Dendrochronological potential of tropical species in the Peruvian Amazon: An analysis in flooded forests of the southeast

Abstract

Research on growth rings in tropical trees within flooded forests has unveiled the complexity of climate change in these ecosystems. However, there has been limited understanding regarding species and their potential for dendrochronology. This study assessed 20 species from 13 botanical families in a flooded forest in southeastern Peru. Wood samples were collected during the dry season using a non-destructive sampling with motorized drill, alongside botanical samples for identification. Growth ring features were described following the IAWA, at a macroscopic level. Thirteen species showed promise for dendrochronological studies, eight of which were previously undocumented. These findings are pivotal for prioritizing species in future dendrochronological investigations in the Peruvian Amazon.

Full Article

Dendrochronological Potential of Tropical Species in the Peruvian Amazon: An Analysis in Flooded Forests of the Southeast

Miguel A. Ranilla-Huamantuco ,^a,f Ingrid A. Cassana-Huáman ,^b,e John Canales-Ramirez ,^c Javier Navio-Chipa ,^d Kerly J. Macedo-Enriquez ,^d Robert F. Farfan-Huanca ,^d and Leif A. Portal-Cahuana ^b,e,*

Research on growth rings in tropical trees within flooded forests has unveiled the complexity of climate change in these ecosystems. However, there has been limited understanding regarding species and their potential for dendrochronology. This study assessed 20 species from 13 botanical families in a flooded forest in southeastern Peru. Wood samples were collected during the dry season using a non-destructive sampling with motorized drill, alongside botanical samples for identification. Growth ring features were described following the IAWA, at a macroscopic level. Thirteen species showed promise for dendrochronological studies, eight of which were previously undocumented. These findings are pivotal for prioritizing species in future dendrochronological investigations in the Peruvian Amazon.

DOI: 10.15376/biores.20.1.1482-1503

Keywords: Growth rings; Madre de Dios; Peruvian woods; Tropical dendrochronology; Tropical forest; Wood anatomy

Contact information: a: Laboratorio Desconcentrado de Peritaje sede Madre de Dios, Ministerio Público, Puerto Maldonado, Perú; b: Instituto de Investigación, Innovación y Desarrollo para el Sector Agrario y Agroindustrial (IIDAA), Facultad de Ingeniería y Ciencias Agrarias (FICA), Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas (UNTRM), Calle Higos Urco 342 – Ciudad Universitaria, Chachapoyas, 01000, Perú; c: Empresa Agroforestal Orgánica R&H, Puerto Maldonado, Perú; d: Consultor independiente, Puerto Maldonado, Perú; e: Xiloteca Gocta, Escuela Profesional de Ingeniería Forestal, Universidad Nacional Toribio Rodríguez de Mendoza de Amazonas, Perú; f: Universidad Nacional San Antonio Abad del Cusco, Cusco, Perú;*Corresponding author: leif.portal@untrm.edu.pe

GRAPHICAL ABSTRACT

INTRODUCTION

Dendrochronology, an expanding field, has ventured into tropical forests, unveiling intriguing secrets and facing unique challenges. The presence of annual rings in over 20 tropical countries, as evidenced by Worbes (2002), highlights synchronized growth linked to extreme climatic events, such as brief droughts and prolonged floods. This technique has enabled an understanding of the influence of phenomena like El Niño on tree growth and the longevity of broadleaf trees in tropical lowlands. Rozendaal and Zuidema (2011) emphasize the relevance of understanding the growth and ecology of tropical trees to conserve and manage these ecosystems.

Despite the challenges posed by less pronounced changes in seasons, techniques, such as dendrometric measurement, cambial injuries, or carbon dating, have allowed us to comprehend the relationship between climate and tree growth in these regions. Meanwhile, Brienen et al. (2016) reveals that ring formation in the tropics is primarily linked to seasonal variations in rainfall or flooding rather than temperature. Despite the moderate sensitivity of tropical trees to climate, the ring studies offer valuable insights into the effects of climate change on these ecosystems.

Many studies have reinforced the importance of annual growth rings in understanding forest ecology and dynamics, as well as their application in projecting timber extraction and sustainable resource management (Schöngart et al. 2017; Marcelo-Peña et al. 2020). In the specific context of Peru, Portal-Cahuana et al. (2023a) highlight how the development of ring chronologies in tropical trees allows for the reconstruction of the neotropical environmental history. They identify sampling gaps and underscore Peru’s high tree diversity as a natural laboratory to better comprehend the growth and functioning of tropical species in interaction with the climate. This specific research provides a detailed insight into dendrochronology in the Peruvian context, contributing to guiding future dendrochronological studies in this region.

