Phenological cycles in the Pantanal woody communities: Responses to climate and soil moisture seasonality

Rainfall remained below 40 mm month-1 from May to September, triggering leaf fall. Most communities presented strong negative correlations between rainfall and leaf fall intensity, with correlation coefficients above 75% for PS, IS, CS, MF, and ASDF (Fig 6). The influence of rainfall on leaf fall intensity occurred up to one month before the event in some communities, such as the IS (r1 = -0.62, p = 0.02) and PS (r1 = 0.63, p = 0.02) communities. Unlike the other meteorological variables, increased insolation positively influenced leaf fall phenology in open canopy communities, such as the IS (r0 = 0.67, p = 0.01), PS (r0 = 0.57, p = 0.04), and CS (r0 = 0.56, p = 0.04) (Fig 6). Leaf fall intensity in the seasonal alluvial forest (ASDF) was also posi- tively correlated with increased insolation in dry months (r0 = 0.57, p = 0.04). There was no significant correlation between the soil moisture content and the leaf fall cycle in any of the communities, although the soil moisture showed expressive seasonal variation in most of the sampled vegetation (Fig 2). Flowering. The influence of seasonality on meteorological variables and its correlation with flowering intensity varied between communities (Fig 6). In the SHB community, annual variability in soil moisture triggered flowering events, with flowering occurring in August and September, when the soil moisture dropped below 5%, and the mean (29˚C) and maximum (37˚C) temperatures reached their peak. The impact of evapotranspiration was observed with

a two-month delay (r1 = -0.62, p = 0.02). Insolation had a positive effect on flowering one month prior to the event (r1 = 0.59, p = 0.03) and a negative effect in the two months preceding it (r2 = -0.61, p = 0.03). In PS and IS, an increase in insolation hours coincided with more intense flowering events (PS: r0 = 0.57, p = 0.04; IS: r0 = 0.56, p = 0.05). The minimum temperature had a strong influ- ence on flowering events in PS (r0 = -0.74, p = 0.004), suggesting that cold fronts can trigger flower blooming. The cyclical change in weather significantly influenced the flowering cycles in the MF, IS, and PS communities, aligning with trends in various meteorological variables (Fig 6). For example, a reduction in evapotranspiration was associated with an increased percentage of flowering in the MF (r0 = -0.86, p < 0.001), IS (r0 = -0.71, p = 0.003), and PS (r0 = -0.70, p = 0.01) stands. Increased insolation enhanced bloom coverage in the IS (r0 = 0.56, p = 0.05) and PS (r0 = 0.57, p = 0.04) communities. Additionally, reduced maximum tem- peratures significantly affected the flowering intensity in the two months preceding the event. In the CS, flowering intensity was weakly correlated with climate seasonality. The decrease in maximum temperature two months prior to the reproductive event was associ- ated with increased flowering intensity (r2 = -0.60, p = 0.03). Notably, the months leading to the peak of flowering were characterized by cooler temperatures of 5˚C cooler. Finally, the peak flowering periods in the ASDF (r2 = 0.62, p = 0.02) and LSDF (r2 = 0.61, p = 0.03) for-

ests were linked to increased rainfall in the two preceding months, marking transitions from wet to dry periods. Fruting. Climate seasonality had limited influences on fruiting phenology. No correlation was found with soil moisture. Only in the SHB and MF, communities subjected to prolonged flooding (> 3 months, Fig 2), did the fruiting intensity correlate with weather variables. In the SHB, the maximum temperature (r0 = 0.56, p = 0.04) during the phenological event and insola- tion one month prior (r1 = 0.64, p = 0.02) were positively correlated with the fruiting percent- age of the Fournier index, whereas insolation two months prior (r2 = -0.55, p = 0.05) was negatively correlated. In the MF treatment, the mean temperature (r0 = 0.57, p = 0.04) posi- tively influenced fruiting, with higher mean temperatures two months earlier triggering high- intensity fruiting events (r2 = 0.57, p = 0.04). PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 17 / 30

Discussion Leaf phenology The leaf-shedding cycles observed in the savanna and forest communities in our study ranged from weakly seasonal to aseasonal, highlighting the diverse phenological responses among woody species. This variation in leaf fall traits reflects the evolutionary and ecological forces linked to environmental filtering that enable species to thrive under different habitat condi- tions [74]. In all the analyzed communities, semideciduous species dominated, comprising more than 50% of the leaf exchange strategies. This aligns with Camargo et al.’s (2018) classification of the woody Cerrado as seasonal semideciduous vegetation [63]. Approximately 29% of the woody species in these communities exhibited complete deciduousness, particularly among those biogeographically associated with the Bolivian Bosque Chiquitano, mesotrophic cer- radão, and other seasonal forests in South America [75]. Notable examples include A. colu- brina, C. fasciculata, H. heptaphyllus, and M. hirtum. In the sandy, low-fertility soils of the PS community, the number of deciduous species doubles compared with those of the other six communities, mirroring the trend observed along the gradient from the Cerrado to seasonally dry tropical forests in northeastern Brazil, where deciduous species increase with aridity [76]. Evergreen species are present in all communities but are more abundant in seasonal seasonal forests, which are typically associated with low to moderate flooding [77]. Convergent responses to leaf shedding within communities were closely tied to climate sea-

sonality. Increased exposure to solar radiation and reduced evapotranspiration during periods of low cloud cover positively influenced leaf fall in the dry winter, particularly in open-canopy communities such as PS and CS. Synchronizing leaf emergence and shedding with solar radia- tion maxima and minima enables trees to maximize water use, contributing to fluctuations in evapotranspiration rates [78]. The increase in evapotranspiration observed at the start of the rainy season corresponds with rising net radiation and temperature [79,80]. Notably, increas- ing maximum monthly temperatures are correlated with increased leaf fall, especially in the SHB community, which experiences several months of inundation during the wet season and very low soil moisture during the driest months [77] C. lanceolatum, the monodominant spe- cies in this community, behaves as an evergreen for most of the year but experiences increased leaf shedding during periods of peak temperatures. In this context, leaf shedding aids in dissi- pating heat during high temperatures and intense solar exposure while also reducing water loss, enhancing leaf cooling, and mitigating the risk of surpassing stress resistance thresholds [81]. Meteorological influences on the vegetative phenological cycle begin approximately two months prior to high-intensity events, underscoring the importance of predictable seasonal cli- mate transitions—such as wet periods following droughts and the arrival of cold air masses— that trigger significant phenological shifts [82].

