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Article

Vegetative and Reproductive Responses from Full Sunlight to Shade of a Pantropical Herbaceous Plant in Caatinga Vegetation

by
Bruno Ayron de Souza Aguiar
1,*,
Elda Simone dos Santos Soares
2,
José Djalma de Souza
3,
Júlia Arruda Simões
2,
Danielle Melo dos Santos
2,
Vanessa Kelly Rodrigues de Araujo
2,
Josiene Maria Falcão Fraga dos Santos
4,
Clarissa Gomes Reis Lopes
1 and
Elcida de Lima Araújo
3
1
Centro de Ciências da Natureza, Universidade Federal do Piauí, Teresina 64049-550, PI, Brazil
2
Departamento de Biologia, Universidade Federal Rural de Pernambuco, Recife 52171-900, PE, Brazil
3
Departamento de Botânica, Universidade Federal de Pernambuco, Recife 50670-420, PE, Brazil
4
Departamento de Biologia, Universidade Estadual de Alagoas, Palmeira do Índios 57076-100, AL, Brazil
*
Author to whom correspondence should be addressed.
Forests 2026, 17(2), 153; https://doi.org/10.3390/f17020153
Submission received: 25 December 2025 / Revised: 19 January 2026 / Accepted: 22 January 2026 / Published: 23 January 2026
(This article belongs to the Section Forest Ecophysiology and Biology)
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Abstract

Herbaceous plants from dry forests respond to different levels of light availability over time and space through strategies that promote their establishment and survival. This study aimed to evaluate the tolerance of the perennial herb Talinum triangulare (Jacq.) Willd., which is pantropically distributed and forms dense populations in the Caatinga, under varying light availabilities. The treatments applied were full sun and 70%, 50%, and 30% light availability, each with 30 replicates. Vegetative, reproductive, and phenological responses were monitored over six months, during which the plant’s reproductive cycle was completed and water availability was higher. In T100, plants showed greater height, diameter, leaf production, flowers, fruits, and seeds. In contrast, reduced light availability led to lower values in these traits but resulted in increased leaf area, seed viability, and higher fruit/flower and seed/fruit ratios as compensatory responses. These findings suggest that higher light availability increases the establishment success of the studied species, although reduced light does not necessarily limit its reproductive success. The study highlights its adaptability to different light conditions and its potential for continued population expansion in dry tropical forests, despite fluctuations in light availability.

1. Introduction

The annual and interannual irregularity of rainfall in dry forests [1,2,3] causes year-to-year variation in leaf area of tree species composing the canopy cover [4,5] and, consequently, in soil shading [6,7]. In response to these changes in light availability, plants may alter their morphology, anatomy, physiology, and phenology [7,8,9,10], adjustments that reflect adaptation strategies to the environment and allow inference of species’ tolerance to high and low light availability [11,12]. In this context, light and water availability jointly regulate vegetative growth and reproductive processes in plants [6,13,14,15].
Despite the well-documented influence of light availability on plant performance across various terrestrial ecosystems, the strategies of native herbaceous plants in semiarid regions to tolerate high and low light availability may differ among species and remain poorly understood. This occurs because studies focusing on herbaceous species in these Neotropical ecosystems receive less attention compared to woody plants [16]. This fact is supported by evidence that, depending on the species, responses to reduced light can be expressed as lower heights, larger leaf areas, lower leaf production, and changes in biomass distribution patterns [17,18,19]. More conservative strategies can be observed, with plants reserving water in leaf tissues rather than accumulating larger amounts of biomass [12]. In addition, reproductive responses may be delayed or anticipated in flowering and fruiting [8,20,21,22], with a reduction or increase in the number of fruits and seeds produced [8,23,24].
In this context, and under current climate change scenarios, predictive models of climate change point to extreme events, with a tendency to reduce rainfall in semi-arid environments [25,26,27,28,29]. As a consequence of this projection, longer periods of leafless conditions are expected, particularly in deciduous species common in the Caatinga [3]. This may reduce canopy overlap, creating light mosaics, that is, different gaps of varying sizes that allow greater or lesser light irradiation in the understory [6,30,31,32]. Consequently, herbaceous plants will face more intense solar exposure, which may prevent many of them from starting the seed germination cycle due to soil desiccation [33]. Towards this evidence, emphasizing plant responses to variations in light availability is important for a better understanding of the functioning of Brazilian seasonally dry tropical forests, such as Caatinga vegetation, which harbors high herbaceous species richness, many of which are favored by increased light intensity [19].
Within this environmental scenario, the vegetation of the Caatinga is rich in endemic species as well as pantropical species that have established and formed large populations in the region over time, influenced by global aridification processes and climate change. The endemic and naturalized flora of the region evolved in response to these conditions, revealing a complex history of adaptation and ecological specialization to aridity levels [34]. Therefore, it is essential to investigate how these species established themselves in the Caatinga, considering their strategies for light tolerance, and to assess whether they will be able to cope with future climate changes. These findings may also enhance our understanding of the global process of species homogenization, which is progressing rapidly due to anthropogenic factors and climate change [35,36].
In light of these considerations, our hypothesis is that greater light availability positively influences the growth and reproduction of Talinum triangulare (Jacq.) Willd. (Talinaceae), a trend observed in many species of the Caatinga. However, reduced light levels may not necessarily limit the reproductive success of this pantropical species, which is adapted to both humid and dry environments [12,15,37,38,39,40,41]. We answer the following questions: Do greater vegetative growth and accumulation of plant biomass occur as light availability increases? Does reducing the light availability delay and limit the production of flowers, fruits, and seeds? Within this study, we investigated both the effects of high light stress and the shade tolerance strategies (low light availability) of the herbaceous plant.

