Evergreen Broad-leaved Trees Research Articles

Forest structure and regeneration of the Tertiary relict Taiwania cryptomerioides in the Gaoligong Mountains, Yunnan, southwestern China

We studied forests containing Taiwania cryptomerioides of various ages and habitats on the eastern slopes of the Gaoligong Mountains in terms of forest structure and composition, population structure (size, age), regeneration patterns, and persistence of the species in relation to their favored habitats. Taiwania thrives in unstable habitats on riverbanks in deep valleys, on steep slopes, on cliffs, on roadsides and by mountain paths at the altitudes of 1175-2500 m a.s.l. All these locations were subject to frequent landslides, whereas Taiwania was very rare at similar altitudes on stable gentle slopes or on mountain ridges free of major disturbances. The maximum age of Taiwania was calculated to be c. 1,872 yr, with 358 cm DBH (diameter at a height of 1.3 m) and 70 m high. The size and age classes of Taiwania in old-growth forests were multimodal, indicating that the regeneration varied by chance, depending on disturbances. In the old-growth forests where above-ground competition for light was intense, shade-intolerant and long-lived coniferous Taiwania became emergent (40–70 m), rising above a forest canopy comprised of more shade-tolerant evergreen broad-leaved trees. The reproduction of the species was mainly by means of minute wind-dispersed seeds falling into rock crevices on cliffs or a rocky forest floor, or on disturbed sites. These populations depended on disturbances or gap regeneration to survive. Taiwania gave way to evergreen broad-leaved tree species of Lithocarpus, Cyclobalanopsis, and Manglietia, and to other conifers such as Tsuga dumosa, where landslides were infrequent. Our results provide insights into the ecological characteristics and survival mechanisms of this East Asian paleoendemic conifer, and contribute to our understanding of the differentiation of forests.

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest.

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as βNRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.

Disentangling long- and short-term changes in perennial organ functions in seasonal environments: A model of foliar chlorophyll and nitrogen in saplings of four evergreen broad-leaved trees

Perennial organ functions of trees living in seasonal environments exhibit temporal changes that can be classified as long-term interannual changes and seasonal fluctuations within single years. However, few studies have separately quantified these changes from longitudinal measurement data or analyzed the relationships between them. We developed a hierarchical Bayesian statistical model consisting of three parts: a long-term interannual change expressed by consecutive annual linear trends, seasonal fluctuations with 26 values for two-week periods in a year, and a random effect for repeated measurements. The model can extract long-term interannual changes and seasonal fluctuations from longitudinal repeated measure data. The pattern of seasonal fluctuation, the amount of seasonal fluctuation, and the net annual change are expressed by the estimated model parameters. We applied our model to foliar chlorophyll (Chl) and nitrogen (N) content measured repeatedly on more than 1-year-old leaves of saplings in four evergreen broad-leaved tree species using nondestructive optical methods. The model successfully explained large variations in the Chl and N content. In general, seasonal fluctuations corresponded to the phenology of current-year leaves; Chl and N tended to decrease from the opening to maturation of new leaves and increased during the rest period. The magnitude of the decrease in the Chl and N content in the growth period of current-year leaves (Δγ) did not decrease noticeably as leaves aged. For the Chl content, Δγ was positively correlated with the maximum value before leaf opening across species. For the N content, Δγ and the maximum value before leaf opening were not clearly correlated across species, but were positively correlated within some species. A model parameter for annual linear trends in Chl and N varied from positive (indicating increasing trends) to negative values (indicating decrease) depending on species and leaf age in years.

Regional-scale directional changes in abundance of tree species along a temperature gradient in Japan.

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad-leaved, deciduous broad-leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad-leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad-leaved trees. Similarly, relative abundance of deciduous broad-leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional-type-level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.

Architectural differences among shaded saplings of four evergreen broad-leaved tree species in Japan.

Architectural differences among shaded saplings of four evergreen broad-leaved tree species in Japan.

Inhibitory Effects of Nano-Extract from Dendropanax morbifera on Proliferation and Migration of Vascular Smooth Muscle Cells.

