Dendrochronology of lianas of the Leguminosae family from the Atlantic Forest, Brazil

Information about plant growth, development and age forms the basis for understanding complex forest ecological processes. Although lianas play an important role in tropical forests, little is known about their growth and development from either climatic or ecological perspectives. Therefore, we studied the growth rings in Legume liana species collected in a mountainous Atlantic Forest in southeastern Brazil. Four of the eight studied species did not show cambial variants, three had a lobed stem, and one had a furrowed xylem. Distinct growth rings were observed in all species. Semi-ring porosity, marginal parenchyma, fibrous zone and radially flattened latewood cells were the main characteristic features of these growth rings. Species without cambial variants, including Dalbergia frutescens, Piptadenia adiantoides, P. micracantha and Senegalia tenuifolia, showed very distinct growth rings visible in macroscopic and microscopic analysis. Ring-width time series and cambial wound assessment were performed to analyze periodicity and dendrochronology. The species with cambial variants, S. grandistipula, S. lacerans, S. martiusiana and S. pedicellata, also showed distinct growth rings, however, sometimes barely detectable or not detected at all. Cambial wounding, cross-dating and climate-growth relationships indicated the annual nature of growth rings in species without cambial variant. Cross-dating between radii within one individual and between individuals was successful, and the synchronized series enabled us to build species chronologies and a mean chronology. Climate-growth analysis revealed significant correlations between chronologies and precipitation, indicating that available moisture is the main factor determining growth rates of lianas in the Atlantic forest.     

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.     

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...     

热带雨林木质藤本植物叶片性状及其关联

热带雨林中木质藤本植物较为丰富。随着全球气候变化加剧,木质藤本植物的丰富度具有不断增加的趋势,有可能对热带森林的结构、功能和动态产生重要影响。然而,目前对木质藤本响应环境变化的机制所知甚少。本研究以13个科20种热带雨林常见木质藤本植物为材料,测定了冠层叶片的17个形态特征及结构性状,并分析了性状间的相互关系。结果表明,叶片相对含水量的种间变异最小（变异系数为5%）,而上表皮厚度的种间变异最大（变异系数为80%）,其它性状的种间变异系数为24%~61%。木质藤本植物的叶脉密度、叶片密度均与气孔密度呈显著正相关,叶片干物质含量与比叶面积呈显著负相关。与相同生境的树木相比,木质藤本的叶面积更小、气孔密度和叶片密度更低、比叶面积更高,但两种植物类群的叶片横切面组织结构厚度无显著差异。研究结果对理解木质藤本植物的生态适应性具有重要意义。     

The negative effect of lianas on tree growth varies with tree species and season

Lianas reduce tree growth, reproduction, and survival in tropical forests. Liana competition can be particularly intense in isolated forest fragments, where liana densities are high, and thus, host tree infestation is common. Furthermore, lianas appear to grow particularly well during seasonal drought, when they may compete particularly intensely with trees. Few studies, however, have experimentally quantified the seasonal effects of liana competition on multiple tree species in tropical forests. We used a liana removal experiment in a forest fragment in southeastern Brazil to test whether the effects of lianas on tree growth vary with season and tree species identity. We conducted monthly diameter measurements using dendrometer bands on 88 individuals of five tree species for 24 months. We found that lianas had a stronger negative effect on some tree species during the wet season compared to the dry season. Furthermore, lianas significantly reduced the diameter growth of two tree species but had no effect on the other three tree species. The strong negative effect of lianas on some trees, particularly during the wet season, indicates that the effect of lianas on trees varies both seasonally and with tree species identity. Abstract in Portuguese is available with online material.     

Do lianas really have wide vessels? Vessel diameter–stem length scaling in non-self-supporting plants

Lianas and other climbing plants are known for their extraordinarily wide vessels. Wide vessels are thought to contribute to the extreme hydraulic efficiency of lianas and to play a part in their ability to dominate many tropical habitats, and even their globally increasing abundance with anthropic disturbance. However, recent hydraulic optimality models suggest that the average vessel diameter of plants generally is the result of tip-to-base vessel widening reflecting the effects of selection buffering conductive path length-imposed hydraulic resistance. These models state that mean vessel diameter should be predicted by stem length, by implication even in lianas. We explore vessel–stem relations with 1409 samples from 424 species in 159 families of both self- and non-self-supporting plants. We show that, far from being exceptional in their vessel diameter, lianas have average natural (not hydraulically weighted) vessel diameters that are indistinguishable for a given stem length from those in self-supporting plants. Lianas do, however, have wider variance in vessel diameter. They have a small number of vessels that are wider than those in self-supporting plants of similar stem lengths, and also narrower vessels. This slightly greater variance is sufficient to make hydraulically weighted vessel diameters in lianas higher than those of self-supporting counterparts of similar stem lengths. Moreover, lianas have significantly more vessels per unit of wood transection than self-supporting plants do. This subtle combination of slightly higher vessel diameter variance and higher vessel density for a given stem length is likely what makes lianas hydraulically distinctive, rather than their having vessels that are truly exceptionally wide.     

