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Article

Leaf Chemistry Patterns in Populations of a Key Lithophyte Tree Species in Brazil’s Atlantic Forest Inselbergs

by
Roberto Antônio da Costa Jerônimo Júnior
1,*,
Ranieri Ribeiro Paula
2,*,
Talitha Mayumi Francisco
3,
Dayvid Rodrigues Couto
3,
João Mário Comper Covre
1 and
Dora Maria Villela
1
1
Programa de Pós-Graduacão em Ecologia e Recursos Naturais, Laboratório de Ciências Ambientais, Universidade Estadual do Norte Fluminense Darcy Ribeiro (UENF), Av. Alberto Lamego, 2000, Campos dos Goytacazes 28013-602, RJ, Brazil
2
Carbone Boréal, Département des Sciences Fondamentales, Université du Québec à Chicoutimi, 555 Boulevard de l’Université, Chicoutimi, QC G7H 2B1, Canada
3
Instituto Nacional da Mata Atlântica (INMA), Av. José Ruschi, 4, Santa Teresa 29650-000, ES, Brazil
*
Authors to whom correspondence should be addressed.
Forests 2025, 16(7), 1186; https://doi.org/10.3390/f16071186
Submission received: 31 May 2025 / Revised: 15 July 2025 / Accepted: 16 July 2025 / Published: 18 July 2025
(This article belongs to the Section Forest Ecophysiology and Biology)
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Abstract

Inselbergs are rocky outcrops with specialized vegetation, including woody species growing in poorly developed soils. We investigated whether populations of the lithophytic tree Pseudobombax petropolitanum A. Robyns (Malvaceae), a key species endemic to Atlantic Forest inselbergs, have convergent or divergent patterns of functional traits related to leaf chemistry. This study was carried out on three inselbergs located in southeastern Brazil. Green and senescent leaves from nine healthy trees and soil samples were collected in each inselberg. The carbon, nitrogen, phosphorus, potassium, calcium, and magnesium concentrations, and the natural abundances of δ13C and δ15N, were measured in leaves and soil, and the C/N, C/P, and N/P ratios were calculated. The specific leaf area (SLA) was measured, and the nutrient retranslocation rate between green and senescent leaves was estimated. Divergences between populations were observed in the concentrations of potassium and magnesium in the green and senescent leaves, as well as in the C/P and N/P ratios in senescent leaves. Our results suggest that nutrient and water dynamics may differ in some inselbergs due to specific nutrients or their relationships, even though there were convergences in most functional traits related to leaf chemistry among the Pseudobombax populations. The divergences among the populations could have important implications for species selection in the ecological restoration context.

Graphical Abstract

1. Introduction

Inselbergs are rocky outcrops consisting mainly of granite and/or gneiss, characterized by large expanses of exposed rock, which are colonized by diverse plant groups, ranging from small herbs, palms, and shrubs to large trees, forming different lithophytic plant communities [1,2,3,4]. These plant communities are home to specialized flora adapted to the restrictive environmental conditions of rocky outcrops, involving high temperatures, water and nutrient restrictions, and full exposure to sun and wind. They also function as ecological refugees for species typical of the surrounding vegetation [4,5]. In the tropics, inselbergs harbor the richest expression of lithophytic vascular flora, with southeastern Brazil standing out as a region of exceptional diversity and as one of the global hotspots for plant species associated with these ecosystems [6]. In the Brazilian Atlantic Forest, inselberg vegetation has been studied in the various types of habitats reported for these ecosystems, including monocot mats, shallow depressions, and vertical rock walls [2,3,4,6]. The inselberg woody vegetation is strongly influenced by the surrounding forest core [4,7,8,9] and plays a vital role in conservation, as it supports both threatened and endemic species, as well as key groups of Neotropical flora, such as vascular epiphytes, which find structural support and protection in the lithophytic trees of these habitats [8,9,10,11]. In this context, the lithophytic tree Pseudobombax petropolitanum A. Robyns (Malvaceae) (henceforth Pseudobombax) stands out, being endemic to the inselbergs of the Atlantic Forest (southeastern Brazil) and considered endangered (EN). This species exhibits high density, dominance, and frequency within this habitat type and is among the most ecologically important species in the woody community based on its importance value index [4].
The woody community growing on the inselbergs of the Atlantic Forest faces different constraints at the soil and microclimate level. These limitations may impose physiological and morphological adaptations on species and populations [12]. This is because the shallow and poorly developed soils, predominantly being sandy and nutrient-poor, are associated with high temperatures in warmer regions of the Atlantic Forest. Overheating is expected in these rocky environments, resulting in high evapotranspiration rates and limiting conditions for most species initially present in the forests [6,12,13,14]. Assuming that inselbergs behave like terrestrial islands in the Atlantic Forest, the populations of endemic species could develop different functional traits due to the influence of factors such as soil chemistry, slope orientation, and microclimate. Whether these distinct groups are becoming more similar (i.e., converging) or different (i.e., diverging) in key functional traits related to leaf chemistry is a relevant question for the ecology and the restoration of inselbergs in the Atlantic Forest. The geographic isolation of inselbergs, associated with the different environmental pressures that may exist in each environment, can lead to the formation of new species [15]. In addition, marked divergences in functional traits could suggest local adaptations, implying the selection of propagules from populations more adapted to each environment in the ecological restoration context. The use of local propagules is recommended to enhance the long-term success of ecological restoration [16].
It is expected that plants adapted to inselbergs will exhibit conservative strategies, characterized by low growth and resource processing rates, greater resource allocation in persistent tissues, as well as enhanced environmental resistance [17,18]. The abundance and dominance of a species such as P. petropolitanum in different inselbergs [4] raises questions about the functional traits related to nutrition and water use. P. petropolitanum is a species that reaches a height of about 15 m and a diameter at breast height (1.3 m) of 117 cm in the Atlantic Forest inselbergs [4]. There is very little known about the functional traits of this species. The species exhibits remarkable phenotypic plasticity in the shape of its leaflets and the size of its petiolules [19], displaying deciduousness during the dry season. To our knowledge, the species has an unknown wood density, but its genus, Pseudobombax, comprises 20 species, and the known members have a low wood density (0.266 g cm−3) [20]. This value of wood density from the genus Pseudobombax may indicate an acquisitive strategy, associated with a fast growth rate and a higher resource processing rate, but lower resistance to environmental stresses [18,21,22]. Thus, the success of a species like P. petropolitanum on inselbergs remains poorly understood.
Depending on environmental conditions, individuals and populations of the same species can adopt different strategies for nutrient absorption and reabsorption [23] and water use regulation [24]. The retranslocation of nutrients, such as nitrogen, phosphorus, potassium, and magnesium, is an important strategy, allowing 60 to 85% of the total content of absorbed nutrients to be conserved, especially when the nutrient availability in the soil is low [25]. The natural abundance of 15N (δ15N) is related to nitrogen sources and preferred acquisition strategies [26]. For example, more significant 15N values in the plant tissues may indicate predominant nitrate uptake or strong ammonia volatilization. In contrast, low 15N values may indicate a preference for ammonium uptake or the occurrence of atmospheric N2 fixation by bacteria in compatible plants [27]. The mycorrhizal association and the diversity of mycorrhizal species can influence the 15N values in plant tissues [27,28]. The C/N, C/P, and N/P ratios are widely used to infer ecological processes, such as the ease of decomposition by microorganisms, imbalance, and nutritional limitation [29,30]. Finally, the natural abundance of 13C (δ13C) serves as a functional indicator of the amount of water used per carbon gain, referred to as the intrinsic water-use efficiency [31,32,33]. Differences in δ13C between individuals and species have been linked to variations in photosynthetic activity, soil water availability, and the growth strategies of plants [22,31,32,33,34]. Plants that grow under water restrictions tend to keep their stomata closed for more extended periods than those in humid environments, leading to differences in the isotopic discrimination of 13C [35]. There is a preference for the lighter isotope 12C over the heavier 13C during CO2 assimilation in photosynthesis [31,34].
This study assessed functional traits related to leaf chemistry in green and senescent leaves of a key species in the inselberg flora, P. petropolitanum (hereafter referred to as Pseudobombax). Our objective was to investigate whether populations of Pseudobombax show convergent or divergent patterns for carbon (C), nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), and magnesium (Mg), as well as for the stoichiometric relationships of C/N, C/P, and N/P; the natural abundance of 13C and 15N; and the rates of nutrient retranslocation. We also showed for the first time the soil chemistry patterns of these environments and the specific leaf area (SLA) of the Pseudobombax trees. This information was related to the chemistry of the leaves. We hypothesize that populations of Pseudobombax exhibit divergent patterns of leaf chemistry due to dissimilarity in the environmental conditions across the studied inselbergs.

