A global review on wood growth rings in lianas

Lianas (woody vines) have a critical role in ecosystem dynamics, especially in the tropics. At present, there is an increasing need to develop reliable methods to measure liana’s growth and age, given their suggested increase in abundance following deforestation events and climatic changes. Dendrochronological analyses offer powerful tools that provide temporal information for ecological studies at different scales, with the first step being the detection of distinct growth rings and compiling a list of species with this feature. Hence, this study offers a thorough list of lianas with distinct growth ring markers across both gymnosperm and angiosperm lianas, with and without cambial variants, from both north and southern hemispheres, based on global surveys, bibliographical references and our own collections. Lianas with distinct growth rings are common in species from both temperate and tropical regions. We found 530 lianas with distinct growth rings belonging to 74 families. Bignoniaceae, Celastraceae, Malpighiaceae, Menispermaceae, and Leguminosae are the families with more species listed. Thick-walled and/or radially flattened latewood fibres, semi-ring-porosity, marginal parenchyma, ring-porosity were the main anatomical markers delimiting growth rings in lianas. Future analyses evaluating the periodicity of growth ring formation, mainly in species with cambial variants, are the next step to advance dendroecological application in lianas.     

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.     

海南霸王岭山地原始林与伐后林中木质藤本对支持木的选择

通过对海南霸王岭热带山地原始林与伐后林中树木及其攀附木质藤本的调查,研究原始林与伐后林中木质藤本对支持木的选择性。结果表明:1)6科优势树木中附藤率最高的是野牡丹科(Melastomataceae),附藤率最低的科,原始林中是山矾科(Symplocaceae),伐后林中是茜草科(Rubiaceae)。2)原始林中,谷木(Memecylon ligustrifolium)与线枝蒲桃(Syzygiumaraiocladum)的附藤比率和每木藤本数均高于样地平均水平;三角瓣花(Prismatomeris tetrandra)和龟背灰木(Symplocosandenophylla)的附藤比率均低于样地平均水平,而每木藤本数与样地平均水平之间没有显著差异。伐后林中,谷木的附藤比率和每木藤本数高于样地平均水平;九节(Psychotria rubra)的附藤比率和每木藤本数低于样地平均水平。3)杜仲藤(Parabariummicranthum)的主要支持木是谷木,夜花藤(Hypserpa nitida)的主要支持木是线枝蒲桃。研究表明,木质藤本对支持木在科和种水平上都具有选择性,因此木质藤本会对树木造成不对称影响,进而影响森林的结构和动态。     

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.     

Climatic signals in tree‐rings of Araucaria angustifolia in the southern Brazilian highlands

Araucaria angustifolia (Bertol.) O. Kuntze (Araucariaceae) is a Neotropical tree, widely distributed in subtropical mountain rain forests and nearby natural grasslands of Southern Brazil. This species produces annual growth rings, but its dendroclimatic potential is barely known. In the present paper, the long‐term growth patterns of A. angustifolia were investigated using annual growth ring time series and association to climate over the last century. Wood cores of A. angustifolia trees growing in forest and grassland habitats were obtained with an increment borer. The cores were surfaced, measured and cross‐dated. The dated ring‐width time series were standardized and submitted to correlation and principal component analysis to verify growth trends among sites and trees. Growth‐climate relationships were investigated using correlation and regression analyses, comparing the ordination axes scores to regional time series of precipitation and temperature. Due to anatomical irregularities, mainly partial rings, only 35 out of 60 trees were cross‐dated. The correlation and ordination analyses showed common tree‐growth trends within and between sites, indicative of a regional environmental force determining inter‐annual cambial activity variation. Despite growing in distinct habitats and disturbance regimes, A. angustifolia trees share a common long‐term growth pattern, which is significantly related to thermal conditions during the current and previous growing seasons. Moreover, site‐specific characteristics may have influenced opposite growth responses and association to climate conditions between forest and grassland trees.     

