Ecology

Basic information of intervieweesResults of the descriptive analysis summarized in Table 2 show that more than half of the respondents were males (69%) and were on average 41.3 years old while more than 32 years of farming experience. The study area is comprised of multiple ethnic groups (Han, Tibetan, Yugur, Mongolian, Hui, etc.). In most cases, the main livelihood activity of the Ethnic Minorities (Tibetan, Yugur, Mongolian, Hui, etc.) is livestock, while Han people main livelihood activity is farming. The majority of respondents (64%) were minority nationality. The vast majority of the agro-pastoralists (86%) have a primary school education or above, even though only 1% of them have Undergraduate education or Above. The results also reveal that 92% of respondents have access to weather information. The average cultivated land Per household is 10.23 Mu and Grassland is 156.21 Mu, respectively. The average per household income is RMB78000, and agricultural income is RMB52000.Table 2 Descriptive statistics of agro-pastoralist characteristics.Full size tableDue to their long-term farming experience, the agro-pastoralists were expected to have a high-level of understanding of local climate knowledge. Also contributing to this could be the information they receive about climate change and for some, the associated training through agro-pastoralists’ associations. Therefore, they also have a propensity to adapt to adverse conditions resulting from climate change impacts. In addition, the high-level of farming experience, the cultivated-land size, grassland size, Credit loan, Insurance, Village cadres all have a positive impact on the level of agro-pastoralists’ adaptation to new climate scenarios.However, the education level and cadres experience may be the major limiting factors for adopting specific long-term adaptation strategies. Ethnicity and gender are also expected to be key factors influencing awareness and adaptation to climate change. There are differences in relative perception intensity between Ethnic Minority and Han because of their cultural ecology (the main livelihood activity of minorities nationality is livestock, while Han main livelihood activity is farming.). In terms of gender, women in rural areas are less mobile and have less access to information and rights. They are also heavily involved in domestic work. However, men may have easier access to information (socializing, going out to work, etc.) Therefore, male headed households are expected to be more likely to adapt to the impact of climate change.Climate change trend in the study areaFigure 2 shows the trend of annual precipitation, annual rainfall and annual snow at different meteorological stations in the study area. As shown in the Fig. 2, precipitation, rainfall and snow show an increasing trend, but the increase range of snow (0.0325–0.375/a) is significantly lower than that of precipitation (1.22–3.1/a) and rainfall (1.04–2.81/a). Similarly, through the inspection, it is found that the multi-collinearity among precipitation, rainfall and snow at each meteorological station is obvious (most R2  > 0.5, and p  More

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Experimental designThis experiment was set up containing two levels of soil biodiversity (high and low soil biodiversity) and seven treatments considering the number of global change factors (GCFs) (0, 1, 2, 4, 6, 8, 10) (Table 1, Supplementary Fig. 1 and Supplementary Data 1). We used the dilution-to-extinction approach to create the high and low soil biodiversity treatments (Supplementary Methods). Soil dilution can lead to a gradual loss of rare soil microbes, which can simulate a realistic loss of soil biodiversity, because rare soil microbes are more sensitive to anthropogenic pressures, e.g., warming, nitrogen addition and drought15. We note that the low soil biodiversity treatment is a subset of the high biodiversity, as many rare species have been eliminated through the dilution; this approach will likely lead to relatively more tolerant microbes in the resulting communities.An increasing number of GCFs was created inspired by the experimental design of the studies on biodiversity-ecosystem function relationships, based on random sampling from a species pool5,6,14. The combination of multiple GCFs was replicated 15 times at each level by randomly selecting GCFs from a pool with 10 GCFs for each replicate (Table 1 and Supplementary Data 1). For each replicate of combined GCFs, there were identical GCF combinations between the high and low soil biodiversity treatments to avoid a confounding effect of GCF combination and soil biodiversity treatments. The pool of 10 GCFs included: warming, nitrogen deposition, drought, heavy metal pollution, plastic mulching film residues, salinity, agricultural fungicide, bactericide