Morphological and anatomical characterization of Qualea dinizii Ducke (Vochysiaceae) applied to the identification of Brazilian Amazon forest species

Caracterização morfológica e anatômica de Qualea dinizii Ducke (Vochysiaceae) aplicada à identificação de espécies florestais na Amazônia Brasileira

Abstract

Knowledge of Amazonian tropical forest species depends on accurate identification and characterization, which are legal requirements for sustainable forest management, traceability, appropriate end use, and control of the forest production chain. However, difficulties in species identification remain common and may compromise the reliability of forest inventories used in Forest Management Plans. This study aimed to describe the morphological and wood anatomical characteristics of Qualea dinizii Ducke. The species presents a trunk with a smooth rhytidome and horizontal striations, with powdery and exfoliating bark. Branches range from exfoliating to puberulous and exhibit intrapetiolar stipules with paired extrafloral glands. Growth rings are distinct and delimited by darker fibrous zones. The wood is characterized by banded axial parenchyma, occasionally confluent and aliform, as well as by traumatic axial canals and the occurrence of macules. The detailed characterization of these macules represents an original contribution of this study. These findings fill an important knowledge gap by providing integrated morphological and wood anatomical descriptions of Q. dinizii in a single document, supporting more reliable identification and contributing to future ecological and technological studies of the species.

Keywords:
growth rings; extrafloral glands; macules; forest management; wood identification

Resumo

O conhecimento sobre as espécies florestais tropicais amazônicas depende de uma identificação e caracterização precisas, fundamentais para o manejo florestal sustentável, a rastreabilidade, o uso adequado da madeira e o controle da cadeia produtiva florestal. No entanto, ainda são frequentes dificuldades na identificação correta das espécies, o que pode comprometer a confiabilidade dos inventários florestais utilizados em Planos de Manejo Florestal. Este estudo teve como objetivo descrever as características morfológicas e anatômicas de Qualea dinizii Ducke. A espécie apresenta tronco com ritidoma liso e estriações horizontais, com casca pulverulenta e exfoliante. Os ramos variam de exfoliantes a pubérulos e apresentam estípulas intrapeciolares com glândulas extraflorais pareadas. Os anéis de crescimento são distintos e delimitados por zonas fibrosas mais escuras. A madeira é caracterizada por parênquima axial disposto em faixas, ocasionalmente confluente e aliforme, bem como pela presença de canais axiais traumáticos e pela ocorrência de máculas. A caracterização detalhada dessas máculas representa uma contribuição original deste estudo. Esses resultados preenchem uma importante lacuna de conhecimento ao reunir, em um único documento, descrições morfológicas e anatômicas integradas de Q. dinizii, contribuindo para uma identificação mais confiável e subsidiando futuras pesquisas ecológicas e tecnológicas sobre a espécie.

Palavras-chave:
anéis de crescimento; glândulas extraflorais; máculas; manejo florestal; identificação da madeira

1. Introduction

The Amazon Forest is the largest continuous tropical forest area in the world and plays an important role in climate regulation and ecosystem functioning. It contains more than 10% of global terrestrial biodiversity and represents approximately 67% of the remaining tropical forests worldwide (Flores et al., 2024; Fernandes et al., 2025).

Due to its ecological relevance and legal status, the Brazilian Federal Constitution of 1988 recognizes the Amazon Forest as part of the national heritage. As a result, its exploitation must follow regulated forest management practices designed to maintain environmental integrity. Forest management includes strategies for the sustainable use of timber and non-timber forest products, aiming to generate social and economic benefits while reducing waste, increasing productivity, and minimizing impacts on the remaining forest (Rodrigues et al., 2020).

Reliable species identification is required for forest inventories, management planning, and law enforcement. It is also necessary to support conservation efforts and reduce illegal logging, which remains a major issue in the Brazilian Amazon (Siam et al., 2023). In this context, CONAMA Resolution No. 406/2009 established the mandatory use of technical and scientific procedures for the botanical identification of managed forest species in Brazil, requiring the recognition of both scientific and vernacular names (Brasil, 2009).