Assessing the utility of direct and indirect methods for estimating tropical tree age in the Western Ghats, India

The knowledge of tree age is important for understanding tree growth and forest dynamics. It may be estimated by ‘direct’ methods involving growth ring counts, or by ‘indirect’ methods involving field measurements of growth rates. Direct methods are considered more accurate, but it is not clear if they are appropriate for all species, notably from the humid tropics. In this paper we assess the occurrence of annual growth rings and their utility for age estimation in three tropical tree species, Acrocarpus fraxinifolius, Dalbergia latifolia (Fabaceae) and Syzygium cumini (Myrtaceae), growing in traditional shade coffee plantations of the southern Western Ghats, India. These species previously were described as having “indistinct or absent” growth rings. We used anatomical studies, field measurements and computational methods to characterise growth rings and assess similarities between directly and indirectly estimated tree ages. Our study revealed that annual growth rings were characterised by different sets of anatomical features per species and were most distinct in the fast-growing deciduous A. fraxinifolius. Growth rates measured in the field showed annual periodicity in all three species, and reflected annual rainfall-drought cycles in D. latifolia and S. cumini. Direct age estimates were most similar to indirect estimates in D. latifolia, and least so in S. cumini. The results of direct age estimation by counting rings are consistent with them being annual in nature in tropical species with distinct and reliable annual growth ring formation. However, for species with poorly defined growth rings, indirect age estimation methods might be more useful.     

Seasonal cambium activity in the subtropical rain forest tree Araucaria angustifolia

Information on the timing and dynamics of tree ring formation is essential to assess the seasonal behavior of secondary wood growth and its associated environmental influences. Araucaria angustifolia is a dominant species in highland pluvial ecosystems of southeastern South America. Previous investigations indicated that their growth rings are formed annually, but no information exists about the timing of growth ring formation and the environmental triggers influencing cambium activity. In this paper we examine inter- and intra-annual cambial activity in A. angustifolia, through anatomical and dendrochronological evidence at two study sites, and model the relationships between regional climate variation and intra-annual tree ring formation. The results confirm the annual growth ring formation in A. angustifolia and indicate that its growth season extends from October to April. Day length and temperature were the main environmental factors influencing the seasonal cambium activity. Our results evidence the dendrochronological potential of A. angustifolia for ecological and climatological studies in southeastern South America.     

Dendrochronology and dendroclimatology of Ceiba speciosa (A. St.-Hil.) Ravenna (Malvaceae) exposed to urban pollution in Rio de Janeiro city,Brazil

Nowadays, the biological monitoring through the growth rings has received increasing attention from ecologists and toxicologists. Structural analysis of these rings allows the incorporation of a time component in the study of plant responses to environmental variation. This allows also to evaluate long time series from the woody plants. In this paper, we assessed the dendrochronological characteristics of Ceiba speciosa growing in forest environment and under urbanization impact. Stem samples were obtained with Pressler probe into trees growing the campus of the Oswaldo Cruz Foundation, adjacent to one of the main urban thoroughfares of the city of Rio de Janeiro (Avenida Brasil), and at Tinguá Biological Reserve, an important remnant of Atlantic Forest. The samples were processed and analyzed following usual dendrochronological methods, with COFECHA and ARSTAN softwares. A negative exponential curve was used for standardization of the series. The residual chronologies were correlated with precipitation and temperature indexes obtained from NOAA weather database. Growth rings are distinct and annual, marked by bands of marginal parenchyma, thick-walled and radially flattened fibres in latewood and distended rays in earlywood. In both sites, the intercorrelation between the trees was above 0.40. Ages ranged from 11 to 41 years in the urban site and from 27 to 64 years in the forest site. In urban area, mean annual increment and cumulative average growth rates were 6 mm/year and 142.62 mm, respectively. At the forest site, these rates were 4 mm/year and 173.07 mm, respectively. The comparison between cumulative radial increment of the two sites revealed that trees of the urban site had higher increment rates beginning at the start of their development and consequently, they showed similar diameters despite lower ages. Correlation analysis between the chronologies and climatic factors revealed a positive association between growth and hot and rainy periods for both study sites. However, there is an immediate response of urban trees in relation to the rains and, a late response of forest trees to the same factor. The dry and hot climate, typical of urban environments, and the absence of natural water reserves in urban soil, may explain this more immediate response of urban tree growth to rainfall and temperature indexes. Our results revealed that Ceiba speciosa is a plastic and stress-tolerant species that is able to survive and adapt to polluted urban conditions. These features, along with its wide natural distribution and frequent planting for city landscaping, make this species an important biomarker for environmental monitoring studies.     

