Tropical Seasonal Rain Forest Research Articles

Savanna Plants Have a Lower Hydraulic Efficiency than Co-Occurring Species in a Rainforest

A plant species can have diverse hydraulic strategies to adapt to different environments. However, the water transport divergence of co-occurring species in contrasting habitats remains poorly studied but is important for understanding their ecophysiology adaptation to their environments. Here, we investigated whole-branch, stem and leaf water transport strategies and associated morphology traits of 11 co-occurring plant species in Yuanjiang valley-type savanna (YJ) with dry–hot habitats and Xishuangbanna tropical seasonal rainforest (XSBN) with wet–hot habits and tested the hypothesis that plants in YJ have a lower water transport efficiency than co-occurring species in XSBN. We found high variation in whole-branch, stem and leaf hydraulic conductance (Kshoot; Kstem and Kleaf) between YJ and XSBN, and that Kstem was significantly higher than Kleaf in these two sites (Kstem/Kleaf: 16.77 in YJ and 6.72 in XSBN). These plants in YJ with significantly lower Kshoot and Kleaf but higher sapwood density (WD) and leaf mass per area (LMA) showed a lower water transport efficiency regarding less water loss and the adaptation to the dry–hot habitat compared to co-occurring species in XSBN. In contrast, these co-occurring plants in XSBN with higher Kshoot and Kleaf but lower WD and LMA tended to maximize water transport efficiency and thus growth potential in the wet–hot habitat. Our findings suggest that these co-occurring species employ divergent hydraulic efficiency across YJ and XSBN so that they can benefit from the contrasting hydraulic strategies in adaptation to their respective habitats.

Elevational patterns of soil organic carbon and its fractions in tropical seasonal rainforests in karst peak-cluster depression region.

Karst ecosystems, especially tropical karst forests, are crucial to the global carbon cycle. In mountainous and hilly areas, elevation-related changes in environment and vegetation often lead to shifts in the accumulation and decomposition of soil organic carbon (SOC). However, the elevational patterns and influencing variables of SOC and its fractions in tropical karst forest ecosystems remain largely unexplored. Here, we analyzed the elevational patterns of SOC and its fractions in the topsoil and subsoil in the tropical seasonal rainforests within typical peak-cluster depression region of Southwest China. Our results indicated that the SOC content was highest at 400 m asl, which was significantly higher than that at 200 m asl (p < 0.05). Overall, SOC content demonstrated an increasing trend with rising elevation. Additionally, SOC content was significantly higher in the topsoil compared to the subsoil (p < 0.05). The majority of SOC fractions exhibited an increase with elevation but decrease with soil depth. Notably, only water-soluble organic carbon (WSOC) displayed a decrease with elevation. Meanwhile, recalcitrant organic carbon (ROC, 54.27%), particulate organic carbon (POC, 30.19%), and easily oxidizable organic carbon (EOC, 16.95%) were the main SOC fractions. Labile organic carbon (LOC) in the karst forest soil was predominantly composed of EOC and POC. Correlation analysis unveiled significant positive correlations between SOC and certain fractions with elevation, soil total nitrogen, and exchangeable magnesium. Conversely, significant negative correlations were observed with soil bulk density (SBD), soil total phosphorus, and litter phosphorus (Litter P). Redundancy analysis indicated that elevation, SBD, and Litter P were the main environmental variables influencing shifts in SOC and its fractions. Structural equation models showed that SOC was primarily directly impacted by soil properties but indirectly impacted by elevation. ROC was mainly associated with the direct effects of soil properties and litterfall, although elevation exerted a substantial impact through indirect pathways. Moreover, LOC was predominantly influenced by the direct impact of soil properties. Therefore, this study demonstrates that SOC and its fractions are strongly influenced by elevation in karst peak-cluster depression regions and have important implications for forest management and sustainable ecosystem development in these regions.

Ants may buffer the Janzen-Connell effect in a tropical forest in Southwest China.

Mutualistic symbioses between ants and plants are widespread in nature. Ants can deter unwanted pests and provide protection for plants in return for food or housing rewards. Using a long-term demographic dataset in a tropical seasonal rain forest in Southwest China, we found that associations with ants positively influenced seedling survival and adult growth, and also, species with extrafloral nectaries experienced weaker conspecific negative density dependence compared with species without extrafloral nectaries. Furthermore, we found strong evidence suggesting that species in our forest experienced conspecific density dependence, which we interpreted as heavy pest pressure that may drive the development of anti-pest symbioses such as the plant-ant relationship. Our findings suggest that ants and conspecific neighbors play important but inverse roles on plant survival and growth and that ants can buffer tree neighborhood interactions in this tropical forest.