The plant Dendropanax morbifera Léveille (D. morbifera), a subtropical broad-leaved evergreen tree, have been used in folk medicine for the treatment of infectious diseases, skin diseases, and other maladies. However, the effect of extracts from D. morbifera in vascuar diseases has not yet been reported. In this study, BrdU assay revealed that extracts from D. morbifera inhibit significantly the proliferation rate of Rat Aortic Smooth Muscle Cells (RAoSMCs) by -40% in treated samples compared to controls. Notably, 2-D wound healing assay and 3-D boyden chamber assay showed the significant reduction of RAoSMCs migration induced by serum in nano extracts treated groups by -50%. We further observed that the phosphorylated levels of Akt and ERK were significantly reduced by 70% in extracts treated RAoSMCs. Moreover, the expression levels of matrix metalloproteinase (MMP) 2 and 9 were significantly reduced by extracts from D. morbifera. Our results suggest that extracts from D. morbifera inhibit proliferation and migration in RAoSMCs via the modulation of phosphorylated levels of Akt and ERK. Subsequently, the reduced MMP2 and 9 expression might result to reduced migration of RAoSMCs.

Developing Optimal Site Prediction Model for Evergreen Broad Leaved Trees, Machilus thunbergii in Warm Temperate Zone of the Korean Peninsula

Developing Optimal Site Prediction Model for Evergreen Broad Leaved Trees, Machilus thunbergii in Warm Temperate Zone of the Korean Peninsula

조경용 상록활엽수로서 종가시나무의 종자발아 및 유묘생장 특성

조경용 상록활엽수로서 종가시나무의 종자발아 및 유묘생장 특성

Responses of the photosynthetic apparatus to winter conditions in broadleaved evergreen trees growing in warm temperate regions of Japan

Photosynthetic characteristics of two broadleaved evergreen trees, Quercus myrsinaefolia and Machilus thunbergii, were compared in autumn and winter. The irradiance was similar in both seasons, but the air temperature was lower in winter. Under the winter conditions, net photosynthesis under natural sunlight (Anet) in both species dropped to 4 μmol CO2 m−2 s−1, and the quantum yield of photosystem II (PSII) photochemistry in dark-adapted leaves (Fv/Fm) also dropped to 0.60. In both species the maximum carboxylation rates of Rubisco (Vcmax) decreased, and the amount of Rubisco increased in winter. A decline in chlorophyll (Chl) concentration and an increase in the Chl a/b ratio in winter resulted in a reduction in the size of the light-harvesting antennae. From measurements of Chl a fluorescence parameters, both the relative fraction and the energy flux rates of thermal dissipation through other non-photochemical processes were markedly elevated in winter. The results indicate that the photosynthetic apparatus in broadleaved evergreen species in warm temperate regions responds to winter through regulatory mechanisms involving the downregulation of light-harvesting and photosynthesis coupled with increased photoprotective thermal energy dissipation to minimize photodamage in winter. These mechanisms aid a quick restart of photosynthesis without the development of new leaves in the following spring.

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits.

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity-Ecosystem Functioning experiment at Jiangxi (BEF-China). Information-theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi-layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi-predictor models for stomatal conductance (gs ) and xylem cavitation vulnerability. In addition, maximum gs was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum gs increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ50 ) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ50 values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.

Modeling qualitative and quantitative elements of branch growth in saplings of four evergreen broad-leaved tree species growing in a temperate Japanese forest

Factors influencing branch growth differed between a qualitative element (whether a branch grows) and a quantitative element (the amount of biomass accretion). We measured branch growth in saplings of four evergreen broad-leaved tree species growing in a temperate Japanese forest. A hierarchical Bayesian approach was used to model two elements of branch growth: a qualitative component [whether branches produce new annual shoots (ASn)] and a quantitative component (total ASn mass). The two components were influenced by somewhat different factors. The probability that a branch would produce ASn was affected by the total mass of previous-year annual shoots (total ASn–1 mass), branch age, relative branch height, canopy openness measured at the branch tip, maximum canopy openness within a single sapling, and the interaction between canopy openness and maximum canopy openness. The total ASn mass was influenced by the total ASn–1 mass, relative branch height, branch inclination, and maximum canopy openness. These two components of branch growth should be considered separately when tree architecture is modeled. In addition, we detected interactive responses among branches within individual saplings. The relative importance of interactive modular response and branch autonomy may differ depending on the status of plants, such as individual age, environmental conditions, and the timing of the measurements. We found considerable similarities in the responses of branch growth to the internal and environmental factors among the species studied. We also found some among-species differences in branch growth responses to the explanatory variables measured.