2. Materials and Methods

2.1. Study Area

The investigation was conducted on three inselbergs south of Espírito Santo, southeastern Brazil (Figure 1). Small fragments of Seasonal Semideciduous forests surround these inselbergs [36], as well as small rural properties with activities ranging from cattle raising and citrus and coffee plantations to commercial forests [4,9,37]. The woody vegetation studied comprises more than 80% of the deciduous species during the dry season, characterized by an open canopy with dispersed individuals and short stature [4]. The predominant climate is the Aw type, according to the Köppen classification, with a dry period from May to October, where the minimum rainfall is generally less than 50 mm per month, and the rainy period from November to April, with an average monthly rainfall of 180 mm, usually accumulated between November, December, and January [10,38]. However, the measurements from the automatic meteorological station located near “Pedra Três Irmãs” inselberg indicated that the total annual rainfall was 1105 mm, and the yearly average temperature was 26 °C in the sampling year (2023/2024). This value exceeds the region’s average temperature, which is 23 °C, by 3 °C, according to [38], indicating warming in the study region. During the execution of the experiment (2023/2024), the average annual minimum temperature was 20 °C, reaching 15 °C in the coldest month (June). The average maximum temperature was 34 °C, reaching values above 40 °C during the rainy season.
The “Pedra da Aliança” inselberg (PA) is located in the municipality of Muqui (20°54′05″ S and 41°21′15″ W), also known locally as “Serra da Aliança,” with an altitude ranging from 200 to 450 m. The Pseudobombax is the most dominant and abundant species [4,9]. The orientation of the sampled slope is west. The “Pedra Três Irmãs” inselberg (TI) is located in the municipality of Jerônimo Monteiro (20°46′19″ S and 41°21′12″ W), within the limits of the “Monumento Natural do Maciço das Andorinhas”, a municipal protected area. It is a rocky complex with an altitude ranging from 120 m to 600 m, where Pseudobombax is the most abundant species [37], and the sampled slope orientation is north. The “Santa Angélica” inselberg (SA) is in the municipality of Alegre (20°43′54″ S and 41°25′58″ W), with an altitude ranging from 200 m to 420 m, where Pseudobombax is the second most abundant species [4], and the orientation of the sampled slope is south.

2.2. Characteristics of the Soil and the Species Studied

The predominant soil in the Atlantic Forest’s inselbergs is the Humic Litholic Neosol [10]. There is no information about soil chemistry in the Atlantic Forest’s inselbergs [4]. Based on a laboratory analysis of soil samples from [39] the TI, PA, and SA inselbergs (Jerônimo Júnior et al., in preparation), we identified distinct patterns in certain surface soil chemistry variables across the inselbergs (Table 1).
The lithophytic tree Pseudobombax is caespitose, with extensive branching and a known height that varies in the southern region of Espírito Santo from 2.5 to 15.3 m [10]. A notable morphological characteristic of this species is its large, superficial roots, which are fixed to exposed rocks, forming natural physical barriers that retain organic debris, rock fragments, and soil [4]. These accumulations are essential not only for the nutrition of the tree but also for the nutrition of the plant communities of the inselbergs [4]. Furthermore, Pseudobombax contributes substantially to aboveground biomass and carbon storage [4], increases the fertility of rocky substrates by recycling nutrients from its biomass [40], and provides critical structural support for the establishment of a diverse vascular epiphyte community [8,11]. Based on the five leaves sampled from each individual of Pseudobombax in each inselberg for this study (see details below), we found that the SLA values in green leaves (Table 2) were situated at the upper end of the SLA spectrum in plant communities [18,41]. In addition, there was a difference between the populations’ SLA values, mainly an apparently higher SLA from individuals of Pseudobombax growing in the SA inselberg compared to TI and PA (Table 2).

