Multiproxy Analysys of Ambers From the Romualdo Formation (Cretaceous), Araripe Basin, Ceará State, Brazil

Multiproxy Analysys of Ambers From the Romualdo Formation (Cretaceous), Araripe Basin, Ceará State, Brazil | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Multiproxy Analysys of Ambers From the Romualdo Formation (Cretaceous), Araripe Basin, Ceará State, Brazil Laura Tomaoka, Alcina MF Barreto, Ludmila AC Do Prado, Eduarda Quadros Machado, and 2 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9349105/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract This study advances the understanding of ambers as important paleobiological and paleoenvironmental archives for the Romualdo Formation of the Araripe Basin and the Lower Cretaceous (Aptian) of Gondwana. Twenty-one (21) ambers collected from the Sobradinho Site, were investigated. Modern and traditional techniques, such as confocal microscopy, petrography, and Laser Ablation ICP-MS (LA-ICPMS), were used, allowing for a detailed analysis of the physical, chemical, and taphonomic characteristics of these fossils. Among the inclusions were fungi, pollen grains of Classopollis and xylem fragments. The analyzed inclusions suggest the presence of coastal forests dominated by gymnosperms with the presence of the Cheirolepidiaceae family. Geochemical analyses revealed a significant enrichment in elements such as sulfur, phosphorus, iron, and zinc, which directly influenced the physical properties and coloration of the samples. From a taphonomic perspective, these ambers represent an allochthonous component that was transported from the forest floor to the coastal marine environment where they were finally deposited. This interpretation was corroborated by the abundance of plant and amber microfragments found in the shales associated with the samples studied. Paleoenvironmental conditions allow to infer a prevailing arid climate, with forests adapted to hydric deficits but potentially sustained by groundwater aquifers. The results offer new perspectives on the evolution of the Araripe Basin ecosystems during the Cretaceous period. Hyphae Aptian Cheirolepidiaceae Classopollis Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 1. INTRODUCTION Resins are natural compounds that are generated by both gymnosperms and angiosperms. They have complex chemical structures containing a variety of elements such as terpenoids (volatile and non-volatile), phenols, acids, and various other substances (Grimaldi, 2019 ; Langenheim,1969, 2003). Resins are essential compounds for plant defense and protection. Its large-scale production is triggered by environmental stress, which can result from abiotic factors, such as variations in temperature, rainfall, atmospheric composition, fires, volcanic activity, changes in sea level, or from biotic factors, such as damage caused by animals (i.e., arthropods) or the action of microorganisms (Delclòs et al., 2023 ; Langenheim, 1969 , 2003 ; Martínez-Delclòs et al., 2004 ; Seyfullah et al., 2018 ). The fossilization of resin, known as amberization (Martínez-Delclòs et al., 2004 ), begins when the resin is exuded and exposed to the atmosphere. The contact with the atmosphere triggers the polymerization of the resin components, leading to gradual hardening and becoming amber (Langenheim, 2003 ; Solórzano-Kraemer et al., 2020 ). The resin can be produced in the upper regions of trees, such as the trunk and branches and when it detaches and falls into the forest ground. The resin can also be covered by a thin layer of sediment (Martínez-Delclòs et al., 2004 ). Secreted by the roots and retained in the soil, it can occur colonization by fungi while the resin is still attached to the tree trunk, as well as once in the ground, the fungi develop on this substance because they feed on the resin components (Álvarez-Parra et al., 2024 ). The oldest records of fossilized resins date back to the Carboniferous Period (Sargent Bray and Anderson, 2009 ), whereas the oldest amber-containing biological inclusions belong to the Triassic Period (Schmidt et al., 2012 ). On a global scale, organic inclusions in amber are quite diverse, including records of invertebrates, such as aquatic arthropods, insects, arachnids (Grimaldi et al., 2018 ; Néraudeau et al., 2017 ), soft parts of insects (Jiang et al., 2022 ) and interactions between insects and their products (p.ex. coprolites) (Qvarnström et al. 2021 ). Records of plants such as leaves, spores, pollen grains (McKellar and Wolfe, 2010 ; Saint Martin et al., 2020 ), coals (Najarro et al., 2010 ) and sap (Lozano et al., 2020 ) have been studied. In addition to fungi (Schmidt et al., 2018 ), bioinclusions of lichens (Schmidt et al., 2022 b), unicellular green algae and diatoms (Schmidt and Dilcher, 2007 ), filamentous microorganisms (Frau et al., 2021 ), bacteria [e.g., flagellated cells, amoebas, ciliates, rotifers (Schmidt and Schäfer, 2005 ), and bacterial filaments (Saint Martin and Saint Martin, 2018 ) have also been described. The places where amber occurs globally are diverse and widely distributed over the last 200 Ma, mostly belonging to the Cretaceous Period (Martínez-Delclòs et al., 2004 ). In the Southern Hemisphere, particularly South America, few occurrences have been described and studied, opening up an interesting field for paleontological research in Brazil. The aim of the research is to improve the knowledge of Brazilian Cretaceous ambers through the study of specimens from the Sobradinho site of the Romualdo Formation. Until 1970 only three amber occurrences were known in Brazil (Fróes de Abreu, 1937; Castro et al., 1970 ; Langenheim and Beck, 1968 ), including the first analysis using infrared spectrometry of Miocene ambers from the Pirabas Formation (state of Acre) (Langenheim and Beck, 1968 ). Decades later, the first gas chromatography-mass spectrometry (GC-MS) was performed on a Brazilian amber (Pereira et al., 2011c ). These pioneers in the amber study in Brazil enabled other researchers to follow their paths, and many more analyses were performed using a few samples of Brazilian amber for a better understanding of this occurrence (Carvalho, 1998 ; Pereira et al., 2009b , 2009c ; Pereira et al., 2011b ). Today in Brazil, amber occurrences are mainly located in five basins: (i) the Araripe Basin (Rio da Batateira and Romualdo Formations), (ii) the Recôncavo Basin (Maracangalha Formation), (iii) the Parnaíba Basin (Itapecuru, Codó and Cabeças), (iv) the Amazonas Basin (Alter do Chão Formation) and (v) the Acre Basin (Pirabas Formation) (Fróes de Abreu, 1937; Castro et al., 1970 ; Langenhein and Beck, 1968; Carvalho, 1998 ; Carvalho et al., 2000 ; Langenhein and Beck, 1968; Paiva and Carvalho, 2021 ; Paula-Freitas et al., 2007 ; Pereira et al. 2009a , 2009b , 2009c , 2011a , 2011b , 2011c , 2020 ; Seyfullah et al., 2020 ). Of these five basins, most were present in the Cretaceous Period (Fig. 1 ), including this study. Other basins belong to the Devonian (Cabeças Formation) and Miocene periods (Pirabas Formation). Organic geochemical analyses conducted to determine botanical affinity indicated that the ambers were products derived from conifers, especially from the families Cheirolepidiaceae and/or Podocarpaceae (Pereira et al., 2009b ). Few macroscopic inclusions have been reported in Brazilian records (Martill et al., 2005 ). Studies on the inclusions found in Brazilian ambers cover ostracods (Piovesan et al., 2022 ) and Ciliophera (Paiva and Carvalho, 2021 ). The occurrence of fossil resin drops associated with plants has also been described, such as dicotyledonous leaves (Langenheim and Beck, 1968 ), pollen grains from Araucariceae, Cupressaceae, and Cheirolepidiaceae (Pereira et al., 2009c , 2011a ), and pollen grains and cones from Erdtmanithecales (Seyfullah et al., 2020 ). Fungal spores have also been found in amber from the Amazon Basin (Pereira et al., 2009c ). 2. GEOLOGY SETTING The basement of the Araripe Basin has as its dominant lithology orthogneisses (quartz diorite to granite) of Precambrian age (from the transverse zone of the Borborema Province (Brito Neves et al., 2000 ; Brito Neves, 2003 )) as well as rocks from the metamorphic suite such as metavolcanics (1.12–0.93 G.a.) (Kozuch et al., 1997 ; Leite, 1997 ; Van Schmus et al., 1995 ) and metagranitoids (980 − 920 Ma) (Leite et al., 2000 ; Brito Neves et al., 2001 ; Kozuch et al., 1997 ; Van Schmus et al., 1995 ). Proterozoic gneisses from the Rio Capibaribe Terrain (Brito Neves et al., 2013 ) calc-alkaline rocks containing minerals such as Ba, Rb, K, Nb, Ta, P and Ti from the Alto Moxotó Terrain (de Lira Santos et al., 2022 ). Orthogneisses, trondjenite, gabbro-diorite and migmatites from the Alto Pajeú Terrain (de Oliveira et al., 2023 ), meta-peraluminous rocks containing Rb, Th, Nb and K from the Cabaceiras Complex (Lages et al., 2009 ) and metagranitoids containing Co, V and Sr from the Caicó Complex (Souza et al., 1996 ). The Araripe Basin, with a sedimentary deposition area of approximately 9,000 km², is the largest inland sedimentary basin in northeastern Brazil (Fambrini et al., 2020 ). The Mauriti Formation was deposited in an interlaced fluvial environment with a lithology of immature sandstone and conglomerates (Assine, 2007 ; Assine et al., 2014 ). This unit was previously called the Cariri Formation, (Beurlen, 1962 , 1963 ) and a new name was proposed by Gaspary and dos Anjos ( 1964 ) and later adopted by Ponte and Appi ( 1990 ). The age of this formation is still under debate, with authors (i.e., Assine, 2007 ; Assine et al., 2014 ) placing it in the Devonian and others arguing for a Jurassic age based on the presence of dinosaur footprints in some of its outcrops (i.e., Carvalho et al., 2024, Carvalho and Leonardi, 2024). In the chronological sequence are the Jurassic-, Brejo Santo-, and Missão Velha formations. The former includes red shales and mudstones of lacustrine origin, whereas the latter refers to a sedimentary sequence of quartz and/or feldspathic sandstones resulting from deposition in an interlaced fluvial environment. The latter are part of the initial stage of the Opening Rift of the Ocean (Assine, 2007 ; Assine et al., 2014 ; Cesero et al., 1997 ), followed by the Abaiara Formation of the Neocomian age, which is characterized by shales, siltstones, and fine sandstones of the fluvial-lacustrine environment (Assine, 2007 ; Assine et al., 2014 ) and is associated with the rifting stage of the opening of the ocean (future Atlantic). All formations belong to the Vale do Cariri Group. The Santana Group (Fig. 2 ) is composed of the Barbalha, Crato, Ipubi and Romualdo Formations (from oldest to youngest) (Assine, 2007 ). The Romualdo Formation, the top portion of the Santana Group, hosts a sedimentary succession that documents a marine transgression that occurred during the fragmentation of the Gondwana supercontinent and the subsequent expansion of the South Atlantic Ocean (Assine et al., 2014 ; Neumann and Assine, 2015 ; Fambrini et al., 2020 ). There is still disagreement about the age of the Romualdo Formation and the perspective that the boundary of Aptian-Albian stands. But it has been dated based on fossils, ostracods, and palynomorphs and considers the entire succession of the Romualdo Formation as the upper Aptian age (Regali, 2001; Rios-Netto and Regali, 2007 ; Teixeira et al., 2018 ; Arai and Assine, 2020 ). Sedimentary beds of this formation crop out in various areas of the Araripe Basin and exhibit different phases of marine transgression resulting from the opening of the Atlantic Ocean. However, the precise extent of these flooding episodes is not well defined. Observations reveal inconsistencies between the sedimentary patterns and macro- and microfossil records found in different geographical locations where the Romualdo Formation crops out (Nascimento et al., 2023 ; Araripe et al., 2025 ). These discrepancies contribute to the fact that the exact configuration of the paleoenvironments remains partially unknown. The fauna of the Romualdo Formation represents a transitional environment between coastal zones and shallow marine waters (see Mabesoone and Tinoco, 1973 ; Arai and Coimbra, 1990 ; Berthou, 1990 ; Kellner et al., 2002 ; Bruno and Hessel, 2006 ; Lima et al., 2012 ; Araripe et al., 2021 ; Melo et al., 2020 ). Recognition of echinoids, calcareous nanofossils, and foraminifera is the main evidence of a marine influence on the sedimentary processes of this formation. These records indicate the occurrence of a transgressive-regressive cycle limited by two regional discordances, which resulted in marine deposition throughout the unit (i.e., Beurlen, 1962 , 1963 ; Prado et al., 2015 , 2018 ; Custódio et al., 2017 ; Melo et al., 2020 ; Araripe et al., 2021 , 2022 ; Pedrosa et al., 2023 ). Regarding macroflora, the Romualdo Formation hosts the remains of the genera Brachyphyllum and Pseudofrenelopsis as the most common fossils, but it can also contain angiosperms and gymnosperms (most abundant) (Duarte, 1993 ; Lima et al., 2012 ). However, fossils of this character are poorly preserved, making taxonomic studies difficult. The Romualdo Formation is mainly composed of sandstones interspersed with dark grey shales, which are rich in organic matter, as well as greenish shales and marls at their base. Most of the upper part of this formation consists of fossiliferous concretions found in green shales, which contain large numbers of vertebrates (i.e., osteichthyes), ichnofossils (i.e., coprolites), ostracods, and, less frequently, plants (Arai and Coimbra, 1990 ; Berthou, 1990 ; Colin and Dépêche, 1997 ; Mabesoone et al., 1999 ; Coimbra et al., 2002 ; Tomé et al., 2014 ). In the upper part, layers of limestone-containing invertebrates are covered by thin layers of sandstone, siltstone, and shale. These sediments host fossils from mixohaline and marine environments, including crustaceans, mollusks and foraminifera, both benthic and planktonic (Beurlen, 1971 ; Assine, 2007 ; Prado et al., 2015 , 2018 ; Araripe et al., 2021 ). The Sobradinho section (Fig. 3 ) is one of the most studied sites in the Romualdo Formation, because it shows the most complete section of this formation, in the eastern portion of the Araripe Basin (Custódio et al., 2017 ; Fursich et al., 2019; Arai e Assine, 2020; Melo et al., 2020 ; Bom et al., 2021 ; Kroth et al., 2021 ; Araripe et al., 2021 , 2022 ).). 