application, surfactant contaminant and soil compaction (Supplementary Methods). These GCFs frequently occur in intensively managed agroecosystems and are treated as anthropogenic pressures10,13,14,15.MicrocosmsThis experiment was conducted using 50 ml conical Mini Bioreactors (Product Number 431720, Corning Inc., NY) as experimental units (Supplementary Fig. 2). Each Mini Bioreactor has four vents in the cap, where a hydrophobic membrane avoids microbial contamination but allows gas exchange. We filled each Mini Bioreactor with 40.0 g (dry weight, d.w.) of soil in total, which received the appropriate treatments.Soil sterilization and inoculum preparationWe collected the field soil from the top 10 cm of an intensive farming system in Berlin (52.466°N, 13.303°E). Field soil was passed through a 2 mm mesh to remove large roots and stones. We sterilized 20 kg of soil for 90 min at 121 °C, and stored 2 kg of fresh soil at 4 °C. The dilution-to-extinction approach38,39,40,41,42 was used to create high and low soil biodiversity (Supplementary Fig. 1). A parent inoculum suspension was prepared by mixing 100 g of fresh soil with 200 ml of sterilized VE water. The sediment settled for 1 min. The upper 200 ml of soil suspension was treated as parent inoculum suspension. 50 ml of parent inoculum suspension was added to 500 g of sterilized soil in a plastic bag, and homogenized by turning the bag up and down 30 times to obtain the inoculum of high soil biodiversity. Another 5 ml of parent inoculum suspension was mixed with 45 ml of sterilized parent inoculum suspension to create the 10-1 dilution. This procedure was repeated five times to reach the 10-6 dilution. 50 ml of the 10-6 dilution was mixed and homogenized with 500 g of sterilized soil in a plastic bag to obtain the inoculum of low soil biodiversity. This whole dilution procedure was repeated five times to obtain 10 bags of soil inoculum (five bags for each soil biodiversity inoculum).Sterile water was added to each plastic bag to reach the water content of the fresh soil in the field. All bags were closed with a sterilized cotton plug and a rubber band to avoid microbial contamination but allow gas exchange42. All bags were incubated in a dark room at 20 °C until similar microbial abundance was observed between the high and low soil biodiversity inoculum. Soil inoculum was homogenized by shaking and turning the bags once a week. After incubation, 2.0 g of soil in each bag was collected and stored at −80 °C for DNA extraction. Quantitative real-time PCR (qPCR) was used to determine fungal and bacterial abundance. In the present study, it took two months to recover soil microbial biomass (Supplementary Fig. 3).The implementation of GCFs and harvestAgroecosystems, some of the most intensively managed ecosystems, are affected by the co-occurrence of multiple GCFs13,14,15. This study focused on GCFs that frequently occur in agroecosystems, including warming, nitrogen deposition, drought, heavy metal pollution, plastic film residues, salinity, agricultural fungicide and bactericide application, surfactant contaminant and soil compaction. We present the rationale for the 10 tested GCFs in the Supplementary Methods.Loading soils were used to achieve an effective mixing of chemical agents into 40.0 g soil in each Mini Bioreactor. We created separate ‘loading soil’ for each GCF with chemical addition by mixing an appropriate dose of a chemical agent with sterilized soil through careful homogenization. This was done to avoid exaggerated effects of more concentrated chemicals when mixed with soil. For each chemical, 1.0 g (d.w.) of loading soil contained an appropriate dose for 40.0 g soil in a Mini Bioreactor. For instance, 1 634 mg of NH4NO3 was mixed with 100 g (d.w.) of sterilized soil, to ensure that there was about 16.34 mg of NH4NO3 in 1.0 g of sterilized loading soil. We weighed 40.0 g (d.w.) of soils, including 1.0 g (d.w.) of each loading soil, 5.0 g (d.w.) of soil inoculum (high or low soil biodiversity), an appropriate amount of film (0 or 0.16 g plastic film) and sterilized soil, according to GCF combination for each experimental unit. We put 40.0 g of mixed soils into a clean and sterilized cup (200 ml) with a cap, and then homogenized it by turning the cup up and down for 5 min using a shaking machine (Heidolph Reax 2, Heidolph Instruments GmbH & CO. KG, Schwabach, Germany). After homogenization, 40.0 g of mixed soils was transferred to a Mini Bioreactor, and a mesh bag containing about 100 mg of dry Medicago lupulina leaves (65 °C for 72 h) was buried 1 cm below the soil surface. We used a stick to press soils in each Mini Bioreactor to simulate an ambient condition (1.3 g cm−3) or mechanical compaction (1.7 g cm−3) in farmland.For the warming treatment with an increment of 5.0 °C over the ambient temperature (20 °C), we wrapped