The climate seasonality associated with an annual flood pulse imposes additional con- straints on woody plants, influencing patterns of deciduousness [83,84]. Anoxic conditions and reduced gas exchange during flooding shape various plant strategies to cope with near- zero oxygen concentrations [9]. At the end of the rainy season, when the maximum soil mois- ture and flood levels are reached, a second peak in leaf fall is observed, with individuals experiencing more than 50% leaf shedding across all communities. This pattern indicates that leaf shedding serves as a strategy to avoid moisture stress caused by both dry-season droughts and flood-season soil hypoxia [9,11] Understanding the relationship between vegetative phe- nology and water stress requires species-specific information on adaptations to drought and flood resistance [11,85]. Jancoski et al. (2022) provided evidence of interspecific variation in hydraulic function regulation influenced by deciduousness [23]. Young individuals of T. PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 18 / 30

aurea, a typical Cerrado species, tolerate flood stress by preventing root system death and leaf abscission [86]. Total or partial deciduousness, combined with strong stomatal regulation, appears to be a common water regulation strategy for both dry and flood season species in the Cerrado and flooded forests [23,81]. L. parvifolius, commonly found in MF, efficiently main- tains water use during the dry season but exhibits stomatal limitations in response to seasonal flooding [81], with more than 50% of its leaves lost during this period. The two peaks in leaf fall intensity during both dry and wet periods have significant implications for carbon cycle dynamics within these communities, as the decomposition of senescing leaves contributes to the release of carbon from the soil back into the atmosphere [87]. Many species identified in the studied floodplain were found across multiple communities, exhibiting intraspecific variation in vegetative activity. This variation is attributed to distinct ontogenetic stages within populations, small-scale environmental heterogeneity, and genetic structure [73,88,89]. Water table depth and fluctuations are considered to modulate plasticity in maximum root depth among species occurring in both the Cerrado and Pantanal. For example, the root depth of T. aurea and C. americana in the Pantanal was found to be half that of their counterparts in the Cerrado [73]. Shallow root systems during waterlogging may lead to reduced transpiration and tree growth, triggering additional leaf-shedding events, a phe-

nomenon also observed in Amazon floodplain trees. Some individuals of A. discolor were observed to partially and shortly shed leaves during the flood season, reinforcing the idea that leaf shedding can occur in evergreen species in the aquatic phase [11,83]. Phenotypic plasticity allows species populations to adapt to various positions along the soil hydrological gradient, facilitating their persistence in savannas and forests [90]. Even without clear evidence that surface soil moisture seasonality affects leaf shedding, water table dynamics may influence the ability of leaf fall to adjust during periods of low atmo- spheric and soil moisture. We speculate that the lack of a significant correlation between soil moisture and leaf shedding intensity may stem from habitat-specific interactions among soil physical attributes, rainfall, and flood dynamics that disrupt the alignment between rainfall timing and soil moisture seasonality [13,91]. These dynamics can lead to prolonged water shortages, reducing the seasonality of leaf shedding, as observed in Lowland Seasonally Dry Forests (LSDF). In these seasonal forests, which are typically found in areas of high soil fertility along flood-free riverbanks and relict dunes [91], the relatively high number of evergreen spe- cies suggests that some trees are buffered against seasonal drought due to access to deep soil layers (2–6 m) or storage in succulent tree trunks [92,93]. In contrast, the floristically similar ASDF may experience significant leaf fall seasonality due to stronger connections between

rainfall, soil surface moisture, and groundwater fluctuations at lower elevations near the river’s watercourse [91]. Access to groundwater reserves is thus a critical factor for the resilience of both forest and savanna ecosystems [73,94]. Reproductive phenology cycle The intricate interaction between climate signals and the reproductive patterns of plant com- munities in diverse ecosystems, particularly in the Pantanal region, highlights the critical role of phenological events in shaping ecological dynamics [22,95–98]. The studied communities exhibited significant seasonal flowering, with notable exceptions in the ASDF. In essence, these results from the Pantanal underscore the crucial role of lower minimum and maximum temperatures, rainfall, and evapotranspiration in triggering flowering events, highlighting dry- ing and cooler conditions as a period of peak flower production [99]. Climatic antecedence emerges as a key determinant of the rhythms of the phenophase and is influenced by the feed- back between hydroclimatic conditions and energy flux with land cover type [100]. PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 19 / 30

The distinct synchronies and varied functions of plant species assemblies have led to some communities exhibiting peak flowering activity during both the dry season and wet phase, which is related to the presence of species with multiple reproductive strategies within the same community. Species associated with ASDF, such as T. catigua and A. cuspa, have individ- uals who flower in opposite hydrological phases in the rainy and dry seasons, respectively, which could be influenced by their distinct floristic affinities with neighboring ecosystems (e.g., Atlantic forest, Seasonal forest, and Cerrado) [75]. In general, the studied species popula- tions exhibited low-intensity and brief-to-intermediate flowering events [101], which aligns with the findings of other studies [22,99,102,103]. Damasceno-Junior (2020) noted that the flowering phenophase in the Pantanal is generally weakly seasonal (111), gaining significance in certain species groups associated with dispersal strategies [104]. The communities with greater synchronisms were those dominated by a few species (SHB, IS, and MF) or that occurred on flood-free ground on ridges or ancient levees (i.e., LSDF) [46]. Species employing seed or fruit dispersal during periods of high river levels, such as V. divergens and M. guianen- sis, presented peak flowering activity at the very end of the wet season (June), when flooding receded, resulting in fruiting in January. T. aurea and H. heptaphyllus, known for their massive and synchronized flowering, produce numerous anemochoric seeds that respond clearly to

variations in climate [51] The flowering phenophase in SHB occurs simultaneously with fruit- ing during the dry season. Synchronized reproductive behavior is influenced by high tempera- tures and consecutive months of exposure to solar radiation, indicating the high tolerance of savanna shrub species to extreme heat and drought [105]. In addition to climatic influence, endogenous forces play a role in explaining low and asyn- chronous flowering within species populations [106,107], as observed in both CS and ASDF, respectively. Endogenous factors, which are crucial for flower production, can manifest through spatial variations in the chemical attributes of the soil [106] and as a life strategy to avoid competition for visitors and pollinators [107]. Notably, the physicochemical aspects of the soil, such as the phosphorus content [108], vary among the distinct vegetation types in the Pantanal [109]. In general, the seasonal synchrony of fruiting among the different species was weak (con- centration coefficient, rc < 0.50) across the various vegetation communities. This phenome- non may arise when coexisting plant species exhibit diverse life cycle modes, resulting in the decoupling of phenophases from climatic seasonality [69,108,110]. At the individual level, the presence of fruits at reduced frequency and intensity (Fournier intensity < 6%) underscores the potential influence of environmentally heterogeneous ecosystems and resource availability on heightened species competition and reduced energy investment in reproduction [111,112].