Seedling dynamics differ between canopy species and understory species in a tropical seasonal rainforest, SW China

We used 11 years of census data from 450 seedling quadrats established in a 20-ha forest dynamics plot to study seedling dynamics in tree species of a tropical seasonal rainforest in Xishuangbanna, southwestern China. We found that overall seedling recruitment rate and relative growth rate were higher in the rainy season than in the dry season. Both the recruitment rate of seedlings from canopy tree species (two species) and the relative growth rate of seedlings from understory species (nine species) were higher in the rainy season than in the dry season. However, in the rainy season, the recruitment rate of seedlings was higher for canopy tree species than for understory tree species. In addition, relative growth rate of seedlings was higher in the canopy species than in understory seedlings in the dry season. We also observed that, in both rainy and dry seasons, mortality rate of seedlings was higher for canopy species than for understory species. Overall, canopy tree species appear to have evolved a flexible strategy to adapt to the seasonal changes of a monsoon climate. In contrast, understory tree species seem to have adopted a conservative strategy. Specifically, these species mainly release seedlings in the rainy season and maintain relatively stable populations with a lower mortality rate and recruitment rate in both dry and rainy seasons. Our study suggests that canopy and understory seedling populations growing in forest understory may respond to future climate change scenarios with distinct regeneration strategies.

Beta Diversity Patterns and Determinants among Vertical Layers of Tropical Seasonal Rainforest in Karst Peak-Cluster Depressions

Karst peak-cluster depressions in tropical China are characterized by high habitat heterogeneity, supporting complex seasonal rainforest communities, and harboring a rich abundance of endemic and endangered plants. However, for these rainforests, species and phylogenetic beta diversity and their limiting factors are poorly understood. In this study, the relationships between the beta diversity of three vertical layers (herb, shrub, and tree), environmental factors, and physical distance in China’s tropical karst seasonal rainforest were studied. The results showed that each layer exhibited high species and beta diversity, with species turnover being the dominant contributing factor. Environmental filtering and dispersal limitations were significant drivers of community assembly. Environmental filtering exerted a strong influence, with slope position, soil availability of phosphorus and potassium, pH, and organic matter being the key factors. These findings elucidate seasonal rainforest species and beta diversity spatial patterns within karst peak-cluster depressions, providing a foundation for developing karst ecosystem forest management and vegetation restoration measures.

A Tree-Shrub Layer Separation Method for MLS LiDAR Point Clouds of Typical Tropical Seasonal Rainforests

A Tree-Shrub Layer Separation Method for MLS LiDAR Point Clouds of Typical Tropical Seasonal Rainforests

A dataset of carbon and water fluxes observed in Xishuangbanna tropical seasonal rain forest from 2011 to 2015

The proposal for China's dual-carbon target is based on the foundation of addressing climate change. It is imperative to enhance carbon sinks and reduce carbon sources to implement climate governance. The terrestrial ecosystem is an important atmospheric carbon sink, while the tropical seasonal rain forest, as a component of the terrestrial ecosystem in China characterized by rich biomass, and strong productivity, plays an essential role in carbon sequestration. As one of the basic observation and research stations of China Flux Research Network (ChinaFLUX), the Xishuangbanna Tropical Seasonal Rainforest Ecological Station built in 2002 has accumulated 20 years of carbon and water flux observation data. Based on the ChinaFLUX data processing system, this dataset processes and collates the tropical seasonal rain forest flux and conventional meteorological data of Xishuangbanna from January 2011 to December 2015. The result is a collection of data products at half-hourly, daily, monthly, and yearly intervals. This dataset can be applied to research fields such as the analysis of the carbon cycle response mechanism of ecosystems and the assessment of carbon sequestration benefits under climate change, and it can also serve as a valuable reference for the optimization of ecosystem model parameters and the verification of results. Through multi-platform architecture, multi-scale integration, and multi-method coupling, the dataset is expected to further provide basic support for analyzing the mechanisms of regional and even global carbon-water-heat cycles in ecosystems, spatiotemporal dynamics and predicting future carbon sink potential, thus providing the scientific basis for achieving China's carbon neutrality goal.