2. Materials and Methods

2.1. Selected Species

Talinum triangulare (Jacq.) Willd. (Talinaceae), synonymous with Talinum fruticosum (L.) Juss., is a perennial herb with a pantropical distribution [37], originating from Africa but widely dispersed across Asia and South America [33,42]. Although not endemic to Brazil, it is considered a native species and occurs across almost the entire Brazilian territory [41]. Despite its African origin, this herbaceous plant forms large populations in the Caatinga vegetation, a Brazilian seasonally dry tropical forest, where it significantly contributes to landscape structure, particularly in the Agreste Caatinga [43]. In addition, the species has developed adaptations that allow its occurrence in a wide range of environments, including humid forests and urban areas worldwide [39,40]. These characteristics make T. triangulare a suitable model species for evaluating plant responses to contrasting light conditions across heterogeneous environments.
The herb is used in many parts of South America (mainly in the North, Northeast, and Midwest of Brazil), Africa, and Asia in traditional medicine (antifungal and antioxidant activity) in human food (non-conventional plant food), and as animal fodder [39,44,45,46]. It is a bulbous geophyte that presents leaf succulence, stems and roots that store nutrients, characteristics that are considered as adaptations to the drought [38,43]. Its reproduction can be crossed or by self-pollination, and it is considered facultatively autogamous with barochoric dispersion [47,48,49]. The herb presents C3 photosynthesis during the rainy season and facultative CAM induced by prolonged droughts [38].

2.2. Experiment Installing

Seeds of Talinum triangulare were collected in July 2016 from a Caatinga vegetation fragment at the Instituto Agronômico de Pesquisa (IPA), Caruaru-PE (8°14′18″ S, 35°55′20″ W, 535 m asl, 30 ha), during the fruit dehiscence period. Searches for populations of the species were conducted in 50 plots of 25 m2 each, previously demarcated within a 1-hectare area of preserved forest. The largest populations of T. triangulare were found in open areas within the forest interior and along edges, which receive higher light availability. A total of 400 seeds were collected from different locations within the fragment, and for each parent plant, a representative number of seeds was selected in a haphazard manner to capture variability among individuals.
In the laboratory, a week after collecting, the seeds were sanitized with 2.5% sodium hypochlorite and submitted to the chemical scarification pre-germinative treatment with 10% sulfuric acid (H2SO4) for 10 min, followed by imbibition in water for 24 h [33]. Afterward, the seeds were placed to germinate in Petri dishes, containing filter paper with cotton previously moistened. The plates were maintained in a germination chamber (BOD) at 25 °C with a 24-h photoperiod. One week after germination, 180 healthy-looking seedlings were selected. The seedlings were carefully transferred to properly labeled polyethylene bags containing 2 kg of soil collected in the same site of seed collection. The collected soil was previously autoclaved to avoid germination of other species and, consequently, competition for water and nutrients. The water retention capacity of 2 kg of soil, equivalent to soil pot capacity (PC), was determined before the seedling transplanting by the gravimetric method proposed by [14,15].
Polyethylene bags containing the seedlings were placed in a greenhouse (average natural daylight: 10,449.06 lux; average temperature: 28.7 °C; relative humidity: 64.8%) and were irrigated daily up to 100% of PC during 15 days for acclimatization of the young plants. On the 16th day, among those 180 plants, 120 were selected and submitted to four treatments of light availability: T100: full sun (11,638.9 lux; control treatment), T70: 70% (8203.5 lux), T50: 50% (5729.5 lux), and T30: 30% (3669.7 lux), with 30 replicates per treatment. Light availability was defined as a proportion of full sunlight, with T100 used as the reference condition. These average light levels were measured as mean illuminance (lux) values using a THAL-300 instrument (Thermo-Hygro-Anemometer-Luximeter, Instrutherm, São Paulo, Brazil) between 11:00 and 13:00, under each nursery shading level. These shading percentages were obtained in nurseries of 2.5 m long by 3 m wide and 2 m high, built with PVC pipes and black polyethylene shading screens (commercially known as Sombrite®, Vonder, Curitiba, Brazil) for each shading percentage (70%, 50%, and 30% shade). The experiment was monitored over a six-month period. During monitoring, the plants were irrigated daily, and the pots were maintained at 100% pot capacity (PC) to avoid water stress that could result from different light incidence percentages. All treatments were conducted in the same greenhouse, ensuring similar temperature and air humidity conditions, and any minor environmental fluctuations occurred uniformly across treatments. Therefore, light availability was the only experimental factor intentionally manipulated.
The experimental light treatments were designed based on four-year (2014–2017) temporal and spatial variations in light availability within the studied Caatinga fragment. Light intensity under natural conditions during the rainy season (March–August) ranged from 4424.6 to 16,240 lux, measured between 11:00 and 13:00 with a THAL instrument (Thermo-Hygro-Anemometer-Luximeter).