Despite this legal framework, botanical identification remains a major challenge for forestry professionals in Brazil and is often restricted to the genus level. Misidentification of species in forest management plans compromises the quality of forest inventories, leads to inaccurate data being entered into forest information systems, and negatively affects decision-making processes related to sustainability (Procópio and Secco, 2008; Botosso, 2009; Souza et al., 2020). Conversely, accurate species identification improves transparency in the timber supply chain and facilitates access to high-value markets.

Qualea dinizii Ducke belongs to the family Vochysiaceae, one of the most species-rich families in the Amazon region, comprising approximately 250 species distributed among eight genera. Six of these genera occur in Brazil, highlighting the representativeness and diversity of the family in the country (Angiosperm Phylogeny Group, 2016; Silva et al., 2021). Although native, these species are not endemic to Brazil (Flora e Funga do Brasil, 2025). Approximately 60 species occur throughout Central and South America (Kawasaki, 2007; Moya and Brea, 2015; Shimizu et al., 2016; Shimizu et al., 2020).

Species of the genus Qualea Aubl. are large trees that may reach heights of 30–35 m (Howe and Smallwood, 1982; Barbosa et al., 2017), and their wood is recognized for its economic potential. These species exhibit similar physical and mechanical properties, good workability, and favorable characteristics for turning, finishing, and gluing, making them suitable for civil construction and the furniture industry (Cardoso et al., 2012; Silva et al., 2021). However, the high diversity of Vochysiaceae and the similarity among species may hinder accurate identification in the field, especially when diagnostic morphological and anatomical information is limited.

In this context, detailed and integrated descriptions of the morphology and wood anatomy of Qualea dinizii remain scarce in the scientific literature, limiting its reliable identification and proper classification in forest inventories and management plans. Therefore, this study aimed to describe the morphological and anatomical characteristics of Qualea dinizii Ducke in the Brazilian Amazon. The results provide original reference data to support species identification and contribute to ecological and technological studies involving this species.

2. Materials and Methods

2.1. Study area and sampling

The study area is located in the municipality of Juína, northwestern Mato Grosso State, Brazil (Figure 1A), at 58°44’21.74” W and 11°22’31.03” S. The area comprises a fragment of primary tropical forest covering a total area of 340.28 ha and managed under a sustainable forest management regime within the Brazilian Legal Amazon region. The regional climate is classified as “Am” according to the Köppen–Geiger system and is characterized as hot and humid tropical, with marked seasonality defined by rainy and dry periods (Alvares et al., 2014). Mean annual precipitation is approximately 2,200 mm, with monthly rainfall below 60 mm during the dry season. The average annual temperature is 24 °C, with maximum temperatures reaching up to 40 °C (IBGE, 2002; Cruz and Drescher, 2025).

Figure 1
(A) Location of the study area; (B) material for morphological characterization; (C) wood sampling and equipment used for the characterization of Qualea dinizii Ducke, showing the different wood regions: A = inner heartwood, B = intermediate heartwood, B′ = outer heartwood, C = sapwood, and D = special anatomical structure observed in the 144-year-old tree.

2.2. Species selection and material collection

The selection of the study species was based on data obtained from the forest inventory of the Sustainable Forest Management Plan (Plano de Manejo Florestal Sustentável – PMFS). Five individuals of Qualea dinizii were selected and felled, representing diameter at breast height (DBH) classes both above and below 50.0 cm. Morphological characterization was performed based on the external characteristics observed in all five sampled trees.

Tree age was determined using the stem analysis methodology (Barusso, 1977; Cruz, 2024). After age determination, two individuals were selected for wood anatomical analysis. The DBH of the sampled trees ranged from 23.23 cm (tree A1, 79 years old) to 70.00 cm (tree A0, 144 years old). The anatomical characterization was based on two trees, which may limit representativeness. Therefore, the anatomical descriptions presented should be interpreted as reference information for the sampled individuals, and future studies with larger sample sizes are recommended to evaluate intraspecific anatomical variability.

Morphological characteristics of the leaves and trunk were documented through photographic records and field notes. The evaluated attributes included the appearance of the rhytidome (outer dead bark) and phloem (inner live bark), bark color, presence of exudates, odor, sapwood color, and trunk base morphology (straight, swollen, buttressed, or fluted). Vegetative material (Figure 1B) was collected for the morphological characterization of leaves and branches.