Mangrove growth rings: fact or fiction?

The analysis of tree rings in the tropics is less straightforward than in temperate areas with a demarcated unfavourable winter season. But especially in mangroves, the highly dynamic intertidal environment and the overriding ecological drivers therein have been a reason for questioning the existence of growth rings. This study aimed at casting light on growth rings in mangroves. In six mangrove species growing in Gazi Bay, Kenya (Sonneratia alba, Heritiera littoralis, Ceriops tagal, Bruguiera gymnorrhiza, Xylocarpus granatum and Lumnitzera racemosa), the occurrence of growth rings was examined. Growth rate of each species was determined based on a 1-year period using the cambial marking technique. The effect of climate was furthermore considered by comparing the results with a number of wood samples originating from contrasting climatic regions. We can conclude that for growth rings to appear in mangroves more than one condition has to be fulfilled, making general statements impossible and explaining the prevalent uncertainty. Climatic conditions that result in a range of soil water salinity experienced over the year are a prerequisite for the formation of growth rings. For species with an anatomy characterized by indistinct ring boundaries, this should be combined with a growth rate of at least 0.3 mm/year. The use of growth rings for age or growth rate determinations should thus be evaluated on a case by case basis.     

天山东部西伯利亚落叶松树轮生长对气候要素的响应分析

天山东部西伯利亚落叶松的树木年轮学研究可以看出：森林上限树轮宽度年表之间相关性较高而下限年表间相关稍低,表明下限小生境要素对树木生长干扰较大。森林上下限树轮年表中样本的总解释量（ESP）和信噪比（SNR）都比较高,说明树木中都含有较多的环境信息;但标准年表中平均敏感度（M．S．）和轮宽指数的标准差（S．D．）都是森林上限数值低于下限,这表明森林上限树木生长对环境变化响应的敏感性降低;相关分析和响应分析也发现森林下限生长的树木对气候因子的响应较为显著。就温度而言,森林上限和下限表现基本一致,树木生长多与温度负相关,其中下限树木生长与春季均温和3．6月份均温显著负相关;降水表现出一定的差别,上限树木生长与春季、夏季及年降水量有较高的负相关,而对下限树木生长影响最大的则是冬季和3—6月份降水。湿润指数与降水基本一致即上限呈负相关而下限正相关,温暖指数全为负相关,寒冷指数下限负相关显著;显然该地区森林上下限树木生长的生态模式存在着一定的差异。研究发现,冬春季节的不同水热组合则是形成树木年轮宽窄的限制因素;同时,前期生长的滞后效应对年轮形成有重要的影响。     

High-fidelity representation of climate variations by Amburana cearensis tree-ring chronologies across a tropical forest transition in South America