3. MATERIALS AND METHODS The samples used in this study and the methodologies adopted are listed in Table 1 . Table 1 Samples and analyses carried out on them. Caption. L. Petrographic slides. C. Confocal microscope. LA-ICP-MS. Laser Ablation Inductively Coupled Plasma Mass Spectrometry. SEM/EDS. Scanning Electron Microscope / X-ray Energy Dispersive Spectroscopy. N° amostra (DGEO-CTG-UFPE) L C LA- ICPMS MEV/EDS Location 8800 1 x Sobradinho, CE - perfil 27 m 8801 1 Sobradinho, CE - perfil 27 m 8803 1 x Sobradinho, CE - perfil 27 m 8804 1 x x Sobradinho, CE - perfil 27 8805 1 Sobradinho, CE - perfil 28 m 9121 1 x x Sobradinho, CE - perfil 28 m 9123 x Sobradinho, CE - perfil 28 m 9124 1 x x Sobradinho, CE - perfil 28 m 9125 1 Sobradinho, CE - perfil 28 m 9126 2 x Sobradinho, CE - perfil 28 m 9128 2 x Sobradinho, CE - perfil 28 m 9129 1 x Sobradinho, CE - perfil 28 m 9131 1 Sobradinho, CE - perfil 28 m ROM-03 1 Sobradinho, CE ROM-04 1 Sobradinho, CE Test 1 Sobradinho, CE Pyrite 2 Sobradinho, CE 3.1 Location of the study area The sample collection site is located in the municipality of Jardim, State of Ceará (CE) (Fig. 2 ), within the stratigraphic sequence of the Sobradinho site (Araripe et al., 2025 ) associated with the shale layers at 28 m at the top of Romualdo Formation (Fig. 3 ). The Sobradinho site belongs to the Romualdo Formation and in the map (Fig. 2 ) the amber site is located on top of Exu Formation, that is due to the fact that the outcrop was exposed in the bed of an intermittent river and has no exposed surface. 3.2 Samples collected for the study A total of 21 complete or fragmented samples were collected during two field expeditions (the first in 2019 and the second in 2022). The ambers were stored in the Scientific Collection of Paleontology at the Department of Geology (DGEO), Center of Technology (CTG), UFPE (Federal University of Pernambuco). Those from the first collection were included under DGEO-CTG-UFPE numbers 8800 to 8806 and those from the second collection were included under DGEO-CTG-UFPE numbers 9120 to 9133. 3.3 Study of the thin sections using a petrographic microscope After preparing the slides, they were analyzed using a Zeiss Axioscope AI Imager petrographic microscope with an attached digital camera to capture images, which allowed the inclusions to be identified, described, and recorded. The images were acquired using the Zen Blue Zeiss program and saved in .tif format. To better characterize and detail the inclusions, thin sections of selected amber fragments were prepared. To achieve this, adopting a safe and effective lamination method was necessary, as this was not a standard procedure (Tomaoka and Ricardi-Branco., 2025). Samples were prepared for lamination at the Palaeohydrogeology Laboratory, part of the Institute of Geosciences, State University of Campinas (UNICAMP). 3.3.1 Making the petrographic thin section The references Corral et al. ( 1999 ), Nascimbene and Silverstein ( 2000 ) and Sadowski et al. ( 2021 ) were used as the basis for creating the slides. They described techniques for preparing amber for lamination such as sanding, polishing, and impregnation in epoxy resin using a vacuum chamber. Therefore, to make the thin sections of an amber sample, it must first be cleaned with distilled water to remove traces of the matrix; then, in the case of thicker and/or opaque samples, thickness was reduced using water sandpaper graded from 150 to 2000 (in ascending order) to achieve a good finish and better visualization. A Stemi V6 ZEISS stereomicroscope was used to select samples with possible bioinclusions and the best place to make thin sections. Twenty-two (22) thin sections of the different samples were made; so therefore, getting a better idea of the occurrences of inclusions in the samples was possible (Table 1 ). All fragments were photographed with a CANON EOS Rebel T5i camera using a light table (Tomaoka and Ricardi-Branco, 2025 ). First, the chosen and prepared material was impregnated with epoxy resin until it was halfway up the sample in a delimited space. After sufficient hardening of the epoxy, a new layer of epoxy was placed on the samples until they were completely covered. They were then placed in a vacuum chamber so that the resin could fill all the gaps, avoiding the generation of reflections by light from the petrographic microscope (Tomaoka and Ricardi-Branco, 2025 ). The last stage was gluing the impregnated sample to a petrographic slide and subsequently sanding and polishing it until it reached the desired thickness, in this case 100 micrometers (Tomaoka and Ricardi-Branco, 2025 ). 3.4 Confocal LASER microscopy imaging The principle of confocal microscopy differs from that of conventional microscopy (Hein et al., 1995 ; González and Halpern, 2007 ; Borlinghaus, 2017 ). Laser confocal microscopy was used to complement the study of the organic inclusions. This technique allows the matrix to be isolated, and the kerogen and organic molecules to be better mapped. This stage was carried out on the premises of the National Institute of Science and Technology for Photonics Applied to Cell Biology (INFABIC) at UNICAMP using the Zeiss LSM780 NLO inverted microscope, and the slides viewed using this equipment can be seen in Table 1 . 3.5 SEM/EDS Scanning electron microscope (SEM) and energy-dispersive spectroscopy (EDS) was used to acquire a better image of micro-inclusions presents in the amber samples. Model Zeiss Leo 430i with EDS detector Oxford model 7059 (10 mm 2 ), software ISIS. The samples were coated with gold using the Quorum Q 150TES. 3.6 Elemental analysis by Laser Ablation (LAICP-MS) Laser Ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) or laser ablation inductively coupled plasma mass spectrometry was used to complete the geochemical characterization of ambers. To accomplish this, samples with the potential for analysis had to be prepared. The samples used to identify the inorganic species were previously cut and sanded (according to the same procedure described above) to obtain thin, smooth, and flat surfaces for ablation. The samples were then placed in a holder and fixed using double-sided tape. Three ablation lines were created based on the size of each sample to cover the entire surface. The samples were selected to cover the widest range of characteristics among the available fossil resins; thus, DGEO-CTG-UFPE samples 8800, 8804, 9121 and 9132 were selected. The elements 13 C, 34 S, 57 Fe, 51 V, 28 Si, 43 Ca, 59 Co, 63 Cu, 45 Sc, 111 Cd, 107 Ag, 31 P, 75 As, 66 Zn and 118 Sn were evaluated to identify the inorganic constituents of the amber samples. The ablation parameters used were: 110 µm spot, 110 µm s-1 scan speed and 60% intensity and plasma conditions of 1200 W RF power, 1.6 L min-1 auxiliary gas and 1.2 L min-1 nebulizer gas. This technique combines plasma ablation and spectrometry to determine the quantity of inorganic elements present in the samples. 4. RESULTS 4.1 Macroscopic characterization In general, the ambers under study range in color from light brown (caramel) to dark brown, or 10YR 2/2, 3/2, 4/4, 3/4, 6/6, 5/6, 4/6, 3/6, 7/8, 6/8, 5/8, 4/8, 7/10, and 6/10 (according to the Munsell color chart), and some samples can even be light brown (caramel) and completely opaque (Fig. 4 ). The shape varies from round to oval, and fractures occur frequently, including those in the cortex. Most samples show more than one resin deposition flow (Fig. 4 ). The inclusions cannot be discerned by the naked eye. The outer surface (cortex) of the samples is more altered than the inner surface. 4.2 Bioinclusions 4.2.1 Analyses under Petrographic and Confocal Microscopy. CONIFEROPHYTA Secondary xylem in transverse view Sample. DGEO-CTG-UFPE 9124 Description. Fragment of secondary xylem in transverse view; tracheids mostly polygonal to sub-radial, with visible walls separated by diffuse parenchyma. Arranged in rows; walls appear lignified (Figs. 5 c, 5 d, 5 e and 5 f). Dimensions: tracheids 25–30 \(\:\mu\:\) m long axis. Secondary xylem radial longitudinal view Sample. DGEO-CTG-UFPE 9124 Description. Fragment of three tracheids in the radial longitudinal view; tracheids with lignified walls and polygonal (hexagonal, pentagonal), bordered pits in an araucarioid pattern. Dimensions: fragment 230 \(\:\mu\:\) m of length, tracheids 40 \(\:\mu\:\) m wide, pits 10 \(\:\mu\:\) m wide (Fig. 5 a, 5 b). POLLENITES H. Potonié, 1993 Anterturma VARIEGERMINANTES Potonié, 1975 Turma SACCITES Erdtman, 1947 Subturma MONISACCITES (Chitaley, 1951) Potonié & Kremp,1954 Infraturma CIRCUMPOLLINI Klaus, 1960 Gênero Classopollis Pflug, 1960 Classopollis spp. Samples. DGEO-CTG-UFPE, 9126 and 9132 Description. Monoporate pollen grains with circular equatorial outline, isolated, exine with a triangular mark, exoexine with a belt. Presence of striations in varying numbers; exine ornamentation ranging from punctate to reticulate (Fig. 6 ). Dimensions. 30–34 µm FUNGI Samples. DGEO-CTG-UFPE 9126, 8805, 9128, 9131, 9132. Description. Mycelium with translucent hyphae showing varying degrees of preservation, ranging from well-preserved to degraded (Figs. 7 a-f). Hyphae in decomposition (Fig. 7 a), continuous hyphae, simple and branched, septate (Fig. 7 b) and non-septate (Fig. 7 d); Spiral hypahes (Fig. 7 e). In general, filament growth occurs from the edges of the amber toward its interior, with the density of hyphae being higher at the periphery Mycellium; (Fig. 7 f). 4.3 Elemental analysis 4.3.1 Scanning Electron Microscope (SEM) and Energy Dispersive Spectrometer (EDS) mapping. SEM and EDS analysis (Fig. 8 ) was carried out in order to better understand the composition of the sample's surface, as we were able to observe the presence of mineral species near the vascular plant structures when observing the slide of sample DGEO-CTG-UFPE 9124 under the SEM (Figs. 8 a), and Figs. 8 b, c and d show the elemental maps made with EDS of chemical elements such as Al and Ca. 4.3.2 Laser Ablation (LAICP-MS) According to the analysis of the DGEO-CTG-UFPE 8800 sample using the three ablation lines (Fig. 9 ), higher signal intensities were obtained for the elements sulphur (S), zinc (Zn), phosphorus (P) and iron (Fe). For the elements S, P, and Zn, the distribution in amber apparently showed a more homogeneous behavior when compared to Fe, which showed an even greater intensity at the extremities. The intensities of the signals recorded for DGEO-CTG-UFPE 8804, which had a redder color and greater homogeneity, are shown in Fig. 9 . The same elements were found in DGEO-CTG-UFPE 8804 as in the previous sample (DGEO-CTG-UFPE 8800), S, P, Zn, and Fe, but with a difference in their distribution. In addition, copper (Cu) was also present over the entire surface of the sample, but it was distributed heterogeneously, as was Zn, with peaks throughout the ablation. For sample DGEO-CTG-UFPE 9121, the same elements, S, P, Zn, and Fe, had higher signal intensities than the other two samples (DGEO-CTG-UFPE 8800 and 8804), but there was also a higher signal intensity for calcium (Ca), as illustrated in Fig. 9 . The distribution of most of the elements on the surface of the sample was more homogeneous, with Fe showing the most pronounced distribution, especially in the darker amber areas. DGEO-CTG-UFPE 9123 was visually more homogeneous in color; however, it had small darker regions, as shown in Fig. 9 , which also shows the signals obtained for the elements evaluated. As in the previous three samples, S, P, and Fe exhibited higher intensities than the other elements explored. In this case, the distribution of the elements at the start of the ablation showed an even greater intensity, drawing attention to the presence of vanadium (V) expressed on ablation line 3, which coincides with the darkest region of the sample. In addition, the distribution of Fe in the sample was heterogeneous on the surface. In all the graphs shown, the 13 C signal was used to identify the beginning and end of sample ablation (in which the signal increases at the beginning and decreases at the end). Si was not expressed in the above graphs because its intensity was higher than that of the other elements. This causes the intensity of the signals to decrease, and they cannot be graphically evaluated together. 