The study of potential species for tropical dendrochronology is crucial for understanding tree growth under diverse climatic conditions. Several researchers have addressed this pertinent area (Brienen et al. 2006; Beltrán and Valencia 2013; Groenendijk et al. 2014; Schöngart et al. 2017; Bauer et al. 2020; Marcelo-Peña et al. 2020). Traditionally, the analysis of growth rings was believed to be feasible only in biomes with marked seasonality, such as temperate forests. However, studies in Peru have revealed the presence of annual rings in various species across altitudinal gradients. These findings debunk the seasonal limitation in the tropics for dendrochronological studies (Marcelo-Peña et al. 2020).

The association between the phylogenetic relationship and the distinctiveness of growth rings, as discussed by Marcelo-Peña et al. (2019), suggests a diversity of responses among deciduous and evergreen species to environmental conditions. Bauer et al. (2020) complement this picture by observing the relationship between leaf phenological patterns and growth rings in subtropical forests, through identifying species with clear anatomical boundaries in their rings. Beltrán and Valencia (2013) delve into the challenges and findings of the anatomical characterization of growth rings in potential species for dendrochronological studies in the Central Selva of Peru. These studies highlight issues, such as difficulty in ring visualization, irregularities, and the presence of parenchyma in bands, emphasizing the importance of understanding variability in cell dimensions to infer the annual formation of rings.

Groenendijk et al. (2014) and other researchers have assessed the potential for applying growth ring analysis in tropical tree species under moist conditions. Despite the lack of seasonal variation in certain tropical forests, distinct boundaries in tree rings were identified, demonstrating the potential for obtaining precise growth data. Collectively, these studies underscore the significance of investigating and understanding potential species for tropical dendrochronology, particularly in the flooded forests of southeastern Peru. The complexity and unique conditions of these ecosystems, characterized by seasonal flooding, highlight the necessity of dendrochronological research in this region. This not only expands knowledge about growth ring formation in diverse tropical conditions but also provides valuable insights for sustainable forest management and understanding tree responses to climate change in these ecosystems. The upland forests exhibit greater species diversity compared to flooded forests (Myster 2015). These forests face annual flooding, raising water levels by over 10 m and submerging seedlings, and trees for up to seven months. This disrupts nutrients, oxygen, and toxins, creating optimal conditions for plant growth, and demanding adaptations (Parolin et al. 2004). Amid current climate changes, understanding the mechanisms of floods and water flows and their impact on the development and alterations of riparian forests becomes essential (Berthelot et al. 2015).

This is one of the first studies assessing the potential of tree species growing in a flooded forest in southeastern Peru. To achieve this, the authors distinguished and characterized the growth rings of 20 tree species. Specifically, the following questions were addressed: (i) How do the growth ring boundaries of 20 tree species growing in a flooded forest differ?; (ii) What are the anatomical characteristics of the growth ring boundaries of 20 tree species in a flooded forest?; (iii) Which tree species show greater potential for dendrochronological studies in this flooded forest? The authors approached these inquiries using 20 tree species from 13 different botanical families.

EXPERIMENTAL

Study Area

The study area focuses on the Low Terrace Forests (LTF) situated along the right bank of the Madre de Dios River in the Madre de Dios Department, Peru. These forests fall under the jurisdiction of the Amazon Producers Association of Camu Camu, known as APAYCNA-MDD, positioned at an altitude of approximately 120 m around coordinates 12°35’26.69″S – 69°18’25.52″W. The sampling sites are located 600 m away from the Madre de Dios River (Fig. 1).

Fig. 1. Location of the study site in the Madre de Dios region, southeastern Peru: A) Terrace Forest (blue circle). The color map illustrates the forest types. B) Red triangles indicate the position of sampled trees. C) The red arrow marked on the tree image indicates the maximum flood level in the area. D) Climate diagram for the period 1990 to 2019 is shown (Walter and Lieth 1960), where darker gray areas represent periods with over 100 mm of precipitation.