This divergence in reproductive schedules establishes a resource availability timetable for fauna, promoting complementarity and potentially sustaining biodiversity throughout a longer part of the year [113]. In most of the monitored communities, species populations consistently produce fruits year- round, although at low frequency and intensity, which can be advantageous for frugivorous ani- mals (e.g., Tapirus terrestris and Ramphastos toco), allowing them to align their fruit consump- tion with the phenology of various fruiting species [114–116]. Conversely, communities characterized by a strong dominance of a few species, notably SHB and MF, presented pro- nounced seasonality in terms of fruit production influenced by increasing temperatures during the austral wetter summer. This finding underscores the significant influence of the flowering season on subsequent fruit dispersal in these environments [104,117]. While leaf development is typically more responsive to changes in water availability than flowering or fruit production is [83], access to groundwater may allow fruiting to occur independently of the usual moisture fluctuations [118]. Additionally, species’ phylogenetic signals can further decouple reproductive PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 20 / 30

cycles from water availability, leading phylogenetically related species to exhibit similar pheno- logical periods, irrespective of moisture conditions or seasonal changes [118]. These fruiting patterns reveal a nuanced strategy mediated by dispersal syndromes and phylogenetic signals in plant phenology [22,119]. While abiotically dispersed species (mostly wind-dispersed) release seeds mainly in the dry season, the wet-season peak is driven almost entirely by biotically dispersed species [22,120]. For example, T. catigua and A. discolor tend to grow in alluvial forests and flood-prone areas [75] during the wettest months (December-Jan- uary; ~24% soil moisture). On the other hand, fructification of species found in CS, such as A. fraxinifolium and A. colubrina, occurred during the driest months (August-September; < 14% soil moisture). The relationships between plant communities and seed dispersers, which include mammals (e.g., bats and primates), birds, and fishes, underscore the critical role of flood pulses in shap- ing patterns of seed deposition. The intricate balance required for ecosystem health and regen- eration and the role of dispersal strategies within each community assembly [108] provide valuable insights into why the onset and end of the reproductive period may not fully coincide across communities. Frugivory by fish and hydrochory are relevant mechanisms for determin- ing lateral dispersal and seed deposition patterns across floodplain communities [121,122]. For example, M. guianensis and O. suaveolens, which undergo fruiting during the rainy season in

the MF and ASDF, serve as examples of species dispersed by fish [96]. Their involvement plays a vital role in the regeneration of forest ecosystems [114]. Concluding remarks: Informing conservation and management initiatives The unique hydrological response of the Pantanal to the climatic cycle sets it apart from other nonflooded savanna vegetation [17,123]. This response involves the intricate interplay between phenological dynamism and alternating drought and wet phases [11,25]. This study revealed the existence of interspecific synchrony in vegetative and reproductive phenology within savanna and forest communities. Differences in seasonal phenological patterns among com- munities were predominantly observed in flowering, with communities with intensities con- centrated from the very end of the wet season (when flooding is receding) to the dry season, differing from those with flowering events more scattered throughout the year (Figs 5A and S1). Interspecific synchronism and seasonal responses in leaf fall were triggered by climate sea- sonality in terms of rainfall, temperature, and evapotranspiration, while the climate signals for flowering and fruiting events were predominantly detected for communities with high domi- nance for few species. The difference in assemblage activity times observed in other Pantanal regions [108] might result from the complex interplay between plant evolution and the current selective habitat fil- ters [51,124,125], resulting in unique strategies for coping with periods of soil or atmospheric

water stress concomitant with high air temperatures [126,127]. The broad array of phenologi- cal responses, which might confer stability to interannual climate fluctuations at the commu- nity level, is further supported by the ability of some species to acclimate to increasing drought or wet conditions [128] and occupy distinct zones of the soil hydrological gradient in the Pan- tanal [17]. Nevertheless, in the context of climate change, the northern Pantanal has experi- enced a shift toward aridity, marked by a 13% increase in the number of days without rain compared with that in the last 60 years. Additionally, there was a 16% reduction in water mass during the drought season compared with that observed a decade ago [129]. This approach tests tolerance limits and alters the dynamics of competitive strategies among shrubs, palms, lianas, and trees while vying for resources in challenging environments. PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 21 / 30

Through repeated cycles of drought and a warming climate, the studied area may be sub- jected to changes in species dominance and potential community declines due to elevated tree mortality across various ontogenetic stages [25,50]. Communities characterized by weak responses to rainfall seasonality, such as the LSDF and SHR, are anticipated to experience less impact amidst ongoing climate change. Conversely, communities that quickly adjust to yearly climate patterns may experience more significant impacts. The trajectory of these changes will depend heavily on the species-specific phenological and physiological strategies that plants adopt to cope with drier and warmer climate conditions, as well as disturbances such as fires [130]. Arau´jo et al. (2021) suggested that deciduousness becomes more likely in forest trees, causing a physiognomic shift toward savanna-like vegetation formation [90]. Species that exhibit leaf drop (deciduous) and employ low investment in fruits and seeds (anemochory and autochory) [108] may perpetuate this phenomenon in the context of extended and intense dry seasons, along with reduced flooding [51,88,131]. The replacement of long-lived trees with pioneer trees and lianas is also expected under such climate change scenarios and after wild- fires [25,131]. Even zoochoric tree species, including the caducifolious and pioneer Cerrado species C. americana, may become dominant under drier climate conditions and intensified fire regimes [132]. The interactions between groundwater recharge and surface water dynam-

ics in communities are critical in determining vegetation shifts under climate change scenarios [133]. Nevertheless, the diminished signals for flowering triggered by low temperatures may lead to decreased flowering or the prevention of flowering events in several species [106]. The potential for long-distance dispersal via zoochory and hydrochory across the floodplain is expected to be constrained, which may lead to a reduction in species richness [25] and a decrease in available resources for the diverse fauna found in the Pantanal [28,96,114]. Understanding the selective pressures that shape species’ adaptations to the environment can provide insight into the mechanisms driving ecosystem change and help in conservation and management efforts [134]. Considering the substantial contribution of vertebrates to woody plant dispersal in the Pantanal, the mortality of fauna during wildfires [28] might pose critical concerns to vegetation regeneration and conservation of the diverse mosaics of plant communities in the Pantanal, reducing seedling recruitment [25,135]. The challenges posed by climate change, wildfires, and the potential loss of large-bodied vertebrates necessitate ongoing research, long-term monitoring, and conservation efforts to safeguard the unique biodiversity and ecosystem services offered by the Pantanal. Since the timing of the phenophase is irregular to some degree between years [69], long-term monitoring of phenological cycles could provide additional insight into future scenarios related to climate variability in Pantanal vegetation.