5. DISCUSSION Owing to their nature, ambers offer unique opportunities for research, especially if modern analysis techniques such as confocal microscopy and laser ablation (LA-ICP-MS) are used. However, to obtain better results, it was important to acquire the ability to polish the amber before lamination, which enabled the prospecting of different inclusions, was necessary. The amber samples studied have a range of colors and can be more opaque or translucent, sometimes weathered, which makes the fragments partially or completely matte and opaque, as is the case with sample DGEO-CTG-UFPE 9122. Therefore, after lamination, the interiors of most of the samples studied could be accessed. The banded pattern of the ambers (Fig. 4 ) is a consequence of the successive flows of resin, as well as their shape. Although the shape is also influenced by gravity as the resin slides down the aerial part of the plant, acquiring an amygdaloid shape (Martínez-Delclòs et al., 2004 ; Labandeira, 2014 ; Lozano et al., 2020 ). Ambers derived from resins secreted by roots, in addition to having an amygdaloid appearance, take longer to solidify because of the underground conditions under which the resin is secreted (Speranza et al., 2015 ). Burial within the soil profile of the still ductile resin or at the final deposition site can also influence changes in the original shape (Labandeira, 2014 ; Lozano et al., 2020 ; Martínez-Delclòs et al., 2004 ). Moreover, the frequent microfractures observed in the ambers are due to the weight of the sedimentary load that was successively deposited above the layer carrying them (Corral et al., 1999 ). 5.1 Bioinclusions In the amber studied, in addition to micro inclusions, there are pseudo-inclusions, according to some authors, (Lozano et al., 2020 ; Quinney et al., 2015 ) these correspond mostly to sap droplets, probably exuded along with the resin because they accompany its flow. Resinous filamentous fungi are the most likely candidates for colonizing non-liquid environments, as is the case with plant resins; resins provide nutrients to fungi (Wu et al., 2018 ). Pseudo-inclusions with sap droplets incorporated into the resin can be more nutritious than the resin, providing a favorable environment for the development of mycelia (Lozano et al., 2020 ). In the studied ambers, the most abundant bioinclusions observed and common to all the samples analyzed were fungi, which were present at various stages of preservation and with a variety of structures, such as mycelia (Fig. 7 ), reproductive structures (i.e., conidiophores), and spores. The fungal populations were denser on the outside of the samples, but they were also observed on the inside, which suggests that they developed after the resin fell from the aerial part of the plant, or that many of the samples studied may have been generated close to the forest floor or even by the roots (Speranza et al., 2015 ), because it would take longer for the resin to become hard (polymerized) enabling the fungus to enter. The presence of spores and pollen grains as inclusions is crucial, especially considering that the genus Classopollis , possibly derived from the Cheirolepidiaceae family, is extremely abundant in the Romualdo Formation shales (Pereira et al., 2009a , 2009c ; Arai and Assine 2020 ; Nascimento et al., 2023 ). Classopollis is, therefore, of parautochthonous origin and is possibly incorporated into the resin near its parent tree during the breeding year (Néraudeau et al., 2020 ; Pereira et al., 2020 ; Piovesan et al., 2020). This genus is paleoenvironmental marker of the Cretaceous (Carvalho et al., 2016 ; Pocock and Jansonius, 1961 ; Taylor et al., 2009 ) suggests a hot and dry climate. Furthermore, the abundant occurrence of Classopollis in the shales of the upper portion of the Romualdo Formation is considered further evidence of a proximal marine environment (Arai and Assine 2020 ). The abundance of continental palynomorphs in the layers where amber occurs contributes to an interpretation of an environment with more continental influence (Teixeira et al.; 2018 ; Ferreira et al. 2025). The anatomical characteristics of the xylem fragment in the longitudinal radial view, showing tracheids with lignified walls and polygonal (hexagonal, pentagonal) bordered pits in an araucarioid pattern, correspond to the characteristics of the wood of the conifer genera described for the Romualdo Formation (Batista et al., 2018 ). These bioinclusions, together with the paleobotanical records of the Romualdo Formation, confirm that the forests associated with the banks of rivers or even the coastal region of a lagoon could have been surrounded by coniferous vegetation, including the Cheirolepidiceae family (Batista et al., 2018 , Teixeira et al. 2018 ; Ferreira et al. 2025). The presence of pollen grains of the genus Classopollis has already been reported in shales at the same level where the ambers were collected (Nascimento et al., 2023 ), and in Cenomanian ambers collected in layers of clays rich in organic matter from the Mayenne region in France (Néraudeau et al., 2020 ) and in various mid Cretaceous deposits from France. 5.2 Elemental analysis When analyzing the results of the elemental composition obtained using the Laser Ablation technique (LA-ICP-MS), various elements were present, the intensity of which varied according to their distribution in each sample. Elements such as V, Cu, Sc, Cd, Sn, and Zn exhibited low intensities in the samples. In contrast, the intensities of the S, P, Fe, and Zn peaks were higher. The elements Fe, Zn, and sometimes V (sample DGEO-CTG-UFPE 9123) seemed to coincide with the darker-colored regions, which seemed to influence the shade of the samples. The intensities of P and S stood out as elements common in the rocks of the Santana Group, as does Si, which also predominated in the SEM/EDS analyses. Fe continues to be a frequent element, together with Mn in the samples analyzed when interpreted with the results obtained from the SEM/EDS elemental analyses. The presence of Ca demarcating the tracheids in sample DGEO-CTG-UFPE 9124 could indicate that calcium was dissolved in the water used by the plant after the inclusion of the xylem fragment deposited in the walls. The composition of ambers varies considerably depending on many external factors to the plant that produced the resin flow, including the area of provenance, paleosol, regional geology, diagenetic changes and paleoclimate (Cockx and McKellar, 2024 ; Martínez-Delclòs et al., 2004 ; Peñalver et al., 2006 ). Thus, the oxidation of Fe, for example, during the early diagenesis of the ambers and the colonization by the fungi that degrade the organic matter (resin) rich in P, C and S (indispensable elements for the development of fungi) may have partly influenced their coloring and the development of the weathering crust that the samples show (Cockx and McKellar, 2024 ; Martínez-Delclòs et al., 2004 ; Peñalver et al., 2006 , Natkaniec-Nowak et al., 2026 ). The elemental composition found in the samples suggests that the elemental variety presented was acquired by resin-secreting plants through groundwater (Aquilina et al., 2013 ), used by the vegetation that inhabited the shores of the marine basin (Melo et al., 2020 ). In addition, according to Aquilina et al. ( 2013 ) no enrichment of typical marine elements such as Na in completely solidified samples was shown. Therefore, enrichment in minerals anomalous to amber may have occurred in two ways: externally, i.e., by water droplets that managed to reach the resin before it was solidified; or internally, before the resin was secreted (Aquilina et al., 2013 ), since the resin is produced inside the plant and, in some cases, in specialized ducts (Langenheim, 2003 ), from elaborate substances that the plant stores in the parenchyma (Aquilina et al., 2013 ). These substances are derived from sap, which in turn is derived from solutions absorbed by the soil; therefore, the composition of the resin is directly influenced by the composition of groundwater and the biological processes related to resin production (Munns, 2002 ). 5.3 Taphonomy The stratigraphic level located approximately 28 m from the composite column of the Sobradinho Site (Araripe et al., 2025 ) was interpreted, based on the of benthic foraminifera, present as stressful conditions associated with a marginal marine environment with variations in salinity; the marine condition was reinforced by the presence of echinoderms and calcareous nannofossils (Prado et al., 2015 , 2018 ; Custódio et al., 2017 ; Melo et al., 2020 ; Araripe et al., 2021 , 2022 , 2025 ; Pedrosa et al., 2023 ). However, the continental origin of the ambers indicates the occurrence of marine ingressions (Custódio et al., 2017 ) that affected the dryland forests or the occurrence of flooding from freshwater streams. Both the first and second possibilities involve removing a portion of the soil in which the ambers were originally deposited and transporting it to the final place of deposition in the marine environment; therefore, they represent an allochthonous portion of the fossiliferous assemblage (Schmidt and Dilcher, 2007 ; Labandeira, 2014 ). At the same levels where the ambers were collected, plant debris and continental palynomorphs are very abundant. Thus continental bioclasts contributed with a decrease in the abundance of marine microfossils (Custódio et al., 2017 ; Teixeira et al.; 2018 ; Fereira et al. 2025; Araripe et al., 2025 ). The conifer-related biological inclusions observed in the ambers and even ambers were possibly derived from resins produced by conifers and indicated the presence of a forest composed of species resistant to the water deficit that would have inhabited near the coast, exhibiting an arid climate (Batista et al., 2018 ; Arai and Assine, 2020 ). However, the presence of groundwater would have enabled the trees to grow and cannot be ruled out. The influence of underground aquifers is most evident in the elemental composition of the amber, where Ca, Si, V, Fe, and Zn ions could have been solubilized from the basin’s basement rocks (Aquilina et al., 2013 ; Cústodio et al., 2017; Fereira et al. 2025). 6. CONCLUSION The studied samples showed diverse colors, transparencies, and weathering influenced by geological and environmental processes over time. The observed banded patterns, amygdaloid and deformed forms, and microfractures reflect the dynamics of resin flows, gravity, burial, and sedimentary pressure. These aspects highlight the importance of depositional and diagenetic contexts in the formation and preservation of ambers. The presence of inclusions such as Classopollis , spores, filamentous fungi, and pseudo-inclusions reinforces the role of ambers as biological and paleoenvironmental archives. The genus Classopollis , associated with the gymnosperm Cheirolepidiaceae, indicates coastal vegetation that tolerated the arid climate that prevailed during the deposition of the upper part of the Romualdo Formation (Aptian). The results of the elemental analyses revealed a variable composition influenced by external factors such as paleosol, regional geology, diagenetic changes, and paleoclimate. Elements such as Fe, P, S, and Zn stand out and are possibly responsible for the coloration. The lack of enrichment in marine elements such as Na suggests that the composition of the resin is directly linked to underground water absorbed by the coastal vegetation that inhabited the margins of the basin, leading to the Romualdo Formation. The combination of innovative methodological approaches and a detailed analysis demonstrates the potential of ambers as natural archives, paving the way for future interdisciplinary research in this field. Declarations Acknowledgments We would like to thank CAPES for the master’s scholarship, FAPESP for funding this research through Processes 2019/16727-3 and 23/16631-1, CNPQ for the researcher grant 307333/2021-3, the Paleohydrogeology Laboratory for the infrastructure that facilitated this research, the Electron Microscopy Laboratory (SEM), the Lamination Laboratory (LAM), Dr. Marco Aurélio Zezzi, the Group of Spectrometry, Sample Preparation and Mechanization (GEPAM), and Eduarda Machado for the analyses. 