heating cables (Exo Terra PT-2012; Hagen Deutschland GmbH & Co. KG, Holm, Germany) around the outside of the bioreactors. A set temperature was maintained by temperature controllers (Voltcraft ETC-902; Conrad Electronic SE, Hirschau, Germany), which can switch off and on heating cables depending on the real-time temperature of the outside surface of Mini Bioreactors. Mini Bioreactors were placed in beakers filled with sand to reduce thermal radiation from neighboring units. At the start of the experiment, we added suitable amounts of sterilized water to each experimental unit to reach 60% of water holding capacity for the non-drought treatment and 30% water holding capacity for the drought treatment.All Mini Bioreactors were incubated at 20.0 °C in the dark for six weeks before the final harvest. Because there was 2.0 g of weight loss on average in each Mini Bioreactor in the first three weeks, we added 2 ml of sterilized water to each Mini Bioreactor on the first day of the fourth week. During the final harvest, soil in each Mini Bioreactor was gently homogenized using a spoon, and then 2.0 g of fresh soil was collected and stored at −80 °C for DNA extraction; 5.0 g was stored at 4 °C for the determination of soil enzyme activity; the leftover was oven-dried at 40 °C for other measurements. DNA of each soil sample was extracted from 250 mg soil, using DNeasy PowerSoil Pro Kit (QIAGEN GmbH, Hilden, Germany), following manufacturer’s instructions. Soil DNA extraction was stored at −80 °C for further analysis.The measurement of response variablesWe measured the following response variables: microbial activity (soil respiration), microbial abundance (bacterial and fungal abundance), nutrient cycling (litter decomposition rate and soil enzyme activity), physical properties (water-stable soil aggregates and soil water repellency), bacterial and fungal biodiversity (richness and microbial network features) (See details in the Supplementary Methods). We measured soil respiration as CO2 concentration (ppm h−1 g−1 soil) in the third and sixth week as an indicator for soil microbial activity. Bacterial and fungal abundance was estimated by qPCR. The proportional loss of litter (Medicago lupulina leaves) dry weight during soil incubation was used as an indication of decomposition rate. We measured the activity of β-glucosidase (cellulose degradation), β-D-celluliosidase (cellulose degradation), N-acetyl-β-glucosaminidase (chitin degradation) and phosphatase (organic phosphorus mineralization) using high throughput microplate assay43,44. A modified protocol by Kemper and Rosenau was used to measure water-stable soil aggregates45. Soil water repellency was measured using the water drop penetration time method46. High throughput sequencing (Illumina MiSeq) was used to measure the taxonomic composition of soil fungal and bacterial communities with the primers fITS7 and ITS4 for fungi and 515F-Y and 806 R for bacteria47,48 (Supplementary Methods).Statistical analysesFor diversity and community composition analysis, we excluded the samples with less than 1% of the observations of the largest sample in the ASV table. For network analysis, we then removed ASVs with low prevalence, which presented less than 20% of samples across all experimental units to reduce the high percentage of zero counts. A co-occurrence network was constructed based on both fungal and bacterial ASV tables. The PLNnetwork function in the R package PLNmodels was employed to infer the network, using a sparse multivariate Poisson log-normal (PLN) model49. According to the Extended Bayesian Information Criterion (EBIC), the best model was extracted with the function getBestModel. The network was compartmentalized into different modules using the cluster_fast_greedy function in the igraph package and visualized with partial correlations with |ρ| > 0.05. We focused on the response of the relative abundance of modules, also known as clusters, which represent the closely associated microbes, e.g., groups of coexisting or co-evolving microbes27. The relative abundance of modules was calculated by summing relative abundances for individual ASVs in modules. We used the package FUNGuildR50 to taxonomically parse fungal trait information, using the FUNGuild database51.For each single GCF treatment, we took the average response from the 10 replicates before analysis. To confirm how the effect of soil biodiversity treatment can change along with the increasing number of GCFs, we quantified the effect size of soil biodiversity treatment for each response variable using Hedges’ g (mean and 95% CIs) at each level of the number of GCFs, using the R package effsize52. Hedges’ g is calculated as the mean difference between the high and low soil biodiversity treatments in units of the pooled standard deviation as a paired-samples because