Supporting information S1 Fig. Frequency of woody individuals undergoing leaf shedding, flowering, and fruiting phenological activities throughout an annual hydroclimatic cycle (July 2017 to June 2018). The color gradient illustrates periods of high activity (intense colors) and low activity (softer colors) among individuals. (TIF) S2 Fig. Frequency of woody species in reproductive activity of, A. Flowering activity and B. Fruiting activity, across the annual hydroclimatic cycle for the seven communities. (TIF) S1 Table. Woody species composition and vegetative and reproductive phenology of seven savanna and forest communities of the northern Pantanal. A total of 59 species from 28 families were recorded, totaling 193 individuals in 7,200 m2. The table presents the occupied PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 22 / 30

communities and the months of the year (from 1 to 12, corresponding to January to Decem- ber) for the activities of leaf fall, flowering, and fruiting of each species. Leaf fall activity is con- sidered when it is above or equal to 50%. PL < 50%: Partial Leaf Fall (less than 50%); LY: Leaf Fall Lost Year-round (� 50%). Communities: Cerrado Savanna (CS), Invaded Savanna (IS), Parkland Savanna (PS), Shrubland (SHB), Lowland Seasonal Semideciduous Forest (LSDF), Alluvial Seasonal Semideciduous Forest (ASDF), and Semi-evergreen Monodominant Forest of Vochysia divergens (MF). (DOCX) S1 Appendix. Dataset containing the original data utilized in the phenological study con- ducted in the Pantanal Mato-grossense, SESC Baı´a das Pedras, Pocone´ municipality, Mato Grosso (Brazil). It encompasses information on reproductive and vegetative phenology, envi- ronmental data, and results from derived analyses. (XLSX) Acknowledgments We dedicate this work to the memory of the UFMT technician He´lio Ferreira for his invalu- able support, which contributed significantly to our research and that of many young scientists in plant ecology at the UFMT. We are thankful for the technical and logistic support provided by the Federal University of Mato Grosso, the SESC Pantanal (Brazilian Social Service of Com- merce), the Pantanal Research Center (CPP), and the National Institute for Science and Tech- nology of Wetlands (INAU). Author Contributions Conceptualization: Julia Arieira, Karl-L. Schuchmann, Marinêz Isaac Marques. Data curation: Julia Arieira. Formal analysis: Julia Arieira.

Funding acquisition: Karl-L. Schuchmann, Marinêz Isaac Marques. Investigation: Julia Arieira, Michelle D. Lanssanova. Methodology: Julia Arieira. Project administration: Julia Arieira, Karl-L. Schuchmann, Marinêz Isaac Marques. Resources: Julia Arieira, Karl-L. Schuchmann, Marinêz Isaac Marques. Software: Julia Arieira. Supervision: Julia Arieira, Karl-L. Schuchmann, Marinêz Isaac Marques. Validation: Julia Arieira. Visualization: Julia Arieira, Ana Silvia O. Tissiani, Osvaldo Borges Pinto Junior. Writing – original draft: Julia Arieira, Michelle D. Lanssanova. Writing – review & editing: Julia Arieira, Karl-L. Schuchmann, Arnildo Pott, Osvaldo Borges Pinto Junior, Marinêz Isaac Marques. References 1. Boulton CA, Lenton TM, Boers N. Pronounced loss of Amazon rainforest resilience since the early 2000s. Nat Clim Chang. 2022; 12: 271–278. https://doi.org/10.1038/s41558-022-01287-8 PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 23 / 30

2. Ricaurte LF, Olaya-Rodrı´guez MH, Cepeda-Valencia J, Lara D, Arroyave-Sua´rez J, Max Finlayson C, et al. Future impacts of drivers of change on wetland ecosystem services in Colombia. Global Environ- mental Change. 2017; 44: 158–169. https://doi.org/10.1016/J.GLOENVCHA.2017.04.001 3. Ramsar Technical Report 3: Valuing wetlands: Guidance for valuing the benefits derived from wetland ecosystem services | Ramsar. [cited 28 Mar 2020]. Available: https://www.ramsar.org/document/ ramsar-technical-report-3-valuing-wetlands-guidance-for-valuing-the-benefits-derived-from 4. Sieben EJJ, Khubeka SP, Sithole S, Job NM, Kotze DC. The classification of wetlands: integration of top-down and bottom-up approaches and their significance for ecosystem service determination. Wetl Ecol Manag. 2018; 26: 441–458. https://doi.org/10.1007/S11273-017-9585-4 5. Finlayson CM, Groot RS de, Hughes FMR, Sullivan CA. Freshwater ecosystem services and func- tions. In: Hughes J, editor. Freshwater ecology and conservation: approaches and techniques. Oxford, University Press.; 2019. p. 464. Available: https://global.oup.com/academic/product/ freshwater-ecology-and-conservation-9780198766421?cc=br&lang=en&# 6. Junk WJ. Wetlands of tropical South America. In: Whigham D, Hejny S, Dykyjova D, editors. Wetlands of the World I. Dordrecht: Kluwer Academic Publishers; 1993. pp. 679–739. 7. Darrah SE, Shennan-Farpo´ n Y, Loh J, Davidson NC, Finlayson CM, Gardner RC, et al. Improvements to the Wetland Extent Trends (WET) index as a tool for monitoring natural and human-made wetlands.

