Annual Growth Ring Patterns in Brachystegia spiciformis Reveal Influence of Precipitation on Tree Growth1

The availability of exactly dated tree‐ring chronologies is limited in tropical regions. However, these chronologies could contribute widely to studies of the influence of natural and human‐induced factors on tropical forests. We examine the potential for building a chronology based on three sites in the miombo woodland of western Zambia. Brachystegia spiciformis Benth., a dominant species from this vegetation type, is used. Response of the chronology to several climatic factors is examined. All specimens showed very clear growth rings, and cross‐dating between radii of a tree was successful for all trees. Site chronologies could be constructed after cross‐dating of growth ring series of individual trees. The mean growth ring curves of the three sites were significantly similar, allowing for the construction of a regional chronology. Correlation function analysis between the tree‐ring chronology and regional climatic variables revealed that climate at the core of the rainy season, in December and January, has an explicit influence on tree growth. Where precipitation and relative humidity in these months influence tree growth positively, temperature correlates in a negative way. Some 20 percent of the variance in the B. spiciformis tree‐ring chronology is accounted for by wet season rainfall. The successful cross‐dating and correlation between a tree‐ring chronology and climate demonstrated in this study indicate annual ring formation in B. spiciformis trees and sensitivity to climatic conditions.     

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).     

Seedling ecology of two miombo woodland trees

The development of seedlings of two miombo trees, Brachystegia spiciformis Benth. and Julbernardia paniculata (Benth.) Troupin, was studied during two growing seasons (December 1989–April 1991) at a Zambian grassland site. Seed germination rates under laboratory and field conditions were not significantly different although germination in the field was delayed by 1–2 weeks due to insufficient rainfall. After one year of storage J. paniculata seed germination had declined from 67% to 17% while germination of B. spiciformis seeds remained at about 83%.Leaf production was confined to the rainy season. Leaf fall occurred during the dry season and in J. paniculata this was followed by shoot die-back during the hot dry period (August–November). Two-thirds of B. spiciformis seedlings experienced shoot die-back but shoot die-back did not necessarily result in seedling mortality. Seedling deaths occurred during the germination period (6–10 weeks after planting) and in the hot dry period (40–50 weeks after planting) during September–November. Survivorship of B. spiciformis seedlings was 74% at the end of the second growing season while this was 46% for J. paniculata.Shoot growth was negligible during the second growing season. In fact mean maximum leaf area of B. spiciformis decreased significantly from 19.7 cm² (SD=5.7) per plant at the end of the first growing season to 13.3 cm² (SD=5.8) at the end of the second growing season (t=3.31, P<0.01). However, root biomass of B. spiciformis seedlings increased 2.8 times during the second growing season.These results suggest that shoot die-back in seedlings of miombo trees is caused by drought and that the slow shoot growth is the result of allocating most of the biomass to root growth during seedling development.     

Annual growth rings, rainfall-dependent growth and long-term growth patterns of tropical trees from the Caparo Forest Reserve in Venezuela

1 Tree-ring analyses and dendrometer measurements were carried out on 37 tree species in a semi-deciduous forest of the Reserva Forestal de Caparo, Venezuela, where the mean annual rainfall is about 1700 mm and there is a dry season from December to March. The main purposes of the investigation were to show the seasonality of cambial growth, and the connection between precipitation patterns and tree-ring curves. Long-term rates of wood increment were also estimated.
2 Cambial markings in consecutive years showed that annual rings were formed by many species.
3 The distinctiveness of growth zones was usually greater in deciduous species than in evergreen species, although not all deciduous species had distinct rings.
4 Dendrometer measurements showed that the annual growth rhythm was related to precipitation patterns. Evergreen species tended to show only a short interruption of wood growth (during the later part of the dry season), whereas deciduous species stopped growth completely at the end of the rainy season.
5 For deciduous species, regression analyses showed close relations between tree-ring width and the sum of precipitation outside the rainy seasons (i.e. November to April). Evergreen species reacted to the total annual amount of precipitation.
6 Variation in longest available ring chronology (for Terminalia guianensis ) showed little correlation with the El Niño–Southern Oscillation effect.
7 On average trees from natural forests showed relatively constant growth over the entire life span. Plantation trees grew fast up to an age of 15–20 years, but annual increments then decreased to values seen in natural forest trees.     

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...     