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Cretaceous Research , 91. https://doi.org/10.1016/j.cretres.2018.05.015 Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 22 Apr, 2026 Reviewers invited by journal 22 Apr, 2026 Editor assigned by journal 10 Apr, 2026 First submitted to journal 08 Apr, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9349105","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":627720189,"identity":"76a19ee7-73f4-40f0-886c-0dec7efe6926","order_by":0,"name":"Laura Tomaoka","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA9UlEQVRIiWNgGAWjYBACPvb+hw8+VLDJMRwA8yXAJDM+LWw8Z5gNZ5zhMyZBi0QOmzBvi1xiwwEkUfxaGHKPMfM2mKX3HT/8gJl3h4U8/7SzB5gL9+DTci7t4dwdabkzz6QZMPOekTCccTsvgXnGMzxaGBvMDd6eOZa74UAOAzNvm0QCw+0cA2aeA3i0MDOYSfC2/U83OP8GokWeoBY2HjNJ3ja2BIMbUFsMCGrhYUsGBjKb4cwbzwwOzm2TMNwI9MvhGXi08Ms/PgiKSnm+88kPH7xtq5OXu5178HEBHi0oAKqOB8YgHvCQqmEUjIJRMAqGOQAAgP9R+zecbmgAAAAASUVORK5CYII=","orcid":"https://orcid.org/0009-0009-1445-0153","institution":"UNICAMP: Universidade Estadual de Campinas","correspondingAuthor":true,"prefix":"","firstName":"Laura","middleName":"","lastName":"Tomaoka","suffix":""},{"id":627720190,"identity":"8254e631-1590-4e7c-9981-017e3d6ad778","order_by":1,"name":"Alcina MF Barreto","email":"","orcid":"","institution":"UFPE: Universidade Federal de Pernambuco","correspondingAuthor":false,"prefix":"","firstName":"Alcina","middleName":"MF","lastName":"Barreto","suffix":""},{"id":627720191,"identity":"2e590a03-53b6-46e2-8f26-196a2c6f68cc","order_by":2,"name":"Ludmila AC Do Prado","email":"","orcid":"","institution":"URCA: Universidade Regional do Cariri","correspondingAuthor":false,"prefix":"","firstName":"Ludmila","middleName":"AC Do","lastName":"Prado","suffix":""},{"id":627720192,"identity":"766f6522-b787-4af1-8c78-6e381eed6204","order_by":3,"name":"Eduarda Quadros Machado","email":"","orcid":"","institution":"UNICAMP: Universidade Estadual de Campinas","correspondingAuthor":false,"prefix":"","firstName":"Eduarda","middleName":"Quadros","lastName":"Machado","suffix":""},{"id":627720193,"identity":"5e02b9b0-da8d-4d32-a2aa-66531af58198","order_by":4,"name":"Marco Aurélio A Zezzi","email":"","orcid":"","institution":"UNICAMP: Universidade Estadual de Campinas","correspondingAuthor":false,"prefix":"","firstName":"Marco","middleName":"Aurélio A","lastName":"Zezzi","suffix":""},{"id":627720194,"identity":"8c914037-8ec7-4b21-ae42-f511ca7c0f97","order_by":5,"name":"Fresia Ricardi-Branco","email":"","orcid":"","institution":"UNICAMP: Universidade Estadual de Campinas","correspondingAuthor":false,"prefix":"","firstName":"Fresia","middleName":"","lastName":"Ricardi-Branco","suffix":""}],"badges":[],"createdAt":"2026-04-07 20:10:17","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9349105/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9349105/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108238009,"identity":"00c4fa99-394c-4961-a946-880ca70ada89","added_by":"auto","created_at":"2026-04-30 19:28:45","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":829391,"visible":true,"origin":"","legend":"\u003cp\u003eLocation map showing Brazilian Cretaceous amber locations.\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/b234595a192232b5b497ce25.png"},{"id":108491475,"identity":"4dc6cf6d-3e81-474e-96d6-cbbe6508651d","added_by":"auto","created_at":"2026-05-05 09:54:05","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":793540,"visible":true,"origin":"","legend":"\u003cp\u003eLocalization map with the Araripe Basin and World map of the Cretaceous (Mod. After Araripe et al., 2025).\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/611a7d8e3edbee2e61a11227.png"},{"id":108238011,"identity":"a3929546-65bf-4b45-b81f-3cfefd630210","added_by":"auto","created_at":"2026-04-30 19:28:45","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":292772,"visible":true,"origin":"","legend":"\u003cp\u003eStratigraphic column from the Sobradinho section, Jardim, CE, Romualdo Formation (Santana Group) showing amber at 28 meters (Mod. Araripe et al., 2025).\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/95e1077a5cf2e36a8475d062.png"},{"id":108491218,"identity":"5c098d07-c522-4df3-b20c-9c91ce4357a7","added_by":"auto","created_at":"2026-05-05 09:52:56","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1562432,"visible":true,"origin":"","legend":"\u003cp\u003eThe selection of samples showing the variety of raw amber collected at the Sobradinho Site, Romualdo Formation. \u003cstrong\u003ea: \u003c/strong\u003eSample DGEO-CTG-UFPE 9121; \u003cstrong\u003eb:\u003c/strong\u003e Sample DGEO-CTG-UFPE 9120; \u003cstrong\u003ec:\u003c/strong\u003eSample DGEO-CTG-UFPE 9132; \u003cstrong\u003ed:\u003c/strong\u003e Sample DGEO-CTG-UFPE 9130; \u003cstrong\u003ee: \u003c/strong\u003eSample DGEO-CTG-UFPE 9129; \u003cstrong\u003ef:\u003c/strong\u003e Sample DGEO-CTG-UFPE 9128; \u003cstrong\u003eg: \u003c/strong\u003eSample DGEO-CTG-UFPE 8805; \u003cstrong\u003eh:\u003c/strong\u003e Sample DGEO-CTG-UFPE 8801; \u003cstrong\u003ei: \u003c/strong\u003eSample DGEO-CTG-UFPE 8800; \u003cstrong\u003ej\u003c/strong\u003e and \u003cstrong\u003ek:\u003c/strong\u003e Amber in situ. Scale bar A-I = 5 mm; J = 4 cm; K = 2 cm.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/aa9b566b60a2c5222fe129b3.png"},{"id":108238013,"identity":"b9331d0d-9722-4a42-bd95-94338b21b42e","added_by":"auto","created_at":"2026-04-30 19:28:45","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":25783183,"visible":true,"origin":"","legend":"\u003cp\u003eFragments of xylem, sample DGEO-CTG-UFPE 9124; \u003cstrong\u003ea:\u003c/strong\u003e Tracheids in radial longitudinal section; \u003cstrong\u003eb: \u003c/strong\u003ePolygonal (hexagonal, pentagonal) bordered pits (yellow arrow) Araucarioid pattern. \u003cstrong\u003ec:\u003c/strong\u003e Fragment of secondary xylem transversal section, highlighting kerogen using confocal microscopy; \u003cstrong\u003ed:\u003c/strong\u003e Detail of the interior of tracheids with differentiated matrix filling obtained by confocal microscopy; \u003cstrong\u003ee:\u003c/strong\u003e Detail of mineral species differing from kerogen in confocal microscopy; F: detail of parenchyma around tracheids (black arrow) in MEV.\u003c/p\u003e\n\u003cp\u003eScale bars: a; b; c = 50 μm; d = 10 μm; e and f= 20 μm\u003c/p\u003e","description":"","filename":"Figure5.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/88722a40845fb4aefeb74338.png"},{"id":108491498,"identity":"e6e33430-4592-47bd-89ca-6e292ce29478","added_by":"auto","created_at":"2026-05-05 09:54:11","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":2269507,"visible":true,"origin":"","legend":"\u003cp\u003ePollen grains of the genus \u003cem\u003eClassopollis \u003c/em\u003espp. \u003cstrong\u003ea:\u003c/strong\u003e Fluorescence image under Confocal Microscopy (DGEO-CTG-UFPE 8801); \u003cstrong\u003eb:\u003c/strong\u003e Transmitted light image generated by petrographic microscopy (DGEO-CTG-UFPE 8801); Scale bars: a, b = 20 μm.\u003c/p\u003e","description":"","filename":"Figure6.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/cad4e7de834327ef8102477d.png"},{"id":108491626,"identity":"3815b377-7eed-4d59-be1a-8106ce620567","added_by":"auto","created_at":"2026-05-05 09:54:48","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":3301491,"visible":true,"origin":"","legend":"\u003cp\u003eSeveral fungal structures in ambers. \u003cstrong\u003ea:\u003c/strong\u003e Branched hypha in the process of decomposition (red arrow) with a conidium (external asexual spore) (black arrow), (DGEO-CTG-UFPE 9126); \u003cstrong\u003eb: \u003c/strong\u003eMycelium, (DGEO-CTG-UFPE 9131); \u003cstrong\u003ec:\u003c/strong\u003e Mycelium, (DGEO-CTG-UFPE 8805); \u003cstrong\u003ed: \u003c/strong\u003eMycelium (DGEO-CTG-UFPE 9131); \u003cstrong\u003ee:\u003c/strong\u003e Spiral hyphae (DGEO-CTG-UFPE 9128); \u003cstrong\u003ef:\u003c/strong\u003eFungi clustering near the sample margin (DGEO-CTG-UFPE 9131). Scale bars: a-f = 50 µm.\u003c/p\u003e","description":"","filename":"Figure7.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/abae261a35a3f225c354d02f.png"},{"id":108491333,"identity":"733e5923-ebf9-47e4-9712-f49c7529c85a","added_by":"auto","created_at":"2026-05-05 09:53:20","extension":"png","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":1220710,"visible":true,"origin":"","legend":"\u003cp\u003eElemental maps and histograms generated via the SEM/EDS analysis (DGEO-CTG-UFPE 9124) of different portions of the xylem fragment in transverse view. \u003cstrong\u003ea:\u003c/strong\u003e SEM image of the Map 3 area; \u003cstrong\u003eb:\u003c/strong\u003e Elemental Map 3 with the distribution of Aluminium in orange and Calcium in yellow; \u003cstrong\u003ec:\u003c/strong\u003e Distribution of Aluminium \u003cstrong\u003ed: \u003c/strong\u003eDistribution of Calcium; \u003cstrong\u003ee:\u003c/strong\u003e Histogram of present elements. Scale bars = 100 µm.\u003c/p\u003e","description":"","filename":"Figure8.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/98564a54f83d37e3444a1761.png"},{"id":108238016,"identity":"42a709db-55f8-4b30-b3df-10db2d179906","added_by":"auto","created_at":"2026-04-30 19:28:45","extension":"png","order_by":9,"title":"Figure 9","display":"","copyAsset":false,"role":"figure","size":1619338,"visible":true,"origin":"","legend":"\u003cp\u003eResults of LAICP-MS analysis. Samples DGEO-CTG-UFPE 8808, 9123, 8804 and 9121, respectively. Scale bars = 10 mm.\u003c/p\u003e","description":"","filename":"Figure9.png","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/bd2550b937a8faceac604e1e.png"},{"id":108494888,"identity":"f96c147d-1b57-4405-9e54-7234e563b00b","added_by":"auto","created_at":"2026-05-05 10:07:54","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":63032525,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9349105/v1/3284f6f2-6c3e-4c91-b699-74276b67ae56.pdf"}],"financialInterests":"","formattedTitle":"\u003cp\u003eMultiproxy Analysys of Ambers From the Romualdo Formation (Cretaceous), Araripe Basin, Ceará State, Brazil\u003c/p\u003e","fulltext":[{"header":"1. INTRODUCTION","content":"\u003cp\u003eResins are natural compounds that are generated by both gymnosperms and angiosperms. They have complex chemical structures containing a variety of elements such as terpenoids (volatile and non-volatile), phenols, acids, and various other substances (Grimaldi, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Langenheim,1969, 2003).\u003c/p\u003e \u003cp\u003eResins are essential compounds for plant defense and protection. Its large-scale production is triggered by environmental stress, which can result from abiotic factors, such as variations in temperature, rainfall, atmospheric composition, fires, volcanic activity, changes in sea level, or from biotic factors, such as damage caused by animals (i.e., arthropods) or the action of microorganisms (Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Langenheim, \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1969\u003c/span\u003e, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Seyfullah et al., \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe fossilization of resin, known as amberization (Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), begins when the resin is exuded and exposed to the atmosphere. The contact with the atmosphere triggers the polymerization of the resin components, leading to gradual hardening and becoming amber (Langenheim, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Sol\u0026oacute;rzano-Kraemer et al., \u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe resin can be produced in the upper regions of trees, such as the trunk and branches and when it detaches and falls into the forest ground. The resin can also be covered by a thin layer of sediment (Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSecreted by the roots and retained in the soil, it can occur colonization by fungi while the resin is still attached to the tree trunk, as well as once in the ground, the fungi develop on this substance because they feed on the resin components (\u0026Aacute;lvarez-Parra et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe oldest records of fossilized resins date back to the Carboniferous Period (Sargent Bray and Anderson, \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), whereas the oldest amber-containing biological inclusions belong to the Triassic Period (Schmidt et al., \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOn a global scale, organic inclusions in amber are quite diverse, including records of invertebrates, such as aquatic arthropods, insects, arachnids (Grimaldi et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; N\u0026eacute;raudeau et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), soft parts of insects (Jiang et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and interactions between insects and their products (p.ex. coprolites) (Qvarnstr\u0026ouml;m et al. \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Records of plants such as leaves, spores, pollen grains (McKellar and Wolfe, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Saint Martin et al., \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), coals (Najarro et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and sap (Lozano et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) have been studied. In addition to fungi (Schmidt et al., \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), bioinclusions of lichens (Schmidt et al., \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2022\u003c/span\u003eb), unicellular green algae and diatoms (Schmidt and Dilcher, \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), filamentous microorganisms (Frau et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), bacteria [e.g., flagellated cells, amoebas, ciliates, rotifers (Schmidt and Sch\u0026auml;fer, \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), and bacterial filaments (Saint Martin and Saint Martin, \u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) have also been described.