there were identical GCFs and GCF combinations for both high and low biodiversity conditions, with the exception of the zero and 10 GCF treatments.To evaluate how each of the response variables changes along with the number of GCFs, we applied a generalized additive model (GAM)53. GAM is a penalized generalized linear model that fits a nonparametric, nonlinear smooth curve54. The degree of smoothness of model terms is estimated as part of fitting, using the generalized cross validation. We reasonably assume that the curve shapes are different between high and low soil biodiversity treatments. Therefore, we included biodiversity conditions (low/high) as the model intercept and as the “factor smooth” smoothing class, where a smooth function is created for each factor level independently55. For GAM modeling, we used the mgcv package55. The dimension of the basis used to represent the smooth term was set as k = 5 so that the model does not overfit to the data. For this, we compared some other values (from 3 to 8) and confirmed that the results are essentially the same within the tested range. The other parameters were set as default.The relationships between soil microbial indices and other soil indices were tested using Spearman correlation in the package microbiome, and the adjustment method “fdr” was employed to control the false discovery rate for multiple testing correction56. For the further multivariate integration of soil functions/properties and composition of modules, the DIABLO (Data Integration Analysis for Biomarker discovery using a Latent component method for Omics studies) was employed to detect correlation (Pearson’s correlation |r| > 0.5) among variables using the package mixOmics57.The Z-scores for each of the eight soil functions (as shown in Fig. 1, with the exception of soil water repellency) were evaluated, and then we computed an improved weighted multifunctionality metric to represent soil multifunctionality (Supplementary Methods)58. Structural equation models (SEMs) were used to reveal the direct and indirect effects of an increasing number of GCFs on soil multifunctionality within each soil biodiversity treatment using the package lavaan59. We assumed that an increasing number of GCFs influences soil multifunctionality by regulating the bacterial and fungal abundance and the relative abundance of modules. All response variables were standardized to the same comparison scale using the z-score transformation before constructing SEMs. Models with optimal fitting indices were reported (Supplementary Fig. 11).The permutational multivariate analysis (ADONIS) and non-metric multidimensional scaling (NMDS) ordination based on the Bray-Curtis distance were conducted to test the effect of soil biodiversity and GCF treatments on the community composition of bacteria and fungi using the R package vegan60. For the data handling, processing, and visualization, we used the packages tidyverse61, reshaping62, cowplot63, RColorBrewer64, qgraph65, igraph66, factoextra67, phyloseq68 and itsadug69. These data manipulation and analyses were conducted using R version 4.1.370. The R script and data are available in a publicly accessible database71.Reporting summaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More

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The pre-evaporitic, evaporitic, and post-evaporitic phases are recognized for the late Aptian. These phases are recorded within the K40–K50 sequences (Fig. 2A), and show an average maximum thickness of approximately 650 m in the studied basins. The pre-evaporitic phase is represented by carbonate and siliciclastic deposits formed in fluvial and lacustrine deltaic environments within a large proto-oceanic gulf28 (Fig. 2A). The peak of the evaporitic deposition is recorded in the K50 sequence, with widespread occurrences in the Brazilian equatorial margin. The origin of these deposits is the heat intensification associated with the widening of the Atlantic Ocean. These conditions caused strong evaporation leading to a wide distribution of evaporites (mainly halite and anhydrite gypsum) in the South Atlantic basins. The eastern continental margin of Brazil contains a restricted marine section characterized by evaporites, which are particularly prominent in thickness and occurrence in the Espírito Santo Basin (Itaúnas Member of the Mariricu Formation) and the Sergipe Basin (the Ibura Member of the Muribeca Formation)28. Evaporites form the most prominent evidence of dry climates in the South Atlantic basins11, with evaporation exceeding precipitation. The post-evaporitic phase is characterized by fully marine conditions evidenced by rich assemblages of marine fossils. During this phase, carbonates were deposited, followed by muddy and sandy sediments in shallow-marine and slope environments.Figure 2Paleoclimatic