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Article

Allometric Equations for Aboveground Biomass Estimation in Natural Forest Trees: Generalized or Species-Specific?

1
Key Laboratory of State Forestry and Grassland Administration On Subtropical Forest Biodiversity Conservation, Co-Innovation Center for Sustainable Forestry in Southern China, College of the Life Sciences, Nanjing Forestry University, Nanjing 210037, China
2
State Environmental Protection Key Laboratory On Biosafety, Research Center for Biodiversity Conservation and Biosafety, State Environmental Protection Scientifc Observation and Research Station for Ecological Environment of Wuyi Mountains, Biodiversity Comprehensive Observation Station for Wuyi Mountains, Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment of China, Nanjing 210042, China
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(7), 493; https://doi.org/10.3390/d17070493
Submission received: 16 June 2025 / Revised: 2 July 2025 / Accepted: 6 July 2025 / Published: 18 July 2025
(This article belongs to the Section Plant Diversity)

Abstract

Accurate estimation of aboveground biomass (AGB) in tree–shrub communities is critical for quantifying forest ecosystem productivity and carbon sequestration potential. Although generalized allometric equations offer expediency in natural forest AGB estimation, their neglect of interspecific variability introduces methodological pitfalls. Precise AGB prediction necessitates resolving two biological constraints: phylogenetic conservation of allometric coefficients and ontogenetic regulation of scaling relationships. This study establishes an integrated framework combining the following: (1) phylogenetic signal detection (Blomberg’s K/Pagel’s λ) across 157 species’ allometric equations, revealing weak but significant evolutionary constraints (λ = 0.1249, p = 0.0027; K ≈ 0, p = 0.621); (2) hierarchical error decomposition of 9105 stems in a Mt. Wuyishan forest dynamics plot (15 species), identifying family-level error stratification (e.g., Theaceae vs. Myrtaceae, Δerror > 25%); (3) ontogenetic trajectory analysis of Castanopsis eyrei between Mt. Wuyishan and Mt. Huangshan, demonstrating significant biomass deviations in small trees (5–15 cm DBH, p < 0.05). Key findings resolve the following hypotheses: (1) absence of strong phylogenetic signals validates generalized models for phylogenetically diverse communities; (2) ontogenetic regulation dominates error magnitude, particularly in early developmental stages; (3) differential modeling is recommended: species-specific equations for pure forests/seedlings vs. generalized equations for mixed mature forests. This work establishes an error hierarchy: ontogeny > taxonomy > phylogeny, providing a mechanistic basis for optimizing forest carbon stock assessments.