A dataset of the litterfall recovery amount and the standing crop dynamics in the tropical seasonal rainforests of Xishuangbanna (2005–2020)

Litterfall refers to the collective organic matter generated by the aboveground parts of plants that accumulates on the surface. It is a key medium linking the ecological processes between the aboveground and the underground in terrestrial ecosystems. Litterfall recovery and the standing crop comprise the dry weight of branches, leaves, flowers and fruits, bark, moss and lichen, and so on. The annual yield of leaf litterfall can provide data support for the assessment of forest productivity, and the falling flowers and fruits are of great value to the research of forest seed rain. The proportion of each component is an important factor affecting nutrient cycling and carbon cycling in the ecosystem. The dataset collects the litterfall recovery amount and the standing crop of the tropical seasonal rainforest in Xishuangbanna during 2005–2020. The establishment and sharing of this dataset can furnish fundamental data for the in-depth examination of vegetation structure and ecosystem function under the global hydrothermal pattern changes, and serve as a foundation for advancing the study of the litterfall and tropical seasonal rain forests.

A dataset of soil moisture in Xishuangbanna tropical seasonal rain forests (2009–2017)

As the carrier of material circulation and energy flow, soil moisture plays an important role in terrestrial forest ecosystems. Observing soil moisture is beneficial for unveiling the hydrological characteristics, water conservation capacity and ecosystem service function of forest ecosystems. The tropical rain forest is among the most biodiverse ecosystems. There are large areas of tropical rain forests in Xishuangbanna, where the tropical seasonal rain forest represents a prominent vegetation type. Conducting continuous, long-term and high-quality ecological monitoring is one of the main tasks of China Ecosystem Research Network (CERN). Soil moisture content is an important indicator in the long-term site-specific water environment observations within the terrestrial ecosystems monitored by CERN. This dataset comprises soil moisture content data collected from January 2009 to December 2017 at the Xishuangbanna Station for Tropical Rainforest Ecosystem Studies of the Chinese Academy Sciences (also known as National Forest Ecosystem Research Station at Xishuangbanna, BNF for short), with a total of 2,879 entries. The determination method used for this dataset is fixed-point continuous observation using Time Domain Reflectometry (TDR) instruments; observations were conducted at multiple, ranging from 0–10 cm, 10–20 cm, 20–30 cm, 30–40 cm, 40–50 cm, 50–70 cm, 70–90 cm and 90–110 cm. The observation frequency consists of 3 times per month, respectively at the beginning, middle and end of each month. The observations were conducted strictly according to the CERN observation and quality control protocols. The dataset is readily available for online sharing and serves as essential foundational data for conducting research on tropical forest hydrology under the global climate change and different vegetation environments.

Hydraulic traits and photosynthesis are coordinated with trunk sapwood capacitance in tropical tree species.

Water stored in trunk sapwood is vital for the canopy to maintain its physiological function under high transpiration demands. Little is known regarding the anatomical properties that contribute to the hydraulic capacitance of tree trunks and whether trunk capacitance is correlated with the hydraulic and gas exchange traits of canopy branches. We examined sapwood capacitance, xylem anatomical characteristics of tree trunks, embolism resistance, the minimal xylem water potential of canopy branches, leaf photosynthesis and stomatal conductance in 22 species from a tropical seasonal rainforest and savanna. The results showed that the mean trunk sapwood capacitance did not differ between the two biomes. Capacitance was closely related to the fiber lumen fraction and fiber wall reinforcement and not to the axial and ray parenchyma fractions. Additionally, it was positively correlated with the theoretical hydraulic conductivity of a trunk and the specific hydraulic conductivity of branches, and showed a trade-off with branch embolism resistance. Species with a high trunk sapwood capacitance maintained less negative canopy water potentials in the dry season, but higher leaf photosynthetic rates and stomatal conductance in the wet season. This study provides a functional link among trunk sapwood capacitance, xylem anatomy, canopy hydraulics and photosynthesis in tropical trees.