During sampling, the species was not in its reproductive phase. In the Amazon region, phenological heterogeneity among tree species often limits the observation of flowering and fruiting, thereby hindering taxonomic identification based solely on reproductive structures (Duarte et al., 2022). In this context, Dendrology, focused on the study and identification of trees, represents a complementary tool that enables accurate species recognition and reduces errors associated with the use of vernacular names.

2.3. Characterization of leaves, branches, trunk, and wood

The morphological characteristics of leaves and branches were analyzed using a Leica M205C stereomicroscope equipped with a camera and image capture system. The description of external characteristics was based on Ribeiro et al. (1999), Obermuller et al. (2011), and Flora e Funga do Brasil (2025). Species identification was confirmed by comparison with reference specimens deposited in virtual herbarium databases (SpeciesLink Network, 2024), including samples IAN158471 and IAN162842 from the Herbarium IAN – Embrapa Amazônia Oriental (Herbário IAN, 2024) and specimen 3340159 from the Missouri Botanical Garden Herbarium (Missouri Botanical Garden Herbarium, 2024).

To characterize the wood of Q. dinizii, discs from two trees were used, one 144 years old and the other 79 years old. The material was collected from the disk at DBH level (Figure 1c), from which a radial sample was taken. Subsequently, test specimens were prepared, measuring 2 × 2 × 2 cm (cross × tangential × radial sections), following the recommendations of COPANT (1974) and IAWA Committee (1989). Three samples were taken from the 79-year-old tree (Figure 1C), corresponding to the inner heartwood (A), intermediate heartwood (B), and sapwood (C). From the 144-year-old tree, five samples were collected, four of which were used for anatomical analysis: inner heartwood (A), intermediate heartwood (B), outer heartwood (B′), and sapwood (C). Additionally, one sample from this tree was used to evaluate a special wood structure (D).

The samples were polished using sandpaper with grit sizes ranging from 80 to 2000 grains·mm⁻² and evaluated for the organoleptic characteristics of the wood (color and distinction between heartwood and sapwood, odor, taste, grain, and shine), according to Latorraca et al. (2018). Macroscopic characterization was performed on the transverse, tangential, and radial sections with a binocular magnifying glass, and the images were recorded using a Leica M205C stereomicroscope. The identification of the wood was confirmed by comparison with samples recorded in the Xylotheque of the Wood Technology Laboratory, Mato Grosso Agricultural Defense Institute (INDEA-MT), according to the identification certificate number: 00009/2024/NLTM/INDEAMT. Microscopic analyses were conducted by microtomy, with softening of the test specimens in an autoclave at 120 °C, obtaining histological sections approximately 25 μm thick in a sliding microtome, decolorization in sodium hypochlorite, staining with safranin, dehydration in an alcohol series (30–99.99%), and fixation in ethyl alcohol/butyl acetate (1:1) and absolute butyl acetate. The macro and microscopic description followed the recommendations of COPANT (1974), IAWA Committee (1989), and Latorraca et al. (2018), evaluating axial parenchyma (visibility and arrangement), vessels (microscopic visibility, porosity, grouping, arrangement, and content), rays (visibility in the cross section, contrast in the radial section, and stratified rays in the tangential section), growth layers, and the presence of special structures. The quantitative microscopic parameters evaluated included pore frequency (Fp), tangential pore diameter (Øp), and ray dimensions (width – Lr, height – Hr, and frequency per mm – Fr).

Fiber morphology was characterized by maceration, according to the specifications of Brazilian Regulatory Standard NBR 15.066 (ABNT, 2004). Longitudinally sectioned samples (~3 mm) were treated with an equimolar (1:1) solution of glacial acetic acid and hydrogen peroxide and incubated at 60 °C for 48 h. After washing and staining with Astra Blue, temporary slides were prepared and analyzed using a ZEISS Primo Star trinocular microscope, coupled with an image acquisition system and operated with Zen 3.1 software (Figure 1C).