1. Introduction

Biomass is an important measure of forest ecosystem productivity [1]. It is also a key indicator of forest ecosystem function, and assessments of forest carbon stocks are based upon the estimation of forest biomass [2]. Forest biomass can be measured at both individual tree and stand level [3], or at both individual and stand level [4]. The biomass of individual trees and forest communities is dynamic over time. The process of forest development and community succession can affect forest biomass. The stand development increases biomass [5], while forest succession links biomass dynamics [6]. In addition to being associated with forest ecological processes, forest biomass is also positively correlated with biodiversity indices such as species richness and functional diversity, though context-dependent exceptions exist in degraded ecosystems [7,8,9].
Forest biodiversity is essential to ecosystem function and can provide a variety of ecosystem services, including pollination, seed dispersal, resistance to wind storms, and others [10]. Aboveground biomass (AGB) of trees and shrubs serves as a fundamental basis for evaluating forest ecosystem functions and the value of ecosystem services such as biodiversity and carbon storage [11]. As a proxy for habitat complexity, AGB mediates biodiversity conservation by (i) providing multi-layered niches for canopy specialists and understory taxa, and (ii) enhancing resource partitioning through size-structured allocation patterns. It directly reflects the productivity and health status of the forest [12]. Higher AGB indicates a more complex forest structure including different layers of canopy and vegetation that provide diverse ecological niches [13]. Hence, quantifying natural forest AGB is crucial for assessing global carbon sinks, as well as for developing effective forest management and conservation strategies [14]. By estimating AGB, we can understand the growth patterns of different tree species under various environmental conditions and analyze the productivity and resource use efficiency of forest ecosystems [15].
An allometric equation for aboveground biomass is a model that describes the relationship between aboveground biomass and factors such as tree species diameter at breast height (DBH) and height. The estimation of AGB can be based on a single factor (DBH), or dual factors (DBH and tree height), e.g., biomass–diameter regression and biomass–diameter–height regression [16]. In the allometric model of Y = bxa, b is a constant, indicating y’s value when x = 1, and a is the allometric coefficient, determining the growth rate of y relative to x, e.g., isometric (a = 1), positive (a > 1), and negative allometry (a < 1) [17,18,19]. Allometric equations are widely applied to different forests or geographical regions, such as various ecosystems in USA [20], Douglas-fir plantation in Italy [21], pine stands in Portugal [22], Omo-Gibe woodland species in Ethiopia [23], dry secondary forest trees in Thailand [24], and Atlantic forest species in Brazil [25]. In China, allometric equations have been developed for tree species in both temperate [26,27] and subtropical forests [28,29,30].
The widespread application of allometric equations across biomass presents both opportunities for standardized carbon accounting and challenges in addressing ecological specificity, particularly in hyper-diverse forests. However, in terms of natural forests (especially tropical and subtropical forests), species highly assemble at a local scale. The species density per unit area in long-term forest dynamics plots (FDPs) often reaches a high level, e.g., 1519 (species)/25 ha in Manaus (Brazil), 1114/50 ha in Yasuni (Ecuador), 468/20 ha in Xishuangbanna (Yunnan, China), 299/50 ha in Barro Colorado Island (Panama), and 153/20 ha in Tiantongshan (Zhejiang, China) [31]. In FDPs with abundant species, the estimation of overall forest biomass faces a potential problem: not all species have a specific allometric equation. On the other hand, for natural forests of different ages, sizes, and species, carbon storage is often estimated using a generalized allometric equation [13,30,32,33]. In this context, a similar question arises: is there sufficient scientific basis for estimating biomass using a genenralized allometric equation? There are few reports on such investigations, although some studies have noted between tree species (phylogenetic) [34,35] and between size classes (ontogenetic) [29,36]. Although allometric equations have been widely studied across taxa, our understanding of how phylogenetic history and ontogenetic development shape these relationships remains incomplete. Phylogenetic comparative methods [37,38] have substantially advanced the field, but their integration into allometric studies is still limited. Most empirical studies focus either on interspecific scaling without accounting for developmental changes, or on within-species ontogenetic patterns without considering evolutionary history. As highlighted by Voje et al. [39] and Pélabon et al. [40], there is a lack of studies that jointly analyze both phylogenetic and ontogenetic influences on trait scaling. Moreover, classic work such as Gould [41] and more recent syntheses like Klingenberg [42] emphasize that separating the effects of growth and descent remains a major challenge. These gaps limit our ability to generalize allometric patterns and predict trait variation across lineages and developmental stages.
In this study, we focus on two potential errors, namely phylogenetic and ontogenetic errors, in estimating the biomass of FDPs using a general biomass–DBH allometric equation. The datasets created by our lab from two FDPs at Mt. Wuyishan [43,44] and Mt. Huangshan [43,45] were used to evaluate the two types of errors. We plan to collect parameters of the biomass–DBH allometric equations from different species around the world for phylogenetic signal detection. Based on these, we propose two hypotheses: (1) Since the growth patterns of trees are influenced by evolutionary history, tree species with different phylogenetic locations have distinct allometric equation parameters. Thus, there may be significant phylogenetic errors when using a general allometric equation to estimate AGB in FDPs, compared to using species-specific equations. (2) The growth patterns of trees are influenced by environmental factors, and local environmental factors vary over time. Thus, using general allometric equations to estimate AGB in FDPs with different size classes may introduce ontogenetic errors. The verification of the two hypotheses not only provides a basis of model selection for the determination of AGB in FDPs, but also lays a scientific foundation for ecosystem function evaluation and biodiversity conservation.