Development of Brachystegia‐Julbernardia woodland after clear‐felling in central Zambia: Evidence for high resilience

Question: Does the development of Brachystegia‐Julbernardia (miombo) woodland after felling, and under a variable fire regime, occur via a serai stage of fire‐tolerant species? Location: Four sites in central Zambia, Africa. Methods: Trees in replicate plots were clear‐cut and stumps and resprouts enumerated. Species recruited into the tree layer (> 2.0 m tall) were monitored for 11 years (1991–2001) and fire occurrence and herbaceous biomass assessed annually to determine fuel loads. Results: Fire frequency was variable at the study sites and fuel loads were generally too low to suppress woodland regeneration after felling. However, at one site a change from low to high fire frequency arrested woodland development and triggered a regression towards a ‘fire‐trap’ vegetation type in which a few fire‐tolerant species survived. There was no evidence to support the hypothesis that miombo woodland regeneration is facilitated by a sere of fire‐tolerant species. All regrowth after felling was from resprouting plants present before felling. Trees with a previous history of felling sprouted more vigorously than trees that had not been felled before. Species richness in the tree layer increased with time since felling because resprout species had different height growth rates. Conclusion: The resilience of miombo trees after clear‐felling is largely due to their capacity to regenerate vegetatively from resprouts and stumps after release from frequent fires. Coppicing is therefore recommended as a suitable management technique for miombo woodland in central southern Africa.     

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.     

Spatial pattern,regeneration and growth rates of Brachystegia spiciformis and Julbernardia globiflora

Past work on tree-to-tree interactions in semi-arid savannas, through the use of nearest neighbour analysis, has shown both a regular dispersion pattern and a positive correlation between the size of a tree and the distance to its nearest neighbour. From these results, the importance of competition has been inferred. In the present study, tree-to-tree interactions in a mesic savanna woodland at Marondera, Zimbabwe, dominated by relatively small-sized individuals of Brachystegia spiciformis and Julbernardia globiflora, were examined. Results show that the regular dispersion pattern is uncommon; that there are few significant positive correlations between size of individual and distance to nearest neighbour; and that growth rate is not consistently correlated with size and distance of nearest neighbour. Results of a study of regeneration relative to canopy cover suggest a possible mechanism for these findings: unlike semi-arid savannas, young plants often grow in under-canopy environments so that early spacing of young trees does not occur. Significant positive correlations between the size of a tree and the distance to its nearest neighbour were found in a stand of larger, presumably older, trees, suggesting that such a relationship only develops in mesic miombo woodland through a thinning process as the trees mature. Familial clumping, which is common in the study area, may mask spatial patterns.Abbreviations RBAI Relative basal area increment     

A tree-ring based comparison of Terminalia superba climate–growth relationships in West and Central Africa

Tropical lowland forests are characterized by humid climate conditions with interannual variations in amount of precipitation, length of dry season, and relative humidity. The African tree species, Terminalia superba Engl. & Diels has a large distribution area and potentially incorporates these variations in its tree rings. Tree ring analysis was performed on 60 plantation trees (increment cores) and 41 natural trees (stem disks) from Ivory Coast and the Congolese Mayombe Forest. Natural forests and old plantations (50–55 years) showed similar growth patterns. Regional chronologies were developed for the two sample regions and showed a long-distance relationship for the period 1959–2008. Growth in the Mayombe was associated with early rainy season precipitation, but no relation was found between tree growth and precipitation in Ivory Coast. Congolese trees possibly show a higher climate-sensitivity than Ivorian trees, because precipitation in the Mayombe is more limiting, and Congolese T. superba trees are found closer to the margins of their distribution. Likewise, tree growth in the Mayombe was also influenced by the SSTs of the Gulf of Guinea and the South Atlantic Ocean during the early rainy season. However, tree growth was influenced by ENSO in both regions. In the Mayombe, La Niña years were associated with stronger tree growth whereas in Ivory Coast, El Niño years corresponded with stronger tree growth. The presented relation between ENSO, precipitation and tree growth is original for equatorial African forests, suggesting an influence of global climate variability on tree growth.     

Dendroecological investigations on Swietenia macrophylla King and Cedrela odorata L. (Meliaceae) in the central Amazon

The width of the increment zones in the xylem of Swietenia macrophylla King and Cedrela odorata L. was investigated by dendroecological methods in a primary forest near Aripuanã, Mato Grosso, Brazil (10°09′S, 59°26′W). The annual period of cambial cell division and its intra-annual variation were determined by dendrometer measurements of 30 trees of each species. Tree-ring width chronologies for Swietenia and Cedrela were developed from cross-dated increment curves of 33 out of 47 Swietenia and 51 out of 64 Cedrela trees. Simple correlations were computed between the radial growth increment and monthly precipitation for the period 1890–2000. In Swietenia, cambium activity occurred throughout almost the whole year, but in Cedrela it was restricted to the rainy period from September of the previous year to June of the current year. Tree-rings were formed annually in the juvenile and adult wood of Cedrela, while in Swietenia the annual formation of tree-rings was restricted to the adult wood. Consequently the age of the Swietenia trees could be dated by the tree-rings in good approximation, while age dating of the Cedrela trees was exact. Correlation analyses revealed a significant relationship between the precipitation at the beginning and at the end of the growth season and the width of the increment zones in the adult xylem of Swietenia. In contrast, the width of the growth increment in the xylem of Cedrela was significantly correlated with the precipitation in March and May of the previous growth period.     

