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    Thermal evolution of light hydrocarbon fingerprints in biodegraded oils from Ordovician reservoirs, Tabei Uplift, Tarim Basin

    AbstractWithin the Tabei Uplift of the Tarim Basin, Ordovician reservoirs in both the northern Halahatang (N-Halahatang) and western Lunnan (W-Lunnan) areas experienced extensive biodegradation during the Late Hercynian (Permian). Subsequent Himalayan (Neogene–Quaternary) tectonism induced divergent burial-thermal histories: the N-Halahatang reservoirs underwent intensive maturation (> 6,500 m depth; 1.02–1.22% Ro), while the W-Lunnan reservoirs experienced milder maturation (< 5,800 m depth; 0.70–0.85% Ro). Despite similar δ13Coil values indicating genetic affinity, the relatively deeply buried biodegraded oils from the N-Halahatang area contain abundant C6–C8 light hydrocarbons (LHs), while the biodegraded oils from the W-Lunnan area exhibit only trace amounts of C6–C8 LHs. To elucidate the evolution of LHs compositions and fingerprints in biodegraded oils under thermal maturation, and to determine whether the more enriched C6–C8 LHs in the N-Halahatang oils can be attributed to enhanced burial-thermal maturation, two relatively shallower-burial biodegraded oils (Well LG40: slight to moderate biodegradation‌; Well LG7: heavy to severe biodegradation) from the W-Lunnan area were artificially pyrolyzed to various maturities. Subsequently, LH parameters of the pyrolyzed oils were compared with those of the naturally matured, deeply buried oils (heavy to severe biodegradation) from the N-Halahatang area. The results indicated that both biodegraded oils generated C6–C8 LHs through thermal cracking, and the more severely biodegraded oil (Well LG7) exhibited a lower LH maximum yield than that from Well LG40. Certain parameters for organic matter type classification (n-C7–DMCP–MCH and 3RP–5RP–6RP diagrams) generally remained applicable during thermal maturation, whereas most parameters for secondary alteration identification and maturity assessment were significantly compromised. Additionally, LH parameters of the N-Halahatang oils (1.02–1.22% Ro) matched those of the LG7 pyrolyzed oils at EasyRo = 1.00–1.20%, confirming that the enriched C6–C8 LHs in the N-Halahatang oils can be attributed to cracking of biodegraded oils (with ‌biodegradation levels equivalent to Well LG7‌) under intense burial-thermal maturation. Furthermore, the potential C6–C13 LHs derived from biodegraded oil cracking constitute 11–16 wt% of N-Halahatang’s liquid hydrocarbon resources.

    Data availability

    Datasets used and analyzed during the current study are available from the corresponding author upon reasonable request.
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    Download referencesAcknowledgementsThe authors thank Dr. Jinzhong Liu, Mr. Yong Li, Dr. Zewen Liao, and Dr. Yankuan Tian for their assistance in laboratory analyses. The authors are also grateful to the anonymous reviewers for their constructive suggestions.FundingThis work was supported by the National Natural Science Foundation of China (Grant Nos. 42173056 and 42572184), the project Theory of Hydrocarbon Enrichment under Multi-Spheric Interactions of the Earth (Grant No. THEMSIE04010104). This is also a contribution to the Special Fund for the Strategic Priority Research Program of the Chinese Academy of Sciences (Grant No. XDA14010103).Author informationAuthors and AffiliationsState Key Laboratory of Deep Earth Processes and Resources, Guangzhou Institute of Geochemistry, Chinese Academy of Sciences, Guangzhou, 510640, ChinaYuwei Yang, Yuhong Liao, Yueyi Huang, Bin Cheng, Huanyu Lin & Yunpeng WangState Key Laboratory of Advanced Environmental Technology, Guangzhou Institute of Geochemistry, Chinese Academy of Sciences, Guangzhou, 510640, ChinaYijun Zheng & Ping’An PengUniversity of Chinese Academy of Sciences, Yuquan Road, Beijing, 100049, ChinaYuwei Yang, Yuhong Liao, Yijun Zheng, Bin Cheng, Huanyu Lin, Yunpeng Wang & Ping’An PengCAS Center for Excellence in Deep Earth Science, Guangzhou, 510640, ChinaYuwei Yang, Yuhong Liao, Yijun Zheng, Bin Cheng, Huanyu Lin, Yunpeng Wang & Ping’An PengBiogas Institute of Ministry of Agriculture and Rural Affairs, Chengdu, 610041, ChinaYueyi HuangAuthorsYuwei YangView author publicationsSearch author on:PubMed Google ScholarYuhong LiaoView author publicationsSearch author on:PubMed Google ScholarYueyi HuangView author publicationsSearch author on:PubMed Google ScholarYijun ZhengView author publicationsSearch author on:PubMed Google ScholarBin ChengView author publicationsSearch author on:PubMed Google ScholarHuanyu LinView author publicationsSearch author on:PubMed Google ScholarYunpeng WangView author publicationsSearch author on:PubMed Google ScholarPing’An PengView author publicationsSearch author on:PubMed Google ScholarContributionsYuwei Yang: Investigation, Methodology, Formal analysis, Writing-Original Draft; Yuhong Liao: Supervision, Conceptualization, Funding acquisition, Validation, Writing-Reviewing and Editing; Yueyi Huang: Data Curation; Yijun Zheng: Writing-Reviewing and Editing; Bin Cheng: Resources; Huanyu Lin: Visualization; Yunpeng Wang: Project administration, Funding acquisition; Ping’An Peng: Funding acquisition.Corresponding authorCorrespondence to
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    Reprints and permissionsAbout this articleCite this articleYang, Y., Liao, Y., Huang, Y. et al. Thermal evolution of light hydrocarbon fingerprints in biodegraded oils from Ordovician reservoirs, Tabei Uplift, Tarim Basin.
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    KeywordsBiodegradationBurial-thermal maturationLight hydrocarbonsOrdovician reservoirsTabei UpliftTarim Basin More

