomical features associated with the delimitation of growth bands were examined to establish the presence of tree rings. Different forest types reflect the precipitation gradients and wood anatomical features vary accordingly. Characteristics of wood structure are closely related to the dominant climatic patterns of each forest, revealing a common pattern of anatomical arrangements in terms of water transport and safety. In the Chaco and transitional forests, ring boundaries are related to marginal parenchyma whereas in montane forests growth ring boundary is mostly associated with the presence of thicker fibers at the end of the ring. The largest proportion of species with clearly marked growth rings occurs in the montane forest type of NW Argentina. Clear growth rings is a requisite for dendrochronological applications, hence the present work represents the first regional attempt to address the potential of subtropical species in South America to be used in dendrochronological studies.     

Drivers of growth variability of Hymenaea stigonocarpa,a widely distributed tree species in the Brazilian Cerrado

Dendrochronology is a valuable tool to understanding climate-growth and growth-age relationships of native tree species from tropical forests. The information obtained from growth rings can elucidate climate responses of tree-growth under the ongoing environmental changes and support the development of sustainable forest management strategies based on species and site conditions. The Cerrado, which is a vast tropical savannah ecoregion of Brazil, has precipitation seasonality capable of inducing the formation of annual tree rings in moisture sensitive woody species. Hymenaea stigonocarpa is the typical tree species in the Cerrado with proven annual tree rings. It is an important commercial species that has been massively exploited for timber causing the considerable reduction of its natural populations. This study provides information about tree age and growth trajectories as well as climatic-growth signals of H. stigonocarpa in southeastern Brazil. We sampled 13 trees for tree-ring analysis. Tree-ring measurement and analysis were conducted using standard dendrochronological techniques. Sampled trees revealed the young successional stage of the stand, with ages varying from 20 to 35 years old. Nine out of 13 trees were used to build the standard chronology (1981 to 2013) that was positively correlated with precipitation at the end of the growing season (March-April). The chronology was able to capture SST anomalies patterns related to the South American Monsoon System. Growth modeling indicated that minimum logging diameter of 10.4cm is achieved at 24 years of age. The results reported here add valuable contribution to the discussion of sustainable management strategies for Cerrado ecoregion species.     

Growth rings, growth ring formation and age determination in the mangrove Rhizophora mucronata

BACKGROUND AND AIMS: The mangrove Rhizophora mucronata has previously been reported to lack annual growth rings, thus barring it from dendrochronological studies. In this study the reported absence of the growth rings was reconsidered and the periodic nature of light and dark brown layers visible on polished stem discs investigated. In addition, the formation of these layers in relation to prevailing environmental conditions, as well as their potential for age determination of the trees, was studied. METHODS: Trees of known age were collected and a 2.5-year cambial marking experiment was conducted to determine the periodic nature of the visible growth layers. KEY RESULTS: Annual indistinct growth rings were detected in R. mucronata and are defined by a low vessel density earlywood and a high vessel density latewood. The formation of these growth rings and their periodic nature was independent from site-specific environmental conditions in two forests along the Kenyan coast. However, the periodic nature of the rings was seriously affected by slow growth rates, allowing accurate age determination only in trees with radial growth rates above 0.5 mm year(-1). The onset of the formation of the low vessel density wood coincided with the onset of the long rainy season (April-May) and continues until the end of the short rainy season (November). The high vessel density wood is formed during the dry season (December-March). Age determination of the largest trees collected in the two studied forests revealed the relatively young age of these trees (+/-100 years). CONCLUSIONS: This study reports, for the first time, the presence of annual growth rings in the mangrove R. mucronata, which offers further potential for dendrochronological and silvicultural applications.     

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.     

Inter-annual and seasonal variability of radial growth, wood density and carbon isotope ratios in tree rings of beech (Fagus sylvatica) growing in Germany and Italy

We investigated the variability of tree-ring width, wood density and ¹³C/¹²C in beech tree rings (Fagus sylvatica L.), and analyzed the influence of climatic variables and carbohydrate storage on these parameters. Wood cores were taken from dominant beech trees in three stands in Germany and Italy. We used densitometry to obtain density profiles of tree rings and laser-ablation-combustion-GC-IRMS to estimate carbon isotope composition (δ ¹³C) of wood. The sensitivity of ring width, wood density and δ ¹³C to climatic variables differed; with tree-ring width responding to environmental conditions (temperature or precipitation) during the first half of a growing season and maximum density correlated with temperatures in the second part of a growing season (July–September). δ ¹³C variations indicate re-allocation and storage processes and effects of drought during the main growing season. About 20% of inter-annual variation of tree-ring width was explained by the tree-ring width of the previous year. This was confirmed by δ ¹³C of wood which showed a contribution of stored carbohydrates to growth in spring and a storage effect that competes with growth in autumn. Only mid-season δ ¹³C of wood was related to concurrent assimilation and climate. The comparison of seasonal changes in tree-ring maximum wood density and isotope composition revealed that an increasing seasonal water deficit changes the relationship between density and ¹³C composition from a negative relation in years with optimal moisture to a positive relationship in years with strong water deficit. The climate signal, however, is over-ridden by effects of stand density and crown structure (e.g., by forest management). There was an unexpected high variability in mid season δ ¹³C values of wood between individual trees (−31 to −24‰) which was attributed to competition between dominant trees as indicated by crown area, and microclimatological variations within the canopy. Maximum wood density showed less variation (930–990 g cm⁻³). The relationship between seasonal changes in tree-ring structure and ¹³C composition can be used to study carbon storage and re-allocation, which is important for improving models of tree-ring growth and carbon isotope fractionation. About 20–30% of the tree-ring is affected by storage processes. The effects of storage on tree-ring width and the effects of forest structure put an additional uncertainty on using tree rings of broad leaved trees for climate reconstruction.     