phases scheme and principal component analysis for paleoclimatic phases. (A) Paleoclimatic phases scheme for the late Aptian and the main depositional environments. (B) Principal component plot of bioclimatic groups. (C) Principal component for the pre-evaporitic phase (N = 92), evaporitic phase (N = 78), and post-evaporitic phase (N = 385); see Supplementary Fig. 9 for individual basins.Full size imagePaleovegetationWe identified a rich plant community with 139 spore and pollen genera/morphotypes representing all plant groups: bryophytes (five genera), ferns (58 genera), lycophytes (18 genera), pteridosperms (one genus), gymnosperms (27 genera), and angiosperms (30 genera) (Supplementary Table 2). The inferred systematic affinities at the family level reached 100% in bryophytes, 56.9% in ferns, 100% in lycophytes, 100% in pteridosperms, 92.6% in gymnosperms, and 40.0% in angiosperms, totaling 67.6% of the recorded genera (Supplementary Table 2). Marine elements (e.g., dinoflagellate cysts and microforaminiferal linings) were identified, in particular from the Sergipe and Araripe basins (Fig. 1). Pollen grains from gymnosperms were most abundant, represented mainly by the conifer families Cheirolepidiaceae, Araucariaceae, and Podocarpaceae, although representing different climatic settings. Classopollis (Cheirolepidiaceae) is the most abundant genus in all sections studied, followed by Araucariacites (Araucariaceae). Gymnosperms showed low diversity. Spore-producing plants are the most diverse in the assemblages of all basins (82 genera) and represented by several families of bryophytes, ferns, and lycophytes (e.g., Sphagnaceae, Anemiaceae, Cyatheaceae, Marsileaceae, Selaginellaceae, and Lycopodiaceae). These plant groups depend on water to reproduce and are therefore associated with humid settings.Cicatricosisporites (Anemiaceae) is the third most abundant palynomorph in all the basins, but especially in the northeastern basins (e.g., Sergipe Basin). Angiosperms are among the least abundant; however, they are diverse and include the most abundant and controversial genus Afropollis, herein attributed to angiosperms. In the most recent publication that addressed this question, ref.29 suggest that Afropollis should be treated as an angiosperm genus, although without more precise systematic assignment. The 30 genera/morphotypes of angiosperms are assigned to 8 families, viz., Arecaceae, Chloranthaceae, Euphorbiaceae, Flacourtiaceae, Illiciaceae, Liliaceae, Solanaceae and Trimeniaceae. The second most abundant genus is Stellatopollis also without precise systematic assignment.Spatio-temporal distribution of bioclimatic groupsOn the basis of their botanical affinities, most taxa were classified into five bioclimatic groups [see “Methods” section and Supplementary information], viz., hydrophytes, hygrophytes, tropical lowland flora, upland flora, and xerophytes (Supplementary Table 2) (Fig. 3).Figure 3Relevant palynomorphs of bioclimatic groups: (1) Aequitriradites sp.; (2) Crybelosporites sp.; (3) Perotriletes sp.; (4) Cicatricosisporites sp.; (5) Echinatisporis sp.; (6) Verrucosisporites sp.; (7) Bennettitaepollenites sp.; (8) Stellatopollis sp.; (9) Afropollis sp.; (10) Dejaxpollenites microfoveolatus; (11) Classopollis classoides; (12) Equisetosporites ovatus; (13) Gnetaceaepollenites jansonii; (14) Regalipollenites sp.; (15) Araucariacites sp.; (16) Callialasporites dampieri; (17) Complicatissacus cearensis; (18) Cyathidites sp.. Scale bar 20 µm.Full size imageOverall, the vegetation is dominated by the xerophytic bioclimatic group on account of the very high abundance of Classopollis (Cheirolepidiaceae) (general mean of 60.5%). However, the stratigraphic distribution of the bioclimatic groups in the sections studied (Supplementary Figs. 1–6) indicates wet phases confirmed by the curves of the other bioclimatic groups (hygrophytes, hydrophytes, tropical lowland flora, and upland flora). We used Pearson correlation analysis (Supplementary Fig. 7) to assess the correlation between the bioclimatic groups. The analysis revealed positive correlations between the bioclimatic groups of hygrophytes, hydrophytes, tropical lowland flora, and upland flora, and a negative correlation between these groups and the xerophyte group (Supplementary Fig. 7). The positive correlation between upland flora and hygrophytes confirms previous studies for the Sergipe Basin6,7, suggesting a relation between these groups and the hot and humid climate. The weak negative correlation between tropical lowland flora and upland flora is presumably related to elevation.The upland flora forms the second most abundant bioclimatic group, with an average of 18.9%. The large number of specimens of Araucariacites (Araucariaceae) in this group is notable. The hydrophytes are the least abundant group, with an average