The Low Terrace Forests (Btb), also known as floodable forests, undergo annual flooding lasting between 3 to 8 months (Poma 2007). The diversity and floristic composition of tree species in these forests differ and are lower compared to upland forests (Normand et al. 2006; Féret and Asner 2014).

In climatic terms, this region is characterized by a warm, humid, and seasonal climate, with an average annual precipitation ranging between 2200 and 2400 mm, and an average annual temperature of 24.2 °C. Maximum temperatures reach 37.9 °C, while minimum temperatures drop to 11.3 °C (Terborgh and Andresen 1998; Román-Dañobeytia et al. 2015; Best et al. 2021).

Species, Processing, and Analysis of Growth Rings

Twenty tree species from a low terrace forest, belonging to 18 genera and 13 different botanical families, representative of the study area, were selected; detailed information regarding the wood samples is provided in Table 1. Botanical samples were collected, and photographs of leaves, flowers, fruits, trunk, roots, and outer and inner bark were recorded. Subsequently, the samples were preserved and sent to the Forest Herbarium (MOL) at the National Agrarian University La Molina (UNALM) in Lima, Peru, for proper classification and deposition.

Wood samples were collected during the dry season (May to August) using a non-destructive method employing the Stihil BT45 gasoline-powered motorized drill with hollow bits (Fig. 3), measuring 2.5 cm in diameter and adjustable in length to fit the tree diameter, ranging from 60 to 110 cm, collected at 1.30 m in a 90° angle from the tree axis (Marcelo-Peña et al. 2019; Aragão et al. 2022). The wound created on the tree trunk was covered with a protective and wound-healing paste and subsequently sealed with a liquid silicone gun to prevent the entry of insects and pathogens (Portal-Cahuana et al. 2023c).

Fig. 2. In the field: A) Non-destructive collection; B) Sample extraction; C) Fungicide paste; and D) Sealed with silicone

The samples were placed on coded wooden mounts and secured with twine. They were then left at room temperature to air dry at the Xiloteca Gocta (specialized environment) of the National University Toribio Rodríguez de Mendoza de Amazonas. Subsequently, they were affixed to the wooden mounts with the cross-section facing upward and sanded and polished using an 80 to 600 grain/cm² sandpaper sequence (Portal et al. 2021; Roquette et al. 2023b) to aid in the precise visualization of ring growth boundaries (Roquette et al. 2023a). The samples were digitized at 1200 dpi using a scanner with a scale and examined under a stereoscopic microscope at various magnifications (Aragão et al. 2019; Menezes et al. 2022).

Finally, the growth rings of the twenty wood samples (Fig. 3) were described following the IAWA guidelines of (IAWA 1989); including their general and macroscopic descriptions. Additionally, the qualitative information was systematized, and a Principal Component Analysis (PCA) was conducted to distinguish and describe the growth ring boundaries using the PAST software (Hammer, Øyvind, version 1.0, Tromsø, Norway) (Hammer 2001).

RESULTS AND DISCUSSION

General and Macroscopic Description of Woods

A comprehensive study was conducted to provide a general and macroscopic description of 20 types of wood from a flooded Forest of Southeast Peru, as detailed in the accompanying Table 1.

Through meticulous observation and analysis, the distinctive and anatomical features of each wood species have been recorded. Additionally, macrophotographs of the transverse cuts of each species are presented for better visual understanding. Furthermore, a PCA analysis was performed, which provides a visual representation of the similarity between forest species, along with clustering to facilitate the identification and characterization of groups of species with similar anatomical characteristics (Appendix, Supplementary Material).

The findings of the study support the importance of investigating wood anatomy as a crucial tool to address the issue of illegal logging in the Peruvian Amazon, by providing a means to accurately identify tree species and their origins, which can help trace and prevent illegal harvesting practices (Portal-Cahuana et al. 2023c). A detailed understanding of the anatomical characteristics of different wood species allows for better identification and classification of wood, playing a fundamental role in combating the trade of illegally extracted timber.

Furthermore, the implementation of precise and reliable wood identification systems, such as the convolutional neural networks mentioned in the article (Ferreira et al. 2020), facilitates the distinction between legal and illegal timber, thereby strengthening efforts to promote sustainable logging practices and protect valuable forest resources. It is necessary to emphasize the importance of monitoring, incentivizing, and monetizing legal and sustainable wood value chains in the Peruvian Amazon, thus highlighting the ongoing need for studies on wood anatomy to address environmental and socio-economic challenges in the region (Da Cunha Soares 2017).