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    Integrated geographical and ecological analysis reveals environmental drivers of Gardenia jasminoides distribution and chemical variation

    AbstractGardenia jasminoides, a widely distributed resource rich in Crocin, has generated substantial market demand due to its potential value as a saffron substitute. This necessitates the exploration of efficient and sustainable cultivation strategies to obtain target compounds for specific purposes. To enhance cultivation efficiency and secure supply chains, we integrated MaxEnt modeling, spatial interpolation, and geodetector analysis. This framework aimed to predict suitable habitats for G. jasminoides across China, map spatial variation in bioactive compounds including Crocin, Gardenia Yellow, and Geniposide, and identify environmental drivers influencing their distribution. MaxEnt achieved high predictive accuracy (AUC = 0.960), identifying Jiangxi, Zhejiang, and Guangdong as key high-suitability regions. Precipitation of the driest month and human population density emerged as dominant factors shaping species distribution. Spatial gradients revealed that Crocin and Gardenia Yellow decrease from southwest to northeast, whereas Geniposide exhibits latitudinal differentiation characterized by higher concentrations in northern regions. Geodetector analysis highlighted vegetation type as the primary driver of compound variation, with q values of 0.618 for Crocin, 0.606 for Gardenia Yellow, and 0.639 for Geniposide. These results indicate that the accumulation of target compounds is strictly modulated by ecological niches, where specific vegetation types drive metabolic differentiation through microclimate regulation and interspecific competition. Based on these findings, we advocate for an industry-oriented divergent cultivation strategy. Southwestern China should be prioritized for Crocin-rich germplasm to support the natural pigment industry, whereas northern regions are designated as premium zones for pharmaceutical-grade Geniposide sourcing. Furthermore, recognizing vegetation type as a critical driver facilitates the implementation of targeted habitat management techniques. These findings provide a direct guide for designating priority cultivation zones and optimizing harvest timing to maximize the yield of target compounds for specific industrial uses.

    Data availability

    The datasets used and/or analysed during the current study available from the corresponding author on reasonable request.
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    Download referencesAcknowledgmentsWe sincerely thank all the scholars and farmers who have helped us during the sample collection process. We would like to express our gratitude to all previous researchers who helped and references for this study.FundingThis research was supported by the National Natural Science Foundation of China (No. 82274052), CACMS Innovation Fund (No.CI2023E002, CI2024E003), Special Project on Survey of Scientific and Technological Basic Resources (No. 2022FY101000), National Key R&D Program: Intergovernmental Cooperation in International Science and Technology Innovation (No. 2022YFE0119300).Author informationAuthors and AffiliationsState Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing, 100700, ChinaMingxu Zhang, Cong Zhou, Suhua Huang, Hui Wang, Tingting Shi, Meng Li, Zhixian Jing & Xiaobo ZhangAuthorsMingxu ZhangView author publicationsSearch author on:PubMed Google ScholarCong ZhouView author publicationsSearch author on:PubMed Google ScholarSuhua HuangView author publicationsSearch author on:PubMed Google ScholarHui WangView author publicationsSearch author on:PubMed Google ScholarTingting ShiView author publicationsSearch author on:PubMed Google ScholarMeng LiView author publicationsSearch author on:PubMed Google ScholarZhixian JingView author publicationsSearch author on:PubMed Google ScholarXiaobo ZhangView author publicationsSearch author on:PubMed Google ScholarContributionsM.Z.: Writing – original draft, Conceptualization, Methodology, Validation, Data curation, Writing – review & editing; C.Z.: Writing – original draft, Conceptualization, Methodology, Validation, Data curation, Writing – review & editing; S.H.: Conceptualization, Writing – review & editing; H.W.: Writing-review and editing, Methodology; T.S.: Writing – review & editing, Data curation; Z.J.: Writing – review & editing, Data curation; M.L.: Writing – review & editing; X.Z.: Writing – review & editing, Conceptualization, Methodology.Corresponding authorCorrespondence to
    Xiaobo Zhang.Ethics declarations

