Growth-ring boundaries of tropical tree species: Aiding delimitation by long histological sections and wood density profiles

Recent methodological advances have opened new perspectives for tropical dendrochonological studies by facilitating the visualization, delimitation, and analyses of tree-rings. One of those improvements was brought by X-ray densitometry, which allows building radial wood density profiles at microscopic scale. Furthermore, recent methods allow for cutting long histological sections to study anatomical variations along the entire radius of trees. These techniques have mainly been applied to low wood-density species from temperate and Mediterranean regions, with only limited applications in the tropics. Here we provide an improved protocol that allows for obtaining long histological sections of tropical woods, apply it to six species with varying wood densities 0.45−0.85 g cm⁻³ (Eucalyptus grandis, Tectona grandis, Acacia mangium, Cedrela fissilis, Hymenaea courbaril, and Copaifera duckei), and explore potential applications for tree-ring analyses. We provide instructions on core-microtome knife adjustments, procedures for softening and sectioning long histological samples of high wood-density species. We also present a multi-proxy approach that combines X-ray density profiles with the histological sections that improve the characterization and distinction of the various and complex tropical growth rings anatomical markers (fibre zone, marginal parenchyma, and ring porosity). This multi-proxy approach also opens the door for obtaining quantitative anatomy and physical parameters of tropical species with (intra-annual resolution. Our proposed approach is thus not only an additional tool to improve ring-boundary delimitation of tropical species, but it also paves the way to more innovative, borderline approaches in tropical dendrochronology.     

Cedrela nebulosa: A novel species for dendroclimatological studies in the montane tropics of South America

Up to now, the development of dendrochronological records from tropical regions in South America has been limited to the lowlands with emphasis in the Amazon basin. In this contribution, we present the first chronology of Cedrela nebulosa, a species that develops in the tropical mountainous regions of South America. We collected samples from trees in Monobamba district in Peru, analysed the anatomical features that determine the growth rings, and processed following the methods commonly used in dendrochronology. The 133-years chronology covering the 1883–2015 period, showed large correlation between series. In order to determine the climatic variables that control tree growth, we performed correlation analyses between tree-growth and local and regional precipitation and temperature records. We found that precipitation triggers tree growth at the beginning of the spring season but temperature seems to be the main control in annual growth. Also, C. nebulosa chronology present coherent variations with Multivariate Enso Index (MEI) and Pacific Ocean sea surface temperatures during summer months. This climate-sensitive tree-ring record indicates good potential for dendroclimatic studies and provides an opportunity to reconstruct climatic variations in montane forests of the tropical Andes.     

Wood day capacitance is related to water content,wood density,and anatomy across 30 temperate tree species

Water released from wood during transpiration (capacitance) can meaningfully affect daily water use and drought response. To provide context for better understanding of capacitance mechanisms, we investigated links between capacitance and wood anatomy. On twigs of 30 temperate angiosperm tree species, we measured day capacitance (between predawn and midday), water content, wood density, and anatomical traits, that is, vessel dimensions, tissue fractions, and vessel–tissue contact fractions (fraction of vessel circumference in contact with other tissues). Across all species, wood density (WD) and predawn lumen volumetric water content (VWC_L-pd) together were the strongest predictors of day capacitance (r²_adj = .44). Vessel–tissue contact fractions explained an additional ~10% of the variation in day capacitance. Regression models were not improved by including tissue lumen fractions. Among diffuse-porous species, VWC_L-pd and vessel–ray contact fraction together were the best predictors of day capacitance, whereas among semi/ring-porous species, VWC_L-pd, WD and vessel–fibre contact fraction were the best predictors. At predawn, wood was less than fully saturated for all species (lumen relative water content = 0.52 ± 0.17). Our findings imply that day capacitance depends on the amount of stored water, tissue connectivity and the bulk wood properties arising from WD (e.g., elasticity), rather than the fraction of any particular tissue.     

A comparison of the wood anatomy of 11 species from two cerrado habitats (cerrado s.s. and adjacent gallery forest)

A comparative study of the secondary xylem (wood) anatomy of 11 species (38 specimens) occurring in cerrado s.s. and the adjacent gallery forest (both cerrado s.l. habitat) was made with the aim of identifying the anatomical characteristics of ecological value and correlating them with the environmental conditions. The anatomical features that vary, in general, between the two habitats are: growth ring distinctness (well or poorly defined); tyloses and deposits (more abundant in cerrado specimens); gelatinous fibres (more evident in cerrado specimens and in different patterns between habitats); variation in paratracheal and banded parenchyma (more abundant in cerrado); and more cells per parenchyma strand in cerrado. In general, gallery forest specimens have wider vessels, fewer vessels per square millimetre and larger intervessel pits, indicating more efficient water conduction, whereas cerrado s.s. specimens are the opposite, with low vulnerability and mesomorphy indices, demonstrating greater safety under conditions of water stress. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ?? , ??–??.     