of only 1.4%. In this group, the highest values are attributed to the genus Crybelosporites (Marsileaceae).Principal component analyses (PCA) were used to reduce the multidimensional dataset, based on the percent abundance of the bioclimatic groups to a smaller number of dimensions for interpretive analysis. For all sections, two components or axes explain 97.6% of the observed variability (Fig. 2B). Hygrophytes, hydrophytes, tropical lowland flora, and upland flora show positive correlation (positive loading, 0.320, 0.029, 0.006, and 0.468, respectively), whereas xerophytes show a negative relationship (negative loading, − 0.823) on the first axis, which alone explains 83.0% of the variability. In summary, the first axis of the PCA reveals a separation of two major climatic conditions (wet and dry) along the axis (Fig. 2B). The wet conditions include the associations of hygrophytes, hydrophytes, tropical lowland flora, and upland flora, with dry conditions associated with taxa from the xerophyte group. The second axis explains 14.6%, in which hygrophytes, hydrophytes, and tropical lowland flora show a positive correlation relationship (positive loading, 0.719, 0.037, 0.036, respectively), whereas upland flora and xerophytes show a negative relationship (negative loading, − 0.684 and − 0.108, respectively). With respect to the second axis, a polarization between the hygrophytes (positive loading, 0.719) and the upland flora (negative loading, − 0.684) can be interpreted as a lowland–upland trend. The same pattern was recorded for all paleoclimatic phases (Fig. 2C) and sections (Supplementary Fig. 8), that is, the first axis is related to humidity vs. aridity, and the second axis to elevation (lowland vs. upland). This suggests that these two factors, particularly the first one, controlled the vegetation distribution in the late Aptian of the region. As all bioclimatic groups occurred in the three evaporitic phases, these trends in abundance reflect expansion and contraction of the recorded vegetation.Parallel increasing trends of bioclimatic groups mark the pre-evaporitic phase: hygrophytes and upland flora in the Bragança-Viseu, São Luís, Parnaíba, Ceará, Potiguar, and Araripe basins (Supplementary Figs. 1–3 and 5), suggesting that there was a certain amount of moisture in these areas. The xerophytes show the lowest average of this phase (44.1%) (Table 1), whereas hygrophytes show the highest average (27.0%). These humid conditions are confirmed by the highest mean of the Fs/X ratio (Fs/X = 0.4), representing the predominance of spore-producing plants [see Methods section and Supplementary information]. Despite the low abundance of hydrophytes in the sections, a prominent feature is the highest average (2.5%) of this group (Table 1), which is assigned to aquatic environments, confirming relatively wet conditions in this phase. There are no pre-evaporitic samples available from the Sergipe and Espírito Santo basins.Table 1 Average abundance of bioclimatic groups, diversity, Fs/X and marine elements for the paleoclimatic phases.Full size tableThe evaporitic phase is characterized by the highest abundance of the xerophyte bioclimatic group (76.4%) (Table 1), represented mainly by Classopollis (Supplementary Figs. 1–6). A high abundance of xerophytes occurred widely distributed in all basins studied. In this phase, tropical lowland flora is notable, showing an average higher than the overall average (3.3%), particularly in the Bragança-Viseu, São Luís, Parnaíba, and Ceará basins (Supplementary Figs. 1 and 2). This result is related to the moderate to high abundance of the genus Afropollis in these basins. The evaporitic phase is also characterized by the lowest average Fs/X ratio (Fs/X = 0.1) (Table 1), confirming the dominance of xerophytes.The post-evaporitic phase is characterized by the upland flora bioclimatic group (mean = 24.4%) (Table 1). The moderate to high abundance of upland flora in this phase is represented, in particular, by pollen grains of Araucariacites, which represent the high-relief family Araucariaceae. This bioclimatic group is associated with more humid conditions, as confirmed by an Fs/X ratio higher than the overall average (Fs/X = 0.2). The upland flora is significant in all basins, except the Espírito Santo Basin, where xerophytes predominate in both studied phases in this basin.Latitudinal biome distributionsBiome change is a fundamental biological response to climate change. In the study area, the predominance of a specific biome is mainly related to humidity, since all five recorded bioclimatic groups are related to a warm climate (Supplementary Table 2) representing two biomes: tropical xerophytic shrubland and tropical rainforest. In the rainforest biome two phytophysiognomies are recognized: lowland and montane rainforest. The tropical xerophytic