2. Materials and Methods

2.1. Site Description

The study site is located near Sixin Village, Xingcun Town, Wuyishan City. This region is characterized by a mid-subtropical monsoon humid climate, with an annual average temperature ranging from 17.0 °C to 18.4 °C, an average relative humidity of 75–84%, annual precipitation of 1800 mm, an average annual sunshine duration of 1910.2 h, and a frost-free period of 227–246 days [43]. The soil type of the sample site is dominated by red soil with pH ranging from 4.5 to 5.0. Evergreen broad-leaved forests represent the zonal forest vegetation of Mt. Wuyishan, covering a vast area and predominantly distributed at elevations between 350 and 1400 m. The plant community is primarily composed of species from families such as Fagaceae, Lauraceae, Ericaceae, Magnoliaceae, Elaeocarpaceae, and Theaceae. Common species include Castanopsis carlesii, Castanopsis fordii, Castanopsis eyrei, and Engelhardia roxburghiana.
A long-term forest dynamics plot (WY) was established at Mt. Wuyishan according to the criteria proposed by the Center for Tropical Forest Sciences (CTFS). All woody plants with DBH (diameter at breast height) ≥ 1 cm in the WY were calibrated and surveyed. WY’s geographic coordinate is 27°35′24.23″ N, 117°45′55.43″ E, with an elevation ranging approximately from 450 to 580 m as it shown in Figure 1. The plot is rectangular, with a projected area of 9.6 ha (400 m × 240 m).

2.2. Data Determination of Biomass

To advance the research, this study collected a total of 157 datasets [Supplementary Materials, Dataset s1: Coefficients of 157 sets of species-specific allometric equations] measured as ln(W) = a + b * ln(DBH) from Web of Science, Google Scholar. The time frame covered the period from 2000 to 2023, and the study objects were limited to woody plants. To ensure data quality, we implemented stringent inclusion criteria. Only peer-reviewed studies with clearly reported ln(W) = a + b × ln(DBH) parameters and associated statistics were included. Studies focusing on herbaceous species, managed forests, or those lacking site metadata were excluded to minimize bias from non-representative conditions. Allometric models without variance estimates were also omitted to maintain statistical integrity [46,47,48,49,50,51,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78,79,80,81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109,110,111,112,113,114,115,116,117,118,119,120,121,122,123,124]. The search was conducted using keyword combinations such as (“allometric equation” or “biomass equation”) and (“tree” or “woody plant”) and (“coefficient” or “parameter”). The equations were required to explicitly provide coefficients a and b, as well as relevant statistical metrics. The coefficients a and b collected from papers were subjected to descriptive statistical analysis and visualization in Python 3.11 to compare the values across different tree species, where
  • W represents the aboveground biomass of the tree species;
  • DBH represents the diameter at breast height of the tree.
Following the methodology outlined by Anhui Agricultural University [125], phylogenetic signal analysis was conducted on the scaling constant (a) of the allometric equation only. The allometric exponent (b), which reflects the proportional relationship between DBH and AGB, was not included in the phylogenetic analysis due to its higher stability and lower sensitivity to phylogenetic constraints. To verify that the 157 sets of coefficients have a phylogenetic signal, we calculated Blomberg’s K statistic [34] and its corresponding p-value in R 4.2.3 using the phylogenetic-K function. Blomberg’s K is a statistical measure that quantifies the strength of trait signal along a given phylogenetic tree. A K-value < 1 suggests that the trait evolution follows the Brownian motion model, indicating a strong phylogenetic signal. A K-value significantly less than 1 implies that the trait is more randomly distributed across the phylogeny. The p-value is used to evaluate the significance of the phylogenetic signal, with values < 0.05 indicating that the signal is statistically significant.
In this study, 9105 individuals from 15 species in the Mt. Wuyishan dynamics plot were selected as the analysis subjects [Supplementary Materials, Dataset s2: AGB errors calculated by generalized and species-specific allometric equations]. AGB was estimated using both generalized and species-specific equations, and error analysis was conducted in Python 3.9.13. For the error analysis with the generalized and species-specific allometric equations, the generalized equation from the study by Zuo et al. [126] on the subtropical evergreen broadleaf forests of China was used. This study site is located in Nanping, Fujian, China, the same region as the Mt. Wuyishan forest dynamics plot. The species overlap is high, and the climatic and soil conditions are similar, making the equation highly relevant for this study:
ln(W) = −1.9783 + 2.3771 × ln(DBH)
To compare the regional differences in biomass of the dominant species, data from 1319 Castanopsis eyrei individuals in Mt. Wuyishan and 14,216 individuals in Mt. Huangshan were selected. The DBH data were classified into 5 cm intervals to analyze the biomass at different size classes. The allometric equation of C. eyrei in the dataset compiled by Luo Y J et al. [127] was used to calculate the AGB of C. eyrei within each DBH interval. Since there is a large amount of data on Mt. Huangshan and the maximum DBH of the C. eyrei is larger than the maximum DBH of the C. eyrei in Mt. Wuyishan, there is an imbalance problem in the data. A downsampling method is used to limit the data volume of Mt. Huangshan to the level of the Mt. Wuyishan data and limit the DBH to 5~60 cm to ensure that comparability of data between the two places. t-test was performed on different DBH intervals to verify whether there were significant ontogenetic differences.