Table 1. General and Macroscopic Description of 20 Wood Species from Flooded Forests in Southeast Peru

Note: MCR: Manual cutting resistance; VIS: Visibility; APO: Apotracheal; DIA: Diffuse in aggregate; PAR: Paratracheal; VAS: Vasicentric; SCA: Scalariform

Distinguishing the Boundaries of the Growth Ring

The distinction of growth ring boundaries is a relevant aspect of wood anatomy, which aids in the identification of tree species. Among the 20 studied species, varied growth ring patterns were observed: three species (15%) displayed highly distinct growth rings, ten species (50%) showed moderately distinct growth rings, seven species (30%) presented growth rings with low distinctiveness, and in one species (5%), the growth rings were indistinct or absent (Table 2).

Table 2. Characterization of Growth Rings of Twenty Trees from the Madre de Dios Region

DLAC = Distinction of growth ring boundaries; MuyD = Very distinct; MD = Moderately distinct; PD = Less distinct; IoA = Indistinct or Absent; DLA = Anatomical limits descriptor; CEPF = Change in fiber wall thickness; PM = Marginal parenchyma; CEPFyPM= Change in fiber wall thickness and Marginal parenchyma

The research on the distinction of growth ring boundaries in Peru has been limited. A previous study conducted in the Central Selva pointed out notable issues in this distinction, highlighting difficulties in visualizing some rings, irregular growth patterns, and the presence of parenchyma in bands (Beltrán and Valencia 2013). These same characteristics identified in the prior research have been reaffirmed in the current study on the distinction of growth ring boundaries, emphasizing the persistence and relevance of these particularities in the tree rings of the region. Additionally, more recent investigations have shown that deciduous species in seasonally dry tropical forests exhibit distinct growth rings, mainly marked by the presence of marginal parenchyma, while evergreen trees in lowland Amazonian and premontane forests show indistinguishable growth rings (Marcelo-Peña et al. 2020). The coherence between the current findings and previous studies underscores the consistency of patterns found in diverse types of tropical forests regarding the formation of growth rings and the relevance of environmental factors in this process.

Fig. 3. Growth ring boundaries in the macroscopic cross-section: A) Brosimun lactescens; B) Calycophyllum spruceanum; C) Clarisia racemosa; D) Diospyros sp.; E) Ficus insipida; F) Guatteria sp; G) Inga sp.; H) Luehea grandiflora; I) NN; J) Pouteria baenhiana; K) Pouteria sp; (1). L) Pouteria sp; (2). M) Pseudolmedia laevis; N) Pterocarpus rorhii; O) Senegalia sp.; P) Spondias mombin; Q) Terminalia oblonga; R) Virola elongata; S) Zanthoxylum acuminatum; T) Zygia sp.; The scale bar is 5 mm. White triangle marks the limit of the growth ring.

The clear identification of growth ring boundaries is essential for dendrochronological studies, as it directly impacts the ability to accurately determine tree age, assess growth rates, and understand how trees respond to environmental variables such as climate fluctuations, flooding, and droughts. This makes the distinction of growth rings a critical tool for exploring past climate conditions and for monitoring long-term environmental changes in tropical forests (Brienen et al. 2009; Marcelo-Peña et al. 2020; Roquette et al. 2023).

Anatomical Boundary Description

In the trees analyzed in this study, the authors identified different criteria to define the growth ring boundaries. In a total of 15 species (75%), this boundary is characterized by changes in fiber wall thickness in latewood. In four species (20%), the boundary is defined by a combination of changes in fiber wall thickness in latewood and the presence of marginal parenchyma. Finally, in one species (5%), the growth ring boundary is exclusively based on the observation of marginal parenchyma (Table 2).

Beltrán and Valencia (2013) and Marcelo-Peña et al. (2020), in their studies on anatomical boundary descriptors of growth rings in Peru, align with these findings by observing that the majority of the studied species were characterized by changes in fiber wall thickness as a boundary. This suggests that tropical trees in Peru commonly exhibit this delineation in their growth rings. However, although changes in the fiber wall thickness can mark the growth ring boundary in some species, this criterion is not always sufficient, as in the case of Spondias mombin.