Ecol Indic. 2019; 99: 294–298. https://doi.org/10.1016/J.ECOLIND.2018.12.032 8. Davidson NC, Dam AA van, Finlayson CM, McInnes RJ, Davidson NC, Dam AA van, et al. Worth of wetlands: revised global monetary values of coastal and inland wetland ecosystem services. Mar Freshw Res. 2019; 70: 1189–1194. https://doi.org/10.1071/MF18391 9. Parolin P. Submerged in darkness: adaptations to prolonged submergence by woody species of the Amazonian floodplains. Ann Bot. 2009; 103: 359–376. https://doi.org/10.1093/aob/mcn216 PMID: 19001429 10. Albert LP, Restrepo-Coupe N, Smith MN, Wu J, Chavana-Bryant C, Prohaska N, et al. Cryptic phenol- ogy in plants: Case studies, implications, and recommendations. Glob Chang Biol. 2019; 25: 3591– 3608. https://doi.org/10.1111/gcb.14759 PMID: 31343099 11. Parolin P, De Simone O, Haase K, Waldhoff D, Rottenberger S, Kuhn U, et al. Central Amazonian floodplain forests: Tree adaptations in a pulsing system. The Botanical Review. 2004; 70: 357–380. https://doi.org/10.1663/0006-8101(2004)070[0357:CAFFTA]2.0.CO;2 12. Correa SB, de Oliveira PC, Nunes da Cunha C, Penha J, Anderson JT. Water and fish select for fleshy fruits in tropical wetland forests. Biotropica. 2018; 50: 312–318. https://doi.org/10.1111/BTP.12524 13. Ivory SJ, McGlue MM, Spera S, Silva A, Bergier I. Vegetation, rainfall, and pulsing hydrology in the Pantanal, the world’s largest tropical wetland. Environmental Research Letters. 2019; 14. https://doi. org/10.1088/1748-9326/AB4FFE 14. Fonseca LDM, Dalagnol R, Malhi Y, Rifai SW, Costa GB, Silva TSF, et al. Phenology and seasonal

ecosystem productivity in an amazonian floodplain forest. Remote Sensing 2019, Vol 11, Page 1530. 2019; 11: 1530. https://doi.org/10.3390/RS11131530 15. De Simone O, Junk WJ, Schmidt W. Central Amazon floodplain forests: Root adaptations to prolonged flooding. Russian Journal of Plant Physiology 2003 50:6. 2003; 50: 848–855. https://doi.org/10.1023/ B:RUPP.0000003285.70058.4C 16. Wittmann F, Scho¨ngart J, Junk WJ. Phytogeography, species diversity, community structure and dynamics of central amazonian floodplain forests. 2010; 61–102. https://doi.org/10.1007/978-90-481- 8725-6_4 17. Nunes da Cunha C, Piedade MTF, Junk WJ. Classificac¸ão e delineamento das a´reas u´midas brasi- leiras e de seus macrohabitats. Cuiaba´: EdUFMT; 2015. 18. Junk WJ, da Cunha CN, Wantzen KM, Petermann P, Stru¨ ssmann C, Marques MI, et al. Biodiversity and its conservation in the Pantanal of Mato Grosso, Brazil. Aquat Sci. 2006; 68: 278–309. https://doi. org/10.1007/s00027-006-0851-4 19. Pott A, Pott VJ. Flora of the Pantanal. In: DAMASCENO-JUNIOR GA, POTT A, editors. Flora and veg- etation of the Pantanal wetland. Switzerland AG: Cham: Springer Nature, Plant and Vegetation; 2021. pp. 39–228. 20. Pott A, Da Silva JSV. Terrestrial and aquatic vegetation diversity of the Pantanal wetland. In: Bergier I, Assine ML, editors. Dynamics of the Pantanal Wetland in South America (Handbook of environmental chemistry serie). Springer Verlag; 2016. pp. 111–131. https://doi.org/10.1007/698_2015_352 21. Eiten G. Brazilian Savannas. B.J. Huntl. In Ecology of tropical savannas. B.J. Huntl. Springer Berlin

Heidelberg; 1982. pp. 25–47. https://doi.org/10.1007/978-3-642-68786-0_3 22. Aoki C, Faria RR, Damasceno-Junior GA, Pott A. Synthesis of the present knowledge on plant phenol- ogy of the Pantanal. 2021; 535–549. https://doi.org/10.1007/978-3-030-83375-6_13 PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 24 / 30

23. Jancoski HS, Schwantes Marimon B, C. Scalon M, de V. Barros F, Marimon-Junior BH, Carvalho E, et al. Distinct leaf water potential regulation of tree species and vegetation types across the Cerrado– Amazonia transition. Biotropica. 2022; 54: 431–443. https://doi.org/10.1111/BTP.13064 24. Wittmann F, Marques MCM, Damasceno Ju´ nior G, Budke JC, Piedade MTF, de Oliveira Wittmann A, et al. The Brazilian freshwater wetscape: Changes in tree community diversity and composition on cli- matic and geographic gradients. Silva LCR, editor. PLoS One. 2017; 12: e0175003. https://doi.org/10. 1371/journal.pone.0175003 PMID: 28394937 25. Marimon BS, Oliveira-Santos C, Marimon-Junior BH, Elias F, de Oliveira EA, Morandi PS, et al. Drought generates large, long-term changes in tree and liana regeneration in a monodominant Ama- zon forest. Plant Ecol. 2020; 221: 733–747. https://doi.org/10.1007/S11258-020-01047-8 26. Marengo J, Alves L, Torres R. Regional climate change scenarios in the Brazilian Pantanal watershed. Clim Res. 2016; 68: 201–213. https://doi.org/10.3354/cr01324 27. Correa SB, Costa-Pereira R, Fleming T, Goulding M, Anderson JT. Neotropical fish–fruit interactions: eco-evolutionary dynamics and conservation. Biological Reviews. 2015; 90: 1263–1278. https://doi. org/10.1111/brv.12153 PMID: 25599800 28. Tomas WM, Berlinck CN, Chiaravalloti RM, Faggioni GP, Stru¨ssmann C, Libonati R, et al. Distance sampling surveys reveal 17 million vertebrates directly killed by the 2020’s wildfires in the Pantanal,

Brazil. Sci Rep. 2021; 11. https://doi.org/10.1038/S41598-021-02844-5 PMID: 34916541 29. Malhi Y, Aragão LEOC, Galbraith D, Huntingford C, Fisher R, Zelazowski P, et al. Exploring the likeli- hood and mechanism of a climate-change-induced dieback of the Amazon rainforest. Proc Natl Acad Sci U S A. 2009; 106: 20610–20615. https://doi.org/10.1073/pnas.0804619106 PMID: 19218454 30. Junk WJ, Nunes De Cunha C. Pantanal: a large South American wetland at a crossroads. Ecol Eng. 2005; 24: 391–401. https://doi.org/10.1016/J.ECOLENG.2004.11.012 31. McGlue MM, Silva A, Assine ML, Stevaux JC, Pupim F do N. Paleolimnology in the Pantanal: Using lake sediments to track quaternary environmental change in the world’s largest tropical wetland. In: Bergier I, Assine ML, editors. Dynamics of the Pantanal wetland In South America, The handbook of environmental chemistry. Springer; 2015. pp. 51–81. https://doi.org/10.1007/698_2015_350 32. Ko¨ppen W. Climatologia: con un estudio de los climas de la tierra. Mexico; 1948. 33. Cadavid Garcia EA. O clima no Pantanal Mato-Grossense. Embrapa Unidade de Execucao de Pes- quisa de Ambito Estadual de Corumba. Corumba, MS: Corumba´ : EMBRAPA-UEPAE Corumba´ , 1984.; 1984. Available: http://www.infoteca.cnptia.embrapa.br/handle/doc/787712 34. Musis CR de, Campelo Ju´nior JHC, Priante Filhoe N. Caracterizac¸ão climatolo´gica da Bacia do Alto Paraguai. Geografia. 1997; 22: 5–21. Available: https://www.periodicos.rc.biblioteca.unesp.br/index. php/ageteo/article/view/14891