2.3. Data Collection

The growth of the 120 plants was measured weekly in height and diameter, using a measuring tape and digital caliper. In addition, the cumulative number of leaves produced per week of each plant was accounted for each treatment. At the end of the experiment, a completely expanded leaf per individual was selected, and these leaves were digitized to measure the total leaf area (TLA), using the Image pro plus 7.0 software. From these obtained values, we determined the physiological indexes of relative growth rate in height (RGRheight) and diameter (RGRdiameter) (unit: mm mm−1 week−1), using the following formula: RGR = (lnW2 − lnW1)/(T2 − T1), where “lnW2” and “lnW1” are the natural logarithms of the measurements per plant at times “T1” and “T2”, with weekly interval [50,51].
To evaluate total leaf production (TLP), we adapted the foliar gain approach proposed by [52]. Foliar gain (FG) was calculated at each weekly sampling interval as the difference between the final and initial leaf number of that interval (FG = LNf − LNi). Total leaf production (TLP) was then obtained by summing the weekly FG values across the entire experimental period, thereby accounting for cumulative leaf production over time rather than a single net difference between the initial and final leaf counts.
To assess the phenological activity index, the occurrence (presence = 1, absence = 0) of the vegetative phenophases (budding, senescence, and leaf abscission) and the reproductive phenophases (flowering and fruiting), were recorded weekly at each replicate. We considered the flowering phenophase as open flowers and the fruiting stage from the initial development of the fruit until its dispersion. According to [15,53], this index represents the percentage of plants that is simultaneously manifesting the phenological events, allowing assessment assess the population synchrony.
In each treatment, 100 flower buds in the pre-anthesis were marked to observe the process of anthesis duration. We considered anthesis to be the moment when the flowers open up until the senescence of the stamens and pistils, and complete retention of the petals [14]. Flowers and fruits produced per plant were counted daily since the beginning of flowering. Total seed production was estimated by counting seed numbers from 60 fruits randomly selected per treatment. In addition, using the production values, the ratio fruit/flower (Fr/Fl) and seeds/fruits (S/Fr) was estimated in each treatment. At the end of each light availability treatment, seeds were selected for a viability experiment, in which 60 seeds per treatment (15 seeds × 4 replicates) were subjected to germination tests to estimate germination percentage. Dormancy-breaking protocols followed [15,16], including seed sanitization and placement in Gerbox® boxes (Acrilplast, São Paulo, Brazil) containing moistened filter paper and cotton. The boxes were maintained in a B.O.D. chamber at 25 °C under a 12-h photoperiod for 30 days.
At the end of the experiment, the parts from each plant (leaves, stem, and root) were separated, packed in paper bags, and placed in the forced air circulation oven at 65 °C until reaching constant weight, for determining dry matter weight of leaves (LDM), stems (SDM), roots (RDM), and total dry matter (TDM) of each treatment. Total dry matter (TDM) was calculated as the sum of vegetative organs only and did not include reproductive structures.