Seasonal variations in the distribution and abundance of the tsetse fly,Glossina morsitans morsitans in eastern Zambia

The seasonal changes in the distribution of Glossina morsitans morsitans Westwood (Diptera: Glossinidae) and its main host, cattle, were examined in a cultivated area of the plateau of eastern Zambia. During four consecutive years, the tsetse and cattle populations were monitored along a fly-round transect traversing the two main vegetation types in the study area. These were miombo, a one-storied open woodland with the genera Brachystegia and Julbernardia dominant, and munga, a one- or two-storied woodland where the principal tree genera were Acacia, Combretum and Terminalia. Concurrently, a capture/mark/release/recapture (CMRR) exercise was conducted along two other transects also traversing both vegetation types. The index of apparent abundance of tsetse (IAA) in miombo increased at the beginning of the rainy season (November), reached its peak at the end of the rainy season (April) and was low during the cold season (May to late August), but especially the hot dry season (September to late October). The IAA of tsetse in munga showed a pattern that was the reverse of that in miombo. The seasonal changes in the IAA of tsetse in both vegetation types were in accordance with changes in the movement patterns of tsetse between the two vegetation type as observed using CMRR. The distribution and abundance of cattle along the transect also showed a seasonal trend. This was especially so in munga, during the first three years of observations, where cattle abundance increased gradually from June onwards, reached a maximum at the end of the hot dry season (October-November) and declined steeply at the start of the rainy season (November-December). In both vegetation types, the monthly mean IAA of tsetse was positively correlated with the abundance of cattle in the previous month. It is concluded that the distribution of tsetse in cultivated area of the eastern plateau of Zambia undergoes substantial seasonal changes, which can partly be attributed to changes in the distribution of cattle. The implications of these observations for the control of tsetse are discussed.     

Seasonal development of secondary xylem and phloem in <Emphasis Type="Italic">Schizolobium parahyba</Emphasis> (Vell.) Blake (Leguminosae: Caesalpinioideae)

The cambial activity and periodicity of secondary xylem and phloem formation have been less studied in tropical tree species than in temperate ones. This paper describes the relationship between seasonal cambial activity, xylem and phloem development, and phenology in Schizolobium parahyba, a fast growing semideciduous seasonal forest tree from southeastern Brazil. From 2002 to 2003, wood samples were collected periodically and phenology and climate were recorded monthly in the same period. S. parahyba forms annual growth increments in wood, delimited by narrow initial parenchyma bands. The reduction of the cambial activity to a minimum correlates to the dry season and leaf fall. The higher cambial activity correlates to the wet season and the presence of mature leaves. In phloem, a larger conductive region was observed in the wet season, when the trees were in full foliage. The secondary phloem did not exhibit any incremental zone marker; however, we found that the axial parenchyma tends to form irregular bands.     

Seasonality and growth rings in lianas of Bignoniaceae

Lianas are one of the most important components of tropical forest, and yet one of the most poorly known organisms. Therefore, our paper addresses questions on the environmental and developmental aspects that influence the growth of lianas of Bignoniaceae, tribe Bignonieae. In order to better understand their growth, we studied the stem anatomy, seasonality of formation and differentiation of secondary tissues, and the influence of the cambial variant in xylem development on a selected species: Tynanthus cognatus. Afterwards, we compared the results found in T. cognatus with 31 other species of Bignonieae to identify general patterns of growth in lianas of this tribe. We found that cambial activity starts toward the end of the rainy season and onset of the dry season, in contrast to what is known for tropical trees and shrubs. Moreover, their pattern of xylem formation and differentiation is strongly influenced by the presence of massive wedges of phloem produced by a variant cambium. Thus, the variant cambium is the first to commence its activity and only subsequently does cambial activity progress towards the center of the regular region, leading to the formation of confluent growth rings. In summary, we conclude that: the cambium responds to environmental changes; the xylem growth rings are annual and produced in a brief period of about 2 months, something that may explain why lianas possess narrow stems; and furthermore, phloem wedges greatly influence cambial activity.     

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.     

Ectomycorrhizae in coastal miombo woodland of Tanzania

Summary Ectomycorrhizae were found in root samples of the treesAfzelia quanzensis Welw. andBrachystegia spiciformis Benth. (Caesalpiniaceae), collected in the coastal miombo type woodland 50 km west of Dar-es-Salaam, Tanzania. Root nodules with a structure resembling that of nitrogen-fixing root nodules of other leguminous plants were observed in theA. quanzensis material. The climate of the locality is rather dry, and strongly seasonal. In the tropics, ectomycorrhizae have previously been found only in humid or rain forest climate zones.     