3. Results

3.1. Analysis of Coefficients of Allometric Equations

Phylogenetic signal detection was conducted on parameters of 157 allometric equations and corresponding phylogenetic tree is presented in Figure 2. Table 1 evaluates the phylogenetic signals and evolutionary trajectories of allometric coefficients through a multi-metric analytical framework. For the empirical dataset (a), Pagel’s Lambda (λ = 0.1249, p = 0.0027) indicates a weak yet statistically significant phylogenetic signal (p < 0.05), suggesting partial dependency of allometric coefficient evolution on phylogenetic history, albeit substantially modulated by non-phylogenetic drivers such as environmental selection pressures. The significant positive spatial autocorrelation (Moran’s I = 0.0764, p = 0.002) further corroborates elevated trait similarity among phylogenetically proximate taxa relative to stochastic expectations. Concurrently, the positive covariance structure (C mean = 0.1106, p = 0.024) reinforces a weak but detectable association between trait variation and phylogenetic distances. Notably, the non-significant Blomberg’s K (K = 0.0427, p = 0.621) implies stochastic evolutionary dynamics, potentially attributable to convergent adaptation or heterogeneous evolutionary rates across clades.

3.2. Analysis of Species Phylogenetic Error

In this study, we applied both generalized and species-specific allometric equations to estimate the AGB of 15 species with a total of 9105 individuals and systematic phylogenetic error analysis was then performed [Supplementary Materials, Dataset s2: AGB errors calculated by generalized and species-specific allometric equations].
In Figure 3, the differences in boxplot width and position between different families indicate that there are certain differences in the error in calculating AGB for each family. The error variation in Figure 3 for Theaceae and Myrtaceae is more pronounced, with some individual errors being extremely large, indicating that these families may be more sensitive to the allometric equations used. In contrast, Pentaphylacaceae and Ericaceae show more concentrated errors, suggesting that AGB estimates for these families are more stable. Theaceae and Symplocaceae exhibit more outliers, meaning that the AGB estimates for these individual plants deviate significantly, which could reflect the unique growth characteristics of these families or limitations in the current equations.

3.3. Analysis of Species Ontogeny Errors

The t-test results (Table A1) show that in the size classes 5–10 cm and 10–15 cm, the biomass difference between C. eyrei in Mt. Huangshan and Mt. Wuyishan is statistically significant; in most size classes, especially 25–30 cm and above, the biomass difference between the two places is not significant. These results show that within the smaller size class of 5–15 cm, there is a large difference (Figure 4) in the AGB of C. eyrei between the two locations. In the larger DBH classes, the aboveground biomass difference becomes smaller and is not significant.