3. Results

3.1. Morphological characterization of leaves, branches, and trunk of Qualea dinizii

The species presents simple leaves with opposite phyllotaxy (Figure 2AD), entire margins, and smooth leaf blades. The leaves are petiolate, with small lateral stipules at the leaf node, glabrous on both surfaces, and exhibit a coriaceous texture. Leaf size ranges from 7.5 to 8.5 cm in length and from 2.5 to 3.0 cm in width. The leaf blades are oblong to elliptical-oblong, with a non-acute acuminate apex and an acute base, and exhibit discoloration between the adaxial and abaxial surfaces (Figure 2A).

Figure 2
Morphological characterization of leaf and branch architecture of Qualea dinizii: (A) Collected and prepared vegetative material used for species identification and confirmation; (B) Simple leaves with entire margins (black arrow), acuminate but non-acute apex (red arrow), acute leaf base (red arrow), prominent primary venation on the abaxial surface (yellow arrows), and discolored adaxial and abaxial surfaces; (C) Secondary veins not reaching the leaf margin; closed reticulate venation with areoles formed by tertiary and quaternary veins (red arrow); (D) Presence of intrapetiolar stipules (brown arrows) associated with extrafloral glands (blue arrows) and opposite phyllotaxy at the same node (blue arrows).

The primary venation is prominent on the abaxial surface, whereas the secondary veins are arcuate, arising from the midrib and without reaching the leaf margin. The venation pattern is closed, with smaller reticulate veins originating from the secondary veins. These give rise to tertiary and quaternary veins that form well-defined areoles with small enclosed spaces (Figure 2B).

Regarding branch morphology, the species exhibits surfaces ranging from exfoliating to puberulous. Intrapetiolar stipules are present and associated with paired extrafloral glands (Figure 2C).

The morphological characteristics evaluated in the trunk of the sampled adult individuals (Figure 3), included predominantly smooth rhytidomes (dead bark), with horizontal striations, a powdery texture, and exfoliating detachment in small flakes, exhibiting a grayish-brown coloration (Figure 3B). The trunk base commonly presented dilated buttresses (Figure 3A). The phloem (living bark) exhibited a pale brown color, non-characteristic exudate, and no perceptible odor. The sapwood exhibited a fibrous texture and yellowish coloration (Figure 3B).

Figure 3
Morphological characterization of the trunk of Qualea dinizii: (A) Predominantly smooth rhytidome (black arrow), horizontal striations (yellow arrow), and buttressed base (red arrows); (B) Inner bark and sapwood with fibrous appearance (black arrow), showing powdery and exfoliating detachment (red arrow).

3.2. Macroscopic characterization of Qualea dinizii wood

Organoleptic characteristics: The heartwood (HW) and sapwood (SW) are distinct, although poorly contrasted (Figure 4A). The heartwood exhibits a light brown color with a slightly pinkish hue (Figure 4B), whereas the sapwood shows a grayish-brown coloration (Figure 4A). The wood is hard when subjected to cutting. Brightness: mirror-like, with low contrast. Texture: smooth to medium. Grain: straight, with elements arranged parallel to the vertical axis of the trunk. Odor and taste: imperceptible. Growth ring boundaries are distinct and visible to the naked eye and under a 10× hand lens, demarcated by darker tangential fibrous zones (Figures 44F). Vessels are distinct and visible to the naked eye without the aid of a 10× lens, and exhibit diffuse porosity. Vessels are predominantly solitary, with occasional multiple vessels arranged in radial chains of two to four, presenting circular to oblique outlines and a nonspecific distribution (Figure 4F). Vessels are predominantly unobstructed, with isolated occurrences of obstructions containing whitish substances (Figure 4G). Axial parenchyma is distinct and visible under a 10× lens, predominantly paratracheal aliform with short wings and confluent aliform patterns (Figure 4F). Radial parenchyma (rays) is thin and sparse, visible under a 10× lens in both tangential and transverse sections (Figures 44F, respectively), non-storied (Figure 4D), and exhibiting poorly contrasted mirroring in the radial plane (Figure 4E). Macules and intercellular canals: Light-colored macules and traumatic axial canals were macroscopically observed in cross section (Figure 4H).