2.3. Tree Selection and Leaf Collection

Nine adult individuals of Pseudobombax were chosen for the evaluation of each inselberg, maintaining a minimum distance of 30 m between individuals. Previous studies conducted in two study areas (PA and SA) revealed that the height of Pseudobombax populations ranged between 2.0 and 7.5 m [4]. In addition, there are at least 190 to 270 individuals per hectare in the PA and SA inselbergs [4]. We assumed that the nine individuals of Pseudobombax sampled on each inselberg may have represented the population [42]. We chose nine individuals due to the sparse distribution of vegetation, and nine individuals represented a balance between logistical viability (access and locomotion with security) and ecological representativeness. We systematically collected three individuals from the highest canopy stratum, three from the middle stratum, and three from the lowest stratum in each inselberg. The separation of the height stratum was performed visually.
Pseudobombax has compound leaves; therefore, each leaflet was used as a leaf unit. Senescent leaves were collected directly from each tree’s branches at the peak of the dry season and during the natural senescence of the species (in August 2023). The senescent leaf was distinguished by the yellow/greenish petiole and the brownish color, in addition to the easy detachment of the branch. The green leaves were collected from the same trees at the peak of the rainy season (January 2024). Due to the large leaf size (Table 1), we sampled nine green and nine senescent leaflets per plant. These leaves were distributed among the canopy’s lower, middle, and upper thirds. We collected leaves in opposite directions, based on the cardinal points, as much as possible. Five of the nine leaves sampled were measured in a leaf area analyzer (LI-3100-LICOR, Inc., Lincoln, NE, USA), air dried (65 °C), and weighed (Table 1). After this, the leaves were dried at 65 °C to a constant weight, ground in a Willey mill (2.0 mm mesh), and one subsample was stored. The equipment was cleaned between each milling using a compressed air jet.

2.4. Chemical and Isotopic Analyses of the Leaves

The concentrations of P, K, Ca, and Mg were determined by inductively coupled plasma emission spectrophotometry (ICP/OES, Varian Australia Pty LTD, Macquarie Park, New South Wales, Australia). For this, 0.2 g of plant material was digested in a solution of concentrated sulfuric acid (175 mL), hydrogen peroxide (100 volumes, 210 mL), selenium (0.21 g), and lithium sulfate (7 g) [43]. Standard apple, replica, and white samples were added to the digester blocks. The percentage of nutrient recovery was calculated through the acid digestion of the international standard of apple leaves [44].
The total C and N concentration and the natural abundance of 13C (δ‰) and 15N (δ‰) were determined in a Thermo Finnigan Delta V Advantage mass spectrometer (IRMS, Thermo Scientific, Milan, Italy) coupled to a Flash 2000 organic elemental analyzer (Thermo Scientific, Milan, Italy). Approximately 2 mg of leaf material was used in each C/13C and N/15N analysis. Pee Dee Belemnite (PDB) and atmospheric nitrogen were the standards for δ13C and δ15N, respectively.

2.5. Nutritional Relationships and Nutrient Retranslocation

C, N, and P concentrations were kept in the same unit (g kg−1) to calculate the stoichiometric relationships between C/N, C/P, and N/P in green and senescent leaves. The retranslocation of N, P, K, and Mg between green and senescent leaves was calculated according to the literature [45]. We assume that nutrient concentrations in senescent leaves at the peak of the dry season represent the inferior limit of the annual leaf nutrient status, as plants perform maximum nutrient retranslocation before leaf senescence occurs in the dry season. In addition, we assume that the nutrient concentrations in the green leaves sampled at the peak of the rainy season represent the maximum annual concentration in the green leaves. Thus, the rate of nutrient retranslocation from senescent leaves to green leaves could indicate the magnitude of nutrient retranslocation throughout the year for this species.

2.6. Statistical Analysis

Principal component analysis (PCA) was employed to examine the relationship between SLA and soil and leaf chemistry, aiming to gain insights into the convergence or divergence patterns of inselbergs. For this, we used the nine soil data points to match the similar number of tree samples on each inselberg. The nine soil data points selected per inselberg were the closest to the trees with sampled leaves. In addition, the effect of the inselbergs on the PCA scores was tested using a linear mixed model (LMM), with the inselbergs as a fixed factor (n = 3) and the repetitions as a random factor (n = 9). Moreover, the effect of the inselbergs on all chemistry variables (i.e., C, N, P, K, Ca, Mg, C/N, C/P, N/P, δ15N, and δ13C, and the nutrient retranslocation rates) in each leaf type (green and senescent) was tested with LMM using the three inselbergs as a fixed factor and the nine individuals as a random factor. Tukey’s Honestly Significant Difference (HSD) post hoc test was used for mean comparisons (p ≤ 0.05). Finally, we used the Spearman rank correlation to evaluate the relationship between soil and leaf chemistry at the local and ecosystem levels. The LMM analyses were performed using JMP® software, version 17 (SAS Institute Inc., Cary, NC, USA 1989–2023). The PCA and the Spearman rank correlation were performed using the R software, version 2024.12.1 (R Core Team, Vienna, Austria, 2024).

3. Results

3.1. SLA, Soil, and Leaf Chemistry Relations

Some tree and soil variables were primarily responsible for the divergences between inselbergs, as highlighted in the first principal component (PC1) (Figure 2A). PC1 and the second principal component (PC2) explained 42% of the data variability. However, only the PC1 score values differed significantly among the three inselbergs (p < 0.0001), whereas no significant differences were observed for the PC2 score (p = 0.4963). PA and SA were the most divergent, with lower and higher PC1 score values. TI had intermediate PC1 score values and was not significantly different from SA, though was significantly different from PA, as indicated by the HSD test. The SLA of green leaves was a notable factor affecting PC1, with a value for squared cosines higher than 0.8 (Figure 2B). In addition, N, K, C, and P stocks in the soil and the C, K, and Ca concentrations in the green leaves were the variables with the highest effect on PC1, as indicated by squared cosine values greater than 0.6. The Mg concentration in green and senescent leaves, the SLA, and the C and N concentrations in the senescent leaves, as well as the 13C in the soil were the other variables with squared cosines higher than 0.4. The Mg and Ca stocks in the soil, the 13C and K concentrations in senescent leaves, and the P concentrations in green leaves showed a modest influence on PC1, with squared cosines higher than 0.3.
A positive and significant correlation was found between soil and green leaves for P, particularly in the TI and SA inselbergs (Table A1, Appendix A). When data from all inselbergs were combined, a positive and significant correlation was observed between soil and green leaves only for K. No significant correlations were detected with the other nutrients.