2.4. Data Analysis

Variation in vegetative and reproductive traits between treatments, as well as the explanatory percentage of variation in light availability, was evaluated using Generalized Linear Models (GLM) with a Gaussian error distribution, incorporating ANOVA and a posteriori Tukey (F) tests in Statistica 7.0 software (StatSoft Inc., Tulsa, OK, USA). Each plant, derived from a different seed, was treated as an independent experimental unit. Seeds were collected from multiple parent plants across the fragment, minimizing pseudo-replication. All analyses were performed at a significance level of α = 0.05, and the Tukey test was applied to control for multiple comparisons among treatments.
To analyze vegetative and reproductive phenology, we used circular statistics following [54,55] with the aid of the Oriana 4.2 software (Kovach Computing Services, Pentraeth, Wales, UK) [56]. To calculate circular statistical parameters, the weeks and days, depending on the observation interval, were converted into angles (vegetative phenology: 0° = 1st week to 355° = 25th week, an interval of 14.4°; Reproductive phenology: 0° = 1st day to 357.8° = 169° day, an interval of 2.13°). The period of greatest phenological activity, which refers to the average date when most individuals produced leaves or reproduced, was defined based on the mean angle (μ). It is important to remember that the mean angle (and the corresponding date) does not necessarily point to the peak date of activity, but indicates the central trend of the data [14,15,55].
We applied the Rayleigh test (Z), to verify the significance of the mean angle (μ), which indicates seasonality in the phenophase, which is, whether the individuals are concentrated around the average date or distributed uniformly during the observation period. Rayleigh tests were considered significant at α = 0.05. The degree of synchrony of phenological events was determined by the length of the mean vector (r), which indicates the concentration of individuals around the mean date. This index has no units and ranges from 0 to 1, where values close to 1 indicate a greater degree of synchrony [55]. Differences between light availability treatments in each phenophase were assessed by the Watson-Williams test (F test) using Oriana 4.2 software [56], adopting a significance level of α = 0.05. Sample dispersion was previously tested by Watson’s U2 Test to verify the presence of Von Mises distribution, which is required for the F test to be performed.

3. Results

3.1. Vegetative Traits

We found significant differences for all vegetative traits among the studied treatments (p < 0.01) (Table 1). Plants submitted to the highest light availability (control—T100) were significantly higher in height (RGRheight, p < 0.01; Figure 1a) compared to other treatments, which, in turn, presented differences only between T70 and T30 (p < 0.01; Figure 1a). The plants submitted to the T100 differed significantly with the increase in diameter growth (RGRdiameter) in relation to the other treatments, which presented similar responses (Figure 1b). The plants submitted in the T100 presented lower TLA compared to the other treatments (Figure 1c). We found that about 10% to 51% of the variations that occurred in vegetative traits were explained by reductions in light availability (Table 1), with total leaf production (TLP) showing the lowest variation at 10% (Table 1).
The higher total production of leaves (TLP) was verified at T100, while the other treatments presented similar reduction when compared to T100 (Figure 2a).
There were significant differences between reduced light availability treatments in total plant biomass accumulation (p < 0.01), leaf biomass (p < 0.01), stem biomass (p < 0.01) and root biomass (p < 0.01) (Figure 3; Table 1). Control plants (T100) presented higher dry matter yield in the leaves, stem, and root compared with the other treatments (p < 0.01), which did not show significant differences between them (p > 0.05) (Figure 3). We found that about 37% to 59% of the changes in plant biomass were explained by reductions in light availability (Table 1).

3.2. Reproductive Traits

There was a significant difference in flower production (p < 0.01; Figure 2b), fruit production (p < 0.01; Figure 2c), and seed production (p < 0.01; Figure 2d) among treatments (Table 1). However, there were no quantitative differences in reproductive responses between T50 and T30 (p < 0.01) (Figure 2b–d). Fruit and seed production decreased with reduced light availability (Table 2).
We observed that the duration of anthesis was maintained over time, even in the face of reduced light availability. The S/Fr ratios, i.e., the average number of seeds produced per fruit, were 65, 66, 68, and 71 for treatments T100, T70, T50, and T30, respectively. The Fr/Fl ratios were high in all treatments (T100 = 0.82, T70 = 0.81, T50 = 0.91, T30 = 0.98), meaning that most of the produced flowers had fruit formation. Seed germination after the end of the experiment was high in all treatments, with mean germination percentages that did not differ statistically among light availability levels (T100 = 85%, T70 = 83%, T50 = 80%, T30 = 81%; Tukey test, α = 0.05), indicating that reduced light availability did not affect the germination capacity of seeds produced under different light conditions. We found that variations from 41% to 54% in reproductive traits were explained by the variation in light availability (Table 1).