Figure 4
Macroscopic anatomical characterization of Qualea dinizii wood. (A) Disc removed from the tree showing the transition zone between heartwood (HW) and sapwood (SW); (B) Tangential section with rectilinear vascular lines arranged in a “V” pattern, visible to the naked eye (white arrow); (C) Cross section highlighting growth ring boundaries demarcated by darker fibrous zones (black arrows); (D) Tangential section showing nonstoried rays; (E) Radial section; (F) Detailed cross section allowing clearer visualization of vessels, axial parenchyma, and growth ring boundaries (black arrows); (G) Cross section detail showing vessel obstruction by whitish substances (white arrows); (H) Macule (blue arrow) and traumatic intercellular canals (red arrow). Scale bar = 1 mm.

3.3. Microscopic characterization of Qualea dinizii wood

Growth ring boundaries were distinct, demarcated by the contrast between fibrous zones at the end of one growth ring and the presence of marginal parenchyma at the beginning of the subsequent ring (Figures 5A, 5E). Vessels were distinct and clearly visible, exhibiting diffuse porosity (Figure 5A). Vessels occurred predominantly solitary, although radial multiples of two to four vessels arranged tangentially, with circular to oblique outlines, were also observed (Figures 5D, 5E). Most vessels were unobstructed, with occasional obstruction by tyloses (tylosis) (Figures 5A, 5D). Vessel elements presented perforation plates and alternate intervessel pits (Figures 5G, 6G, 6H). Axial parenchyma showed a predominantly paratracheal arrangement in bands more than three cells thick, with occasional confluent aliform and diffuse apotracheal patterns (Figures 5A, 5D, 5E). The axial parenchyma occurred in series of five to eight cells. Fibers were libriform, with simple pits and nonseptate walls (Figures 5F, 6G, 6I, 6J). Radial parenchyma (rays) was predominantly multiseriate, with the presence of uniseriate rays ranging from one to four cells in width and exhibiting a non-storied arrangementchan (Figure 5B). Rays were heterocellular, composed of procumbent cells with marginal rows of square and/or upright cells (Figure 5C). Radial secretory ducts were also observed (Figure 5F). Included phloem was absent. Special structures included mineral inclusions, with prismatic crystals occurring in axial parenchyma cells (Figure 5H) and druses present within vessels (Figure 5I). Traumatic axial canals composed of parenchymatous tissue were observed in cross section, appearing as irregular structures arranged tangentially along the trunk axis, with few associated fibers and embedded within the wood (Figures 5J, 5K). These structures were located in the cambial region of the 144-year-old tree. Additionally, the presence of cambial macule was recorded (Figures 6A, 6B, 6C).

Figure 5
Microscopic characterization of the anatomical structure of Qualea dinizii wood. (A) Cross section highlighting distinct growth ring boundaries (black arrows); (B) Tangential section showing nonstoried rays; (C) Radial section with heterocellular rays. Details of the cross section illustrating; (D) vessel obstruction by tyloses; (E) paratracheal axial parenchyma. Details of the tangential section showing; (F) a radial secretory canal (white arrow) and the presence of libriform, nonseptate fibers; (G) Vessel elements with simple perforation plates and alternate intervessel pits. Details of the radial section showing; (H) prismatic crystals in axial parenchyma cells; (I) deposits within vessels; (J, K) Traumatic intercellular canals (white arrows). Scale bars as indicated in the images.

Figure 6
Microscopic analysis of the abnormal anatomical structure of Q. dinizii in wood regions with macules. (A) Cross section of macule formed by fibrous tissue and rays, with few vessels and axial parenchyma; (B) Radial section of macule. (C, D) Detailed cross-sectional and radial sections of macule, respectively; (E, F) Tangential section of macule. Maceration of normal wood regions: (G) vessel element with simple perforation plate and libriform fiber; (H) cell wall of the vessel element, with alternating pits; (I, J) detailing of the cell wall of the libriform fibers. Maceration of wood regions with macules: (K) abnormal fibers with differentiated structures; (L) detailing of the cell walls of abnormal fibers. Scale bar as indicated in the images.