3.2. Carbon and Nutrients in Leaves

Only the K (Figure 3B) and Mg (Figure 3F) concentrations in both green and senescent leaves were statistically different between inselbergs. K was significantly lower in the green (8.1 g kg−1) and senescent (7.4 g kg−1) leaves of individuals growing in inselberg PA compared to those growing in TI and SA, which ranged from 11.7 to 13.6 g kg−1 and 9.0 to 9.2 g kg−1, respectively. Individuals from PA and TI also showed significantly lower Mg concentrations in green (2.63 and 3.10 g kg−1) and senescent leaves (1.98 and 2.50 g kg−1) than in inselberg SA, which had concentrations of 4.14 and 3.49 g kg−1, respectively. However, the C (Figure 3A), N (Figure 3C), P (Figure 3E), and Ca (Figure 3D) levels in both leaf types were statistically similar across the Pseudobombax populations in the inselbergs. The minimum and maximum concentrations in the leaves ranged from 424.1 to 477.1 g kg−1 for C, from 6.0 to 24.5 g kg−1 for N, from 0.2 to 1.4 g kg−1 for P, from 5.4 to 24.2 g kg−1 for K, from 3.7 to 20.9 g kg−1 for Ca, and from 1.0 to 5.1 g kg−1 for Mg, respectively.

3.3. Stoichiometric Relationships

The C/N ratio in green and senescent leaves was statistically equal among the inselbergs, with mean values of 61.2 and 24.9, respectively (Figure 4A). The C/P ratio in senescent leaves was statistically different between inselbergs, but not for the green leaves (Figure 4B). In the senescent leaves, the C/P ratio was significantly higher in the inselberg TI (2150.5) than in the inselberg PA (1387.1), while in the inselberg SA, the plants presented intermediate values (1676.7). The N/P ratio was also significantly different in the senescent leaves (Figure 4C). Lower N/P ratio values were found in the inselberg PA (22.3) compared to SA (29.2) and TI (32.7).

3.4. Natural Abundance of δ13C and δ15N

The values of the natural abundance of δ13C (‰) and δ15N (‰) in green and senescent leaves were not statistically different between populations of Pseudobombax (Figure 5). The minimum and maximum values of δ13C in the green and senescent leaves were −30.1 and −26.2‰, while they ranged from 0.6 to 5.3‰ for δ15N. The mean values in the green and senescent leaves between the populations of Pseudobombax were −28.1 and −28.5‰ for δ13C (Figure 5A) and 3.2 and 2.3‰ for δ15N (Figure 5B), respectively.

3.5. Nutrient Retranslocation Rate

No significant differences were found in the retranslocation rate of N, P, K, and Mg among Pseudobombax populations (Figure 6). The nutrients with the highest translocation rates were P (68 to 76%) and N (53 to 61%). The retranslocation rate ranged from 8% to 32% for K and from 15% to 24% for Mg.