Detecting invisible growth rings of trees in seasonally dry forests in Thailand: isotopic and wood anatomical approaches

We measured radial variation of carbon isotope composition and vessel traits in tree species in seasonally dry forests of Northeast Thailand to explore a more reliable and amenable method of tropical dendrochronology for trees that lack visually detectable and consistent growth rings. Six Dipterocarpaceae species (3 Shorea, 2 Dipterocarpus, and 1 Hopea species) with indistinct or irregular growth rings and teak (Tectona grandis), a species which forms distinct growth rings, were examined. The δ¹³C value variations in all species showed annual cyclicity. Dipterocarpaceae species usually marked the lowest values of δ¹³C in the middle of the growing season, whereas teak had the lowest values at nearly the end of the growing season. Since the growing season of the species examined almost corresponds to the rainy season in the study area, the δ¹³C variation was likely caused by the change in moisture availability. The different variation pattern of teak was attributable to its stronger dependence on ¹³C-enriched reserved material early in the growing season. Changes in tree vessel traits for all species examined also showed annual cyclicity. Dipterocarpaceae species showed significant correlation between δ¹³C values and vessel measurements. Vessel lumen (mean area, tangential and radial diameter, and proportion of total area) had a negative correlation, whereas vessel frequency showed a positive correlation. The correlations indicated that changes in vessel traits were caused by the seasonal variation of moisture available to the trees. Thus, we concluded that methods using wood anatomy, as well as δ¹³C, have great potential for use as tools in tropical dendrochronology within the context of seasonal climate.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Cambial Growth Season of Brevi-Deciduous Brachystegia spiciformis Trees from South Central Africa Restricted to Less than Four Months

We investigate cambial growth periodicity in Brachystegia spiciformis, a dominant tree species in the seasonally dry miombo woodland of southern Africa. To better understand how the brevi-deciduous (experiencing a short, drought-induced leaf fall period) leaf phenology of this species can be linked to a distinct period of cambial activity, we applied a bi-weekly pinning to six trees in western Zambia over the course of one year. Our results show that the onset and end of cambial growth was synchronous between trees, but was not concurrent with the onset and end of the rainy season. The relatively short (three to four months maximum) cambial growth season corresponded to the core of the rainy season, when 75% of the annual precipitation fell, and to the period when the trees were at full photosynthetic capacity. Tree-ring studies of this species have found a significant relationship between annual tree growth and precipitation, but we did not observe such a correlation at intra-annual resolution in this study. Furthermore, a substantial rainfall event occurring after the end of the cambial growth season did not induce xylem initiation or false ring formation. Low sample replication should be taken into account when interpreting the results of this study, but our findings can be used to refine the carbon allocation component of process-based terrestrial ecosystem models and can thus contribute to a more detailed estimation of the role of the miombo woodland in the terrestrial carbon cycle. Furthermore, we provide a physiological foundation for the use of Brachystegia spiciformis tree-ring records in paleoclimate research.     

Climate/growth relationships of Brachystegia spiciformis from the miombo woodland in south central Africa

We present five Brachystegia spiciformis Benth. (BrSp) tree-ring chronologies from the seasonally dry miombo woodland in south central Africa. Between 9 and 34 stem discs were collected from three dry and two wet miombo sites. All samples showed distinct growth rings, which were marked by terminal parenchyma bands. Site chronologies varied in length between 43 and 149 years. An increase in the number of growth ring anomalies in older trees, however, resulted in an increase in dating error and a decrease in between-tree correlations with increase in the chronology length. Annual precipitation variability accounted for some 28% of the common variance in the BrSp chronologies and we found no difference in climate sensitivity between wet and dry miombo sites. The influence of climate, and of precipitation in particular, on tree growth was strongest at the core of the rainy season (December–February). This is also the time of the year when ENSO peaks in amplitude and ENSO effects on precipitation variability in southern Africa are the strongest. We found a negative response of tree growth to ENSO throughout most of the growth year, suggesting that the development of longer chronologies from the miombo region would allow for the investigation of temporal ENSO variability. A spatial extension of the miombo tree-ring network should therefore focus on regions where ENSO effects are the strongest (e.g., southeastern Africa).     

Reproductive phenology, life-forms, and habitats of the Venezuelan Central Plain

Reproductive phenology of 171 plant species belonging to 57 families of angiosperms was studied according to life-forms in four habitat types in a savanna-forest mosaic on the Venezuelan Central Plain. Flowering, unripe fruit, and mature fruit patterns were affected significantly according to life-forms and habitats respectively. Production of flowers, unripe fruits, and mature fruits showed marked seasonality for all habitats except for the forest. Flowering peaked during the rainy season, and fruiting peaked toward the end of the rainy season. The savanna and the disturbed area had similar proportions of species that flowered over the year. The percentage of species with unripe fruits produced throughout the year was more seasonal for the disturbed area than for the other habitats. Mature fruit patterns showed an increase during the late rainy season for the ecotone and savanna. A large number of herbaceous (annual and perennial) and liana species flowered during the wet season, and a smaller fraction flowered during the dry season; and trees, shrubs, and epiphytes increased flowering activity during the dry season. Unripe fruit patterns were similar to those of flowering for all life-forms, however, tree species were less seasonal. Mature fruit production by shrubs peaked in the period of maximum rainfall, while the peak for perennial herbs was in the late rainy season and the peak for annual herbs was during the transition between the rainy season and the dry season. The largest proportion of tree and liana species with ripe fruits occurred during the dry season. Differences among phenological patterns in habitats were caused mainly by life-forms and promote a wider distribution of reproductive events in habitats and overall community in the Venezuelan Central Plain.