Parrotiaflower blooming in Miocene rainforest

AbstractParrotiaC. A. Meyer (Hamamelidaceae) is a relictual genus with only two extant species disjunctly distributed in the subtropical forests of East China and temperate forests of North Iran. Fossil records suggest thatParrotiawas widespread in Europe and Asia during the Miocene, but its fossils are predominantly based on pollen and leaves. In this paper we describe the first fossil flower ofParrotiabased on an exceptionally well‐preserved amber inclusion from the middle Miocene of Zhangpu, Southeast China. The fossil flower was investigated with light microscope and microcomputed tomography techniques.Parrotia zhiyaniisp. nov. is a small apetalous staminate flower subtended by a pair of prominent bracts. The androecium consists of 12 stamens, and each stamen consists of a short, slender filament and a prominent, elongated anther. The anthers with short simple trichomes on the outer surface and a prominent apical connective extension are opened by longitudinal slits. Unlike its living relatives, the newParrotiafrom Zhangpu grew in a Miocene seasonal tropical rainforest.

Revealing the relative importance among plant species, slope positions, and soil types on rhizosphere microbial communities in northern tropical karst and non-karst seasonal rainforests of China.

Rhizosphere microbes have an extremely close relationship with plants and the study on the relationship between rhizosphere microorganisms and their influencing factors is conducive to the protection of vegetation and the maintenance of biodiversity. Here we investigated how plant species, slope positions and soil types affect the rhizosphere microbial community. Slope positions and soil types were collected from northern tropical karst and non-karst seasonal rainforests. The results indicated that soil types played a predominant role in the development of rhizosphere microbial communities (28.3% of separate contribution rate), more than plant species identity (10.9% of separate contribution rate) and slope position (3.5% of separate contribution rate). Notably, environmental factors closely related to soil properties were the major influence factors that controlling the rhizosphere bacterial community structure in the northern tropical seasonal rainforest, especially pH. Additionally, plant species also influenced the rhizosphere bacterial community. In low nitrogen content soil environments, rhizosphere biomarkers of dominant plant species were often nitrogen-fixing strains. It suggested that plants might have a selective adaptation mechanism to rhizosphere microorganisms to obtain the advantages of nutrient supply. Overall, soil types exerted the biggest influence on rhizosphere microbial community structure, followed by plant species and finally slope positions.

Vessel dimorphism and wood traits in lianas and trees among three contrasting environments.

Determining how xylem vessel diameters vary among plants and across environments gives insights into different water-use strategies among species and ultimately their distributions. Here, we tested the vessel dimorphism hypothesis that the simultaneous occurrence of many narrow and a few wide vessels gives lianas an advantage over trees in seasonally dry environments. We measured the diameters of 13,958 vessels from 15 liana species and 10,430 vessels from 16 tree species in a tropical seasonal rainforest, savanna, and subtropical evergreen broadleaved forest. We compared differences in mean and hydraulically weighted vessel diameter (MVD and Dh ), vessel density (VD), theoretical hydraulic conductivity (Kt ), vessel area fraction (VAF), and wood density (WD) between lianas and trees and among three sites. Nine liana species and four tree species had dimorphic vessels. From the tropical seasonal rainforest to the savanna, liana MVD, Dh and Kt decreased, and VD and WD increased, while only tree WD increased. From the tropical seasonal rainforest to the subtropical forest, six wood traits remained unchanged for lianas, while tree MVD, Dh and Kt decreased and VD increased. Trait space for lianas and trees were more similar in the savanna and more divergent in the subtropical forest compared to the tropical seasonal rainforest. These results suggest that lianas tend to possess greater vessel dimorphism, which may explain how lianas grow well during seasonal drought, influencing their unique distribution across tropical rainfall gradients.

Effects of topography and historical disturbance on canopy height structure of tropical forests in Menglun, Xishuangbanna, China.

With the combination of airborne Lidar and panchromatic images in 1981 and 2021, we investigated the canopy height structure of tropical forests in Menglun sub-reserve in the Xishuangbanna National Nature Reserve of Yunnan Province, and analyzed its relationship with environmental factors by using multiple regression tree (MRT) method. The results showed that forests in the Menglun sub-reserve could be clustered into seven types based on canopy height structures, with tropical rainforest, monsoon evergreen broad-leaved forest, secondary forest, and flood plain forest as the main types. The potential solar radiation, altitude, terrain profile curvature, slope and the brightness value of imageries in 1981 and 2021 were main factors that drove the classification. The tropical seasonal rainforest dominated by Pometia pinnata occupied the largest area in valley and low-land. The monsoon evergreen broad-leaved forest dominated by Castanopsis echinocarpa mainly distributed in the ridge and disturbed areas. The secondary forests had homogeneous canopy surface, which was significantly different from the primary forests. The activities of swidden agriculture about three decades ago had legacy impacts on the physiognomy of secondary forests.