4. Discussion

Our general hypothesis was that the populations of Pseudobombax petropolitanum A. Robyns (Malvaceae) would present divergent patterns of leaf chemistry due to the dissimilarity in the environmental conditions among the studied inselbergs. The inselbergs are separated by at least 10 km and show significant differences in several soil chemistry factors, notably the soil C, N, P, K, δ13C, and Mg (Table 1, Figure 2), and the tree SLA (Table 2, Figure 2). From a soil perspective, the SA and PA inselbergs were the most divergent. For example, the soil in the SA inselberg exhibited higher stocks of C, N, and P, whereas the soil in the PA inselberg exhibited lower stocks of C and K, besides the tendency towards a lower stock of Mg (Table 1). In addition, the natural abundance of 13C was highest in the soil of PA (−25.3‰) and lowest in the soils of SA (−27.6‰). Furthermore, Pseudobombax growing in the inselbergs SA and PA presented higher and lower SLA values. High SLA is associated with better resource availability (e.g., solar radiation, soil water, and nutrients) for plant growth; conversely, lower SLA values are expected to have the opposite effect [18,41]. The soil in the SA inselberg, with higher amounts of C, N, and P than the soil from the PA inselberg, may explain the divergences in SLA. In addition, based on our field observations, the higher SLA values in the SA inselberg are likely associated with humidity, given the southward orientation of the sampled slope. Both the soil and leaves of Pseudobombax populations showed lower δ13C values in the SA than in the PA inselberg, suggesting a potentially greater water availability for populations in SA than for those in PA. All this could suggest that the environmental conditions are less severe in the SA than the PA inselberg, with TI assuming intermediate conditions. It is recognized that environmental factors, such as water and nutrient availability, have a greater influence on the formation of leaf morphofunctional characteristics, being more important than genetic factors [46]. Therefore, further studies could investigate the effect of solar radiation on SLA in our study sites, as the difference in exposure faces among the inselbergs may impact SLA [41]. In addition, the causes of the soil differences in these environments need to be investigated in more detail, as they could be related, for example, to the geology of the inselbergs, the vegetation composition, and nutrient cycling. These soils are acidic (pH < 5), and the amounts of nutrients are limiting for the development of cultivated plants, as is the case in most soils in the state of Espírito Santo [47].
The populations of Pseudobombax in the three inselbergs studied diverged in terms of the K and Mg concentration in the leaves, as well as in the C/P and N/P stoichiometry of the senescent leaves, corroborating the hypothesis for these variables. The divergent patterns in SLA, K, and Mg imply that populations of Pseudobombax are developing distinct characteristics, which could be due to local adaptation to different environmental conditions (e.g., soil properties, water availability, solar radiation), genetic drift, or different selective pressures acting on each population. However, the three Pseudobombax populations exhibited no significant divergences in most leaf chemistry variables, as determined by our one-year exploratory assessment. Thus, our results suggest convergence in most leaf chemistry variables investigated among Pseudobombax populations, despite geographic isolation or variations in local conditions. Several factors may explain these patterns, including species-specific physiological constraints that limit variation, similar selective pressures across different sites related to nutrient acquisition and water use, and phenotypic plasticity resulting in similar outcomes under certain conditions. Nevertheless, we observed patterns at a specific point in time, and although these patterns suggested convergence or divergence, definitively proving the process often requires field data associated with genetic evaluations.
A variety of factors can influence the nutrient levels present in leaves, including edaphic conditions, floristic composition, nutrient absorption, utilization, and the retranslocation capacity of each species, as well as tree age [48,49,50]. The soil nutrient levels in each inselberg could explain part of the differences in leaf chemistry between populations, particularly in terms of K and Mg. However, when we examined the relationship between elements in soil and leaves within populations, we found that most of them were non-significant (Table A1). The lack of a significant correlation for most nutrients within the populations could be explained by various factors, in addition to those mentioned above. For example, the uptake of elements from sources other than soil, such as stemflow and throughfall, and associations with mycorrhizae. Microbial activity in the soil and at the rhizosphere level also plays a crucial role in nutrient availability and can differentially function in species with acquisitive versus conservative strategies [51]. In this first study, we consider that, at the global level, the factors affecting K and Mg concentrations in leaves, which explain the differences between populations, are the amount of K and Mg in the soil, as well as the SLA and microclimate conditions. In inselbergs, we can find different types of habitats with growing sites that are climatically and edaphically dry, supporting highly specialized vegetation [52], which in turn influences access to water and nutrients. It is well established that geographic factors (e.g., latitude and altitude) and environmental pressures (e.g., water stress, solar radiation, and microclimatic variations, including temperature and precipitation) significantly influence the nutrient dynamics and allocation of nutrients in plants [53]. Thus, several factors may affect foliar nutrient levels, and the specific processes interfering with each population need to be investigated in further studies. The K amounts in soils are related to water availability, and K uptake is more related to water uptake than N and P [54]. Differences in K concentrations in leaves are related to both plant responses to drought and plant growth, playing a key role, along with N and P, in plant responses to climate gradients to improve their growth capacities and adaptation to water stress along environmental gradients and over time (seasons) [54]. In addition, K concentration can explain trees’ long-term adaptation to different ecological lifestyles [54] and could likely explain the significant relationship between soil and leaf K across the inselbergs (Table A1). Therefore, the lower K concentrations in the leaves of the Pseudobombax population growing on the inselberg PA compared to others may be due to the environment presenting a lower K availability and greater water restrictions than the others, as well as other factors. The leaf Mg concentration was higher in the population of Pseudobombax growing on the SA inselberg compared to those of the PA and TI inselbergs. The greater soil Mg and water availability may result in a higher number of chloroplasts in the leaves of the SA population, as Mg is a central component of chlorophyll [55]. In addition, the structural characteristics of the leaves can also influence this variation, as plants subjected to different light, water, and nutrient availability generally develop a different SLA, chloroplast density, and concentration of chlorophyll molecules per unit area [41,56]. The higher levels of K and Mg, in addition to Ca, are associated with the better ability of plants to regulate water use [57]. Ca levels were statistically similar between soils and in leaves. However, the tendency for lower Ca concentrations in leaves of the population growing in SA suggests a nutritional imbalance, which may be caused by the K and Mg stocks in the soil of the SA inselberg. Despite the better plant growth conditions in the SA inselbergs, as indicated by the higher SLA values, the species Pseudobombax is not the most abundant in this ecosystem [4]. Future studies could assess other functional traits, such as growth rates in terms of diameter and height, in addition to the phenology of Pseudobombax in each inselberg, to determine whether these differences depend on the environmental conditions within each inselberg. This could provide new insights into the adaptation and survival strategies of species with low wood density in harsh environments such as inselbergs.
The differences in stoichiometric relations were significant only for senescent leaves. These relations between C, N, and P could play an important role in each inselberg. They relate to decomposition and nutrient release because soil microorganisms tend to be limited by C and co-limited by other elements, such as N and P, needing to decompose sources of C [58,59]. The C/N ratio observed in the senescent leaves of Pseudobombax populations may indicate greater recalcitrance, resulting in lower nitrogen availability. A C/N ratio higher than 30 indicates that immobilization is higher than mineralization [60,61]. Plant residues with C/P ratios greater than 300 tend to favor microbial P immobilization [62], decreasing the P availability in the soil [59]. In the present study, the values of the C/P ratio in senescent leaves were higher in TI (2150.5), intermediate in SA (1676.7), and lower in PA (1387.1), indicating that the Pseudobombax leaf material has a strong tendency to microbially immobilize P in all inselbergs. The observed patterns for the C/P ratio in senescent leaves appear to follow the soil C/P ratio, particularly regarding the TI inselberg (Figure 4; Table 1). These patterns may suggest a greater influence of senescent leaves from Pseudobombax populations on the soil in the TI inselberg. In addition, the higher C/P ratio in both senescent leaves and the soil in the TI inselberg may suggest adaptations that improve the retranslocation rate of P in the Pseudobombax population, explaining the tendency for higher P retranslocation in Pseudobombax populations from TI compared to SA or PA. Plants adapted to poor soils generally exhibit a higher C/P ratio because they have evolved to be more efficient in utilizing phosphorus and other essential nutrients [48,63,64]. The N/P ratio followed a similar trend to the C/P ratio, particularly in terms of the N/P values in the senescent leaves and soil from the TI inselberg. Both the C/P and N/P ratios may help in understanding the adaptation of Pseudobombax populations to different inselbergs, especially in more phosphorus-poor soils. Thus, the higher C/P and N/P ratios in the TI inselberg could induce changes in the P use efficiency because plants tend to increase their retranslocation efficiency to minimize losses, and these adaptations are crucial for survival in restrictive environments [65,66,67]. Additionally, the role of changes in N/P ratios on vegetation performance and species composition [68] should be investigated in the studied inselbergs, given the differences in structure and diversity between PA and SA inselbergs [4]. The N/P ratio may determine whether a species or ecosystem is limited by P or N. The N/P ratios observed in the present study (PA = 22.3, SA = 29.2, and TI = 32.7) indicate a substantial limitation by phosphorus [48,69,70]. The positive and significant relationship between soil and leaf P in SA and TI is likely indicative of stronger soil–plant feedback in SA and TI than in PA regarding P limitations (Table A1). Further studies should assess these questions using a larger number of trees and a more specific separation of niches for access to nutrients and water within each inselberg.
We used 15N and 13C as proxies for soil N preferences [27,71] and intrinsic water-use efficiency [22,31,32,33,34]. Our results indicate that the populations exhibited convergent patterns in these aspects, with no significant differences. However, some tendencies were observed in the δ15N and δ13C values between tree leaves and soil. The natural abundance of 15N in the soil followed a similar trend to that of 15N in the green leaves. It was lower in the inselberg TI (5.05 and 2.7‰), intermediate in the PA (6.02 and 3.4‰), and higher in the SA (6.59 and 3.5‰). Conversely, the δ13C values suggest that populations have the same water-use strategies. In addition, the leaf δ13C values reflect the same trends observed between soil and leaf. The natural abundance of soil δ13C was higher in PA (−25.09 and −27.96‰), intermediate in TI (−26.92 and −28.12‰), and lower in SA (−27.46 and 28.33‰). These patterns suggest an intricate relationship between Pseudobombax populations and the soil of these environments, implying differences in the forms of nitrogen and water available in the soil and the photosynthetic activity.
We observed convergent patterns of foliar nutrient retranslocation among populations of Pseudobombax growing on inselbergs. Nutrient retranslocation can vary according to species, age, edaphic and climatic conditions, and management practices adopted [72,73]. Nutrients bound to organic compounds (such as N and P) have higher retranslocation rates from senescent to green leaves [74], as observed in the present study for N and P. The retranslocation dynamics of phosphorus (P) in leaves have been shown to exhibit greater variability compared to those of nitrogen (N) [75]. In the present study, the low soil P contents, mainly in the TI inselberg, may explain the general tendency of there being a higher retranslocation rate of leaf P in TI (76%) compared to SA (68%) and PA (66%). The rates of N (53%–61%) and P (68%–76%) retranslocation found here are consistent with values found in other studies, such as 62% for N and 65% for P [76] and 44%–53% for N and 50%–62% for P [77]. Broad-leaved, deciduous, and woody plants reabsorb more P before leaf senescence when compared to coniferous, perennial, leguminous, and herbaceous plants [53]. Low K retranslocation rates were observed in the three populations of Pseudobombax compared to other studies [78]. When we compared the K retranslocation rates of our study (8%–33%) with the global average of 70.1% [76], we inferred that Pseudobombax populations have a low K retranslocation efficiency. This fact can be explained by the high mobility and leaching of this nutrient, which are likely accentuated by the large SLA. Although both Mg and K are considered leachable nutrients, K often correlates with SLA, but Mg does not. This suggests that Mg is regulated differently in the leaf, possibly due to its role in structural components (such as chlorophyll and cell walls) and not just in metabolic processes [79]. In this sense, leaf K leaching could have been more significant in the SA, which also presented the highest SLA values. In addition, the higher retranslocation of K compared to Mg in the inselberg SA has been observed in the opposite direction in inselberg PA, which could suggest an antagonistic relationship, leaving preferences for K or Mg retranslocation in the populations growing in SA and PA, respectively.
We compared three populations of a key species to investigate whether these populations are becoming more similar (converging) or more different (diverging) in their essential functional traits related to leaf chemistry. Our results suggest that there was divergence in the K and Mg leaf concentrations and C/P and N/P ratios in senescent leaves among the inselbergs, although Pseudobombax had a considerable convergence in most leaf chemistry functional traits. The results suggest that the environmental conditions among the three inselbergs studied—namely, soil chemistry, slope direction, and microclimate—were generally not strong enough to drive divergence in these traits. The implications for ecosystem restoration are significant if we suppose no strong divergence exists between populations. However, some divergence in the nutritional leaf traits of Pseudobombax must be considered, particularly in the K and Mg levels in leaves. The convergence of several traits does not preclude the divergence of other traits. Moreover, convergence could also mean that the Pseudobombax tree lacks adaptive plasticity in these traits, potentially making all populations vulnerable if environmental conditions at restoration sites shift beyond the species’ tolerance range, as defined by the conserved trait values [80]. Significant divergences may indicate strong local adaptation or high environmental plasticity, suggesting that populations are specifically adapted to their local conditions (e.g., soil nutrient levels and water availability). Our results suggest that local differences in the soil nutrient stocks and water availability, as well as in tree SLA, primarily cause the observed divergences. When considering population divergence in leaf K, Mg, and SLA, for example, restoration practitioners should prioritize local provenance that is adapted to environmental conditions closely matching the restoration site [81]. Using non-local sources adapted to different nutrient and water regimes could lead to poor survival, growth, or overall fitness. However, some strategies may be employed in cases where local provenances are unavailable. For example, in sites such as PA (low K), it would be preferable to choose provenances with a higher retranslocation efficiency of K, in addition to a lower tree SLA, to reduce K leaching and water stress. For sites such as TI (low P), the provenances with higher retranslocation efficiency for P may be preferred.