35. Hamilton S, McClain M. Hydrological controls of ecological structure and function in the Pantanal wet- land (Brazil). 2002. 36. Junk WJ, Bayley PB, Sparks RE. The flood-pulse concept in river-floodplain systems. Proceedings of the International Large River Symposium. Can. Spec. Publ. Fish. Aquat. Sci. Canadian Special Publi- cations for Fisheries and Aquatic Sciences.; 1989. pp. 110–127. 37. Fantin-Cruz I, Pedrollo O, Castro NMR, Girard P, Zeilhofer P, Hamilton SK. Historical reconstruction of floodplain inundation in the Pantanal (Brazil) using neural networks. J Hydrol (Amst). 2011; 399: 376– 384. https://doi.org/10.1016/j.jhydrol.2011.01.014 38. Assine ML, Soares PC. Quaternary of the Pantanal, west-central Brazil. Quaternary International. 2004; 114: 23–34. https://doi.org/10.1016/S1040-6182(03)00039-9 39. Alvarenga SM, Brasil AE, Pinheiro R, Kux HJH. Estudo geomorfolo´ gico aplicado à Bacia do alto Rio Paraguai e Pantanais Matogrossenses. Boletim Te´ cnico Projeto RADAM/ BRASIL Se´ rie Geomorfolo- gia Salvador. 1984; 187: 89–183. 40. Brasil. Folha SE. 21 Corumba e parte da folha SE.20: geologia, geomorfologia, pedologia, vegetac¸ão, uso potencial da terra / Projeto RADAMBRASIL [v. 27]. Rio de Janeiro; 1982. 41. Couto EG, Corrêa GR, Oliveira VA, do Nascimento AF, Vidal-Torrado P, Beirigo R, et al. Soils of Panta- nal: The Largest Continental Wetland. 2023; 239–267. https://doi.org/10.1007/978-3-031-19949-3_9 42. Coringa EDAO, Couto EG, Torrado PV. Geoquı´mica de solos do pantanal norte, Mato Grosso. Rev

Bras Cienc Solo. 2014; 38: 1784–1793. https://doi.org/10.1590/S0100-06832014000600013 43. Assine ML, Merino ER, Pupim F do N, Macedo H de A, Santos MGM dos, Assine ML, et al. The Qua- ternary alluvial systems tract of the Pantanal Basin, Brazil. Brazilian Journal of Geology. 2015; 45: 475–489. https://doi.org/10.1590/2317-4889201520150014 44. Thielen D, Schuchmann KL, Ramoni-Perazzi P, Marquez M, Rojas W, Quintero JI, et al. Quo vadis Pantanal? Expected precipitation extremes and drought dynamics from changing sea surface temper- ature. PLoS One. 2020; 15. https://doi.org/10.1371/journal.pone.0227437 PMID: 31910441 PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 25 / 30

Pantanal of Mato Grosso, Brazil. J Biogeogr. 2000; 27: 159–168. https://doi.org/10.1046/J.1365-2699. 2000.00357.X 53. Pott VJ, Pott A, Lima LCP, Moreira SN, Oliveira AKM. Aquatic macrophyte diversity of the Pantanal wetland and upper basin. Brazilian Journal of Biology. 2011; 71: 255–263. https://doi.org/10.1590/ s1519-69842011000200004 PMID: 21537598 54. Nunes da Cunha C, Bergier I, Tomas WM, Damasceno-Junior GA, Santos SA, Assunc¸ão VA, et al. Hydrology and Vegetation Base for Classification of Macrohabitats of the Brazilian Pantanal for Policy- Making and Management. 2021; 365–391. https://doi.org/10.1007/978-3-030-83375-6_7 55. Ferreira-Ju´nior WG, Schaefer CEGR, Cunha CN, Duarte TG, Chieregatto LC, Carmo FMS. Flood regime and water table determines tree distribution in a forest-savanna gradient in the Brazilian panta- nal. An Acad Bras Cienc. 2016; 88: 719–731. https://doi.org/10.1590/0001-3765201620150341 PMID: 27142551 56. Arieira J, Nunes da Cunha C. Fitossociologia de uma floresta inunda´vel monodominante de Vochysia divergens Pohl (Vochysiaceae), no Pantanal Norte, MT, Brasil. Acta Bot Brasilica. 2006; 20: 569–580. https://doi.org/10.1590/S0102-33062006000300007 57. Damasceno-Junior GA, Semir J, Dos Santos FAM, Leitão-Filho HDF. Structure, distribution of species and inundation in a riparian forest of Rio Paraguai, Pantanal, Brazil. Flora. 2005; 200: 119–135. https://doi.org/10.1016/j.flora.2004.09.002 58. Morellato LPC, Rodrigues RR, Leitao Filho HF, Joly CA. Estudo comparativo da fenologia de espe´ cies

arbo´reas de floresta de altitude e floresta mesofila semidecidua na serra do Japi, Jundiaı´, São Paulo. Revista Brasileira de Botaˆ nica. 1989; 12: 85–98. Available: https://repositorio.usp.br/item/000796966 59. Bencke CSC, Morellato LPC. Comparac¸ão de dois me´ todos de avaliac¸ão da fenologia de plantas, sua interpretac¸ão e representac¸ão. Brazilian Journal of Botany. 2002; 25: 269–275. https://doi.org/10. 1590/S0100-84042002000300003 60. Fournier LA. Un me´todo cuantitativo para la medicio´ n de caracterı´sticas feno´ logicas en a´ rboles. Tur- rialba. 1974; 24: 422–423. 61. Talora DC, Morellato PC. Fenologia de espe´ cies arbo´reas em floresta de planı´cie litoraˆ nea do sudeste do Brasil. Brazilian Journal of Botany. 2000; 23: 13–26. https://doi.org/10.1590/S0100- 84042000000100002 62. Morellato LPC, Alberti LF, Hudson IL. Applications of circular statistics in plant phenology: A case stud- ies approach. In: Hudson I, Keatley M, editors. Phenological Research. Dordrecht: Springer Nether- lands; 2010. pp. 339–359. https://doi.org/10.1007/978-90-481-3335-2_16 63. Camargo MGG de, Carvalho GH de, Alberton B de C, Reys P, Morellato LPC. Leafing patterns and leaf exchange strategies of a cerrado woody community. Biotropica. 2018; 50: 442–454. https://doi. org/10.1111/BTP.12552 64. Oliveira PO. Fenologia e biologia reprodutiva das espe´cies de cerrado. In: Sano SM, Almeida SP, edi- tors. Cerrado: ambiente e flora. Planaltina: Embrapa-CPAC; 1998. pp. 169–192. 65. Zar JH. Bioestatistical Analysis. New Jersey: Prentice Hall; 1999. PLOS ONE

Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 26 / 30

66. Batalha MA, Martins FR. Reproductive phenology of the cerrado plant community in Emas National Park (central Brazil). Aust J Bot. 2004; 52: 149–161. https://doi.org/10.1071/BT03098 67. Kovach WL. Oriana for Windows. Pentraeth, UK: Kovach Computer Services.; 1994. 68. Bendix J, Homeier J, Cueva Ortiz E, Emck P. Seasonality of weather and tree phenology in a tropical evergreen mountain rain forest Plant-Soil Systems: Interactions and Feedbacks View project Nature 4.0 | Sensing Biodiversity View project. Article in International Journal of Biometeorology. 2006 [cited 16 Aug 2022]. https://doi.org/10.1007/s00484-006-0029-8 PMID: 16598482 69. Pires JPA, Marino NAC, Silva AG, Rodrigues PJFP, Freitas L. Tree community phenodynamics and Its relationship with climatic conditions in a lowland tropical rainforest. Forests. 2018; 9: 114. https:// doi.org/10.3390/F9030114 70. Perina BB, Liboni AP, Montanher DR, Messetti AVL, Pimenta JA, Bianchini E. Phenology of the tree community in a seasonal forest in southern Brazil. Brazilian Journal of Botany 2019 42:3. 2019; 42: 477–490. https://doi.org/10.1007/S40415-019-00545-Y 71. Agência Nacional De A´ guas–ANA. Hidroweb: Sistemas de informac¸ões hidrolo´gicas. In: ANA [Inter- net]. 26 Nov 2018 [cited 18 Nov 2018]. Available: http://hidroweb.ana.gov.br/ 72. INSTITUTO NACIONAL DE METEOROLOGIA—INMET. Normais Climatolo´ gicas do Brasil. [cited 3 Aug 2024]. Available: https://portal.inmet.gov.br/ normais 73. Salis SM, Lehn CR, Mattos PP, Bergier I, Crispim SMA. Root behavior of savanna species in Brazil’s

Pantanal wetland. Glob Ecol Conserv. 2014; 2: 378–384. https://doi.org/10.1016/j.gecco.2014.10.009 74. Shivaprakash KN, Ramesh BR, Umashaanker R, Dayanandan S. Functional trait and community phy- logenetic analyses reveal environmental filtering as the major determinant of assembly of tropical for- est tree communities in the Western Ghats biodiversity hotspot in India. For Ecosyst. 2018; 5: 1–16. https://doi.org/10.1186/S40663-018-0144-0/FIGURES/5 75. Pott A, Pott JV. Vegetac¸ão do Pantanal: fitogeografia e dinaˆ mica. Anais do 2o Simpo´ sio de Geotecno- logias no Pantanal. Corumba´ : Embrapa Informa´tica Agropecua´ ria/INPE; 2009. pp. 1065–1076. 76. Neves SPS, Mendes dos Santos MG, Vito´ ria AP, Rossatto DR, de Miranda L d AP, Funch LS. The roles of functional traits in canopy maintenance along a savanna/seasonally dry tropical forest gradient in northeastern Brazil. Flora: Morphology, Distribution, Functional Ecology of Plants. 2022; 292. https://doi.org/10.1016/J.FLORA.2022.152090 77. Nunes da Cunha C, Junk WJ, Leitão-Filho HF. Woody vegetation in the Pantanal of Mato Grosso, Bra- zil: a preliminary typology. Amazoniana. 2007; 19: 159–184. 78. Wagner FH, He´rault B, Rossi V, Hilker T, Maeda EE, Sanchez A, et al. Climate drivers of the Amazon forest greening. PLoS One. 2017; 12: e0180932. https://doi.org/10.1371/journal.pone.0180932 PMID: 28708897 79. Cabral OMR, da Rocha HR, Gash JH, Freitas HC, Ligo MAV. Water and energy fluxes from a wood- land savanna (cerrado) in southeast Brazil. J Hydrol Reg Stud. 2015; 4: 22–40. https://doi.org/10. 1016/J.EJRH.2015.04.010

80. da Silva JB, Valle Junior LCG, Faria TO, Marques JB, Dalmagro HJ, Nogueira JS, et al. Temporal vari- ability in evapotranspiration and energy partitioning over a seasonally flooded scrub forest of the Bra- zilian Pantanal. Agric For Meteorol. 2021; 308–309: 108559. https://doi.org/10.1016/J.AGRFORMET. 2021.108559 81. Dalmagro HJ, de A. Lobo F, Vourlitis GL, Dalmolin Aˆ C, Antunes MZ, Ortı´z CER, et al. Photosynthetic response of a wetland- and an upland-adapted tree species to seasonal variations in hydrology in the Brazilian Cerrado and Pantanal. Acta Physiologiae Plantarum 2016 38:5. 2016; 38: 1–10. https://doi. org/10.1007/S11738-016-2125-7 82. Wright CL, de Lima ALA, de Souza ES, West JB, Wilcox BP. Plant functional types broadly describe water use strategies in the Caatinga, a seasonally dry tropical forest in northeast Brazil. Ecol Evol. 2021; 11: 11808–11825. https://doi.org/10.1002/ece3.7949 PMID: 34522343 83. Scho¨ ngart J, Piedade MTF, Ludwigshausen S, Horna V, Worbes M. Phenology and stem-growth peri- odicity of tree species in Amazonian floodplain forests. J Trop Ecol. 2002; 18: 581–597. https://doi.org/ 10.1017/S0266467402002389 84. Parolin P, Wittmann F. Struggle in the flood: tree responses to flooding stress in four tropical floodplain systems. AoB Plants. 2010; 2010. https://doi.org/10.1093/aobpla/plq003 PMID: 22476061 85. Parolin P, Lucas C, Piedade MTF, Wittmann F. Drought responses of flood-tolerant trees in Amazo- nian floodplains. Ann Bot. 2010; 105: 129. https://doi.org/10.1093/aob/mcp258 PMID: 19880423