4. Discussion and Conclusion

4.1. Phylogenetic Signal Analysis of Allometric Equations

To the best of our knowledge, it remains unclear how the phylogenetic history affects large-scale variation in tree allometry [128]. The analysis of phylogenetic signals provides a good platform in studies on tree allometry variation. The interpretation of the results in phylogenetic signal analysis of traits (or parameters) is based on the following rules: significant phylogenetic signals indicating trait phylogenetic conservatism, conversely, insignificant phylogenetic signals revealing traits are mainly influenced by the environment [35]. Significant phylogenetic signals in 50 species demonstrated phylogenetic conservation of the crown-to-stem biomass ratio (Pagel’s λ = 0.359, p < 0.05), demonstrating that biomass allocation of crown-to-stem, rather than stem biomass is phylogenetic conservation [35]. However, shoot–root biomass allocation in 20 dry-grassland species varies greatly among closely related species, implying that this biomass allocation pattern is not phylogenetically conservative [129]. Furthermore, in a study of 894 allometric biomass models of 276 tree species, the model parameters of AGB-DBH regressions are not significantly constrained by phylogeny [34]. It was also demonstrated that the scaling relationships for annualized biomass production rates and different measures of body size were indifferent to phylogenetic affiliation [130]. Therefore, based on existing empirical studies, there is no statistically significant impact of phylogenetics on the allometric equation of biomass estimation.
Will there be significant errors in estimating biomass using a generalized allometric equation established with DBH as a single factor? Several investigation results have confirmed that the error is not significant. Based on both generalized and species-specific allometric equations, the correlation coefficients between aboveground biomass and DBH were >0.85 for subtropical Fagaceae species [131] and > 0.80 for understory woody species in northeastern China forest ecosystems [132], and despite the improved fit of biomass after integrating tree height and crown spread data. DBH is an effective estimator in the process of aboveground biomass estimation, and equations considering only DBH can explain component biomass and total biomass with more than 80% variability for tree species in Tibetan Plateau, China [133].
A phylogenetic signal detection was conducted on the parameters a and b of 157 allometric equations, and no significant phylogenetic signal was found, indicating that phylogeny does not impose constraints on allometry. Phylogenetic signal is present but weak, as evidenced by a statistically significant yet low-magnitude λ (λ = 0.1249, p = 0.0027), indicating that allometric coefficients exhibit only partial dependency on phylogenetic relationships. Environmental selective pressures likely drive trait divergence, thereby attenuating phylogenetic conservatism. The non-significant K value (K ≈ 0, p = 0.621) suggests stochastic trait evolution, with patterns indistinguishable from random processes. Notably, comparative analysis with the Brownian motion model (bm) reveals that the empirical K value (0.0427) is significantly lower than the Brownian motion expectation (0.6239; p < 0.05), further supporting non-conservative evolutionary dynamics dominated by environmental filtering or convergent adaptation. More intriguingly, within the range of tree species in the families Theaceae and Myrtaceae, the general allometric equations exhibited substantial errors, suggesting that allometry exhibits intra-familial or intra-generic differentiation rather than being consistent. Despite the quality screening, we acknowledge that site-specific variables such as soil fertility, microclimate, and tree age were not consistently reported in all sources. This introduces potential unexplained variance, a limitation common in meta-analytical studies on allometry [134].