제주도 곶자왈 상록활엽수 종가시나무의 생물량 추정을 위한 상대생장식

This study developed allometry equation and estimated the aboveground-biomass of Quercus glauca, a warm-temperature, evergreen broad-leaved tree, growing in Kotzawal wetland located on Jeju Island. The allometric equations between DBH(diameter at breast height) and dry weights of stems (Ws), branches (Wb), leaves (Wl) and aboveground biomass (Wab) of Q. glauca were as follows: logWs

Changes in the potential habitats of 10 dominant evergreen broad-leaved tree species in the Taiwan-Japan archipelago

Ecosystem vulnerability to climate change remains elusive in the species-rich Taiwan-Japan archipelago. We predicted potential habitats (PHs) of ten dominant evergreen broad-leaved tree species by using the current and twenty potential climate change scenarios using generalised additive models. The presence/absence records of each species, extracted from vegetation database, were used as response variables. Four climatic and one spatial variables were used as explanatory variables. The results showed that the interaction terms of spatial variable, indicating historical range shifts or species interactions, restricted the distribution of all the target species as much as that by the each climatic variable. The PHs of all the target species were predicted to consistently increase, and in particular, to expand northward and upward to the cool temperate zone. However, the PHs were predicted to decrease within the range of 23.6–38.1 % in the Ryukyu Islands for Castanopsis sieboldii and Elaeocarpus japonica, respectively, and within the range of 32.4–42.3 % in Taiwan for Camellia japonica and Distylium racemosum, respectively. These findings suggest that the four species will be vulnerable at the southern range limits; however, the remaining six species will potentially increase within the PH areas in the future at all regions.

Transpiration and stomatal control: a cross-species study of leaf traits in 39 evergreen and deciduous broadleaved subtropical tree species

Key message Using an extensive dataset for 39 subtropical broadleaved tree species, we found traits of the leaf economics spectrum to be linked to mean stomatal conductance but not to stomatal regulation.

Genetic diversity and historical migration patterns of an endemic evergreen oak, Quercus acuta, across Korea and Japan, inferred from nuclear microsatellites

Forests dominated by broad-leaved evergreen trees cover a narrow zone of the southern coast of Korea as well as warm-temperate and subtropical regions of Japan. However, little is known about their establishment history, especially in Korea. Endemic Quercus acuta grows in Korea and Japan. We evaluated its genetic diversity, population structure, and degree of lineage admixture in terms of its distribution history. Analyses utilized seven nuclear microsatellites that were genotyped from 330 individuals of 16 populations. Genetic diversity was highest for the Japanese Kyushu populations. The allelic diversity of populations was similar between Jeju Island and Kyushu, implying that they are putative refugia. Although the mean FST value was very low, patterns for isolation-by-distance were strong and significant. Results from Bayesian clustering and Monmonier’s algorithm indicated that populations are roughly partitioned from west (Korea and Kyushu) to east (Japanese Honshu). Therefore, the two divisions within this species are demographically independent and may have arisen due to past fragmentation. In considering the mixed genetic structure between Korea and Kyushu populations, and the higher levels of genetic diversity in mainland Korea relative to Honshu, we hypothesized that this species has been able to survive within Jeju Island and Kyushu. Consequently, mainland Korea populations might have been shaped by range expansion that kept them well mixed from a continuous and large genetic resource.

Coldness index does not indicate the upper limit of evergreen broad-leaved forest on a subtropical island

Many ecologists in East Asia consider that Kira's coldness index (CI) of −10 °C month agrees well with the northward and upward limits of evergreen broad-leaved forest (EBLF). It is of interest to establish whether CI is an effective indicator of the upper limit of EBLF in a tropical-to-temperate transitional island. A case study in Taiwan was conducted through a comparison of CI layers with field vegetation information. The results from spatial analysis revealed that the location of CI = −10 isotherm was approximately 3,381 m a.s.l., within the Abies zone. The analysis of plots showed that vegetation near CI = −10 isotherm was evergreen coniferous pure forest with 99.9 % relative dominance. The CI = 0 isotherm lay at approximately 2,594 m a.s.l., close to the boundary between the Tsuga–Picea and upper Quercus zones, with the CI = 0 isotherm in the transformation zone between evergreen coniferous and broad-leaved trees. However, the vegetation near CI = 0 isotherm tended more toward coniferous forest showing a total relative dominance of 78.26 %. The two-way cluster dendrogram of vegetation plots revealed identical results. The preceding evidence, combined with the spatial pattern of 3,564 plots along CI and elevation gradients, made it clear that CI did not indicate the upper limit of EBLF in Taiwan. Other information on the CI criterion are discussed. To explore the differentiation of altitudinal vegetation, further research should be made to connect ecoclimatic factors such as effective warmth index with information from vegetation plots.