    Competing interests
    The authors declare no competing interests.

    Ethics
    Information on the voucher specimens, including the deposition location, deposition number, and specimen identifier, is provided in Additional Table 1. We confirm that all research involving field studies and the collection of Gardenia jasminoides was conducted in strict compliance with relevant institutional, national, and international guidelines and legislation. Given the Least Concern status of Gardenia jasminoides and that collection occurred outside of protected areas, specific collection licenses were not required; all collection adhered strictly to local regulations. Furthermore, we adhere to the principles outlined in the IUCN Policy Statement on Research Involving Species at Risk of Extinction and the Convention on the Trade in Endangered Species of Wild Fauna and Flora. We note that Gardenia jasminoides was most recently assessed for The IUCN Red List of Threatened Species in 2023 and is listed as Least Concern. This classification is consistent with its national assessment in China, where the species is also categorized as Least Concern.

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    Reprints and permissionsAbout this articleCite this articleZhang, M., Zhou, C., Huang, S. et al. Integrated geographical and ecological analysis reveals environmental drivers of Gardenia jasminoides distribution and chemical variation.
    Sci Rep (2025). https://doi.org/10.1038/s41598-025-32876-0Download citationReceived: 01 September 2025Accepted: 12 December 2025Published: 24 December 2025DOI: https://doi.org/10.1038/s41598-025-32876-0Share this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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    Keywords
    Gardenia jasminoides
    Suitable distributionSpatial differentiationQuality variationEnvironmental drivers More

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    Structure and community assembly of rare bacterial community in sediments of Sancha Lake

    AbstractTo explore the structure and assembly of the rare bacterial community within sediment samples, as well as their responses their responses to environmental influencing factors, we collected surface sediment and overlying water samples from Sancha Lake across four seasons. MiSeq high-throughput sequencing was applied to the V3-V4 hypervariable regions of the 16 S rRNA genes, and the β – Nearest Taxon index (βNTI) was utilized to analyze the bacterial community assembly in the sediment samples. Our findings uncovered abundant bacterial diversity within the sediment samples of Sancha Lake, with 9314 operational taxonomic units (OTUs) identified, encompassing 59 phyla, 198 classes, 279 orders, 447 families, and 758 genera of bacteria. Proteobacteria and Chloroflexi were the dominant rare bacteria at the phylum level, whereas Coxiella and hgcl_clade were the principal rare bacteria at the genus level. The variety index of rare communities across diverse seasons was notably higher than that of abundant ones (P < 0.01). Bacterial community structure differed between spring and other seasons, and the rare bacterial community exhibited substantial seasonal alterations during non-spring periods. pH, dissolved oxygen (DO), total phosphorus (TP), and soluble reactive phosphorus (SRP) were the predominant environmental factors, exerting an even greater influence on rare bacteria. Within the co-occurrence network, rare bacteria constituted the majority of nodes and connections and were the dominant key species throughout all seasons. The assembly of their community was chiefly deterministic in autumn and random in other seasons. This study indicated that rare bacteria in Sancha Lake were diverse. They were keystone taxa for maintaining community interactions and stable operation, and their assembly process was influenced by both stochastic and deterministic factors.