\u003c/p\u003e \u003cp\u003eThe places where amber occurs globally are diverse and widely distributed over the last 200 Ma, mostly belonging to the Cretaceous Period (Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). In the Southern Hemisphere, particularly South America, few occurrences have been described and studied, opening up an interesting field for paleontological research in Brazil. The aim of the research is to improve the knowledge of Brazilian Cretaceous ambers through the study of specimens from the Sobradinho site of the Romualdo Formation.\u003c/p\u003e \u003cp\u003eUntil 1970 only three amber occurrences were known in Brazil (Fr\u0026oacute;es de Abreu, 1937; Castro et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Langenheim and Beck, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1968\u003c/span\u003e), including the first analysis using infrared spectrometry of Miocene ambers from the Pirabas Formation (state of Acre) (Langenheim and Beck, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1968\u003c/span\u003e). Decades later, the first gas chromatography-mass spectrometry (GC-MS) was performed on a Brazilian amber (Pereira et al., \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2011c\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThese pioneers in the amber study in Brazil enabled other researchers to follow their paths, and many more analyses were performed using a few samples of Brazilian amber for a better understanding of this occurrence (Carvalho, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Pereira et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2009b\u003c/span\u003e, \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2009c\u003c/span\u003e; Pereira et al., \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eToday in Brazil, amber occurrences are mainly located in five basins: (i) the Araripe Basin (Rio da Batateira and Romualdo Formations), (ii) the Rec\u0026ocirc;ncavo Basin (Maracangalha Formation), (iii) the Parna\u0026iacute;ba Basin (Itapecuru, Cod\u0026oacute; and Cabe\u0026ccedil;as), (iv) the Amazonas Basin (Alter do Ch\u0026atilde;o Formation) and (v) the Acre Basin (Pirabas Formation) (Fr\u0026oacute;es de Abreu, 1937; Castro et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1970\u003c/span\u003e; Langenhein and Beck, 1968; Carvalho, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Carvalho et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Langenhein and Beck, 1968; Paiva and Carvalho, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Paula-Freitas et al., \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Pereira et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2009a\u003c/span\u003e, \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2009b\u003c/span\u003e, \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2009c\u003c/span\u003e, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e, \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2011b\u003c/span\u003e, \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2011c\u003c/span\u003e, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Seyfullah et al., \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOf these five basins, most were present in the Cretaceous Period (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), including this study. Other basins belong to the Devonian (Cabe\u0026ccedil;as Formation) and Miocene periods (Pirabas Formation).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eOrganic geochemical analyses conducted to determine botanical affinity indicated that the ambers were products derived from conifers, especially from the families Cheirolepidiaceae and/or Podocarpaceae (Pereira et al., \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2009b\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFew macroscopic inclusions have been reported in Brazilian records (Martill et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Studies on the inclusions found in Brazilian ambers cover ostracods (Piovesan et al., \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and Ciliophera (Paiva and Carvalho, \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The occurrence of fossil resin drops associated with plants has also been described, such as dicotyledonous leaves (Langenheim and Beck, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1968\u003c/span\u003e), pollen grains from Araucariceae, Cupressaceae, and Cheirolepidiaceae (Pereira et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2009c\u003c/span\u003e, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2011a\u003c/span\u003e), and pollen grains and cones from Erdtmanithecales (Seyfullah et al., \u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Fungal spores have also been found in amber from the Amazon Basin (Pereira et al., \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2009c\u003c/span\u003e).\u003c/p\u003e"},{"header":"2. GEOLOGY SETTING","content":"\u003cp\u003eThe basement of the Araripe Basin has as its dominant lithology orthogneisses (quartz diorite to granite) of Precambrian age (from the transverse zone of the Borborema Province (Brito Neves et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Brito Neves, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2003\u003c/span\u003e)) as well as rocks from the metamorphic suite such as metavolcanics (1.12\u0026ndash;0.93 G.a.) (Kozuch et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Leite, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Van Schmus et al., \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and metagranitoids (980\u0026thinsp;\u0026minus;\u0026thinsp;920 Ma) (Leite et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Brito Neves et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Kozuch et al., \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Van Schmus et al., \u003cspan citationid=\"CR114\" class=\"CitationRef\"\u003e1995\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eProterozoic gneisses from the Rio Capibaribe Terrain (Brito Neves et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) calc-alkaline rocks containing minerals such as Ba, Rb, K, Nb, Ta, P and Ti from the Alto Moxot\u0026oacute; Terrain (de Lira Santos et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Orthogneisses, trondjenite, gabbro-diorite and migmatites from the Alto Paje\u0026uacute; Terrain (de Oliveira et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), meta-peraluminous rocks containing Rb, Th, Nb and K from the Cabaceiras Complex (Lages et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) and metagranitoids containing Co, V and Sr from the Caic\u0026oacute; Complex (Souza et al., \u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e1996\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Araripe Basin, with a sedimentary deposition area of approximately 9,000 km\u0026sup2;, is the largest inland sedimentary basin in northeastern Brazil (Fambrini et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The Mauriti Formation was deposited in an interlaced fluvial environment with a lithology of immature sandstone and conglomerates (Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Assine et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This unit was previously called the Cariri Formation, (Beurlen, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1962\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1963\u003c/span\u003e) and a new name was proposed by Gaspary and dos Anjos (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1964\u003c/span\u003e) and later adopted by Ponte and Appi (\u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). The age of this formation is still under debate, with authors (i.e., Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Assine et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) placing it in the Devonian and others arguing for a Jurassic age based on the presence of dinosaur footprints in some of its outcrops (i.e., Carvalho et al., 2024, Carvalho and Leonardi, 2024).\u003c/p\u003e \u003cp\u003eIn the chronological sequence are the Jurassic-, Brejo Santo-, and Miss\u0026atilde;o Velha formations. The former includes red shales and mudstones of lacustrine origin, whereas the latter refers to a sedimentary sequence of quartz and/or feldspathic sandstones resulting from deposition in an interlaced fluvial environment. The latter are part of the initial stage of the Opening Rift of the Ocean (Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Assine et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Cesero et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), followed by the Abaiara Formation of the Neocomian age, which is characterized by shales, siltstones, and fine sandstones of the fluvial-lacustrine environment (Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Assine et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and is associated with the rifting stage of the opening of the ocean (future Atlantic). All formations belong to the Vale do Cariri Group.\u003c/p\u003e \u003cp\u003eThe Santana Group (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) is composed of the Barbalha, Crato, Ipubi and Romualdo Formations (from oldest to youngest) (Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The Romualdo Formation, the top portion of the Santana Group, hosts a sedimentary succession that documents a marine transgression that occurred during the fragmentation of the Gondwana supercontinent and the subsequent expansion of the South Atlantic Ocean (Assine et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Neumann and Assine, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Fambrini et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThere is still disagreement about the age of the Romualdo Formation and the perspective that the boundary of Aptian-Albian stands. But it has been dated based on fossils, ostracods, and palynomorphs and considers the entire succession of the Romualdo Formation as the upper Aptian age (Regali, 2001; Rios-Netto and Regali, \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Teixeira et al., \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Arai and Assine, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSedimentary beds of this formation crop out in various areas of the Araripe Basin and exhibit different phases of marine transgression resulting from the opening of the Atlantic Ocean. However, the precise extent of these flooding episodes is not well defined. Observations reveal inconsistencies between the sedimentary patterns and macro- and microfossil records found in different geographical locations where the Romualdo Formation crops out (Nascimento et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). These discrepancies contribute to the fact that the exact configuration of the paleoenvironments remains partially unknown.\u003c/p\u003e \u003cp\u003eThe fauna of the Romualdo Formation represents a transitional environment between coastal zones and shallow marine waters (see Mabesoone and Tinoco, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Arai and Coimbra, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Berthou, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Kellner et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Bruno and Hessel, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Lima et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Melo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Recognition of echinoids, calcareous nanofossils, and foraminifera is the main evidence of a marine influence on the sedimentary processes of this formation. These records indicate the occurrence of a transgressive-regressive cycle limited by two regional discordances, which resulted in marine deposition throughout the unit (i.e., Beurlen, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1962\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1963\u003c/span\u003e; Prado et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Cust\u0026oacute;dio et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Melo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Pedrosa et al., \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eRegarding macroflora, the Romualdo Formation hosts the remains of the genera \u003cem\u003eBrachyphyllum\u003c/em\u003e and \u003cem\u003ePseudofrenelopsis\u003c/em\u003e as the most common fossils, but it can also contain angiosperms and gymnosperms (most abundant) (Duarte, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Lima et al., \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). However, fossils of this character are poorly preserved, making taxonomic studies difficult.