5. Conclusions

Our one-year evaluation of Pseudobombax petropolitanum A. Robyns (Malvaceae) populations growing in three inselbergs in the Atlantic Forest, located at least 10 km apart, investigated whether these populations were converging or diverging in several functional traits related to leaf chemistry. Although there were no significant differences among populations in most leaf chemistry traits, with convergence in the N, P, Ca, C:N, δ13C, and δ15N in the green and senescent leaves, and the C/P and N/P in green leaves, besides the retranslocation rate of N, P, K, and Mg, the populations diverged in the concentrations of K and Mg in green and senescent leaves, as well as in the stoichiometric relationships of C/P and N/P in the senescent leaves, corroborating our hypothesis for those variables. The divergences may be explained by differences in the availability of resources and plant functional traits, such as soil chemistry, water availability, and SLA, suggesting that nutrient and water dynamics may differ in some inselbergs, particularly concerning specific elements or processes. The divergences in Pseudobombax populations were related to specific nutrients or their relationships, although there were convergences mostly in functional traits related to leaf chemistry. Furthermore, the divergences among populations could have significant implications for species selection in inselbergs, particularly for conservation and restoration purposes. They should be considered, especially when evaluating the retranslocation efficiency of certain nutrients and the SLA in specific populations, which may be crucial for species survival and adaptation.

Author Contributions

Conceptualization, R.A.d.C.J.J., D.M.V. and R.R.P.; Methodology, R.A.d.C.J.J., D.R.C., R.R.P. and D.M.V.; Formal Analysis, R.R.P. and T.M.F.; Investigation, R.A.d.C.J.J., D.M.V. and J.M.C.C.; Data Curation, R.A.d.C.J.J.; Writing—Original Draft Preparation, R.A.d.C.J.J. and R.R.P.; Writing—Review and Editing, R.A.d.C.J.J., R.R.P., T.M.F., D.R.C., J.M.C.C. and D.M.V.; Supervision, D.M.V. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The data that support the findings of this study are available from the corresponding author or the first author upon reasonable request.