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    Data availability

    The datasets analysed during the current study are available in the NCBI repository (https://www.ncbi.nlm.nih.gov/). The BioProject accession number is PRJNA1336117.
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    Download referencesAcknowledgementsThis research was funded by Student Research Training Program(242005).FundingThis research was funded by Student Research Training Program(242005).Author informationAuthors and AffiliationsSchool of Environmental Science and Engineering, Southwest Jiaotong University, Chengdu, 610031, ChinaYong Li, Yajie Li, Yihan Wu, Zuguang Liu, Shiqi Luo & Sidan GongAuthorsYong LiView author publicationsSearch author on:PubMed Google ScholarYajie LiView author publicationsSearch author on:PubMed Google ScholarYihan WuView author publicationsSearch author on:PubMed Google ScholarZuguang LiuView author publicationsSearch author on:PubMed Google ScholarShiqi LuoView author publicationsSearch author on:PubMed Google ScholarSidan GongView author publicationsSearch author on:PubMed Google ScholarContributionsY.L. writing – review & editing, investigation, conceptualization. Y.L. and Y.W. writing – original draft, visualization, project administration. Z.L. and S.L. writing – original draft, visualization. S.G. and Y.L. methodology, investigation. All authors read and approved the final manuscript.Corresponding authorCorrespondence to
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    Range-wide assessment of habitat suitability for jaguars using multiscale species distribution modelling

    AbstractJaguars (Panthera onca) are highly sensitive to persecution, habitat loss, and fragmentation, making the identification of suitable habitat critical for conservation planning. Using GPS telemetry data from 172 individuals across seven countries – the largest jaguar dataset to date – we developed multiscale Resource Selection Functions (RSFs) incorporating 15 environmental covariates to model habitat suitability across the species’ historic range. Jaguars selected productive habitats near water and strongly avoided human-modified landscapes, including areas with high human population density and livestock presence. The resulting habitat suitability surface showed strong predictive performance (AUC = 0.88; Boyce Index = 0.91) and correlated with known density estimates and distribution models. Jaguar Conservation Units (JCUs) and Protected Areas (PAs) contained 68.7% and 53.9% of predicted suitable habitat, respectively, while occupying only a third of the range. Non-designated lands, though comprising just 4% of the range, held nearly 10% of total suitability. The Amazon and Mayan Forests were identified as core strongholds, while ecoregion-based modelling revealed additional areas of high suitability in the Pantanal, Gran Chaco, Cerrado, and coastal Mexico. While Brazil encompassed the largest extent of highly suitable habitat, countries such as Paraguay, Argentina, and the United States gained conservation relevance under the ecoregion-stratified scenario.