\u003c/p\u003e \u003cp\u003eThe Romualdo Formation is mainly composed of sandstones interspersed with dark grey shales, which are rich in organic matter, as well as greenish shales and marls at their base. Most of the upper part of this formation consists of fossiliferous concretions found in green shales, which contain large numbers of vertebrates (i.e., osteichthyes), ichnofossils (i.e., coprolites), ostracods, and, less frequently, plants (Arai and Coimbra, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Berthou, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Colin and D\u0026eacute;p\u0026ecirc;che, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Mabesoone et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Coimbra et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Tom\u0026eacute; et al., \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). In the upper part, layers of limestone-containing invertebrates are covered by thin layers of sandstone, siltstone, and shale. These sediments host fossils from mixohaline and marine environments, including crustaceans, mollusks and foraminifera, both benthic and planktonic (Beurlen, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1971\u003c/span\u003e; Assine, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Prado et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Sobradinho section (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) is one of the most studied sites in the Romualdo Formation, because it shows the most complete section of this formation, in the eastern portion of the Araripe Basin (Cust\u0026oacute;dio et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Fursich et al., 2019; Arai e Assine, 2020; Melo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Bom et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Kroth et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"3. MATERIALS AND METHODS","content":"\u003cp\u003eThe samples used in this study and the methodologies adopted are listed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSamples and analyses carried out on them. Caption. L. Petrographic slides. C. Confocal microscope. LA-ICP-MS. Laser Ablation Inductively Coupled Plasma Mass Spectrometry. SEM/EDS. Scanning Electron Microscope / X-ray Energy Dispersive Spectroscopy.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eN\u0026deg; amostra (DGEO-CTG-UFPE)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eL\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eLA-\u003c/p\u003e \u003cp\u003eICPMS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMEV/EDS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8800\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 27 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8801\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 27 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8803\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 27 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8804\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8805\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9121\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9123\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9124\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9125\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9126\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9128\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9129\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9131\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE - perfil 28 m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eROM-03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eROM-04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTest\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePyrite\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSobradinho, CE\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Location of the study area\u003c/h2\u003e \u003cp\u003eThe sample collection site is located in the municipality of Jardim, State of Cear\u0026aacute; (CE) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), within the stratigraphic sequence of the Sobradinho site (Araripe et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) associated with the shale layers at 28 m at the top of Romualdo Formation (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The Sobradinho site belongs to the Romualdo Formation and in the map (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) the amber site is located on top of Exu Formation, that is due to the fact that the outcrop was exposed in the bed of an intermittent river and has no exposed surface.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Samples collected for the study\u003c/h2\u003e \u003cp\u003eA total of 21 complete or fragmented samples were collected during two field expeditions (the first in 2019 and the second in 2022). The ambers were stored in the Scientific Collection of Paleontology at the Department of Geology (DGEO), Center of Technology (CTG), UFPE (Federal University of Pernambuco). Those from the first collection were included under DGEO-CTG-UFPE numbers 8800 to 8806 and those from the second collection were included under DGEO-CTG-UFPE numbers 9120 to 9133.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e3.3 Study of the thin sections using a petrographic microscope\u003c/h2\u003e \u003cp\u003eAfter preparing the slides, they were analyzed using a Zeiss Axioscope AI Imager petrographic microscope with an attached digital camera to capture images, which allowed the inclusions to be identified, described, and recorded. The images were acquired using the Zen Blue Zeiss program and saved in .tif format.\u003c/p\u003e \u003cp\u003eTo better characterize and detail the inclusions, thin sections of selected amber fragments were prepared. To achieve this, adopting a safe and effective lamination method was necessary, as this was not a standard procedure (Tomaoka and Ricardi-Branco., 2025). Samples were prepared for lamination at the Palaeohydrogeology Laboratory, part of the Institute of Geosciences, State University of Campinas (UNICAMP).\u003c/p\u003e \u003cdiv id=\"Sec7\" class=\"Section3\"\u003e \u003ch2\u003e3.3.1 \u003cem\u003eMaking the petrographic thin section\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eThe references Corral et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1999\u003c/span\u003e), Nascimbene and Silverstein (\u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) and Sadowski et al. (\u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) were used as the basis for creating the slides. They described techniques for preparing amber for lamination such as sanding, polishing, and impregnation in epoxy resin using a vacuum chamber.\u003c/p\u003e \u003cp\u003eTherefore, to make the thin sections of an amber sample, it must first be cleaned with distilled water to remove traces of the matrix; then, in the case of thicker and/or opaque samples, thickness was reduced using water sandpaper graded from 150 to 2000 (in ascending order) to achieve a good finish and better visualization. A Stemi V6 ZEISS stereomicroscope was used to select samples with possible bioinclusions and the best place to make thin sections.\u003c/p\u003e \u003cp\u003eTwenty-two (22) thin sections of the different samples were made; so therefore, getting a better idea of the occurrences of inclusions in the samples was possible (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All fragments were photographed with a CANON EOS Rebel T5i camera using a light table (Tomaoka and Ricardi-Branco, \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFirst, the chosen and prepared material was impregnated with epoxy resin until it was halfway up the sample in a delimited space. After sufficient hardening of the epoxy, a new layer of epoxy was placed on the samples until they were completely covered. They were then placed in a vacuum chamber so that the resin could fill all the gaps, avoiding the generation of reflections by light from the petrographic microscope (Tomaoka and Ricardi-Branco, \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The last stage was gluing the impregnated sample to a petrographic slide and subsequently sanding and polishing it until it reached the desired thickness, in this case 100 micrometers (Tomaoka and Ricardi-Branco, \u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e3.4 Confocal LASER microscopy imaging\u003c/h2\u003e \u003cp\u003eThe principle of confocal microscopy differs from that of conventional microscopy (Hein et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Gonz\u0026aacute;lez and Halpern, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Borlinghaus, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eLaser confocal microscopy was used to complement the study of the organic inclusions. This technique allows the matrix to be isolated, and the kerogen and organic molecules to be better mapped. This stage was carried out on the premises of the National Institute of Science and Technology for Photonics Applied to Cell Biology (INFABIC) at UNICAMP using the Zeiss LSM780 NLO inverted microscope, and the slides viewed using this equipment can be seen in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.5 SEM/EDS\u003c/h2\u003e \u003cp\u003eScanning electron microscope (SEM) and energy-dispersive spectroscopy (EDS) was used to acquire a better image of micro-inclusions presents in the amber samples.\u003c/p\u003e \u003cp\u003eModel Zeiss Leo 430i with EDS detector Oxford model 7059 (10 mm\u003csup\u003e2\u003c/sup\u003e), software ISIS.\u003c/p\u003e \u003cp\u003eThe samples were coated with gold using the Quorum Q 150TES.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.6 Elemental analysis by Laser Ablation (LAICP-MS)\u003c/h2\u003e \u003cp\u003eLaser Ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) or laser ablation inductively coupled plasma mass spectrometry was used to complete the geochemical characterization of ambers. To accomplish this, samples with the potential for analysis had to be prepared. The samples used to identify the inorganic species were previously cut and sanded (according to the same procedure described above) to obtain thin, smooth, and flat surfaces for ablation. The samples were then placed in a holder and fixed using double-sided tape. Three ablation lines were created based on the size of each sample to cover the entire surface. The samples were selected to cover the widest range of characteristics among the available fossil resins; thus, DGEO-CTG-UFPE samples 8800, 8804, 9121 and 9132 were selected.\u003c/p\u003e \u003cp\u003eThe elements \u003csup\u003e13\u003c/sup\u003eC, \u003csup\u003e34\u003c/sup\u003eS, \u003csup\u003e57\u003c/sup\u003eFe, \u003csup\u003e51\u003c/sup\u003eV, \u003csup\u003e28\u003c/sup\u003eSi, \u003csup\u003e43\u003c/sup\u003eCa, \u003csup\u003e59\u003c/sup\u003eCo, \u003csup\u003e63\u003c/sup\u003eCu, \u003csup\u003e45\u003c/sup\u003eSc, \u003csup\u003e111\u003c/sup\u003eCd, \u003csup\u003e107\u003c/sup\u003eAg, \u003csup\u003e31\u003c/sup\u003eP, \u003csup\u003e75\u003c/sup\u003eAs, \u003csup\u003e66\u003c/sup\u003eZn and \u003csup\u003e118\u003c/sup\u003eSn were evaluated to identify the inorganic constituents of the amber samples. The ablation parameters used were: 110 \u0026micro;m spot, 110 \u0026micro;m s-1 scan speed and 60% intensity and plasma conditions of 1200 W RF power, 1.6 L min-1 auxiliary gas and 1.2 L min-1 nebulizer gas. This technique combines plasma ablation and spectrometry to determine the quantity of inorganic elements present in the samples.\u003c/p\u003e \u003c/div\u003e"},{"header":"4. RESULTS","content":"\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Macroscopic characterization\u003c/h2\u003e \u003cp\u003eIn general, the ambers under study range in color from light brown (caramel) to dark brown, or 10YR 2/2, 3/2, 4/4, 3/4, 6/6, 5/6, 4/6, 3/6, 7/8, 6/8, 5/8, 4/8, 7/10, and 6/10 (according to the Munsell color chart), and some samples can even be light brown (caramel) and completely opaque (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The shape varies from round to oval, and fractures occur frequently, including those in the cortex. Most samples show more than one resin deposition flow (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The inclusions cannot be discerned by the naked eye. The outer surface (cortex) of the samples is more altered than the inner surface.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e4.2 Bioinclusions\u003c/h2\u003e \u003cdiv id=\"Sec14\" class=\"Section3\"\u003e \u003ch2\u003e4.2.1 \u003cem\u003eAnalyses under Petrographic and Confocal Microscopy.\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eCONIFEROPHYTA\u003c/p\u003e \u003cp\u003eSecondary xylem in transverse view\u003c/p\u003e \u003cp\u003eSample. DGEO-CTG-UFPE 9124\u003c/p\u003e \u003cp\u003eDescription. Fragment of secondary xylem in transverse view; tracheids mostly polygonal to sub-radial, with visible walls separated by diffuse parenchyma. Arranged in rows; walls appear lignified (Figs.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec, \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ed, \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ee and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ef). Dimensions: tracheids 25\u0026ndash;30 \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\mu\\:\\)\u003c/span\u003e\u003c/span\u003em long axis.