3.3. Vegetative and Reproductive Phenology

The reproductive phenological responses for all treatments were seasonal (Z test, p < 0.01), but with low synchrony or low concentration around the average date for both flowering and fruiting (Table 3). Only T100 showed high activity with 100% of flowering individuals from the 135th day to the 145th day of the study, unlike fruiting, in which all treatments reached 100% activity (Figure 4). This fact occurred because there is a daily dynamic in the flower openings, different from the fruits that lasted about 2 to 3 weeks until their dispersion. We verified that only T50 and T30 synchronized the average date of flowering and fruiting (F test, p < 0.05), with the other treatments being different from each other.
Our results showed that T. triangulare presented a flowering beginning delay of 7 days in conditions of T70 light availability, 13 days in T50, and 15 days in the treatment of lower light availability (T30) when compared to the control treatment. Consequently, there was a fruiting delay of 9 days in the treatments of T70 and T50, and 15 days in T30 (Figure 4).
In vegetative phenophases, we found that, although budding seasonality is observed in all treatments through the significance of the mean angle (Z test, p > 0.05), our results are not reliable due to the low concentration of data, indicating uniform tendency or multimodal distribution (r ≤ 0.2), being more evident at T30 (r ≤ 0.08) (Table 3). During almost the entire observed period, the treatments were at 100% activity for this phenophase, however, there was a synchronous drop in budding, from the 13th week on in all treatments, reaching 33% in T100, 31% in T70, 25% in T50 and 22% in T30 (Figure 4).
Leaf senescence was seasonal (Z test, p < 0.01), with high synchrony around the average date for all treatments (r ≥ 0.6) starting at the 12th week at T100 and T70 and the 14th week at T50 and T30 (Table 3). The highest indices of activity for leaf senescence reached 100% at T100, 86% at T70, 61% at T50, while at T30 it did not exceed 57% activity during the monitored period (Figure 4). We found that leaf abscission was seasonal (Z test, p < 0.01), with high synchrony of individuals (r ≥ 0.7), starting at the 13th week at T100 and T70, at the 16th week at T30 and the 18th week at T50 (Table 3). During monitoring, individuals reached 100% activity in leaf senescence at T100, 93% at T70, 61% at T50, with a 43% decline at T30 (Figure 4).
In the comparisons between the angular averages of the budding, the values were not reliable due to the low concentration. We found that only T100 and T70, as well as T50 and T30, synchronized the average date of leaf senescence (F test, p > 0.05), different from leaf abscission in which all treatments showed significant differences (F test, p < 0.05).
Among the phenological phases analyzed, only fruiting was influenced by variations in light availability, which explained 11% of the observed variation (Table 1).