shrubland biome predominates in the three paleoclimatic phases, with a wide latitudinal range from the Bragança-Viseu, São Luís, and Parnaíba basins (1° S) to the Espírito Santo Basin (20° S). This wide distribution is compatible with a predominantly arid climate in South America in the late Aptian, as indicated by paleoclimatic maps8,9,15 (Fig. 4A). Most arid and semi-arid ecosystems are mainly controlled by precipitation. Other climate parameters are less important, a condition that simplifies cause-effect interpretations. The PCA (Fig. 2B) demonstrated that the wet–dry trend, which reflects high–low precipitation, was the main determinant in the distribution of the biomes. However, considering all phases, an increasing trend in humidity was observed from the southeast (Espírito Santo Basin) to the northeast (e.g., Potiguar Basin) (Fig. 4B), coinciding with the hot and wet belt attributed to the ITCZ (Fig. 4A)15. The latitudinal distribution of diversity also follows this trend. Diversity increased significantly towards in the basins near the equator. Diversity indices (Shannon – H’) peaked in the Sergipe Basin (H’ = 3.5, CL-47 section) at 11° S. Conversely, the lowest average diversity is recorded in the Espírito Santo Basin (H’ = 1.1) at 20° S. Additionally, there is a clear correlation between high diversity (H’) and humidity (Fs/X ratio) (r = 0.691), regardless of paleoclimatic phase, as evidenced by the synchronicity of the H’ and Fs/X curves (Fig. 5). After data normalization between humidity (Fs/X) and marine elements (dinoflagellate cysts and microforaminifer linings), we performed linear correlation analyses, which showed a weak but positive correlation (r = 0.137). This is due to the fact that pre- evaporitic deposits contain only 19 occurrences of dinoflagellate cysts in 90 samples. Despite this, the curves of Fs/X, marine elements and diversity are synchronous (Fig. 5), suggesting a relation between humidity, diversity, and marine incursions.Figure 4Latitudinal changes in late Aptian biomes from southeast to center-north. (A) Paleoclimatic belts of the late Aptian in South America (climatic belts modified from refer.14). Reconstruction map at 116 Ma modified from ODSN Plate Tectonic Reconstruction Service. The Reconstruction map at 116 Ma was generated by ODSN Plate Tectonic Reconstruction Service (https://www.odsn.de/odsn/services/paleomap/paleomap.html). (B) Late Aptian latitudinal distribution of the tropical xerophytic biome in Brazil. (C) Stratigraphic distribution of biomes for individual basins. (D) Relative Importance of biomes for paleoclimatic phases.Full size imageFigure 5Biome trends in relation to paleoclimatic phases. Change in biomes, diversity, Fs/X ratio and marine elements shown by changepoint analysis plotted against paleoclimatic phases.Full size imageThe pre-evaporitic phase is marked by a certain balance between the biomes (Fig. 4C,D). In the lowlands, the tropical xerophytic shrubland biome predominated in the Bragança- Viseu, São Luís, Parnaíba, and Ceará basins, but in the Potiguar Basin it is co-dominant with the lowland rainforest. The montane rainforest was relatively extensive in this phase, although with several areal changes, and reached its widest extent in the Araripe (7° S) and Potiguar (5° S) basins in response to the deterioration of the tropical xerophytic shrubland biome. These conditions demonstrate that humidity was relatively high at this stage. The pre-evaporitic deposits were characterized by the highest diversity average (H’ = 1.8).The method of indicator species analysis (IndVal) was used to identify the key species of each paleoclimatic phase (Supplementary Table 15). The species identified for the pre-evaporitic phase, Deltoidospora spp. (Cyatheaceae-Dicksoniaceae) related to the montane rainforest, are indicator species for the Bragança-Viseu, São Luís, Parnaíba, and Ceará basins. The Gnetaceaepollenites spp. (Gnetaceae) of the Potiguar Basin and Equisetosporites spp. (Ephedraceae) of the Araripe Basin are related to the tropical xerophytic shrubland biome (Supplementary Table 15). Even for the pre-evaporitic phase, a progressive increase in the tropical xerophytic shrubland biome was observed and interpreted as the start of a climatic deterioration stage (Fig. 4C), which culminated in the evaporitic phase. Shifts in vegetation types may occur when precipitation reaches a threshold value, which means that a regionally synchronous gradual climate change can cause abrupt vegetation shifts. The change from humid to warm and arid conditions (evaporitic phase) is directly related to a decrease in precipitation. This aridization process coincides with the appearance of marine elements (e.g., dinoflagellate cysts). The threshold effect (intense evaporation) is reflected in an abrupt decrease in the abundance of lowland