Does future climate change facilitate expansion of evergreen broad-leaved tree species in the human-disturbed landscape of the Korean Peninsula? Implication for monitoring design of the impact assessment

To quantitatively assess future change of evergreen broad-leaved tree species' distributions in human-disturbed landscapes of the Korean Peninsula under climate change, potential habitats (PHs) were projected for four important evergreen broad-leaved tree species (Quercus acuta, Castanopsis sieboldii, Machilus thunbergii, and Neolitsea sericea) by species distribution models (SDMs). The distribution data (presence/absence) of the target species in Korea and Japan were used as response variables for SDMs, and climatic data were used as explanatory variables. Three general circulation models under A2 emission scenarios were used as future climate scenarios for the years 2070–2099. Potential habitats masked by land-use data (PHLUs) were projected to assess the impact of anthropogenic activities. Highly accurate SDMs were obtained for all the target species. The current PHs were decreased to 21–35 % by the anthropogenic activities. Future PHLUs for all the target species were projected to increase by 2.0–18.5 times of current PHLUs. These results suggest that all the target species are applicable as indicator species for monitoring in the Korean Peninsula, even if anthropogenic effects are incorporated. Variation of the increasing rate was caused by the differences in the response to temperature changes. M. thunbergii responded sensitively to the increase of minimum temperature of coldest month and had a largest increase in PHLUs under future climate. Therefore, M. thunbergii is considered to be most appropriate species for monitoring the changes of horizontal distributions above all focal evergreen broad-leaved tree species.

Genetic Differentiation and Genetic Diversity of Castanopsis (Fagaceae), the Dominant Tree Species in Japanese Broadleaved Evergreen Forests, Revealed by Analysis of EST-Associated Microsatellites

The broadleaved evergreen forests of the East Asian warm temperate zone are characterised by their high biodiversity and endemism, and there is therefore a need to extend our understanding of its genetic diversity and phylogeographic patterns. Castanopsis (Fagaceae) is one of the dominant tree species in the broadleaved evergreen forests of Japan. In this study we investigate the genetic diversity, genetic structure and leaf epidermal morphology of 63 natural populations of C. sieboldii and C. cuspidata, using 32 Expressed Sequence Tag associated microsatellites. The overall genetic differentiation between populations was low (G ST = 0.069 in C. sieboldii and G ST = 0.057 in C. cuspidata). Neighbor-joining tree and Bayesian clustering analyses revealed that the populations of C. sieboldii and C. cuspidata were genetically clearly differentiated, a result which is consistent with the morphology of their epidermal cell layers. This suggests that C. sieboldii and C. cuspidata should be treated as independent species, although intermediate morphologies are often observed, especially at sites where the two species coexist. The higher level of genetic diversity observed in the Kyushu region (for both species) and the Ryukyu Islands (for C. sieboldii) is consistent with the available fossil pollen data for Castanopsis-type broadleaved evergreen trees during the Last Glacial Maximum and suggests the existence of refugia for Castanopsis forests in southern Japan. Within the C. sieboldii populations, Bayesian clustering analyses detected three clusters, in the western and eastern parts of the main islands and in the Ryukyu Islands. The west-east genetic differentiation observed for this species in the main islands, a pattern which is also found in several plant and animal species inhabiting Castanopsis forests in Japan, suggests that they have been isolated from each other in the western and eastern populations for an extended period of time, and may imply the existence of eastern refugia.

Genetic homogeneity between Korean and Japanese populations of the broad-leaved evergreen tree Machilus thunbergii (Lauraceae): A massive post-glacial immigration through the Korea Strait or something else?

The warm-temperate vegetation of Korea, currently limited to southern coastal areas, shifted during the Last Glacial Maximum (LGM) towards glacial refugia, putatively located in southern Japan. We hypothesized two scenarios of post-glacial re-colonization of warm-temperate species in relation to current levels of genetic diversity within their populations. If extant Korean populations originated from a single source (a single glacial refugium), we expect significantly lower levels of genetic diversity relative to those from Japan due to founder effects. Alternatively, if they were derived from multiple source populations, levels of genetic diversity within Korean populations will not be significantly reduced compared to those of Japanese ones. To test which of these scenarios is more likely, we investigated the patterns of genetic diversity in 14 populations (seven from southern Korea and seven from southern Japan) of the broad-leaved evergreen tree Machilus thunbergii, employing 11 allozyme loci. High levels of genetic variation in M. thunbergii were found both in southern Korea and southern Japan, with a considerable genetic homogeneity not only between the two regions but also between populations within the two regions. These results suggest a pattern of re-colonization after the LGM fitting the second scenario (immigration from multiple refugia), probably through multiple waves and/or with large founder populations.

常緑広葉樹から分離されたLachnum nipponicumなど菌類3種の形態的・培養的特徴

常緑広葉樹から分離されたLachnum nipponicumなど菌類3種の形態的・培養的特徴