Changes in the structure and composition of miombo woodlands mediated by elephants (Loxodonta africana) and fire over a 26-year period in north-western Zimbabwe

Changes in structure and composition of miombo woodlands mediated by elephants and fire were studied in 26-year-old permanent transects established in 1972 in north-western Zimbabwe. Elephants caused 48% decline in proportions of large trees (>11 cm diameter), significant reductions (30.9–90.9%) in tree heights, reductions in stem areas (43.5%) and densities (2.5%) of all trees. There were increases in proportions of small trees (64.8%), shrub canopy volumes (271%) and shrub densities (172%). These increases are attributed to natural recruitment because of longer fire-free periods and reduction of tree suppression effects on lower strata as a result of elephant-induced tree declines. Frequencies of occurrence of most species dropped by 28–89.6%. Brachystegia boehmii was replaced by Pseudolachnostylis maprouneifolia as the most dominant tree, largely because of high elephant preference for Brachystegia boehmii . A new suite of species, dominated by Combretaceae, increased in dominance resulting in local floristic changes. Reductions in old elephant (33.4%), old unknown (89.9%) and new elephant (13.7%) damage suggest that elephant occupancy of miombo woodlands has declined, possibly because of limited availability of preferred browse species. This study clearly shows that elephants and fire have contributed significantly to the changes in miombo woodlands in the area.     

Phenology and nutrition of miombo woodland trees in Zambia

Seasonality in nitrogen (N) and phosphorus (P) concentration in soil and shoots of five Brachystegia-Julbernardia (miombo) woodland trees was studied from September 1991 to March 1993 at two regrowth miombo sites in central Zambia. Shoot growth started in the dry season (September–November) and lasted until April during the 1991/92 season but had virtually ceased by January 1993 during the 1992/93 season. The shoot growing season was associated with low foliar N/P ratios. These ratios were much lower (<5) during the 1991/92 season than in the 1992/93 season (12–15). The increase in foliar N/P ratios after the shoot growing season was caused by a sharp drop in foliar P concentration, apparently due to reabsorption before leaf fall. There were no annual variations in biomass N concentration in contrast to P. During the 1992/93 growing season P concentrations in foliage and wood were a quarter and a third, respectively, of the 1991/ 92 levels. Since the short shoot growing season observed during the 1992/93 season is typical of savanna woodland trees in southern Africa, the high biomass P concentration and longer growing season in 1991/92 season were exceptional and may have been related to reduced competition by shallow rooting herbaceous plants caused by the severe drought of that season.     

Phenology of Tree Species in Bolivian Dry Forests

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub–canopy forest tree species were recorded monthly over a two–year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub–canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub–canopy tree species produced larger animal– or gravity–dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerio received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry forests.     

Influence of seasonal variation on the phenology and liriodenine content of Annona lutescens (Annonaceae)

Annona lutescens Saff. (Annonaceae) grows as a native tree in Chiapas, Mexico in Tropical Dry Forest habitat. Like most Annonaceae, it biosynthesizes benzylisoquinoline alkaloids, mostly liriodenine. To determine the influence of seasonal changes in the accumulation of liriodenine, the monthly variation of liriodenine content in roots, stems and leaves of mature and young trees was observed. These parts of young and mature A. lutescens trees were collected monthly over a 1 year period and the alkaloids were extracted; the liriodenine was quantified by high-resolution liquid chromatography. The phenological stages of the species were also assessed (leaf development, flowering and fruiting) using the Biologische Bundesanstalt, Bundessortenamt und Chemische Industrie (BBCH) scale. The analysis of both young and mature trees showed a significant increase in the liriodenine concentration occurs within roots during the dry season, which coincides with leaf fall. A significant decrease also occurred at the beginning of the rainy season (the period of leaf growth); the liriodenine content for the next rainy season did not reach the levels of the previous dry season. The climatic variation induced phenological and physiological changes in this species.     

Shading delays bud break in Brachsyegia spiciformis

Whole tree manipulation experiments were performed in the common southern African tree species, Brachystegia spiciformis to test a novel hypothesis that decreasing Total nonstructural carbohydrates (TNC) in the stem could cause bud break in Brachystegia spiciformis. The experimental treatments included fertilization, canopy defoliation, shading and stem heating to decrease stem carbohydrates. None of the treatments significantly decreased mean stem TNC. Likewise the heating, fertilization and defoliation treatments did not significantly affect the date of bud break. However, shading significantly delayed bud break. This delay in bud break could not be attributed to changes in leaf level photosynthetic traits, stem water content, leaf predawn water potential or delayed leaf fall. These results question widely accepted hypotheses about the mechanisms controlling bud break and suggest a carbohydrate homeostatic mechanism.