4. Discussion

Our findings indicate that Talinum triangulare exhibits high phenotypic plasticity across light availability gradients, allowing this species to establish and persist under contrasting light conditions. Although reductions in light availability explained part of the variation in vegetative and reproductive traits (e.g., TLP = 10%, fruiting phenology = 11%; Table 1), the residual variance remained high. This is likely due to inherent individual variability among plants, microclimatic differences within the greenhouse, and the species’ intrinsic phenotypic plasticity, which can buffer responses to light gradients.
Increased light availability, particularly under full sun conditions, promoted vegetative growth, as evidenced by greater height, stem diameter, and leaf production, thereby confirming our expectations. Similar patterns have been reported for other herbaceous species [7,10,19,20,57], emphasizing the importance of high light availability during the early stages of plant growth [58]. In contrast, increasing shade levels have been shown to enhance stem height and diameter in T. triangulare populations from humid tropical regions [40], suggesting evolutionary adjustments that enable survival in distinct environments [2]. Together, these results reinforce the capacity of this species to respond flexibly to different light gradients, supporting its wide distribution across tropical regions [59]. It is important to highlight that the effects of drought were not the variable analyzed in this study, yet it represents a limiting factor for the growth of herbaceous plants in the Caatinga forests [14,15]. According to [60], these effects of drought can be ameliorated under conditions of intermediate irradiance but become more severe at higher or lower light levels.
The increase in leaf area as light availability decreases is a response observed in T. triangulare [12] and well-documented among herbaceous plants, as described in the literature [10], corroborating our expectations. While the specific biochemical processes influenced by leaf size might not be explicit, changes in leaf area are commonly associated with plant adaptations to optimize resource utilization. For instance, the reduction in leaf area under conditions of high light availability typically corresponds to a strategy aimed not only at decreasing water loss but also at minimizing the photo-oxidation of organic compounds. This reduction naturally enhances photosynthetic efficiency, reduces oxidative stress, and mitigates damage to cell membranes, as indicated by previous studies [11,58,59,61,62]. Conversely, leaf area expansion enhances light absorption but may also elevate transpiration, leading to increased water loss and heightened susceptibility to stress [7,18,21,63]. However, in semiarid forests, under combined drought and increased shading, these effects can be attenuated [60].
Increased leaf production under full sun conditions likely compensates for reduced leaf area [23,58], providing photoassimilates necessary for height and diameter growth [18,61,64]. However, leaf trade-offs were insufficient to compensate for the limiting effects of reduced light on plant reproduction and growth, as evidenced by the low values observed for these traits.
Biomass production of T. triangulare was higher in full sun (T100), as recorded for other herbaceous species [18,20], with an emphasis on biomass accumulation on the stems and leaves. The preferential distribution of resources to the aerial system (stems and leaves) observed in T. triangulare can be limited and reversed to the roots under water and nutrient stress conditions [14,61,64]. Although in this study the plants did not undergo water stress, we can draw comparisons with studies that isolated water stress in the same environment [14]. Thus, the reduced availability of light, along with the decrease in soil water availability, can together explain much of the reduction in the total plant biomass, which remains concentrated in the aboveground portion of the plant.
Our results suggest that a longer duration for vegetative phenophases allows the plant to obtain sufficient resources to invest in its reproduction [8,65] a pattern verified only under low light availability. Thus, we believe there is a trade-off between investing in the time of vegetative growth and the beginning of reproduction in T. triangulare. In contrast, phenological imbalances in herbaceous vegetative growth can generate large losses in biomass, affecting plant growing seasons [66].
In our study, we identified notable patterns in the reproductive responses of T. triangulare, which could potentially become more frequent as the availability of light increases in the open areas of the Caatinga: (a) anticipation of phenophases; (b) greater production of flowers, fruits, and seeds; (c) floral anthesis maintained over time; (d) lower fruit/flower ratios and number of seeds per fruit.
In general, variation in light availability can anticipate, delay, or inhibit flowering and fruiting, as well as reduce or increase flower and fruit production, depending on the biological characteristics of each species and interactions with other environmental factors [6,8,18,20,22,65].
Under full sun conditions in dry forests, herbaceous species anticipate their flowering and tend to have a high production of flowers, fruits, and seeds [21,23]. Additionally, some perennial species may not show differences in their reproductive responses to light heterogeneity [67]. Late flowering is often considered a strategy to avoid or tolerate shade. However, this results in consequences such as low fruit production, which directly affects the reproductive success of the plant [6,8]. Possibly, delays in flowering generate a lack of synchrony with the life cycle of animals actively involved in pollination processes [66], thus explaining part of the reduction in production when cross-pollination occurs.
We emphasize that the scarcity of comprehensive studies investigating the relationship between light availability and seed/fruit ratio, alongside an array of critical reproductive variables in herbs from semi-arid environments, as highlighted by [6] presents a significant challenge in establishing direct correlations among these factors. Despite this scarcity, our observations indicate that allocating resources to enhance the average number of seeds per fruit appears to be an advantageous strategy in compensating for the negative effects of shading, thereby contributing to the species’ persistence in the environment. Nonetheless, it became evident that this strategy alone did not fully counteract the reduction in seed production, highlighting the complexity of factors influencing reproductive success under varying light conditions.
Therefore, the projected reduction in canopy overlap due to climatic extremes and anthropogenic actions [25,27,29,32] may increase light infiltration, potentially expanding the presence of T. triangulare in forest soil and facilitating the growth of its populations in dry tropical forests. Such projections can be made considering only variations in low or high light levels, without taking into account extreme drought or water stress conditions. Since soil water content was maintained at 100% of pot capacity throughout the experiment, light availability was the only intentionally manipulated experimental factor. Therefore, the projections and statements regarding the species’ future performance or expansion reflect this limitation. However, its phenotypic plasticity regarding shading conditions has also enabled its establishment in more humid forests [39,40]. This observed fact contributes to the understanding of how plant communities are undergoing homogenization over the years due to the naturalization of species beyond their native distribution areas through anthropogenic actions, potentially altering ecosystem structures and functions [35,36].

5. Conclusions

Our findings indicate that the greater the light availability in the environment, the greater the vegetative and reproductive success of T. triangulare. However, to compensate for the limiting effect of lower light availability, some tolerance strategies were verified, ensuring the completion of their reproductive cycle and yielding seeds with high viability. Our results suggest that, specifically, an increase in light availability, as expected under extreme climatic conditions, could potentially enhance the presence of T. triangulare in the soil and facilitate the expansion of its populations in dry tropical forests. Although we did not assess the isolated or combined effects of water deficit, which may elicit different responses and should be investigated in future studies, its phenotypic plasticity regarding shading conditions will allow its persistence in humid forests, which is already observed. It is possible that the naturalization of the studied species beyond its native distribution area occurred through anthropogenic actions and was facilitated by the region’s climatic conditions. Future studies are encouraged to investigate this further. Based on the observed responses, this frequency may increase over time in environments with both low and high light availability, contributing to the homogenization of the plant biota and potentially altering ecosystem structures and functions. We further highlight that our study is pioneering in attempting to understand the establishment of this pantropical herbaceous plant in dry tropical forests through its ecophysiological strategies in varying light conditions.

Author Contributions

D.M.d.S., J.M.F.F.d.S. and E.d.L.A. were responsible for the conceptualization. B.A.d.S.A. and E.S.d.S.S. wrote and prepared the original draft. B.A.d.S.A., E.S.d.S.S., J.D.d.S. and J.A.S. collected the data. V.K.R.d.A. and C.G.R.L. wrote, reviewed, and edited the draft. J.M.F.F.d.S. and D.M.d.S. also contributed to writing and preparing the original draft. E.d.L.A. supervised the project. All authors have read and agreed to the published version of the manuscript.