Ethnobotany, Availability, and Use of Lianas by the Kaingang People in Suburban Forests in Southern Brazil

Ethnobotany, Availability, and Use of Lianas by the Kaingang People in Suburban Forests in Southern Brazil. Lianas are important components of tropical forests and also non-timber forest products (NTFPs). In this study we 1) record the species of lianas exploited by indigenous Kaingang people and their availability in the remnant forest patches around the city of Porto Alegre, Southern Brazil, 2) describe the harvesting and crafting techniques, and 3) estimate the amount of lianas sold as handicrafts in the main local market and the income obtained. Lianas are exploited exclusively for the handicrafts trade. Seven liana species, out of 31 found in forest remnants, are used by Kaingangs. Two of them account for 90% of all handicrafts sold and 40.5% of all liana stems in secondary growth remnants—Forsteronia glabrescens Müll.Arg. and Amphilophium paniculatum (L.) Kunth. Another three species are scarce and are exploited less. Raw material must be used within a few weeks and be continuously harvested. Kaingang harvesting strategy includes mechanisms to avoid overexploitation: the division of exploitation areas into plots that are left to rest for one year before a new harvesting campaign, and the family’s control of information about potential harvesting sites. The entire process, from collection to sale, is carried out by the Kaingang families without intermediaries, contrary to the most common pattern of NTFP market chains. Trading occurs only locally. Each family earns from USD 98.15 to USD 371.12 per month (30% to 115% of the minimum Brazilian salary). In one year each family consumed on average 10,328.40 meters of lianas in the manufacture of handicrafts for sale in the major fair in the city, which represents about 50% of the demand for lianas. We estimated that one hectare of unexploited forest can have on average a standing crop of 10,165.40 m of the most desired species. Market pressure regulates exploitation, a common pattern in NTFP use, but in this case leading to a mix of usually opposite trends—manufacture with increasingly sophisticated techniques, but using a few liana species and an increasing number of alternative materials.     

Liana communities exhibit different species composition,diversity and community structure across forest types in the Congo Basin

Lianas are poorly characterized for central African forests. We quantify variation in liana composition, diversity and community structure in different forest types in the Yangambi Man and Biosphere Reserve, Democratic Republic of Congo. These attributes of liana assemblages were examined in 12 1-ha plots, randomly demarcated within regrowth forest, old growth monodominant forest, old growth mixed forest and old growth edge forest. Using a combination of multivariate and univariate community analyses, we visualize the patterns of these liana assemblage attributes and/or test for their significant differences across forest types. The combined 12 1-ha area contains 2,638 lianas (≥2 cm diameter) representing 105 species, 49 genera and 22 families. Liana species composition differed significantly across forest types. Taxonomic diversity was higher in old growth mixed forests compared to old growth monodominant and regrowth forests. Trait diversity was higher than expected in the regrowth forest as opposed to the rest of forest types. Similarly, the regrowth forest differed from the rest of forest types in the pattern of liana species ecological traits and diameter frequency distribution. The regrowth forest was also less densely populated in lianas and had lower liana total basal area than the rest of forest types. We speculate that the mechanism of liana competitive exclusion by dominant tree species is mainly responsible for the lower liana species diversity in monodominant compared to mixed forests. We attribute variation in liana community structure between regrowth and old growth forests mostly to short development time of size hierarchies.     

Thirteen years of change in trees and lianas in a Gabonese rainforest

Inventories of trees and lianas were performed at a 13-year interval (1979–1992) in an evergreen forest in north-east Gabon. The number of trees decreased while their basal area and species richness increased. In contrast, both the richness and density of lianas decreased. Lianas with active climbing mechanisms were the main types to disappear.     