Estimating the gross primary productivity based on VPM correction model for Xishuangbanna tropical seasonal rainforest

ABSTRACT Due to the unique climatic characteristics and vegetation features of tropical regions, the correlation R2 of gross primary productivity (GPP) estimation in tropical regions using remote sensing models was generally lower than 0.3. Therefore, for the cloudy and rainy tropical regions, the influence brought by clouds on remote sensing images needed to be considered in GPP estimation. This paper developed a corrected vegetation photosynthesis model (VPM) for estimating GPP under cloudy conditions. It mainly corrected the two parameters, W scalar and Enhanced Vegetation Index (EVI), which were obtained from remote sensing images and were therefore greatly influenced by clouds in the model. First, the water stress factor W scalar was replaced by Evaporation Fraction (EF). Secondly, using the good correlation between near surface temperature and EVI, the conversion coefficient between near surface temperature and EVI was fitted to achieve the effective reconstruction of EVI contaminated by clouds. The correction of the two factors improved the estimation accuracy of the VPM model, and the comparison with the observed values of the GPP site in 4 years showed that the correction of EVI had a better improvement, with an increase of 0.22 in R2 compared with the pre-correction, and the correction of W scalar was increased by 0.11 in R2. To verify the proposed method, the in-situ observation data of Xishuangbanna flux site from 2007 to 2010 were used. The results showed that the proposed method effectively improved the accuracy of GPP estimation by VPM model, especially in 2007 it was strongly influenced by clouds, and the improvement was significant, with R2 increasing from 0.2 to 0.82. In general, the accuracy of GPP estimation by the proposed method had been significantly improved, with RMSE (gC·m−2·8 day−1) decreasing from 15,14.4, 18.1, 14.2 to 8.07, 6.56, 10.33, 11.44, respectively. Therefore, the proposed method can be used to estimate the GPP for tropical seasonal rain forests in Xishuangbanna.

Erratum to: Partial mutual exclusion of ectomycorrhizal and saprobic fungi in a tropical seasonal rainforest

The original publication contains the following errors: Tables 1 and 2 of the original research article contained the macrofungi collected from all the substrates (soil, fallen branches, twigs and other woody material) in plots that should have been contained with names of macrofungi collected only from soil and woody material burried in the soil

Sapling root ecological stoichiometric characteristics of dominant species in three forests in Yunnan, China, and their relationship with soil physicochemical factors

AbstractQuestionThis study aimed to explore the nutrient‐cycling patterns, supply mechanisms and limiting characteristics of elements in saplings of tropical and subtropical forest ecosystems.LocationTropical seasonal rainforests, tropical montane rainforests and mid‐montane moist evergreen broad‐leaved forests in Yunnan, China.MethodsWe sampled sapling roots from 21 vascular plant species that are dominant in the upper and lower canopies of three forest types. We assessed stoichiometric contents of carbon (C), nitrogen (N) and phosphorus (P), and investigated their relationships with soil nutrients.ResultsThe C content ranged from 449.67 to 483.00 mg/g in the roots of the dominant species, the N content ranged from 1.98 to 9.67 mg/g and the P content ranged from 0.32 to 0.53 mg/g. The C:N ratio was highest in the tropical seasonal rainforest, whereas the C:P and N:P ratios were highest in the tropical montane rainforest. The root average C:N and C:P ratios were higher than Chinese plant averages, which indicated that the forest ecosystems in southern Yunnan had higher absorption and utilization efficiency of C. The average N:P ratio was &gt;12 and &lt;14, which indicated that plant root growth was restricted by N and P availability. In the saplings of upper canopy species, the C:N ratio was higher, the C:P ratio was similar and N:P ratio was lower compared with those of lower canopy species. The N:P ratio in the upper canopy was &lt;12, whereas that in the lower canopy was &gt;14, which indicated that growth in the upper canopy was mainly N‐limited and growth in the lower canopy was mainly P‐limited. Nutrient absorption and utilization efficiency in the upper canopy was slightly higher than in the lower canopy.ConclusionsThe sapling roots of the dominant species in Yunnan had the characteristics of high C:N, high C:P and low N:P, and the growth and development of plants is mainly restricted by N and P availability. The C, N and P contents and their stoichiometric characteristics in roots were affected mainly by the soil P content. This study provides a scientific basis for further revealing the ecological strategies of plants in response to changes in the external environment.