Funding

This study was funded by the following agencies, which supported the provision of scholarships and the execution of the project: FACEPE (Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco; APQ-0083-2.05/15; IBPG-1680-2.03/16); CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico; grants 133758/2017-6 and 167901/2022-2); and FAPEPI (Fundação de Amparo à Pesquisa do Estado do Piauí; grant 8104.UNI291.57805.19092022).

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Acknowledgments

We thank the funding agencies FACEPE, CNPq, and FAPEPI for their financial support. We also thank the IPA (Instituto Agronômico de Pernambuco) and UFRPE, UFPE, and UFPI for their logistical support. In addition, we are grateful to all researchers from the Natural Ecosystems Plant Ecology Laboratory (LEVEN) and the Ecophysiology and Conservation Biology Laboratory (LEBCon) for their assistance with data collection and analysis.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
TLATotal Leaf Area
TLPTotal Leaf Production
RGRRelative Growth Rate
RGRheightRelative Growth Rate in Height
RGRdiameterRelative Growth Rate in Diameter
TDMTotal Dry Matter
LDMLeaf Dry Matter
SDMStem Dry Matter
RDMRoot Dry Matter
PCPot Capacity
FrFruit
FlFlower
SSeeds

References

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Forests 17 00153 g001
Figure 1. Effect of light availability on vegetative traits of Talinum triangulare (Jacq.) Willd. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test. (a) RGRheight (Relative growth rate in height); (b) RGRdiameter (Relative growth rate in diameter); (c) Total leaf area (TLA).
Figure 1. Effect of light availability on vegetative traits of Talinum triangulare (Jacq.) Willd. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test. (a) RGRheight (Relative growth rate in height); (b) RGRdiameter (Relative growth rate in diameter); (c) Total leaf area (TLA).
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Figure 2. Total production of leaf (a), flowers (b), fruits (c), and seeds (d) of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test.
Figure 2. Total production of leaf (a), flowers (b), fruits (c), and seeds (d) of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test.
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Figure 3. Biomass of roots, stem, and leaves from Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test.
Figure 3. Biomass of roots, stem, and leaves from Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Error bars represent the mean ± 95% confidence interval. Distinct letters among light variations denote significant differences by the Tukey a posteriori test.
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Figure 4. Circular distribution of percentage activity of vegetative and reproductive phenophases of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Bars around the circle refer to the weekly or daily percentages of phenological activity for each treatment. The direction of the arrow indicates the average date (μ) and refers to the period of highest activity. The length of the arrow indicates the degree of synchrony (r).
Figure 4. Circular distribution of percentage activity of vegetative and reproductive phenophases of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments of different light availability: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux). Bars around the circle refer to the weekly or daily percentages of phenological activity for each treatment. The direction of the arrow indicates the average date (μ) and refers to the period of highest activity. The length of the arrow indicates the degree of synchrony (r).
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Table 1. Generalized linear models (GLMs) to assess the influence of light availability on the vegetative, reproductive, and phenological traits of Talinum triangulare (Jacq.) Willd., with a Gaussian error distribution and Tukey post hoc tests highlighting significant differences (α = 0.05).
Table 1. Generalized linear models (GLMs) to assess the influence of light availability on the vegetative, reproductive, and phenological traits of Talinum triangulare (Jacq.) Willd., with a Gaussian error distribution and Tukey post hoc tests highlighting significant differences (α = 0.05).
CategoryTraitsSSTSSErrorSMFpη2
Vegetative traitsRGRh0.010.050.0390.00516.46<0.010.28
RGRd0.0020.010.0070.00212.79<0.010.22
TLA2714.810,157.97443904.914.10<0.010.51
TLP15,468.9117,599.1102,130.25156.35.85<0.010.10
Total biomass alocation89.74256.69166.929.9156419.71<0.010.59
Biomass leaves (LDM)7.2251.8744.62.45.93<0.010.37
Biomass stem (SDM)21.4475.0653.653.614.66<0.010.53
Biomass roots (RDM)4.9528.0323.081.657.86<0.010.42
Reproductive traitsFlowers production166,075558,257392,18155,35815.66<0.010.54
Fruits production65,864359,415293,55221,9558.30<0.010.42
Seeds production256,591,3001.51 × 1091.25 × 10985,530,4337.57<0.010.41
Flowering37,971.9711,651749,62312,657.311.95<0.010.22
Phenological traitsFruiting16,747.61,327,2681,310,5215582.52.860.030.11
Leaf budding1673.456,657.754,984.3557.80.970.40-
Leaf senescence6875.993,614.586,738.52291.92.530.06-
Leaf abscission3769.557,922.954,153.31256.52.220.08-
SS: squares sum; TSS: total squares sum; Error: error value; SM: Squares Mean; F: Fisher test; p < 0.05—significant differences; η2: Explanatory Percentage; RGRh: relative growth in height; RGRd: relative growth in diameter; TLP: total leaf production; TLA: total leaf area.
Table 2. Total production of flowers, fruits, and seeds of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux).
Table 2. Total production of flowers, fruits, and seeds of Talinum triangulare (Jacq.) Willd. under different levels of light availability. Treatments: T100 = full sun (11,638.9 lux/control); T70 = 70% (8203.5 lux); T50 = 50% (5729.5 lux); T30 = 30% (3669.7 lux).
TreatmentFlowersFruitsSeeds
tpMean ± SEtpMean ± SEtpMean ± SE
T1006.652221.7 ± 75.45.493183.1 ± 61.3357.04511,864.8 ± 3974.3
T705.242180.7 ± 58.64.287147.8 ± 46.7282.9429815.7 ± 3106.3
T504.059144.9 ± 54.23.697132 ± 51.2251.3968080.5 ± 3134.8
T303.409121.7 ± 43.73.358119.9 ± 44238.4188454.9 ± 3104
tp: total production per treatment, SE: standard deviation.
Table 3. Circular analysis of vegetative and reproductive phonological patterns of Talinum triangulare (Jacq.) Willd. under different levels of light availability.
Table 3. Circular analysis of vegetative and reproductive phonological patterns of Talinum triangulare (Jacq.) Willd. under different levels of light availability.
PhenophasesVariableTreatments
T100T70T50T30
FloweringObservations9922854371616691
Mean angle (μ)224.8° *235.2° *249.2° *247.7° *
Circular standard deviation88.8°78.4°71.7°70°
Length of mean vector (r)0.30.390.450.47
FruitingObservations12,30311,31110,10110,372
Mean angle (μ)228.5° *239° *251.6° *248.6° *
Circular standard deviation86.7°77.2°69.5°71.8°
Length of mean vector (r)0.310.40.470.45
Leaf buddingObservations2090186219172086
Mean angle (μ)72.3° *81.2° *71.9° *92.9° *
Circular standard deviation116.7°99.7°104.6°128.4°
Length of mean vector (r)0.120.220.180.08
Leaf senescenceObservations918629454383
Mean angle (μ)270.6° *270.6° *278.9° *278.3° *
Circular standard deviation49.7°46.5°43.9°49.8°
Length of mean vector (r)0.6860.7190.7450.684
Leaf abscissionObservations646423229339
Mean angle (μ)271.5° *264.4° *288.7° *274.6° *
Circular standard deviation45.6°36.4°34.3°38.6°
Length of mean vector (r)0.720.810.830.79
* Significant mean angles (μ) by Rayleigh test (p < 0.05).
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MDPI and ACS Style