These investigations into the delineation and descriptors of growth rings in tropical forests can be linked to phenology (Bauer et al. 2020) and cambial activity (Marcati et al. 2006, 2008; Lisi et al. 2008). Phenological events, such as seasonal variations in temperature and rainfall, influence the timing of cambial activity, which in turn affects the formation of distinct anatomical boundaries in the wood (Marcati et al. 2006). Cambial activity, driven by these environmental cues, leads to the differentiation between earlywood and latewood, creating clear boundaries in species with pronounced growth rings (Lisi et al. 2008). Understanding these processes is essential for interpreting the relationship between tree growth and climatic conditions, especially in tropical forests where such patterns may vary.

The principal component analysis yielded notable outcomes in distinguishing the growth ring boundaries (Fig. 4) and describing the anatomical limits (Fig. 5). These findings prove highly valuable for species identification and understanding the potential of growth rings in future dendrochronological studies in a low terrace forest located in the Madre de Dios region, Peru.

In particular, certain species with high potential for dendrochronological studies were identified, such as Terminalia oblonga, Zygia sp., and Zanthoxylum acuminatum, due to the pronounced distinction in their growth rings, as well as variations in fiber wall thickness and marginal parenchyma. These two descriptors play a crucial role in the accurate delimitation of growth rings.

In contrast, it was observed that certain species have potential for dendrochronological studies: Pouteria sp. (1), NN, Calycophyllum spruceanum, Brosimum lactescens, Luehea grandiflora, Pterocarpus rohrii, Virola elongata, Pouteria sp., Diospyros sp., and Ficus insipida exhibited moderately distinct rings with changes in fiber wall thickness in latewood, while only Virola elongata showed marginal parenchyma.

In contrast, a group of species displayed rings that were slightly distinct, indistinct, or even absent, suggesting a limited potential for dendrochronological studies. This group includes Clarisia racemosa, Pouteria baehniana, Pseudolmedia laevis, Inga sp., Senegalia sp., Guatteria sp., and Spondias mombin.

Fig. 4. Principal component analysis on the distinction of growth ring boundaries among the twenty studied tree species

Fig. 5. Principal component analysis on the anatomical boundary descriptor of growth ring limits among the twenty studied tree species

The authors report that 65%, equivalent to 13 species found in a low terrace forest located in the Madre de Dios region, Peru, show potential for dendrochronological studies. Among these, 12 species have had their growth ring potential analyzed in other types of tropical forests (Worbes 2002; Schöngart et al. 2007, 2017; Beltrán and Valencia 2013; Marcelo-Peña et al. 2020).

Regarding the potential of forest species for dendrochronological studies, growth rings have been studied in the states of Cajamarca, La Libertad, Pasco, Ucayali, and Junín (Beltrán and Valencia 2013; Marcelo-Peña et al. 2020). However, there has been limited exploration in the Southeastern region of Peru, especially in forests partially submerged in water for parts of the year. It is valuable to collect wood samples in tropical areas experiencing seasonal changes to obtain precise data on tree rings. For instance, in areas affected by seasonal flooding, where cambial activity diminishes, these changes might be evident in wood cells, revealing variations in tree growth rings and highlighting their patterns.

In terms of using forest species in dendrochronological studies, growth rings have been examined across various regions in Peru such as Cajamarca, La Libertad, Pasco, Ucayali, and Junín (Beltrán and Valencia 2013; Marcelo-Peña et al. 2020). However, exploration in the southeastern part of the country and in forests partially submerged seasonally remains limited. It is crucial to collect wood samples in tropical areas with seasonal changes to obtain precise information about tree rings (Rozendaal and Zuidema 2011; Blagitz et al. 2019) For instance, in temporarily flooded forests where cambial activity reduces growth, this is reflected in wood cells, showcasing variations in growth rings and highlighting them as relevant indicators (Wimmer 2002; Schöngart et al. 2005). Finally, regarding species with high potential for dendrochronological studies, there have been investigations on growth rings within the genera Terminalia and Zanthoxylum. Studies have been conducted on species like Terminalia amazonica, T. catappa, T. guyanensis, T. quintalata, Zanthoxylum caribaeum, Z. kellermanii, and Z. rhoifolium (Schöngart et al. 2017; Portal-Cahuana et al. 2023b). However, for Zygia sp., no research with species from this genus has been found.