4.2. Ontogeny and Allometric Equations

Evolution and development jointly shape plant architecture. In theory, plant architecture may be subject to both phylogenetic constraints and ontogenetic genetic control. The formation trajectory of plant architecture, growth, branching, axial differentiation, and reproductive organ development is a dynamic and multi-level process of ontogeny, and 23 architectural models and their names were proposed and coined by Halle and his colleagues [135]. As a support of biomass production and partitioning at organ scale, architectural models can simulate individual tree development and stand dynamics, while thinning treatment can alter tree architectural [136]. Thus, morphological transformation and developmental bias exist in plants, although they are modular organisms [137].
Complex ontogenetic rules, e.g., leaf neoformation or polycyclism, result in tree architecture, and both inherent and environmental factors affect biomass formation and allocation of a tree [138]. In a tropical rainforest, wood density explains specific architectural differentiation, light capture strategy and species co-existence. Interestingly, wood density correlates negatively with stem diameter and positively with stem biomass [139]. Interspecific variation in architectural traits (tree diameter, height and crown dimensions) is related to the functional traits (light requirements and wood density) in a Central Africa tropical forest [140]. In the forest, the canopy’s obstruction of light has a substantial impact on the understory small trees. Stem height growth is limited in understory trees, and following canopy release events diameter growth responds more rapidly than tree height, indicating that tree growth is driven by diameter instead of height [141]. In a forest environment with low light and interference from neighboring trees, the biomass allocation of trees is size-dependent, with major allocation resources to leaves for small-diameter trees (DBH < 8 cm), to stems for medium-diameter trees (8 cm < DBH < 20 cm), and branches for large-diameter trees (DBH > 20 cm) [142]. Ontogenetic variation in tree biomass allocation is influenced by several ecological and biophysical factors: (i) Soil Fertility: In nutrient-rich environments, trees tend to allocate more biomass to aboveground parts (e.g., stems), leading to a more pronounced relationship between DBH and AGB. Conversely, in nutrient-poor soils, root allocation may be prioritized, causing deviations in allometric models for small trees. (ii) Light Availability: For understory trees or those in shaded environments, biomass allocation may favor leaf development for light capture, particularly in the early stages of growth. This alters the DBH-to-AGB relationship in small trees. (iii) Climate: Variability in temperature and precipitation regimes influences tree growth and biomass allocation. In drier or harsher climates, trees may allocate more biomass to roots, whereas in more temperate conditions, aboveground growth is often more pronounced. (iv) Anthropogenic Disturbance: Disturbances such as logging or land conversion can modify growth patterns, often resulting in altered biomass allocation strategies. In disturbed environments, trees may focus more on regrowth or survival, further influencing allometric relationships.
Tree architecture or allometry is also influenced by developmental time, e.g., tree age or size [143]. For a given tree species, the allometric model varies with developmental stages or size classes [144,145,146], and even if the stages differ by two years, their allometric coefficients may also be inconsistent [147]. Branches and leaves are more sensitive to environmental changes and are the tree components most influenced by stand age. The intercepts and slopes differ significantly across different stand age classes, indicating that stand age affects allometric growth models [28]. The allometry of Norwegian spruce exhibits an ontogenetic trend, and as age increases (2, 8, and 21 years), the biomass allocated to the stem elevates [148]. Due to the influence of the ontogenetic stage on biomass–diameter relation, allometric equations established based on different age groups or size classes may underestimate or overestimate biomass [149]. Different allometric models should be adopted to estimate the biomass of large and small trees, and applying equations for large trees to saplings results in biased and/or inaccurate estimates [16,132].
Analysis results on Castanopsis eyrei confirm that ontogenetic deviation of biomass estimation occurs among the individuals with different size classes. We also demonstrate that developing species-specific allometric equations for individuals of different ages can provide more robust estimates of aboveground biomass. Although using a generalized allometric equation for aboveground biomass estimation is convenient and effective for mature trees or forests, species-specific allometric equations should be used to reduce individual ontogenetic errors for small trees or young forests.