Body size determines multitrophic soil microbiota community assembly associated with soil and plant attributes in a tropical seasonal rainforest.

To understand soil biodiversity we need to know how soil communities are assembled. However, the relationship between soil community assembly and environmental factors, and the linkages between soil microbiota taxonomic groups and their body sizes, remain unexplored in tropical seasonal rainforests. Systematic and stratified random sampling was used to collect 243 soil and organism samples across a 20-ha plot in a tropical seasonal rainforest in southwestern China. High-throughput sequencing, variation analysis and principal coordinates of neighbourhood matrices were performed. Soil community composition, spatial distribution and assembly processes based on propagule size (including archaea, bacteria, fungi and nematodes) were investigated. The results showed that: (i) the community assembly of small soil microorganisms (bacteria, fungi) was mostly influenced by stochastic processes while that of larger soil organisms (nematodes) was more deterministic; (ii) the independent effects of habitat (including soil and topographic variables) and its interaction with plant attributes for community structure significantly decreased with increasing body size; and (iii) plant leaf phosphorus directly influenced the spatial distribution of soil-available phosphorus, which indicates their indirect impact on the assembly of the soil communities. Our data suggest that the assembly of multitrophic soil communities can be explained to some extent by changes in above-ground plant attributes. This highlights the importance of above- and below-ground linkages in influencing multitrophic soil microbiota community assembly.

First fossil record of Canarium (Burseraceae) from the middle Miocene of Fujian, southeastern China and its paleoecological implications

The pili nut genus Canarium is an important element in tropical rainforests and seasonal rainforests of the northern and southern hemispheres. The genus contains some economically important species widely cultivated as food crops worldwide. Fossil evidence suggests that Canarium had occurred in Africa, Asia, Europe and North America during the Paleogene, with the earliest fossil record from the early Eocene of eastern North America and southern England. Here we describe the three-dimensionally preserved fruits of Canarium from the middle Miocene Zhangpu flora in Fujian, southeastern China as a new species, Canarium haominiae n. sp. The internal structures and anatomical details of the fossil fruits were investigated using X-ray micro-computed tomography and scanning electron microscope. Endocarps of C. haominiae formed by three connate pyrenes are ovoid to spindle-shaped in lateral view and rounded triangular in cross section, with acute to obtuse base and slightly acuminate to acute apex. Each pyrene is unilocular and dehiscent by an operculate germination valve. The germination valve is ovate and opens from the top of the pyrene to approximate two thirds of its length towards the base. The occurrence of C. haominiae confirms that the middle Miocene Zhangpu flora represents a megathermal seasonal rainforest, and also indicates that Canarium might have been an important component in the tropical seasonal rainforest since the middle Miocene.

Quantifying the vertical microclimate profile within a tropical seasonal rainforest, based on both ground- and canopy-referenced approaches

Quantifying the microclimate of a tall and complex tropical forest is quite challenging because of the difficulty in accessing the canopy. Traditional ground-referenced methods may underestimate the contribution of canopy heterogeneity to structuring the vertical profiles of forest microclimate. The present study examined how the reference height affects vertical variation of microclimate in a tropical rainforest in southwest China, based on both ground- and canopy-referenced approaches. The results show that the canopy-referenced approach yielded a higher model fit than did the ground-referenced method, and only canopy-referenced method could detect two thresholds in the leaf area index at approximately -22.6 ± 2.7 m and -36.6 ± 6.6 m below the canopy top; the higher threshold is consistent with thresholds of the annual mean temperature, the diurnal ranges of air temperature, and the relative humidity in the vertical profile; while the lower threshold is similar with the breakpoints of annual mean relative humidity, the annual ranges of air temperature and the relative humidity along the profile. The discontinuous variance in the microclimatic factors was due to the canopy structure in the vertical profile. Selecting the top of the canopy as the reference height could be a better approach for quantifying the microclimatic profiles in the studied forest, and this approach can improve our understanding of the effects of the vertical stratification of microclimates on species composition and diversity in this forest.