Liana Abundance,Diversity, and Distribution on Barro Colorado Island,Panama

Lianas are a key component of tropical forests; however, most surveys are too small to accurately quantify liana community composition, diversity, abundance, and spatial distribution – critical components for measuring the contribution of lianas to forest processes. In 2007, we tagged, mapped, measured the diameter, and identified all lianas ≥1 cm rooted in a 50-ha plot on Barro Colorado Island, Panama (BCI). We calculated liana density, basal area, and species richness for both independently rooted lianas and all rooted liana stems (genets plus clones). We compared spatial aggregation patterns of liana and tree species, and among liana species that varied in the amount of clonal reproduction. We also tested whether liana and tree densities have increased on BCI compared to surveys conducted 30-years earlier. This study represents the most comprehensive spatially contiguous sampling of lianas ever conducted and, over the 50 ha area, we found 67,447 rooted liana stems comprising 162 species. Rooted lianas composed nearly 25% of the woody stems (trees and lianas), 35% of woody species richness, and 3% of woody basal area. Lianas were spatially aggregated within the 50-ha plot and the liana species with the highest proportion of clonal stems more spatially aggregated than the least clonal species, possibly indicating clonal stem recruitment following canopy disturbance. Over the past 30 years, liana density increased by 75% for stems ≥1 cm diameter and nearly 140% for stems ≥5 cm diameter, while tree density on BCI decreased 11.5%; a finding consistent with other neotropical forests. Our data confirm that lianas contribute substantially to tropical forest stem density and diversity, they have highly clumped distributions that appear to be driven by clonal stem recruitment into treefall gaps, and they are increasing relative to trees, thus indicating that lianas will play a greater role in the future dynamics of BCI and other neotropical forests.     

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.     

Rapid Liana Colonization along a Secondary Forest Chronosequence

Lianas (woody vines) can have profound effects on tree recruitment, growth, survival, and diversity in tropical forests. However, the dynamics of liana colonization soon after land abandonment are poorly understood, and thus it is unknown whether lianas alter tree regeneration early in succession. We examined the liana community in 43 forests that ranged from 1 to 31 yr old in central Panama to determine how fast lianas colonize young forests and how the liana community changes with forest succession. We found that lianas reached high densities early in succession, commonly exceeding 1000 stems/ha within the first 5 yr of forest regeneration. Lianas also increased rapidly during early succession in terms of basal area but did not show evidence of saturation within the 30 yr of our chronosequence. The relative contribution of lianas to total woody plant community in terms of basal area and density increased rapidly and reached a saturation point within 5 yr (basal area) to 15 yr (density) after land abandonment. Our data demonstrate that lianas recruit early and in high density in tropical forest regeneration, and thus lianas may have a large effect on the way in which secondary forests develop both early and throughout succession.     

Cambial dormancy induced growth rings in <Emphasis Type="Italic">Heritiera fomes</Emphasis> Buch.- Ham.: a proxy for exploring the dynamics of Sundarbans,Bangladesh

Key message

Cambial marking experiment and cambial activity analysis offer strong evidence on existence of annual growth rings in Heritiera fomes and revealing the potential of dendrochronological applications in Bangladesh mangroves.

Abstract

Despite enormous significance in coastal protection, biodiversity conservation and livelihood support to the local communities, mangrove ecosystems have been continuously degrading mainly due to anthropogenic disturbances and climate change. Time series based on dated tree ring is an option to identify the causes of forest dilapidation. In this study, we investigated the structure and periodicity of the growth ring in Heritiera fomes, the flagship tree species of the Bangladesh Sundarbans, combining cambial marking experiment and cambial activity analysis. Distinct growth rings were found which are delineated by a band of marginal parenchyma, predominantly one cell wide but up to three and occasionally interrupted with fiber. Of the 13 trees with cambium marking experiment, one growth ring was found in each tree during a year. The dormant cambium was characterized by the abrupt boundary between xylem and cambial zone, absence of enlarging or differentiating cambial derivatives, lower number of cambial cells and thicker radial walls in cambial cells. Growth ring anomalies, i.e., wedging and partially missing rings were also found. In most of the cases, the lower part of the eccentric discs had low radial increment (<0.75 mm) and therefore the growth ring in that area merged with previous one and produced wedging or partially missing ring. However, the existence of annual rings suggests its great potential for future dendrochronological applications to reveal the dynamics of vegetation and climate in Sundarbans.

     

Dendrochronological potential of <Emphasis Type="Italic">Millettia stuhlmannii</Emphasis> in Mozambique

Key message

This study demonstrates that Millettia stuhlmannii produces annual growth rings responsive to seasonal climate and should be useful for dendrochronology.