86. Oliveira AKM, Gualtieri SCJ. Trocas gasosas em plantas jovens de Tabebuia aurea (Bignoniaceae Juss.) submetidas a estresse por alagamento. Revista Arvore. 2016; 40: 39–49. https://doi.org/10. 1590/0100-67622016000100005 PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 27 / 30

87. Manzoni S, Vico G, Thompson S, Beyer F, Weih M. Contrasting leaf phenological strategies optimize carbon gain under droughts of different duration. Adv Water Resour. 2015; 84: 37–51. https://doi.org/ 10.1016/J.ADVWATRES.2015.08.001 88. Chaturvedi RK, Tripathi A, Raghubanshi AS, Singh JS. Functional traits indicate a continuum of tree drought strategies across a soil water availability gradient in a tropical dry forest. For Ecol Manage. 2021; 482: 118740. https://doi.org/10.1016/J.FORECO.2020.118740 89. Lacerda DMA, Rossatto DR, Ribeiro-Novaes E´ KMD, de Almeida EB. Reproductive phenology differs between evergreen and deciduous species in a Northeast Brazilian savanna. Acta Bot Brasilica. 2018; 32: 367–375. https://doi.org/10.1590/0102-33062017ABB0343 90. Arau´ jo I, Marimon BS, Scalon MC, Fauset S, Marimon BH, Tiwari R, et al. Trees at the Amazonia-Cer- rado transition are approaching high temperature thresholds. Environmental Research Letters. 2021; 16: 034047. https://doi.org/10.1088/1748-9326/ABE3B9 91. Girard P, Da Silva CJ, Abdo M. River–groundwater interactions in the Brazilian Pantanal. The case of the Cuiaba´ River. J Hydrol (Amst). 2003; 283: 57–66. https://doi.org/10.1016/S0022-1694(03)00235-X 92. Borchert R. Soil and stem water storage determine phenology and distribution of tropical dry forest trees. Ecology. 1994; 75: 1437–1449. https://doi.org/10.2307/1937467 93. Bordiert R. Water status and development of tropical trees during seasonal drought. Trees. 1994; 8: 115–125. https://doi.org/10.1007/BF00196635/METRICS

94. Chen S, Stark SC, Nobre AD, Cuartas LA, de Jesus Amore D, Restrepo-Coupe N, et al. Amazon forest biogeography predicts resilience and vulnerability to drought. Nature 2024 631:8019. 2024; 631: 111– 117. https://doi.org/10.1038/s41586-024-07568-w PMID: 38898277 95. Van Schaik CP, Terborgh JW, Wright SJ. The phenology of tropical forests: Adaptive significance and consequences for primary consumers. 2003; 24: 353–377. https://doi.org/10.1146/ANNUREV.ES.24. 110193.002033 96. Araujo JM, Correa SB, Anderson J, Penha J. Fruit preferences by fishes in a Neotropical floodplain. Biotropica. 2020; 52: 1131–1141. https://doi.org/10.1111/BTP.12790 97. Bentos T V., Mesquita RCG, Williamson GB. Reproductive phenology of central Amazon pioneer trees. Trop Conserv Sci. 2008; 1: 186–203. https://doi.org/10.1177/194008290800100303 98. Morellato LPC, Camargo MGG, Gressler E. A review of plant phenology in south and central america. Phenology: An Integrative Environmental Science. Springer Netherlands; 2013. pp. 91–113. https:// doi.org/10.1007/978-94-007-6925-0_6 99. Araujo MM V, Lobo FA. Phenology of copernicia alba in flooded and not flooded environments. Flor- esta e Ambiente. 2020; 27: 0–0. https://doi.org/10.1590/2179-8087.097917/PDF/FLORAM-27-1- E20170979.PDF 100. Caballero CB, Ruhoff A, Biggs T. Land use and land cover changes and their impacts on surface- atmosphere interactions in Brazil: A systematic review. Sci Total Environ. 2022; 808. https://doi.org/ 10.1016/j.scitotenv.2021.152134 PMID: 34864033

101. Lopezaraiza-Mikel M, Quesada M, A´ lvarez-Añorve M, A´ vila-Cabadilla L, Marte´ n-Rodrı´guez S, Calvo- Alvarado J, et al. Phenological patterns of tropical dry forests along latitudinal and successional gradi- ents in the neotropics. Tropical Dry Forests in the Americas: Ecology, Conservation, and Manage- ment. 2013; 101–128. https://doi.org/10.1201/B15417-10/Phenological-patterns-tropical-dry-forests- along-latitudinal-successional-gradients-neotropics-martha-lopezaraiza-mikel-mauricio-quesada- mariana- 102. Morellato LPC, Talora DC, Takahasi A, Bencke CC, Romera EC, Zipparro VB. Phenology of Atlantic rain forest trees: A comparative study. Biotropica. 2000; 32: 811–823. https://doi.org/10.1111/j.1744- 7429.2000.tb00620.x 103. Fava WS, Covre W da S, Sigrist MR. Attalea phalerata and Bactris glaucescens (Arecaceae, Arecoi- deae): Phenology and pollination ecology in the Pantanal, Brazil. Flora—Morphology, Distribution, Functional Ecology of Plants. 2011; 206: 575–584. https://doi.org/10.1016/J.FLORA.2011.02.001 104. Ferreira LV, Parolin P. Tree phenology in Central Amazonian floodplain forests: Effects of water level fluctuation and precipitation at community and population level. Pesquisas, Botaˆ nica. 2007; 58: 139– 156. 105. Shen JX, Zhang YJ, Maenpuen P, Zhang S Bin, Zhang L, Yang L, et al. Response of four evergreen savanna shrubs to an incidence of extreme drought: high embolism resistance, branch shedding and maintenance of nonstructural carbohydrates. Tree Physiol. 2022; 42: 740–753. https://doi.org/10. 1093/treephys/tpab150 PMID: 35020937

106. Numata S, Yamaguchi K, Shimizu M, Sakurai G, Morimoto A, Alias N, et al. Impacts of climate change on reproductive phenology in tropical rainforests of Southeast Asia. Communications Biology 2022 5:1. 2022; 5: 1–10. https://doi.org/10.1038/s42003-022-03245-8 PMID: 35449443 PLOS ONE Phenological cycles in the Pantanal woody communities PLOS ONE | https://doi.org/10.1371/journal.pone.0316011 February 7, 2025 28 / 30