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSecondary xylem radial longitudinal view\u003c/p\u003e \u003cp\u003eSample. DGEO-CTG-UFPE 9124\u003c/p\u003e \u003cp\u003eDescription. Fragment of three tracheids in the radial longitudinal view; tracheids with lignified walls and polygonal (hexagonal, pentagonal), bordered pits in an araucarioid pattern. Dimensions: fragment 230 \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\mu\\:\\)\u003c/span\u003e\u003c/span\u003em of length, tracheids 40\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\mu\\:\\)\u003c/span\u003e\u003c/span\u003em wide, pits 10 \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\mu\\:\\)\u003c/span\u003e\u003c/span\u003em wide (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea, \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003ePOLLENITES H. Potoni\u0026eacute;, 1993\u003c/p\u003e \u003cp\u003eAnterturma VARIEGERMINANTES Potoni\u0026eacute;, 1975\u003c/p\u003e \u003cp\u003eTurma SACCITES Erdtman, 1947\u003c/p\u003e \u003cp\u003eSubturma MONISACCITES (Chitaley, 1951) Potoni\u0026eacute; \u0026amp; Kremp,1954\u003c/p\u003e \u003cp\u003eInfraturma CIRCUMPOLLINI Klaus, 1960\u003c/p\u003e \u003cp\u003eG\u0026ecirc;nero \u003cem\u003eClassopollis\u003c/em\u003e Pflug, 1960\u003c/p\u003e \u003cp\u003e \u003cem\u003eClassopollis\u003c/em\u003e spp.\u003c/p\u003e \u003cp\u003eSamples. DGEO-CTG-UFPE, 9126 and 9132\u003c/p\u003e \u003cp\u003eDescription. Monoporate pollen grains with circular equatorial outline, isolated, exine with a triangular mark, exoexine with a belt. Presence of striations in varying numbers; exine ornamentation ranging from punctate to reticulate (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eDimensions. 30\u0026ndash;34 \u0026micro;m\u003c/p\u003e \u003cp\u003eFUNGI\u003c/p\u003e \u003cp\u003eSamples. DGEO-CTG-UFPE 9126, 8805, 9128, 9131, 9132.\u003c/p\u003e \u003cp\u003eDescription. Mycelium with translucent hyphae showing varying degrees of preservation, ranging from well-preserved to degraded (Figs.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea-f). Hyphae in decomposition (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ea), continuous hyphae, simple and branched, septate (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003eb) and non-septate (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ed); Spiral hypahes (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ee). In general, filament growth occurs from the edges of the amber toward its interior, with the density of hyphae being higher at the periphery Mycellium; (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003ef).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e4.3 Elemental analysis\u003c/h2\u003e \u003cdiv id=\"Sec16\" class=\"Section3\"\u003e \u003ch2\u003e4.3.1 Scanning Electron Microscope (SEM) and Energy Dispersive Spectrometer (EDS) mapping.\u003c/h2\u003e \u003cp\u003eSEM and EDS analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003e) was carried out in order to better understand the composition of the sample's surface, as we were able to observe the presence of mineral species near the vascular plant structures when observing the slide of sample DGEO-CTG-UFPE 9124 under the SEM (Figs.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003ea), and Figs.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e8\u003c/span\u003eb, c and d show the elemental maps made with EDS of chemical elements such as Al and Ca.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section3\"\u003e \u003ch2\u003e4.3.2 \u003cem\u003eLaser Ablation (LAICP-MS)\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eAccording to the analysis of the DGEO-CTG-UFPE 8800 sample using the three ablation lines (Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e), higher signal intensities were obtained for the elements sulphur (S), zinc (Zn), phosphorus (P) and iron (Fe). For the elements S, P, and Zn, the distribution in amber apparently showed a more homogeneous behavior when compared to Fe, which showed an even greater intensity at the extremities.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe intensities of the signals recorded for DGEO-CTG-UFPE 8804, which had a redder color and greater homogeneity, are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e. The same elements were found in DGEO-CTG-UFPE 8804 as in the previous sample (DGEO-CTG-UFPE 8800), S, P, Zn, and Fe, but with a difference in their distribution. In addition, copper (Cu) was also present over the entire surface of the sample, but it was distributed heterogeneously, as was Zn, with peaks throughout the ablation.\u003c/p\u003e \u003cp\u003eFor sample DGEO-CTG-UFPE 9121, the same elements, S, P, Zn, and Fe, had higher signal intensities than the other two samples (DGEO-CTG-UFPE 8800 and 8804), but there was also a higher signal intensity for calcium (Ca), as illustrated in Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e. The distribution of most of the elements on the surface of the sample was more homogeneous, with Fe showing the most pronounced distribution, especially in the darker amber areas.\u003c/p\u003e \u003cp\u003eDGEO-CTG-UFPE 9123 was visually more homogeneous in color; however, it had small darker regions, as shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig9\" class=\"InternalRef\"\u003e9\u003c/span\u003e, which also shows the signals obtained for the elements evaluated. As in the previous three samples, S, P, and Fe exhibited higher intensities than the other elements explored. In this case, the distribution of the elements at the start of the ablation showed an even greater intensity, drawing attention to the presence of vanadium (V) expressed on ablation line 3, which coincides with the darkest region of the sample. In addition, the distribution of Fe in the sample was heterogeneous on the surface.\u003c/p\u003e \u003cp\u003eIn all the graphs shown, the \u003csup\u003e13\u003c/sup\u003eC signal was used to identify the beginning and end of sample ablation (in which the signal increases at the beginning and decreases at the end). Si was not expressed in the above graphs because its intensity was higher than that of the other elements. This causes the intensity of the signals to decrease, and they cannot be graphically evaluated together.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"5. DISCUSSION","content":"\u003cp\u003eOwing to their nature, ambers offer unique opportunities for research, especially if modern analysis techniques such as confocal microscopy and laser ablation (LA-ICP-MS) are used. However, to obtain better results, it was important to acquire the ability to polish the amber before lamination, which enabled the prospecting of different inclusions, was necessary.\u003c/p\u003e \u003cp\u003eThe amber samples studied have a range of colors and can be more opaque or translucent, sometimes weathered, which makes the fragments partially or completely matte and opaque, as is the case with sample DGEO-CTG-UFPE 9122. Therefore, after lamination, the interiors of most of the samples studied could be accessed.\u003c/p\u003e \u003cp\u003eThe banded pattern of the ambers (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e) is a consequence of the successive flows of resin, as well as their shape. Although the shape is also influenced by gravity as the resin slides down the aerial part of the plant, acquiring an amygdaloid shape (Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Labandeira, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Lozano et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Ambers derived from resins secreted by roots, in addition to having an amygdaloid appearance, take longer to solidify because of the underground conditions under which the resin is secreted (Speranza et al., \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBurial within the soil profile of the still ductile resin or at the final deposition site can also influence changes in the original shape (Labandeira, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Lozano et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Moreover, the frequent microfractures observed in the ambers are due to the weight of the sedimentary load that was successively deposited above the layer carrying them (Corral et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1999\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003e5.1 Bioinclusions\u003c/h2\u003e \u003cp\u003eIn the amber studied, in addition to micro inclusions, there are pseudo-inclusions, according to some authors, (Lozano et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Quinney et al., \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) these correspond mostly to sap droplets, probably exuded along with the resin because they accompany its flow.\u003c/p\u003e \u003cp\u003eResinous filamentous fungi are the most likely candidates for colonizing non-liquid environments, as is the case with plant resins; resins provide nutrients to fungi (Wu et al., \u003cspan citationid=\"CR115\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Pseudo-inclusions with sap droplets incorporated into the resin can be more nutritious than the resin, providing a favorable environment for the development of mycelia (Lozano et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the studied ambers, the most abundant bioinclusions observed and common to all the samples analyzed were fungi, which were present at various stages of preservation and with a variety of structures, such as mycelia (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e7\u003c/span\u003e), reproductive structures (i.e., conidiophores), and spores. The fungal populations were denser on the outside of the samples, but they were also observed on the inside, which suggests that they developed after the resin fell from the aerial part of the plant, or that many of the samples studied may have been generated close to the forest floor or even by the roots (Speranza et al., \u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), because it would take longer for the resin to become hard (polymerized) enabling the fungus to enter.\u003c/p\u003e \u003cp\u003eThe presence of spores and pollen grains as inclusions is crucial, especially considering that the genus \u003cem\u003eClassopollis\u003c/em\u003e, possibly derived from the Cheirolepidiaceae family, is extremely abundant in the Romualdo Formation shales (Pereira et al., \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2009a\u003c/span\u003e, \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2009c\u003c/span\u003e; Arai and Assine \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Nascimento et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). \u003cem\u003eClassopollis\u003c/em\u003e is, therefore, of parautochthonous origin and is possibly incorporated into the resin near its parent tree during the breeding year (N\u0026eacute;raudeau et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Pereira et al., \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Piovesan et al., 2020). This genus is paleoenvironmental marker of the Cretaceous (Carvalho et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Pocock and Jansonius, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e1961\u003c/span\u003e; Taylor et al., \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) suggests a hot and dry climate. Furthermore, the abundant occurrence of Classopollis in the shales of the upper portion of the Romualdo Formation is considered further evidence of a proximal marine environment (Arai and Assine \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The abundance of continental palynomorphs in the layers where amber occurs contributes to an interpretation of an environment with more continental influence (Teixeira et al.; \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Ferreira et al. 2025).\u003c/p\u003e \u003cp\u003eThe anatomical characteristics of the xylem fragment in the longitudinal radial view, showing tracheids with lignified walls and polygonal (hexagonal, pentagonal) bordered pits in an araucarioid pattern, correspond to the characteristics of the wood of the conifer genera described for the Romualdo Formation (Batista et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). These bioinclusions, together with the paleobotanical records of the Romualdo Formation, confirm that the forests associated with the banks of rivers or even the coastal region of a lagoon could have been surrounded by coniferous vegetation, including the Cheirolepidiceae family (Batista et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2018\u003c/span\u003e, Teixeira et al. \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Ferreira et al. 2025).\u003c/p\u003e \u003cp\u003eThe presence of pollen grains of the genus \u003cem\u003eClassopollis\u003c/em\u003e has already been reported in shales at the same level where the ambers were collected (Nascimento et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and in Cenomanian ambers collected in layers of clays rich in organic matter from the Mayenne region in France (N\u0026eacute;raudeau et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and in various mid Cretaceous deposits from France.