The observed macules exhibited a macroscopic appearance as light-colored spots, visible both to the naked eye (Figure 4H) and under microscopic analysis (Figure 6). These structures were located near the vascular cambium and were characterized by distinct coloration and alterations in xylem fiber density. Anatomically, the macules consisted predominantly of fibrous tissue and multiseriate rays, with a marked reduction in the number of vessels and axial parenchyma (Figures 6A6F).

Maceration analyses of the normal wood regions (Figures 6G, 66J) and the regions containing cambial macules (Figures 66L) revealed clear differences in fiber structure. In the normal regions, libriform fibers exhibited regular morphology, with uniform cell walls and simple pits, which are typical features of unaltered woody tissue. In contrast, fibers associated with the cambial macules displayed abnormal morphology, differing markedly from the libriform fibers observed in the surrounding wood. These structural alterations indicate that the presence of macules is directly associated with localized anatomical modifications in wood cells.

3.4. Quantitative characterization of Qualea dinizii wood

The quantitative characterization of the anatomical parameters of Qualea dinizii wood included pore frequency, tangential vessel diameter, and ray width, height, and frequency. Samples were obtained along the radial direction (pith–heartwood–sapwood) from trees A0 (144 years old) and A1 (79 years old). The values of the analyzed parameters are presented in Tables 1 and 2.

Table 1
Radial variation of quantitative pore parameters in Qualea dinizii wood.

Tree/ Age (years)	Radial Position	Fr (vessels/mm−2)					Øp (μm)
		Descriptive statistics				Clas.	Descriptive statistics				Clas.
		Mean	Min	Max	sd		Mean	Min	Max	sd
A1/ 79y	Inner HW (A)	12.69	9.61	19.22	3.92	Few	86.45	25.85	150.22	24.12	Small
	Intermediate HW (B)	8.27	7.31	9.27	0.86	Few	140.94	31.58	203.27	34.32	Medium
	SW (C)	6.34	5.70	6.97	0.49	Few	152.66	42.07	212.11	34.68	Medium
A0/ 144y	Inner HW (A)	9.91	7.99	17.01	3.97	Few	118.98	45.16	199.23	32.76	Medium
	Intermediate HW (B)	6.28	4.68	8.16	1.36	Few	171.54	50.98	234.94	33.97	Medium
	Outer HW (B')	6.90	5.02	7.82	1.09	Few	148.47	36.82	240.06	43.03	Medium
	SW (C)	7.70	6.12	9.52	1.26	Few	182.79	40.14	275.89	40.30	Medium

• Legend: HW = heartwood; SW = sapwood; Fr = pore frequency (number mm⁻²); Øp = tangential pore diameter (µm); Clas.: Classification; sd = standard deviation. A, B, B′ and C correspond to the ordered radial positions of specimen removal (pith–bark direction) in trees A1 (79 years old) and A0 (144 years old).

Table 2
Quantitative rays’ parameters of Qualea dinizii wood.

Tree/ Age (years)		A1 - 79 years old			A0 - 144 years old
Parameter		Fr (rays/mm)	Lr (μm)	Hr (μm)	Fr (rays/mm)	Lr (μm)	Hr (μm)
Descriptive statistics	Mean	5.50	27.32	465.41	4.07	42.03	353.20
	Min.	4.64	8.12	18.58	3.57	14.28	120.14
	Máx.	6.07	50.59	1,161.31	4.64	70.70	1,046.64
	sd	0.60	5.93	204.61	0.41	10.95	128.10
Classification		Few	Fine	Low	Few	Fine	Low

• Legend: Fr = ray frequency (number mm); Lr = ray width (µm); Hr = ray height (µm); sd = standard deviation.

4. Discussion

The integrated morphological and anatomical characterization of Qualea dinizii provides reference information for the identification of this Amazonian timber species. In sustainable forest management, species identification supports forest inventories, improves traceability, and reduces errors in technical documents. Although morphological and anatomical information on Q. dinizii has been reported in previous studies, the present work compiles external morphology, macroscopic wood description, microscopic anatomy, and biometric parameters into a single document, which may facilitate the recognition of diagnostic traits under field and laboratory conditions (Pinheiro, 2014).