4.3. Selection of Allometric Equations

In order to reduce the model bias of AGB measurement and improve accuracy, a model needs to be validated before the selection allometric equations [150]. The selection of allometric equations often comes with shortcomings or defects. It is not advisable to replace regional models with national models directly, or vice versa [151]. Integrating more models and complete tree size classes and using a weighted function method is beneficial for selection of the optimal allometric equation [152].
Model selection requires multidimensional thinking, such as phylogenetics, ontogeny, and forest types. Phylogenetically, a reliable method for model selection is to group species based on phylogenetic relationships and determine allometric equations [35]. Genus-specific allometric equations, known as mixed models, can improve the accuracy of tree biomass estimates [153]. In addition, the biomass allometric model established based on the method of phylogenetically weighted regression (PWR) can avoid phylogenetic stationarity and improve the accuracy of biomass measurement [154]. Ontogenetically, the AGB aboveground biomass may be underestimated or overestimated when applying the allometric equations of large trees to small trees [16,29,155,156]. For a single tree species, there are significant differences in the allometric equations between large and small trees [157,158]. The generalized allometric equations for large trees should not be simply applied to understory shrubs, subcanopy small trees, and alpine dwarf forests, and need to be independently developed [158,159,160,161]. From the perspective of forest management, the selection of models needs to consider the diversity of tree species and the values of correlation coefficients [162]. In terms of the biomass allometric equations of Picea abies stands, specific models for pure stands cannot be applied to mixed stands (and vice versa); otherwise it may lead to significant bias [163]. This discrepancy in allometry between pure and mixed stands may be species-dependent. For instance, allometric patterns of biomass in mixed stands are significantly different from those in monospecific stands for Quercus petraea, while those in Pinus sylvestris are similar [164]. In addition, site effects [165,166] and climate effects [167,168] on biomass allometric models were documented in a quite a number of reports.
In the choice of biomass estimation models, there are often methodological mistakes and pitfalls, such as the arbitrary choice of analytical methods, model dredging and inadequate model diagnosis, and ignoring collinearity [134]. Based on the above discussion, if the differences in biological estimation objects and purposes are ignored, the selection of allometric models will also result in errors. Our data confirms that in the cases of forest stands with less species, in a young state, or for yield estimation, species-specific models should be applied as much as possible, while in the cases of forest stands with many tree species, in an old state, or for general biomass estimation, general models cannot be excluded.
This study highlights the need for dynamic AGB modeling strategies. Future research could focus on the following: (i) developing size-class-specific biomass equations to address ontogenetic variance; (ii) applying phylogenetically weighted regression in taxa showing consistent deviations; (iii) integrating functional trait data for more biologically informed allometry; (iv) compiling regional biomass databases with site metadata to improve model transferability.

4.4. Conclusions

This study provides critical insights into the interplay between phylogenetic constraints and allometric variation in forest biomass estimation. Theoretical implications emerge from the decoupling of trait evolution from phylogenetic history: the absence of strong phylogenetic signals (λ = 0.1249, p = 0.0027; K ≈ 0, p = 0.621) challenges the assumption of phylogenetic conservatism in allometric scaling, thereby validating the empirical rationale for employing generalized allometric models in phylogenetically diverse forest plots. Ecological significance lies in the hierarchical differentiation patterns, while family-level disparities (e.g., Theaceae vs. Myrtaceae) reflect divergent ecological strategies, and ontogenetic shifts across size classes highlight the necessity of incorporating life-stage dynamics into biomass prediction frameworks.
Methodologically, the reduced error magnitude when applying generalized equations to large individuals (vs. small individuals) suggests a strategic priority: resource-constrained monitoring programs could optimize accuracy by developing size-specific models for smaller individuals while retaining generalized equations for mature trees. These findings directly address the proposed hypotheses of ontogenetic sensitivity: The detectable size-class differentiation underscores a critical limitation-broad-scale equations may systematically misrepresent biomass allocation trajectories, necessitating error-correction protocols when extrapolating across developmental stages. Using generalized allometric equations to estimate AGB in FDPs with different size classes can results in ontogenetic errors. This study not only provides reference for the development of AGB allometric models, but also helps with AGB estimation in different scenarios.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17070493/s1, Dataset s1: Coefficients of 157 sets of species-specific allometric equations; Dataset s2: AGB errors calculated by generalized and species-specific allometric equations.