Aguiar, B.A.d.S.; Soares, E.S.d.S.; Souza, J.D.d.; Simões, J.A.; Santos, D.M.d.; Araujo, V.K.R.d.; Santos, J.M.F.F.d.; Lopes, C.G.R.; Araújo, E.d.L. Vegetative and Reproductive Responses from Full Sunlight to Shade of a Pantropical Herbaceous Plant in Caatinga Vegetation. Forests 2026, 17, 153. https://doi.org/10.3390/f17020153

AMA Style

Aguiar BAdS, Soares ESdS, Souza JDd, Simões JA, Santos DMd, Araujo VKRd, Santos JMFFd, Lopes CGR, Araújo EdL. Vegetative and Reproductive Responses from Full Sunlight to Shade of a Pantropical Herbaceous Plant in Caatinga Vegetation. Forests. 2026; 17(2):153. https://doi.org/10.3390/f17020153

Chicago/Turabian Style

Aguiar, Bruno Ayron de Souza, Elda Simone dos Santos Soares, José Djalma de Souza, Júlia Arruda Simões, Danielle Melo dos Santos, Vanessa Kelly Rodrigues de Araujo, Josiene Maria Falcão Fraga dos Santos, Clarissa Gomes Reis Lopes, and Elcida de Lima Araújo. 2026. "Vegetative and Reproductive Responses from Full Sunlight to Shade of a Pantropical Herbaceous Plant in Caatinga Vegetation" Forests 17, no. 2: 153. https://doi.org/10.3390/f17020153

APA Style

Aguiar, B. A. d. S., Soares, E. S. d. S., Souza, J. D. d., Simões, J. A., Santos, D. M. d., Araujo, V. K. R. d., Santos, J. M. F. F. d., Lopes, C. G. R., & Araújo, E. d. L. (2026). Vegetative and Reproductive Responses from Full Sunlight to Shade of a Pantropical Herbaceous Plant in Caatinga Vegetation. Forests, 17(2), 153. https://doi.org/10.3390/f17020153

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