Abstract

Millettia stuhlmannii is a highly valuable and potentially overexploited timber species indigenous to southeastern Africa. It is of particular economic importance in Mozambique though relatively little is known about its growth rate or response to climate. This study investigates whether M. stuhlmannii is potentially useful for dendrochronology—that is whether this species forms annual growth rings that are responsive to external forcing such as climate. Five methods were used to determine whether M. stuhlmannii growth rings are indeed annual in nature, including analysis of ring anatomy, dating trees of known age, cambial wounding, classical cross-dating, and comparison of annual growth to climate variables. Growth rings of Millettia stuhlmannii are distinct and well formed, young trees from plantations of known age formed an appropriate number of distinct annual rings, trees showed distinct wood reaction to cambial wounding, adding exactly one complete ring in one calendar year, cross-dating within and between trees was somewhat successful, and annual growth is significantly correlated with wet season precipitation. Results of this study indicate that M. stuhlmannii is a potentially useful species for dendrochronology. These findings should allow a better understanding of this species’ growth dynamics and ecology, as well as its response to climate variability in the past and potentially to future climate change.

     

元江干热河谷木质藤本的多样性及其与宿主树木的关系

木质藤本是森林生态系统的重要组分。本研究在元江干热河谷地区随机设置了30个20 m×20 m的样方,调查样方中胸径≥0.5 cm的木质藤本多样性及其与宿主树木之间的关系。结果显示:30个样方中记录到胸径≥0.5 cm的木质藤本植物共945株(隶属于22种20属11科),其中,豆科木质藤本的丰富度和多度最高;胸径≤2 cm的木质藤本占个体总数的63.7%;茎缠绕类木质藤本的个体数最多。样方中胸径≥5 cm的树木共有1060株(隶属于38种31属16科),36.0%的树木上至少附藤1株。不同径级和不同树皮粗糙度的树木被木质藤本侵扰的百分比之间存在极显著差异(P0.001)。随着宿主树木平均枝下高的增加,附藤率呈下降趋势。76.5%的木质藤本选择离其根生长点最近的树木进行攀援。表明元江干热河谷中的木质藤本以小径级占优势,树木胸径、枝下高、树皮粗糙度和木质藤本根生长点到树木的距离是影响木质藤本侵扰树木的重要因素,支持木质藤本对宿主树木的侵扰具有选择性的假说。研究结果对中国西南干热河谷退化植被的恢复与物种多样性保护具有重要意义。     

Interspecific variation in clonality in temperate lianas revealed by genetic analysis: Do clonal proliferation processes differ among lianas?

Lianas are important components of forest communities with a considerable impact on overall forest structure and function. Lianas are characterized by extensive clonal reproduction on the forest floor, which is important for their establishment and growth. Previous studies have suggested that clonal reproduction strategies vary substantially among liana species; however, few studies have quantitatively evaluated the clonality strategy in multiple co-occurring liana species. The primary objective of the present study was to evaluate the relative contribution of clonal reproduction and to understand the clonal proliferation processes in co-occurring liana species by assessing both small stems on the forest floor and mature stems that climbed trees. The clonal reproduction strategy in four common liana species (stem twiner: Wisteria floribunda; root climbers: Schizophragma hydrangeoides, Euonymus fortunei and Rhus ambigua) in a 6-ha plot and a belt transect within an old-growth temperate forest in central Japan was evaluated using genetic analysis. The contribution of clonal reproduction was smaller in root climbers than in W. floribunda. All W. floribunda genets with small ramets in the understory had genetically identical ramets that climbed trees, whereas few such ramets were found in root climbers. This indicates that W. floribunda mature ramets laterally produce small ramets via clonal proliferation, whereas seedlings of root climbers grow horizontally to climb trees. The results indicate that the clonal reproduction processes differ greatly among lianas and the clonal growth in lianas plays a contrasting role in their life-history strategy.     

Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

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质藤本植物的多样性与空间分布

木质藤本植物是热带、亚热带山地森林重要的组分之一, 在森林动态、生态系统过程和森林生物多样性形成与维持等方面具有重要作用。本文调查了哀牢山中山湿性常绿阔叶林木质藤本植物的多样性及其在垂直和水平空间上的分布规律。在20个20 m × 50 m的样地中共调查到DBH≥0.2 cm的木质藤本植物1,145株, 隶属于19科25属29种, 其中物种最丰富的科为菝葜科(4种)和蔷薇科(3种), 但多度最高的科为葡萄科(363株, 占总株数的31.7%)。研究发现林下木质藤本(通常DBH < 1 cm)拥有较高的物种丰富度和多度, 对木质藤本植物多样性具有较大的贡献。有55.7%的个体分布在林下层, 林冠层占28.8%, 亚冠层只有15.5%。木质藤本的垂直空间分布在不同径级、不同攀援类型之间具有明显的差异。 从水平空间分布来看, 地形是影响木质藤本的一个重要因素: 沟谷木质藤本的物种丰富度、多度和基面积分别是坡面的171%, 420%和606%; 有12个物种只分布在沟谷生境。这表明哀牢山中山湿性常绿阔叶林木质藤本植物对生境具有偏好性。