and montane rainforest and a sharp increase to a very high abundance of the tropical xerophytic shrubland biome (Supplementary Figs. 4C and 5). The threshold effect was not detected in the Espírito Santo Basin, where the arid conditions remained stable with minimal shift (expansion and contraction) of the biome. The main representatives of this biome are conifers of the family Cheirolepidiaceae (Classopollis), which were most abundant in lagoons and coastal environments and are often associated with evaporates30,31,32,33,34,35. Even under xeric or water-stressed conditions there was a slight increase in biomes related to a humid climate (lowland and montane rainforest phytophysiognomies) towards the equatorial region, suggesting influence of the ITCZ (Fig. 4A,B).The evaporitic phase was characterized by the lowest diversity average (H’ = 1.2). With modest rainfall, arid regions are generally characterized by fewer species than moister biomes36. However, diversity indices peaked in the Bragança-Viseu, São Luís, and Parnaíba basins (H’ = 2.6, RL-01 section) and along the equatorial margin (2° S) (Supplementary Fig. 1).IndVal emphasizes the xeric conditions in the evaporitic phase by association with the species from the tropical xerophytic shrubland biome: Classopollis spp. (Ceará and Potiguar basins), Classopollis classoides (Sergipe Basin), Classopollis intrareticulatus (Araripe Basin), and Gnetaceaepollenites spp. (Espírito Santo Basin). For the Bragança-Viseu, São Luís, Parnaíba, and Ceará basins, where xeric restrictions are milder, the indicator taxon is Afropollis spp. from the lowland rainforest. This genus shows the weakest negative correlation with xerophytes.After the end of evaporite deposition, all sections indicate climatic stability, which kept the climate hot and arid even in the post-evaporitic phase, although the response was not linear.The shift in the biomes, especially the tropical xerophytic shrubland in the Bragança-Viseu, São Luís, Parnaíba, Ceará, and Araripe basins, occurred in the transition between the evaporitic and post-evaporitic phases, whereas in the Potiguar and Sergipe basins it occurred within the post-evaporitic phase. As indicated in the dendrograms of each section (Supplementary Figs. 1–6), the shift occurred abruptly in all basins, except the Espírito Santo Basin. The tropical rainforest biome (lowland and montane rainforests) replaced the tropical xerophytic shrubland in almost all basins (Fig. 4C). Even the Espírito Santo Basin, far from the influence of the ITCZ, shows a slight increase in lowland rainforest. The changes in the biomes are attributable to threshold effects caused by gradual climate change related to the ITCZ intensification shift and progressive increase in marine influence, indicated by an increase in marine microplankton from an average of 3.9% in the evaporitic phase to 44.1%. The increase in marine influence is reflected in the first major flooding surface observed in the Cretaceous succession27. Thus, a climate amelioration stage was established in the post- evaporitic phase (Fig. 5). In combination with published paleotopographic information25, the bioclimatic groups associated to the humid conditions (hygrophytes, hydrophytes, tropical lowland flora, and upland flora) were combined and visualized to create Fig. 6.Figure 6Reconstruction of the transitional gradient between marine to terrestrial environment (uplands) under ITCZ influence. The illustration is based on paleoflora and environmental information from palynological data from studied sections. Original size illustration: 18 × 24 cm, by Julio Lacerda.Full size imageAccording to refs.7,37, arid conditions are characterized by sea-level lowstands, whereas warm and humid conditions are correlated with sea levels rise, which explains the increase in the tropical rainforest biome (lowland and montane rainforests). The more intense humidity is supported by the results of IndVal for the post-evaporitic phase, with all species related to humid climate: Deltoidospora spp. (Bragança-Viseu, São Luís and Parnaíba basins), Araucariacites limbatus (Ceará Basin), Cicatricosisporites spp. (Potiguar Basin), Cicatricosisporites spp. and Araucariacites australis (Sergipe Basin), Inaperturopollenites spp. (Araripe Basin) and Inaperturopollenites simplex (Espírito Santo Basin).Our results show that the ITCZ combined with the opening of the South Atlantic Ocean during the late Aptian altered vegetation dynamics. As today, the ITCZ influence is stronger in the northeastern and north-central regions of South America. It is notable that the late Aptian climate evolution in the South Atlantic, culminating in higher humidity, was accompanied by an intrinsic relation between plant diversity, humidity, and marine influence. More

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