Numerous ring-width chronologies from different species have recently been developed in diverse tropical forests across South America. However, the temporal and spatial climate signals in these tropical chronologies is less well known. In this work, annual growth rings of Amburana cearensis, a widely distributed tropical tree species, were employed to estimate temporal and spatial patterns of climate variability in the transition from the dry Chiquitano (16–17°S) to the humid Guarayos-southern Amazon (14–15°S) forests. Four well-replicated chronologies (16–21 trees, 22–28 radii) of A. cearensis were compared with temperature and precipitation records available in the region. The interannual variations in all four A. cearensis tree-ring chronologies are positively correlated with precipitation and negatively with temperature during the late dry-early wet season, the classic moisture response seen widely in trees from dry tropical and temperate forests worldwide. However, the chronologies from the dry Chiquitano forests of southern Bolivia reflect the regional reduction in precipitation during recent decades, while the chronologies from the tropical lowland moist forests in the north capture the recent increase in precipitation in the southern Amazon basin. These results indicate that A. cearensis tree growth is not only sensitive to the moisture balance of the growing season, it can also record subtle differences in regional precipitation trends across the dry to humid forest transition. Comparisons with previously developed Centrolobium microchaete chronologies in the region reveal a substantial common signal between chronologies in similar environments, suggesting that regional differences in climate are a major drivers of tree growth along the precipitation gradient. The difficulty of finding A. cearensis trees over 150-years old is the main limitation involved in the paleoclimate application of this species. The expansion of monocultures and intensive cattle ranching in the South American tropics are contributing to the loss of these old growth A. cearensis trees and the valuable records of climate variability and climate change that they contain.     

Reconstruction of pre-monsoon weather conditions in northwestern Thailand from the tree-ring widths of Pinus merkusii and Pinus kesiya

Two pine species, Pinus merkusii and Pinus kesiya, native in Thailand were studied for their potential to reconstruct past weather conditions from their tree rings. Altogether, cores from 209 Merkus pines and from 205 Khasi pines were sampled at 16 sites. Standard methods were applied to assemble tree-ring chronologies for each site and tree species. The longest site chronology of Merkus pine and Khasi pine covered 314 and 183 years, respectively. Principal component analysis (PCA) illustrated a close clustering of all, except two, site chronologies on the PC1 axis, indicating a strong signal common for both pine species throughout NW Thailand. However, along the PC2 axis, there was a distinct separation between the Merkus pine and the Khasi pine site chronologies, which was possibly due to the species-specific differences in site altitudes. Simple correlations between the PC1 time series and a regionalized climate data set showed a highly negative association with temperature during the pre-monsoon period from March to May. Since, pre-monsoon temperature and rainfall were negatively correlated with each other, we used the PC1 as a direct proxy for pre-monsoon weather conditions (warm/dry vs. cool/wet) back to 1834 AD.     

Radial stem variations of Tabebuia chrysantha (Bignoniaceae) in different tropical forest ecosystems of southern Ecuador

Stem diameter increments of the broadleaved deciduous tree species Tabebuia chrysantha were measured with high-resolution dendrometers in a tropical lower montane forest and in a dry forest in southern Ecuador, the latter showing a distinct dry season. Those analyses were complemented by wood anatomical studies on regularly collected microcores to determine the season of active cambial growth and the time of formation of annual growth boundaries. The length of the cambial active period varied between 3 and 7 months at the tropical lower montane forest and 2 and 4 months in the dry forest, respectively. During dry days, amplitudes of daily stem diameter variations correlated with vapour pressure deficit. During October and November, inter-annual climate variations may lead to dry and sunny conditions in the tropical lower montane forest, causing water deficit and stem diameter shrinkage in T. chrysantha. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall as well as vapour pressure deficit in certain periods of the year, indicating that rainfall plays a major role for tree growth.     

Stem growth rhythms in trees of a tropical rainforest in Southern Brazil

Key message

We demonstrate that tropical trees growing in wet climates can have a marked seasonality in cambium activity and stem growth associated with high temperature and day length of summer.

Abstract

Monitoring the rhythm of tree growth associated with the growth rings can contribute substantially to understanding forest dynamics and the management of tropical forests. In this study, we monitored the girth increment rhythm and described the wood characteristics (anatomy of growth rings, wood specific gravity) in 10 tropical tree species (103 individuals) naturally occurring in a wet and weakly seasonal region of Atlantic Forest in southern Brazil. We aimed to verify whether tree growth dynamics are associated with climate and woody anatomy in tropical trees with contrasting ecological characteristics. We installed permanent dendrometer bands and monthly assessed the girth increment for 22 months. We collected wood samples (non-destructive method), measured wood specific gravity and prepared permanent slides to characterize the growth ring markers. We found growth rings in all species (distinct in six species); deciduous species produced more distinguishable tree rings compared with semi-deciduous and evergreen tree species. Species varied in their accumulated girth growth (in average, from 1.83 to 62.64 mm), growth rates (1–15 %), and annual radial increment (0.16–5.44 mm). Girth increment was positively related to temperature and day length in five out of ten tree species, indicating the possible effects of these climatic variables in triggering cambial activity in these species. The growth pattern varied among species and was marginally associated to the tree deciduousness. We concluded that even in wet and less seasonal climates, there can be an association in the cambium activity and stem growth with the hotter and longer days of summer months.