Acknowledgments

The first author thanks the Carlos Chagas Filho Foundation for Research Support of the State of Rio de Janeiro (FAPERJ) for the doctoral scholarship (grants: E-26/200.250/2021). D.M. Villela was supported by the National Council for Scientific and Technological Development (CNPq; 406244/2021-9; 315112/2020-4 scholarship). T.M. Francisco and D.R. Couto thank the CNPq (Programa de Capacitação Institucional—PCI/INMA; grants: TMF #300906/2022-6 and DRC #301141/2022) of the Brazilian Ministry of Science, Technology and Innovation (MCTI) for the research grants provided. We thank Thais Arão Feletti, Rita de Cássia Freire Carvalho, Lucas Batista Vargas, Fábio da Silva Simão “Binho”, Mariana Alves Faitanin, and Caroline Pessanha da Silva for their support during the fieldwork; Henrique Machado Dias, from the Department of Forest Sciences at the Federal University of Espírito Santo (UFES), for his logistical support during the execution of the experiment; José Eduardo Macedo Pezzopane, from the Laboratory of Forest Meteorology and Ecophysiology (UFES, Brazil), for providing the meteorological data; and the Environmental Sciences Laboratory (LCA/UENF) technicians for their support with the analyses.

Conflicts of Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Appendix A

Table A1. Spearman correlation test between soil and leaf chemistry in Pseudobombax populations in each inselberg studied (TI = “Três Irmãs”, PA = “Pedra da Aliança”, and SA = “Santa Angélica”) and among all (All) Atlantic Forest inselbergs studied in southeast Brazil.
Table A1. Spearman correlation test between soil and leaf chemistry in Pseudobombax populations in each inselberg studied (TI = “Três Irmãs”, PA = “Pedra da Aliança”, and SA = “Santa Angélica”) and among all (All) Atlantic Forest inselbergs studied in southeast Brazil.
VariableGreen LeavesSenescent Leaves
TIPASAAllTIPASAAll
C0.520.600.130.06−0.150.100.170.02
N0.20−0.420.570.16−0.120.08−0.020.20
P0.68 *−0.400.82 *0.330.000.13−0.200.07
K0.050.050.320.52 *−0.02−0.42−0.430.22
Ca−0.30−0.22−0.22−0.20−0.13−0.670.500.03
Mg−0.27−0.17−0.270.26−0.50−0.38−0.270.12
Mn−0.05−0.02−0.48−0.080.05−0.230.500.13
δ13C−0.450.27−0.100.100.200.02−0.320.26
δ15N0.25−0.30−0.40−0.050.00−0.150.02−0.01
The values in the table correspond to the Spearman correlation coefficient. The symbol * indicates a significant correlation (p < 0.05), which is either positive or negative.