The morphological traits of branches and trunk observed in this study agree with taxonomic and floristic descriptions available for Q. dinizii (Ribeiro et al., 1999; Obermuller et al., 2011; Martins-da-Silva et al., 2014; Almeida and Almeida, 2018; Shimizu et al., 2020; Flora e Funga do Brasil, 2025). The species can be distinguished from other members of the genus by a combination of characteristics, including oblong to elliptical-oblong leaves, non-acute acuminate apices, acute leaf bases, discolored leaf surfaces, closed venation with the formation of areoles, exfoliating to puberulous branches, intrapetiolar stipules, and paired extrafloral glands. These traits may reduce misidentification in forest inventories, particularly in areas with high floristic similarity among sympatric species.

The organoleptic properties and macroscopic qualitative features described here are consistent with previous reports for Q. dinizii (Lopes et al., 1983; Reis et al., 2014; Queiroz et al., 2020; Silva et al., 2021; SFB, 2024). This agreement supports the use of these macroscopic descriptors in timber inspection routines and anatomical screening.

The presence of well-defined growth rings delimited by darker tangential fibrous zones confirms previous descriptions for Q. dinizii (Lopes et al., 1983; Reis et al., 2014; Queiroz et al., 2020; Silva et al., 2021). This feature indicates seasonal cambial activity and supports the interpretation of annual growth periodicity. The low frequency of false rings observed may be associated with irregular cambial stimulation and discontinuous ring formation, which tend to occur more frequently in specific stem portions (Fritts, 1971; Husch et al., 2003). Although many tropical species do not form regular chronological growth rings, the ring structure observed in Q. dinizii indicates a defined growth pattern that may support future ecological and dendrochronological studies (Boninsegna et al., 2009).

Diffuse porosity and the predominance of solitary vessels, followed by multiples of two, are common traits in Amazonian hardwoods (Higuchi et al., 2019). The vessels observed in this study presented alternate intervessel pits, radiovascular pits, and occasional tyloses, which is consistent with previous anatomical reports (Moya and Brea, 2015; Moya, 2018; Silva et al., 2021). Tyloses may contribute to vessel occlusion and may limit fungal spread through conductive tissues. In addition, their presence may affect permeability and influence preservative treatment efficiency, since vessel blockage can reduce fluid penetration (Coradin and Camargos, 2002). In Amazonian trees, tyloses contribute to partial sapwood deactivation by reducing water and nutrient transport in the active xylem. Their distribution may be heterogeneous along the stem (Bamber, 1976; Kitin et al., 2010). Therefore, tyloses may reflect physiological processes associated with heartwood formation and defense mechanisms.

Axial parenchyma in Q. dinizii was predominantly paratracheal, arranged in bands more than three cells thick, and more evident toward the pith–heartwood region. In some areas, it appeared as tangentially confluent aliform parenchyma or as diffuse apotracheal parenchyma closer to the sapwood–bark interface, as previously reported (Lopes et al., 1983; Moya and Brea, 2015; Moya, 2018). The absence of included phloem corroborates previous studies (Reis et al., 2014) and is consistent with the distribution of this trait within Vochysiaceae, in which included phloem occurs mainly in Erisma and Erismadelphus (Kawasaki, 2007).

Mineral inclusions were observed, including prismatic crystals arranged in chains within axial parenchyma, which agrees with reports for Qualea species (Quirk, 1980) and for Q. dinizii (Moya and Brea, 2015; Moya, 2018; Silva et al., 2021). Druses were also identified, as well as isolated prismatic crystals in ray cells. Although mineral inclusions are not always treated as diagnostic characters, they are frequently reported in anatomical studies and may support species differentiation and wood utilization assessments (Vasconcellos et al., 1995).