Fossil wood of the Juglandaceae: Some questions of taxonomy,evolution, and phylogeny in the family based on wood anatomy

Some problems in the taxonomy of the Juglandaceae are discussed based on wood anatomy; the identification of fossil juglandaceous wood is considered. Data on fossil wood of the Juglandaceae are summarized; a key for identification of wood anatomy in modern and fossil Juglandaceae is compiled. Wood anatomical characters in members of the family are discussed in the light of major evolutionary trends in the secondary xylem of dicots, and a comparative characterization of members of the family is developed. A hypothesis is proposed that the subfamily Engelhardioideae is the most primitive member of the Juglandaceae based on wood anatomy, the tribe Juglandeae and subfamily Platycaryoideae are slightly more highly specialized, and the tribe Hicorieae is the most advanced. Evolutionary relationships between the members of the Juglandaceae are reviewed based on wood anatomy.     

Asteropeia and Physena (Caryophyllales): A case study in comparative wood anatomy

Previous analyses ofAsteropeia andPhysena have not compared the wood anatomy of these genera to those of Caryophyllales s.l. Molecular evidence shows that the two genera from a clade that is a sister group of the core Caryophyllales. Synapomorphies of theAsteropeia-Physena clade include small circular alternate pits on vessels, presence of vasicentric tracheids plus fiber-tracheids, presence of abaxial-confluent plus diffuse axial parenchyma, and presence of predominantly uniseriate rays. These features are analyzed with respect to habit and ecology of the two genera. Solitary vessels, present in both genera, are related to the presence of vasicentric tracheids. Autapomorphies in the two genera seem related to adaptations byPhysena as a shrub of moderately dry habitats (e.g., narrower vessel elements, abundant vasicentric tracheids, square to erect cells in rays) as compared to alternate character expressions that seem related to the arboreal habit and humid forest ecology ofAsteropeia. The functional significance of vasicentric tracheids and fiber-tracheids in dicotyledons is briefly reviewed in the light of wood anatomy of the two genera.     

广义锦葵科火绳树属4个种的枝条木材解剖及其分类学意义

研究了广义锦葵科火绳树属4个种枝条的木材解剖。火绳树属枝条为散孔至半环孔材,管孔主要为单管孔和2～3个管孔组成的径列复管孔;导管间纹孔式和射线导管间纹孔式互列、小;侵填体和螺纹加厚缺如。射线主要为单列射线,2～3列射线较多;射线细胞多为方形,射线组织主要为异型,边缘直立细胞常1行;射线组织稀为同型;鞘细胞和瓦形细胞缺如。轴向薄壁组织傍管和离管型,主要为带状。晶体丰富,主要在射线、纤维和薄壁组织中。研究的4个种可以通过枝条木材解剖特征加以区分。     

Wood anatomy and hydraulic architecture of stems and twigs of some Mediterranean trees and shrubs along a mesic-xeric gradient

Woody species populating the major Mediterranean ecosystems in the world are characterized by different levels of adaptation to the seasonal Mediterranean climate conditions. Many species of these ecosystems show wood features that allow high efficiency of transport when water is available, while maintaining hydraulic safety during drought periods. This study focuses on the anatomy of juvenile and mature wood of some species representative of continuous sequences of Mediterranean vegetation formations according to gradients of water availability, from xeric to relatively mesic: Cistus monspeliensis L., Rhamnus alaternus L., Myrtus communis L., Pistacia lentiscus L., Olea europaea L., Quercus ilex L., Fraxinus ornus L. and Ostrya carpinifolia L. Twigwood collected in Southern Italy was anatomically compared with the stemwood of the same species represented in the reference slide collection of the National Herbarium of the Netherlands (Lw). The “hydraulic distance” between the wood of main stems and twigs was estimated on the basis of suites of anatomical features related to water efficiency/safety. Although some attributes (i.e. porosity and type of imperforate tracheary elements) were similar in young twigs and older rings, other traits (i.e. vessel frequency and size) evidenced the different hydraulic properties of twig and stemwood. The difference between juvenile and mature structures was large in the species of the mesic end of the gradient while it was relatively small in those more xeric. This tendency is in agreement with the habit gradient from medium-sized trees to small evergreen/drought deciduous shrubs according to decreasing water availability in Mediterranean vegetation types.     

Low-temperature resistance in Polylepis tarapacana, a tree growing at the highest altitudes in the world

The Polylepis tarapacana forests found in Bolivia are unique with respect to their altitudinal distribution (4200–5200 m). Given the extreme environmental conditions that characterize these altitudes, this species has to rely on distinct mechanisms to survive stressful temperatures. The purpose of this study was to determine low‐temperature resistance mechanisms in P. tarapacana. Tissue was sampled for carbohydrate and proline contents and micro‐climatic measurements were made at two altitudes, 4300 and 4850 m, during both the dry cold and wet warm seasons. Supercooling capacity (?3 to ?6 °C for the cold dry and ?7 to ?9 °C for the wet warm season) and injury temperatures (?18 to ?23 °C for both seasons), determined in the laboratory, indicate that P. tarapacana is a frost‐tolerant species. On the other hand, an increase in supercooling capacity, as the result of significant increase in total soluble sugar and proline contents, occurs during the wet warm season as a consequence of higher metabolic activity. Hence, P. tarapacana, a frost‐tolerant species during the colder unfavourable season, is able to avoid freezing during the more favourable season when minimum night‐time temperatures are not as extreme.     