Author Contributions

Conceptualization, H.D. and Y.F.; methodology, Y.F.; software, Y.S. and Y.X.; formal analysis, Y.S.; investigation, Y.X., Y.S., X.Z. and X.R.; data curation, Y.X., X.R. and X.Z.; writing—original draft preparation, Y.S. and Y.F.; writing—review and editing, Y.F.; visualization, X.R.; supervision, Y.F.; project administration, Y.F. and H.D. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The original contributions presented in this study are included in the article/Supplementary Materials. Further inquiries can be directed to the corresponding author.

Conflicts of Interest

The authors declare no conflicts of interest.

Appendix A

Table A1. t-test results.
Table A1. t-test results.
DBH Class (cm)T-Statisticp-Value
(0–5]1.9600.050 *
(5–10]−6.059<0.001 ***
(10–15]−3.5880.0004 ***
(15–20]−0.5490.584
(20–25]0.5410.589
(25–30]−1.3200.190
(30–35]1.0170.312
(35–40]1.1810.244
(40–45]1.3200.198
(45–50]0.0330.974
(50–55]1.1730.271
(55–60]−1.6340.141
Note: * indicate statistically significant results at the corresponding significance level. *: p < 0.05; ***: p < 0.001.

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Figure 1. Topographic map of Mt. Wuyishan dynamics plot.
Figure 1. Topographic map of Mt. Wuyishan dynamics plot.
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Figure 2. Phylogenetic tree of the studied 157 species.
Figure 2. Phylogenetic tree of the studied 157 species.
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Figure 3. AGB error distribution by family.
Figure 3. AGB error distribution by family.
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Figure 4. AGB distribution across DBH classes. Note: a and b represent the DBH classes that show significant differences in the t-test.
Figure 4. AGB distribution across DBH classes. Note: a and b represent the DBH classes that show significant differences in the t-test.
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Table 1. Statistical results of allometric coefficient analysis across species groups (a, random, bm).
Table 1. Statistical results of allometric coefficient analysis across species groups (a, random, bm).
GroupC meanIKK.starλ
a0.1106 *0.0764 **0.04270.04710.1249 **
(p = 0.024)(p = 0.002)(p = 0.621)(p = 0.749)(p = 0.003)
random−0.0317−0.00640.04700.05760.00007
(p = 0.674)(p = 0.477)(p = 0.475)(p = 0.476)(p = 1.000)
bm0.5201 **0.2709 **0.6239 **0.6012 **1.0010 **
(p = 0.001)(p = 0.001)(p = 0.001)(p = 0.001)(p = 0.001)
Note: Values represent estimated parameters and associated p-values. Significant results are bolded with asterisks (*: p < 0.05, **: p < 0.01). C mean: Mean effect size of covariates; K: Kruskal–Wallis statistic for median comparisons; K.star: Adjusted K-value; λ: Regularization parameter; a: Empirical dataset; bm: Brownian motion model; random: Random null model.
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Shang, Y.; Xia, Y.; Ran, X.; Zheng, X.; Ding, H.; Fang, Y. Allometric Equations for Aboveground Biomass Estimation in Natural Forest Trees: Generalized or Species-Specific? Diversity 2025, 17, 493. https://doi.org/10.3390/d17070493

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Shang Y, Xia Y, Ran X, Zheng X, Ding H, Fang Y. Allometric Equations for Aboveground Biomass Estimation in Natural Forest Trees: Generalized or Species-Specific? Diversity. 2025; 17(7):493. https://doi.org/10.3390/d17070493

Chicago/Turabian Style

Shang, Yuxin, Yutong Xia, Xiaodie Ran, Xiao Zheng, Hui Ding, and Yanming Fang. 2025. "Allometric Equations for Aboveground Biomass Estimation in Natural Forest Trees: Generalized or Species-Specific?" Diversity 17, no. 7: 493. https://doi.org/10.3390/d17070493

APA Style

Shang, Y., Xia, Y., Ran, X., Zheng, X., Ding, H., & Fang, Y. (2025). Allometric Equations for Aboveground Biomass Estimation in Natural Forest Trees: Generalized or Species-Specific? Diversity, 17(7), 493. https://doi.org/10.3390/d17070493

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