     

Tree-ring distinctness,dating potential and climatic sensitivity of laurel forest tree species in Tenerife Island

Macaronesian laurel forests are the only remnants of a subtropical palaeoecosystem dominant during the Tertiary in Europe and northern Africa. These biodiverse ecosystems are restricted to cloudy and temperate insular environments in the North Atlantic Ocean. Due to their reduced distribution area, these forests are particularly vulnerable to anthropogenic disturbances and changes in climatic conditions. The assessment of laurel forest trees’ response to climate variation by dendrochronological methods is limited because it was assumed that the lack of marked seasonality would prevent the formation of distinct annual tree rings. The aims of this study were to identify the presence of annual growth rings and to assess the dendrochronological potential of the most representative tree species from laurel forests in Tenerife, Canary Islands. We sampled increment cores from 498 trees of 12 species in two well-preserved forests in Tenerife Island. We evaluated tree-ring boundary distinctness, dating potential, and sensitivity of tree-ring growth to climate and, particularly, to drought occurrence. Eight species showed clear tree-ring boundaries, but synchronic annual tree rings and robust tree-ring chronologies were only obtained for Laurus novocanariensis, Ilex perado subsp. platyphylla, Persea indica and Picconia excelsa, a third of the studied species. Tree-ring width depended on water balance and drought occurrence, showing sharp reductions in growth in the face of decreased water availability, a response that was consistent among species and sites. Inter-annual tree-ring width variation was directly dependent on rainfall input in the humid period, from previous October to current April. The four negative pointer years 1995, 1999, 2008 and 2012 corresponded to severe drought events in the study area. This study gives the first assessment of dendrochronological potential and tree-ring climate sensitivity of tree species from the Tenerife laurel forest, which opens new research avenues for dendroecological studies in Macaronesian laurel forests.     

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.     

森林木质藤本数量过度增长的机制与生态效应

木质藤本是维持森林物种多样性的组成部分。近年来，热带、亚热带森林尤其是次生林内的木质藤本数量的过度增长给森林的恢复和健康发展造成了威胁。至今，国内尚少有研究对森林木质藤本过多的现象、增长机制与生态效应进行综合认识。该文梳理国内外相关文献，从木质藤本数量增长的机制与生态效应进行分析和总结，综合相关研究认为：(1)木质藤本数量的增长与干旱化加剧、大气CO₂浓度上升、自然干扰和森林破碎化有关，在环境变化的情况下，木质藤本在形态、行为、生理等方面比树木更具优势，表现为更快的生长速率、更强的繁殖力和可塑性以及资源利用效率。(2)木质藤本主要通过遮荫胁迫、资源竞争和机械压力与损伤等方式对树木造成影响。(3)木质藤本过度增长在个体水平上会阻碍树木生长、生殖并引起树木死亡，在群落水平上会改变物种组成、降低多样性，在生态系统水平上会降低森林碳储量，改变碳、矿质养分和水分循环过程等。因此，建议结合野外长期监测与控制实验开展木质藤本数量动态与环境变化关系、森林干扰对木质藤本生长的影响、木质藤本对环境变化的响应及适应机理、木质藤本数量过度增长的生态效应评价研究。同时，应积极探索合理的森林...     

Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (A _a) of the woody species in the SF ranged from 11.2 to 34.5 μmol m⁻² s⁻¹, similarly to pioneer tree species from literature data in southeast Asia. The A _a and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas A _a in the MF ranged between 9 to 21 μmol m⁻² s⁻¹, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.