The occurrence of macules represents one of the main findings of this study. Macules were observed as light-colored tangential stains in the cross section, characterized by a very low frequency of vessels and axial parenchyma. To our knowledge, this is the first report describing these structures in Q. dinizii. This observation expands the anatomical information available for the species and may support future comparative studies. In addition, traumatic axial canals were identified as dark, irregular tangential structures. Traumatic axial canals are described as characteristic of Qualea species (Quirk, 1980; Alves et al., 2025) and have been previously reported for Q. dinizii (Lopes et al., 1983; Reis et al., 2014). Their presence is generally associated with wound responses caused by mechanical injury or biological attack. The occurrence of macules and traumatic canals in the same specimens suggests that both structures may be linked to localized disturbances during wood formation. Morris et al. (2016) reported that xylem parenchyma consists of living cells involved in defense and repair and that these cells may divide and differentiate in response to injury. In this context, the reduced vessel frequency observed in macule regions may indicate modifications in xylem differentiation associated with stress responses.

Fiber morphology in macule regions showed alterations in cell wall structure when compared with unaffected wood. In normal regions, fibers presented characteristics typical of the species, including libriform fibers with simple, nonseptate pits, as described by Silva et al. (2021). The abnormal structure observed in macule zones suggests changes in wood formation, possibly associated with stress or pathogen activity. These regions may introduce heterogeneity in wood structure, which may influence mechanical performance and processing depending on the frequency and extent of macule formation.

The biometric parameters obtained in this study are consistent with those reported for Q. dinizii (Lopes et al., 1983; Moya and Brea, 2015; Moya, 2018; Silva et al., 2021). Vessel distribution patterns may reflect anatomical plasticity, since wood structure can vary according to environmental conditions and adaptive responses (Silva and Melo Júnior, 2017; Silva et al., 2021). In the younger tree (A1, 79 years old), vessel frequency was higher near the pith. A similar pattern was observed in the older tree (A0, 144 years old), with higher vessel concentration near the pith, a reduction in the intermediate zone, and increase toward the bark. Similar radial variation has been described in other tropical species (Chagas et al., 2007; Zanon et al., 2009; Lima et al., 2011; Longui et al., 2012).

In contrast, vessel diameter increased toward the bark, which has also been reported for other native tropical species, in which vessel size is smaller near the pith and increases radially until stabilization in mature wood (Lobão et al., 2012; Portal Cahuana, 2017). Despite this radial variation, Q. dinizii wood can be characterized by few vessels with small to medium diameters and by rays that are few, thin, and low. These traits appear consistent across the analyzed samples and remain useful for anatomical identification.

Future studies should increase sampling to evaluate intra-specific anatomical variation and determine whether macules are frequent in Q. dinizii or associated with site-specific stress conditions. Additional investigation of the relationship between macules, traumatic canals, and potential biotic or abiotic stress factors may clarify their origin and implications for wood quality. Studies addressing durability, mechanical properties, and processing behavior may also support the technological use of Q. dinizii.

5. Conclusions

This study provided an integrated morphological and anatomical characterization of Qualea dinizii Ducke, generating reference information to support reliable species identification in the Brazilian Amazon. The species can be recognized by its smooth and horizontally striated rhytidome with exfoliating bark, buttressed trunk base, and oblong to elliptical-oblong leaves with discolored surfaces, intrapetiolar stipules, and paired extrafloral glands.

The wood anatomy is characterized by distinct growth rings delimited by darker fibrous zones, banded axial parenchyma (sometimes confluent and aliform), and traumatic axial canals. A major original contribution of this study was the detailed documentation of macules, described as tangential light-colored zones with reduced vessel and axial parenchyma frequency.

By compiling external morphology, macroscopic and microscopic wood features, and biometric parameters into a single document, this work strengthens the technical basis for Qualea dinizii identification and reduces the risk of misidentification in forest inventories, thereby supporting future ecological, taxonomic, and technological studies in the Amazonian Forest.

Acknowledgements

The authors would like to thank the Federal University of Mato Grosso – UFMT, the Forest Management Laboratory – FENF/UFMT, the Wood Technology Laboratory – FENF/UFMT, the Scarabeoidology Laboratory – IB/UFMT and the Wood Identification Laboratory – INDEA – MT.

Data Availability Statement

The research data generated and analyzed that support the findings of this study have been published in the manuscript itself and are available from the corresponding author upon reasonable request.

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Edited by

• Editor:
  Takako Matsumura Tundis

Publication Dates

• Publication in this collection
  03 Aug 2026
• Date of issue
  2026

History

• Received
  01 Feb 2026
• Accepted
  21 May 2026