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003e5.2 Elemental analysis\u003c/h2\u003e \u003cp\u003eWhen analyzing the results of the elemental composition obtained using the Laser Ablation technique (LA-ICP-MS), various elements were present, the intensity of which varied according to their distribution in each sample. Elements such as V, Cu, Sc, Cd, Sn, and Zn exhibited low intensities in the samples. In contrast, the intensities of the S, P, Fe, and Zn peaks were higher. The elements Fe, Zn, and sometimes V (sample DGEO-CTG-UFPE 9123) seemed to coincide with the darker-colored regions, which seemed to influence the shade of the samples. The intensities of P and S stood out as elements common in the rocks of the Santana Group, as does Si, which also predominated in the SEM/EDS analyses. Fe continues to be a frequent element, together with Mn in the samples analyzed when interpreted with the results obtained from the SEM/EDS elemental analyses. The presence of Ca demarcating the tracheids in sample DGEO-CTG-UFPE 9124 could indicate that calcium was dissolved in the water used by the plant after the inclusion of the xylem fragment deposited in the walls.\u003c/p\u003e \u003cp\u003eThe composition of ambers varies considerably depending on many external factors to the plant that produced the resin flow, including the area of provenance, paleosol, regional geology, diagenetic changes and paleoclimate (Cockx and McKellar, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Pe\u0026ntilde;alver et al., \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThus, the oxidation of Fe, for example, during the early diagenesis of the ambers and the colonization by the fungi that degrade the organic matter (resin) rich in P, C and S (indispensable elements for the development of fungi) may have partly influenced their coloring and the development of the weathering crust that the samples show (Cockx and McKellar, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Mart\u0026iacute;nez-Delcl\u0026ograve;s et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Pe\u0026ntilde;alver et al., \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, Natkaniec-Nowak et al., \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2026\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe elemental composition found in the samples suggests that the elemental variety presented was acquired by resin-secreting plants through groundwater (Aquilina et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), used by the vegetation that inhabited the shores of the marine basin (Melo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In addition, according to Aquilina et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) no enrichment of typical marine elements such as Na in completely solidified samples was shown. Therefore, enrichment in minerals anomalous to amber may have occurred in two ways: externally, i.e., by water droplets that managed to reach the resin before it was solidified; or internally, before the resin was secreted (Aquilina et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), since the resin is produced inside the plant and, in some cases, in specialized ducts (Langenheim, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2003\u003c/span\u003e), from elaborate substances that the plant stores in the parenchyma (Aquilina et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). These substances are derived from sap, which in turn is derived from solutions absorbed by the soil; therefore, the composition of the resin is directly influenced by the composition of groundwater and the biological processes related to resin production (Munns, \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2002\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e5.3 Taphonomy\u003c/h2\u003e \u003cp\u003eThe stratigraphic level located approximately 28 m from the composite column of the Sobradinho Site (Araripe et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2025\u003c/span\u003e) was interpreted, based on the of benthic foraminifera, present as stressful conditions associated with a marginal marine environment with variations in salinity; the marine condition was reinforced by the presence of echinoderms and calcareous nannofossils (Prado et al., \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Cust\u0026oacute;dio et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Melo et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Araripe et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2022\u003c/span\u003e, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Pedrosa et al., \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, the continental origin of the ambers indicates the occurrence of marine ingressions (Cust\u0026oacute;dio et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) that affected the dryland forests or the occurrence of flooding from freshwater streams.\u003c/p\u003e \u003cp\u003eBoth the first and second possibilities involve removing a portion of the soil in which the ambers were originally deposited and transporting it to the final place of deposition in the marine environment; therefore, they represent an allochthonous portion of the fossiliferous assemblage (Schmidt and Dilcher, \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Labandeira, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). At the same levels where the ambers were collected, plant debris and continental palynomorphs are very abundant. Thus continental bioclasts contributed with a decrease in the abundance of marine microfossils (Cust\u0026oacute;dio et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Teixeira et al.; \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Fereira et al. 2025; Araripe et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2025\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe conifer-related biological inclusions observed in the ambers and even ambers were possibly derived from resins produced by conifers and indicated the presence of a forest composed of species resistant to the water deficit that would have inhabited near the coast, exhibiting an arid climate (Batista et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Arai and Assine, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, the presence of groundwater would have enabled the trees to grow and cannot be ruled out. The influence of underground aquifers is most evident in the elemental composition of the amber, where Ca, Si, V, Fe, and Zn ions could have been solubilized from the basin\u0026rsquo;s basement rocks (Aquilina et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; C\u0026uacute;stodio et al., 2017; Fereira et al. 2025).\u003c/p\u003e \u003c/div\u003e"},{"header":"6. CONCLUSION","content":"\u003cp\u003eThe studied samples showed diverse colors, transparencies, and weathering influenced by geological and environmental processes over time. The observed banded patterns, amygdaloid and deformed forms, and microfractures reflect the dynamics of resin flows, gravity, burial, and sedimentary pressure. These aspects highlight the importance of depositional and diagenetic contexts in the formation and preservation of ambers.\u003c/p\u003e \u003cp\u003eThe presence of inclusions such as \u003cem\u003eClassopollis\u003c/em\u003e, spores, filamentous fungi, and pseudo-inclusions reinforces the role of ambers as biological and paleoenvironmental archives. The genus \u003cem\u003eClassopollis\u003c/em\u003e, associated with the gymnosperm Cheirolepidiaceae, indicates coastal vegetation that tolerated the arid climate that prevailed during the deposition of the upper part of the Romualdo Formation (Aptian).\u003c/p\u003e \u003cp\u003eThe results of the elemental analyses revealed a variable composition influenced by external factors such as paleosol, regional geology, diagenetic changes, and paleoclimate. Elements such as Fe, P, S, and Zn stand out and are possibly responsible for the coloration. The lack of enrichment in marine elements such as Na suggests that the composition of the resin is directly linked to underground water absorbed by the coastal vegetation that inhabited the margins of the basin, leading to the Romualdo Formation.\u003c/p\u003e \u003cp\u003eThe combination of innovative methodological approaches and a detailed analysis demonstrates the potential of ambers as natural archives, paving the way for future interdisciplinary research in this field.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eWe would like to thank CAPES for the master\u0026rsquo;s scholarship, FAPESP for funding this research through Processes 2019/16727-3 and 23/16631-1, CNPQ for the researcher grant 307333/2021-3, the Paleohydrogeology Laboratory for the infrastructure that facilitated this research, the Electron Microscopy Laboratory (SEM), the Lamination Laboratory (LAM), Dr. Marco Aur\u0026eacute;lio Zezzi, the Group of Spectrometry, Sample Preparation and Mechanization (GEPAM), and Eduarda Machado for the analyses.\u003c/p\u003e \u003cp\u003eFurther image data are available at Reposit\u0026oacute;rio de Dados de Pesquisa da Unicamp. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.25824/redu/TYR2HP\u003c/span\u003e\u003cspan address=\"10.25824/redu/TYR2HP\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003e\u0026Aacute;lvarez-Parra, S., Buenocebollada, C. 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NE Brazil.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWu, H., Li, L., \u0026amp; Ding, M. (2018). The first cyclaxyrid beetle from Upper Cretaceous Burmese amber (Coleoptera: Cucujoidea: Cyclaxyridae). \u003cem\u003eCretaceous Research\u003c/em\u003e, 91. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.cretres.2018.05.015\u003c/span\u003e\u003cspan address=\"10.1016/j.cretres.2018.05.015\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"palaeobiodiversity-and-palaeoenvironments","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pbpe","sideBox":"Learn more about [Palaeobiodiversity and Palaeoenvironments](https://www.springer.com/journal/12549)","snPcode":"12549","submissionUrl":"https://www.editorialmanager.com/pbpe/default2.aspx","title":"Palaeobiodiversity and Palaeoenvironments","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Hyphae, Aptian, Cheirolepidiaceae, Classopollis","lastPublishedDoi":"10.21203/rs.3.rs-9349105/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9349105/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis study advances the understanding of ambers as important paleobiological and paleoenvironmental archives for the Romualdo Formation of the Araripe Basin and the Lower Cretaceous (Aptian) of Gondwana. Twenty-one (21) ambers collected from the Sobradinho Site, were investigated. Modern and traditional techniques, such as confocal microscopy, petrography, and Laser Ablation ICP-MS (LA-ICPMS), were used, allowing for a detailed analysis of the physical, chemical, and taphonomic characteristics of these fossils. Among the inclusions were fungi, pollen grains of \u003cem\u003eClassopollis\u003c/em\u003e and xylem fragments. The analyzed inclusions suggest the presence of coastal forests dominated by gymnosperms with the presence of the Cheirolepidiaceae family. Geochemical analyses revealed a significant enrichment in elements such as sulfur, phosphorus, iron, and zinc, which directly influenced the physical properties and coloration of the samples. From a taphonomic perspective, these ambers represent an allochthonous component that was transported from the forest floor to the coastal marine environment where they were finally deposited. This interpretation was corroborated by the abundance of plant and amber microfragments found in the shales associated with the samples studied. Paleoenvironmental conditions allow to infer a prevailing arid climate, with forests adapted to hydric deficits but potentially sustained by groundwater aquifers. The results offer new perspectives on the evolution of the Araripe Basin ecosystems during the Cretaceous period.\u003c/p\u003e","manuscriptTitle":"Multiproxy Analysys of Ambers From the Romualdo Formation (Cretaceous), Araripe Basin, Ceará State, Brazil","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-30 19:28:39","doi":"10.21203/rs.3.rs-9349105/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2026-04-22T11:54:06+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-22T11:52:01+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-10T04:10:58+00:00","index":"","fulltext":""},{"type":"submitted","content":"Palaeobiodiversity and Palaeoenvironments","date":"2026-04-08T11:34:27+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"palaeobiodiversity-and-palaeoenvironments","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pbpe","sideBox":"Learn more about [Palaeobiodiversity and Palaeoenvironments](https://www.springer.com/journal/12549)","snPcode":"12549","submissionUrl":"https://www.editorialmanager.com/pbpe/default2.aspx","title":"Palaeobiodiversity and Palaeoenvironments","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"4b1178ca-dd35-499c-9207-83befaa6bd97","owner":[],"postedDate":"April 30th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-30T19:28:40+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-30 19:28:39","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9349105","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9349105","identity":"rs-9349105","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}