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Figure 1. Location of the study sites of three Atlantic Forest inselbergs in Espírito Santo State, southeastern Brazil: Três Irmãs (TI); Santa Angélica (SA), and Aliança (PA).
Figure 1. Location of the study sites of three Atlantic Forest inselbergs in Espírito Santo State, southeastern Brazil: Três Irmãs (TI); Santa Angélica (SA), and Aliança (PA).
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Figure 2. Principal component analysis (A) relating chemistry elements in the 0–10 cm soil depth (identified by _S and brown color), the SLA, and the chemistry of the green (identified by _GL and green color) and senescent (identified by _SL and orange color) leaves of Pseudobombax trees growing on three inselbergs in the Atlantic forest: “Três irmãs” (TI, square), “Aliança” (PA, circle), and “Santa Angélica” (SA, triangle); and the values of the squared cosines (B) for each soil and tree variable affecting PC1.
Figure 2. Principal component analysis (A) relating chemistry elements in the 0–10 cm soil depth (identified by _S and brown color), the SLA, and the chemistry of the green (identified by _GL and green color) and senescent (identified by _SL and orange color) leaves of Pseudobombax trees growing on three inselbergs in the Atlantic forest: “Três irmãs” (TI, square), “Aliança” (PA, circle), and “Santa Angélica” (SA, triangle); and the values of the squared cosines (B) for each soil and tree variable affecting PC1.
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Forests 16 01186 g003
Figure 3. Mean values (±SE, n = 9) for C (A), K (B), N (C), Ca (D), P (E) and Mg (F) concentrations in green (dark green) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil. Inselbergs: TI = “Pedra das três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselberg populations for green or senescent leaves separately, and different letters indicate the differences between populations. “ns” indicates that the differences were not significant (p value > 0.05).
Figure 3. Mean values (±SE, n = 9) for C (A), K (B), N (C), Ca (D), P (E) and Mg (F) concentrations in green (dark green) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil. Inselbergs: TI = “Pedra das três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselberg populations for green or senescent leaves separately, and different letters indicate the differences between populations. “ns” indicates that the differences were not significant (p value > 0.05).
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Figure 4. Mean values (±SE, n = 9) for C/N (A), C/P (B), and N/P (C) ratios in green (dark green) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselbergs for green and senescent leaves separately, and different letters indicate the differences between populations. “ns” indicates that the differences were not significant (p value > 0.05).
Figure 4. Mean values (±SE, n = 9) for C/N (A), C/P (B), and N/P (C) ratios in green (dark green) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselbergs for green and senescent leaves separately, and different letters indicate the differences between populations. “ns” indicates that the differences were not significant (p value > 0.05).
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Forests 16 01186 g005
Figure 5. Mean values (±SE, n = 9) of the natural abundances of δ13C (‰) (A) and δ15N (‰) (B) in the green (dark) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil: TI = “Pedra das três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselbergs for green and senescent leaves. “ns” indicates that the differences were not significant (p value > 0.05).
Figure 5. Mean values (±SE, n = 9) of the natural abundances of δ13C (‰) (A) and δ15N (‰) (B) in the green (dark) and senescent (light green) leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) in three Atlantic Forest inselbergs in southeast Brazil: TI = “Pedra das três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences between the inselbergs for green and senescent leaves. “ns” indicates that the differences were not significant (p value > 0.05).
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Figure 6. Mean values (±SE, n = 9) of the retranslocation rate (%) of N, P, K, and Mg among green and senescent leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) trees in three Atlantic Forest inselbergs in southeast Brazil: TI = Três Irmãs”, PA = “Aliança,” and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences. “ns” indicates that the differences were not significant (p value > 0.05).
Figure 6. Mean values (±SE, n = 9) of the retranslocation rate (%) of N, P, K, and Mg among green and senescent leaves of Pseudobombax petropolitanum A. Robyns (Malvaceae) trees in three Atlantic Forest inselbergs in southeast Brazil: TI = Três Irmãs”, PA = “Aliança,” and SA = “Santa Angélica”. p values less than 0.05 indicate significant differences. “ns” indicates that the differences were not significant (p value > 0.05).
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Table 1. p values and the average values (±SE, n = 15) of soil pH, nutrient stock, the natural abundance of 13C and 15N, and the stoichiometric relationships of C/N, C/P, and N/P at the 0–10 cm soil layer in three Atlantic Forest inselbergs in southeastern Brazil. Inselbergs: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. Different letters indicate statistical differences in soil chemistry between inselbergs (p ≤ 0.05) *.
Table 1. p values and the average values (±SE, n = 15) of soil pH, nutrient stock, the natural abundance of 13C and 15N, and the stoichiometric relationships of C/N, C/P, and N/P at the 0–10 cm soil layer in three Atlantic Forest inselbergs in southeastern Brazil. Inselbergs: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. Different letters indicate statistical differences in soil chemistry between inselbergs (p ≤ 0.05) *.
Variablesp ValueTIPASA
pH0.02404.86 ± 0.083 a4.56 ± 0.089 ab4.53 ± 0.074 b
C (kg ha−1)0.000462.65 ± 3.44 b56.47 ± 5.26 b82.69 ± 5.91 a
N (kg ha−1)<0.00014.78 ± 0.25 b4.81 ± 0.40 b6.94 ± 0.43 a
P (kg ha−1)<0.00011.07 ± 0.11 b1.61 ± 0.28 b3.13 ± 0.12 a
K (kg ha−1)0.0001106.62 ± 9.28 a59.99 ± 11.44 b106.11 ± 6.06 a
Ca (kg ha−1)0.9804975.90 ± 157.13 a923.08 ± 211.36 a943.87 ± 186.49 a
Mg (kg ha−1)0.0800111.13 ± 17.64 a58.14 ± 60.34 a178.49 ± 11.64 a
δ13C (‰)<0.0001−27.04 ± 0.12 b−25.33 ± 0.24 a−27.63 ± 0.18 b
δ15N (‰)<0.00015.04 ± 0.16 b6.02 ± 0.21 a6.59 ± 0.22 a
C/N ratio0.006615.27 ± 0.23 a13.77 ± 0.36 b13.85 ± 0.42 b
C/P ratio<0.000165.11 ± 6.64 a38.74 ± 2.34 b28.18 ± 4.63 b
N/P ratio0.00025.02 ± 0.54 a3.35 ± 0.18 b2.35 ± 0.41 b
* A linear mixed model was used to test the effect of inselbergs as a fixed factor and the soil samples as a randomized factor. Tukey’s HSD post hoc test was used for mean comparisons.
Table 2. Average (±SE; n = 9) leaf area (cm2), leaf mass (g), and specific leaf area (SLA) in the Pseudobombax petropolitanum A. Robyns (Malvaceae) trees growing in three Atlantic Forest inselbergs in Espírito Santo State, southeastern Brazil. Inselbergs: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. Specifically for SLA values, the different letters indicate statistical differences among inselbergs (p ≤ 0.05) *.
Table 2. Average (±SE; n = 9) leaf area (cm2), leaf mass (g), and specific leaf area (SLA) in the Pseudobombax petropolitanum A. Robyns (Malvaceae) trees growing in three Atlantic Forest inselbergs in Espírito Santo State, southeastern Brazil. Inselbergs: TI = “Três Irmãs”, PA = “Aliança”, and SA = “Santa Angélica”. Specifically for SLA values, the different letters indicate statistical differences among inselbergs (p ≤ 0.05) *.
InselbergLeaf Area (cm2)Leaf Mass (g)SLA (cm2 g−1)
GreenSenescentGreenSenescentGreenSenescent
TI686.9 ± 39.3531.5 ± 48.68.6 ± 0.67.2 ± 0.581.2 ± 4.8 b73.8 ± 3.4 ab
PA683.6 ± 33.5544.6 ± 49.89.0 ± 0.57.6 ± 0.876.4 ± 2.5 b71.7 ± 1.4 b
SA760.6 ± 48.1504.9 ± 46.17.5 ± 0.76.1 ± 0.8104.3 ± 4.8 a82.8 ± 5.0 a
* A linear mixed model was used to evaluate the differences in SLA, with the inselbergs as a fixed factor and the individuals as a random factor. Tukey’s HSD post hoc test was used for mean comparisons.
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Jerônimo Júnior, R.A.d.C.; Paula, R.R.; Francisco, T.M.; Couto, D.R.; Covre, J.M.C.; Villela, D.M. Leaf Chemistry Patterns in Populations of a Key Lithophyte Tree Species in Brazil’s Atlantic Forest Inselbergs. Forests 2025, 16, 1186. https://doi.org/10.3390/f16071186

AMA Style

Jerônimo Júnior RAdC, Paula RR, Francisco TM, Couto DR, Covre JMC, Villela DM. Leaf Chemistry Patterns in Populations of a Key Lithophyte Tree Species in Brazil’s Atlantic Forest Inselbergs. Forests. 2025; 16(7):1186. https://doi.org/10.3390/f16071186

Chicago/Turabian Style

Jerônimo Júnior, Roberto Antônio da Costa, Ranieri Ribeiro Paula, Talitha Mayumi Francisco, Dayvid Rodrigues Couto, João Mário Comper Covre, and Dora Maria Villela. 2025. "Leaf Chemistry Patterns in Populations of a Key Lithophyte Tree Species in Brazil’s Atlantic Forest Inselbergs" Forests 16, no. 7: 1186. https://doi.org/10.3390/f16071186

APA Style

Jerônimo Júnior, R. A. d. C., Paula, R. R., Francisco, T. M., Couto, D. R., Covre, J. M. C., & Villela, D. M. (2025). Leaf Chemistry Patterns in Populations of a Key Lithophyte Tree Species in Brazil’s Atlantic Forest Inselbergs. Forests, 16(7), 1186. https://doi.org/10.3390/f16071186

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