Contracting montane cloud forests: a case study of the Andean alder (Alnus acuminata) and associated fungi in the Yungas

Alnus acuminata is a keystone tree species in the Yungas forests and host to a wide range of fungal symbionts. While species distribution models (SDMs) are routinely used for plants and animals to study the effects of climate change on montane forest communities, employing SDMs in fungi has been hindered by the lack of data on their geographic distribution. The well‐known host specificity and common biogeographic history of A. acuminata and associated ectomycorrhizal (ECM) fungi provide an exceptional opportunity to model the potential habitat for this symbiotic assemblage and to predict possible climate‐driven changes in the future. We (1) modeled the present and future distributions of suitable habitats for A. acuminata; (2) characterized fungal communities in different altitudinal zones of the Yungas using DNA metabarcoding of soil and root samples; and (3) selected fungi that were significant indicators of Alnus. Fungal communities were strongly structured according to altitudinal forest types and the presence of Alnus. Fungal indicators of Alnus, particularly ECM and root endophytic fungi, were also detected in Alnus roots. Current and future (year 2050) habitat models developed for A. acuminata predict a 25–50 percent decrease in suitable area and an upslope shift of the suitable habitat by ca. 184–380 m, depending on the climate change scenario. Although A. acuminata is considered to be an effective disperser, recent studies suggest that Andean grasslands are remarkably resistant to forest invasion, and future range contraction for A. acuminata may be even more pronounced than predicted by our models.     

Carbon and water balance in Polylepis sericea,a tropical treeline species

Polylepis sericea trees grow well above the continuous forest line in the Venezuelan Andes. In these environments, extreme daily temperature ranges can occur at any time of the year and trees experience a 4 month dry period. The purpose of this work was to study carbon and water relations of this species in the field during wet and dry seasons in order to understand this species' success at such high altitudes. Leaf gas exchange (portable system in open mode) and leaf water potential (pressure chamber) were measured at 1–2 h intervals during several daily courses at 4000 m elevation in the Páramo de Piedras Blancas. CO₂ assimilation versus leaf temperature curves were also obtained for this species in the laboratory. Clear differences in the measured parameters were observed between seasons. For a wet season day, maximum CO₂ assimilation rate was 7.4 mol m^-2 s^-1 and leaf conductance was relatively constant (approximately 100 mmol m^-2 s^-1)In the dry season day, maximum CO₂ assimilation rate was 5.8 molm^-2 s^-1 and leaf conductance was close to 60 mmolm^-2 s^-1. Minimum leaf water potentials measured were -1.3 MPa for the wet and -2.2 MPa for the dry season. The CO₂ assimilation-leaf temperature relationship showed a 13.4°C leaf temperature optimum for photosynthesis with maximum and minimum compensation points of 29.5 and -2.8°C, respectively. Maximum night-time respiration was relatively high (2.7 (imol) m^-2 s^-1)Our results show thatP. sericea maintains a highly positive carbon balance through all daily courses, even though there is a slight water stress effect during the dry season; this suggests that its carbon assimilation machinery is well adapted to the low temperatures and seasonal water stress found in the high tropical mountains.     

Robinia pseudoacacia as a surrogate for native tree species for saproxylic beetles inhabiting the riparian mixed forests of northern Italy

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Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.     

Studying global change through investigation of the plastic responses of xylem anatomy in tree rings

Variability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, sub-annual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that low-cost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.     

Latitudinal trends in wood anatomy within species and genera: case study in Cornus s.l. (Cornaceae)

Latitudinal trends in wood anatomical characters in three Asiatic species of Cornus sensu lato (s.l.) were studied and compared with those for the whole genus based on an extensive sampling covering the specific distribution ranges and the generic data from a previous study. We studied 124 specimens of C. controversa growing between 31.5° and 45.3° N, 54 of C. kousa between 24.4° and 40.5° N, and 64 of C. macrophylla between 27.8° and 41.0° N. Characters studied were vessel element length, fiber length, vessel frequency, tangential vessel diameter, and vessel grouping index. At the species level no latitudinal trends were detected throughout the distribution ranges of the species. Neither tree size, altitude, nor climatic factors had a significant correlation with wood anatomical characters. In contrast, at the genus level, latitudinal trends were significant not just for the whole genus, but for both New and Old World species groups. At the genus level, latitude and three climatic factors all had a significant correlation with wood anatomical characters, but correlation coefficients with latitude were markedly high. The difference in latitudinal trends between the genus and species levels may be due to the radiation of Cornus along paleoclimatic gradients in the early Tertiary.