CONCLUSIONS

1. The authors delineated the growth ring patterns in 20 tree species from a lowland terrace forest in Madre de Dios, Peru. Notably, eight previously undocumented species were discovered. These species exhibited a range of growth ring patterns, with 15% showing highly distinct rings, 50% displaying moderately distinct rings, 30% having rings with low differentiation, and 5% presenting indistinct or absent rings.
2. Criteria for defining growth ring boundaries varied: in 75% of the species, changes in the fiber wall thickness in latewood defined the boundary, while in 20%, it was a combination of these changes and the presence of marginal parenchyma. In the remaining 5%, the boundary was exclusively defined by observing marginal parenchyma.
3. Principal component analysis showcased noticeable differentiation in growth ring boundaries, as well as in anatomical descriptions of these limits. These outcomes are crucial for identifying species with high potential for dendrochronological studies, such as Terminalia oblonga, Zygia sp., and Zanthoxylum acuminatum, which exhibited distinct rings.
4. Conversely, some species showed rings with low differentiation or were absent, limiting their suitability for dendrochronological studies. These findings are essential for future dendrochronological investigations in the region and a better understanding of the ecological history of these forests.
5. This study contributes to a broader understanding of growth ring diversity in tropical forests in the Madre de Dios region, Peru. The findings offer valuable insights for future dendrochronological research and underscore the importance of environmental factors in influencing growth ring formation in these ecosystems.

ACKNOWLEDGMENTS

The author acknowledges the Universidad Nacional San Antonio Abad del Cusco – UNSAAC, for the funding provided for the research work of the Master’s thesis in Ecology and Environmental Management.

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Article submitted: October 28, 2024; Peer review completed: December 9, 2024; Revised version received and accepted: December 10, 2024; Published: December 17, 2024.

DOI: 10.15376/biores.20.1.1482-1503

APPENDIX

Supplementary Information

Dendrochronological Potential of Tropical Species in the Peruvian Amazon: An Analysis in Flooded Forests of the Southeast

Fig. S1. Macroscopic sections of the species. A) Brosimun lactescens (Moore) C.C.Berg. B) Calycophyllum spruceanum Benth. C) Clarisia racemosa R.et P. D) Diospyros sp. Image 1: Cross-section, 2: Radial section, and 3: Tangential section. Note: The bar measures 20 mm in the cross-section, and 10 mm in the radial and tangential sections.

Fig. S2. Macroscopic sections of the species. E) Ficus insipida Willd. F) Guatteria sp. G) Inga sp. H) Luehea grandiflora C. Martius cf. Image 1: Cross-section, 2: Radial section, and 3: Tangential section. Note: The bar measures 20 mm in the cross-section, and 10 mm in the radial and tangential sections.

Fig. S3. Macroscopic sections of the species. I) NN J) Pouteria baenhiana Monachino. K) Pouteria sp. (1). L) Pouteria sp. (2). Image 1: Cross-section, 2: Radial section, and 3: Tangential section. Note: The bar measures 20 mm in the cross-section, and 10 mm in the radial and tangential sections.

Fig. S4. Macroscopic sections of the species. M) Pseudolmedia laevis (R. et P.) J.F. N) Pterocarpus rorhii Vahl. O) Senegalia sp. P) Spondias mombin L. Image 1: Cross-section, 2: Radial section, and 3: Tangential section. Note: The bar measures 20 mm in the cross-section, and 10 mm in the radial and tangential sections.

Fig. S5. Macroscopic sections of the species. Q) Terminalia oblonga (R.et P) Steud. R) Virola elongata (Benth.) Warb. S) Zanthoxylum acuminatum (Sw.) Sw. T) Zygia sp. Image 1: Cross-section, 2: Radial section, and 3: Tangential section. Note: The bar measures 20 mm in the cross-section, and 10 mm in the radial and tangential sections.

Table S1. Popular Name and Main Uses of the Wood of the 20 Forest Species in the Flooded Forest of Southeastern Peru

^a [1]; ^b [2]; ^c http://www.tropicaltimber.info/es/; ^d [3]; e [4].

Fig. S6. A) Principal Component Analysis. B) Clustering of the twenty tree species in the flooded forest of southeastern Peru.

In Fig. S6, it is evident that PCA analysis provides a visual representation of the similarity among tree species in a reduced coordinate space, while clustering allows the identification and characterization of groups of species with similar anatomical features. Combining these approaches offers a more comprehensive understanding of the diversity and variability of tree species based on their wood anatomy.

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