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    Data availability

    GPS telemetry data for 117 of the 172 jaguar individuals used in this study are publicly available via Morato et al. (2018)82. The remaining 55 individuals were provided by collaborators and remain under the stewardship of their respective research groups; these data are not publicly available due to ongoing research use and data-sharing agreements. However, access to these data may be granted upon reasonable request to the corresponding authors, pending approval from the original data providers. The resulting habitat suitability surface generated by this study will be made openly available on the Zenodo repository upon manuscript acceptance (currently accessible for peer review at: https://zenodo.org/records/15824344?token=eyJhbGciOiJIUzUxMiJ9.eyJpZCI6IjY3YTJjNzhjLTVmMzItNDFhZi04YmY1LTk0NTQzZmFkYjgyZSIsImRhdGEiOnt9LCJyYW5kb20iOiJiYTI3YTBjNDRhOGRkNjk3NWI1ZGI1OWEyMDRkYWU3NCJ9._5XjPvxWOw4azyxXv4Ww-eaoHm1FG54BexND5TEEsmBnBTahFRBpbdScnwD_8McXtH-eNHyVaqA6hoWbieufCQ).
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    Download referencesAcknowledgementsGCA doctorate is supported by a grant to David Macdonald at WildCRU from the Robertson Foundation. We are extremely grateful to the Alianza WWF-Fundación, Telmex/Telcel, the Universidad Nacional Autónoma de México (project DGAPA, PAPIT IN208017), and Amigos de Calakmul A.C. for their financial support. We extend special thanks to the ejidos Caobas and Laguna Om, as well as the Calakmul Biosphere Reserve, for granting permission to conduct our research on their lands. The Instituto Homem Pantaneiro is grateful to ICMBio/CENAP, Panthera Brasil, and Onçafari for their valuable partnerships. The Mamirauá Institute thanks CNPq for the scholarships, and the communities of Mamirauá Sustainable Development Reserve for their essential field support – especially Lázaro Pinto dos Santos (Lazinho) and Railgler Gomes dos Santos (Raí), in memoriam. ACSA thanks the Espírito Santo Research and Innovation Support Foundation (FAPES) for funding the project (FAPES 510/2016), as well as for the Capixaba Researcher Fellowship (FAPES 404/2022). SAC thanks NASA for the project grant 80NSSC25K7244. All figures were created by GCA using QGIS v3.36.0 (https://qgis.org). Editorial assistance was provided by ChatGPT (OpenAI) to improve clarity and language use.FundingGCA doctorate is supported by a grant to David Macdonald at WildCRU from the Robertson Foundation. ACSA was supported by the Espírito Santo Research and Innovation Support Foundation (FAPES) for funding the project (FAPES 510/2016), as well as for the Capixaba Researcher Fellowship (FAPES 404/2022). SAC was supported by a NASA project grant number 80NSSC25K7244.Author informationAuthors and AffiliationsWildlife Conservation Research Unit (WildCRU), Department of Biology, University of Oxford, Life and Mind Building, South Parks Road, Oxford, OX1 3EL, UKGuilherme Costa Alvarenga, Caroline C. Sartor, Samuel A. Cushman, Alexandra Zimmermann & David W. MacdonaldGrupo de Pesquisa em Ecologia e Conservação de Felinos na Amazônia, Mamirauá Institute for Sustainable Development (MISD), Estrada do Bexiga, nº 2584, Tefé, AM, BrazilGuilherme Costa Alvarenga, Diogo Maia Gräbin, Emiliano E. Ramalho & Marcos Roberto Monteiro de BritoDepartment of Biological Sciences, Northern Arizona University, Flagstaff, AZ, USAŻaneta KasztaPrograma de Pós-Graduação em Ciência Animal e Programa de Pós-Graduação em Ecologia de Ecossistemas, Universidade Vila Velha, Av. Comissário José Dantas de Melo, 21, Boa Vista, Vila Velha, Espírito Santo, 29102-920, BrazilAna Carolina Srbek-AraujoLaboratório de Ecologia e Zoologia de Vertebrados (LABEV/ICB), Universidade Federal do Pará, Belém, PA, BrazilAna Cristina Mendes-Oliveira & Leonardo SenaPanthera, 104 West 40th Street, 5th Floor, New York, NY, 10018, USABart Harmsen, Rebecca J. Foster & Ronaldo G. MoratoInstituto de Ciencias de la Tierra, Biodiversidad y Ambiente (IBCIA), Universidad Nacional de Río Cuarto and National Scientific and Technical Research Council (CONICET), Ruta Nacional 36 Km 601, Río Cuarto, ArgentinaCarlos De AngeloInstitute for the Conservation of Neotropical Carnivores, Avenida Horácio Neto, 1030, Parque Edmundo Zanoni, Atibaia, SP, BrazilCarolina Franco Esteves, Claudia B. de Campos, Daiana Jeronimo Polli, Emiliano E. Ramalho & Fernando C. C. AzevedoPrograma de Pós-Graduação em Ecologia e Conservação, Instituto de Biociências – Inbio, Universidade Federal de Mato Grosso do Sul, Campo Grande, MS, 79070-900, BrazilDiego F. Passos VianaFundación Rewilding Argentina, Scalabrini Ortiz 3355, 1425, Buenos Aires, ArgentinaEmiliano DonadioWCS Big Cat Program, New York, USAEsteban PayánDepartamento de Ciências Naturais, Universidade Federal de São João del Rei, São João del Rei, MG, 36301-160, BrazilFernando C. C. AzevedoDepartment of Conservation Biology, Doñana Biological Station, CSIC, Avda. Américo Vespucio 26, 41092, Isla de la Cartuja, Seville, SpainFrancisco PalomaresWildlife Protection Solutions, 2501 Welton Street, Denver, CO, 80205, USAGeorge V. N. PowellLaboratorio de Ecología y Conservación de Fauna Silvestre, Instituto de Ecología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Coyoacán, Ciudad de México, MexicoGerardo CeballosInstituto Homem Pantaneiro, Corumbá, Mato Grosso do Sul, BrazilGrasiela Porfirio & Wener Hugo Arruda MorenoDepartamento de Ciencias Ambientales, Universidad Autónoma Metropolitana, Unidad Lerma, CBS, Lerma de Villada, MexicoHeliot ZarzaPrimero Conservation, Box 158885935, Pinetop, AZ, USAIvonne CassaigneCentro de Pesquisa de Limnologia, Biodiversidade e Etnobiologia do Pantanal; Laboratório de Mastozoologia; Programa de Pós-graduação em Ciências Ambientais, Universidade do Estado de Mato Grosso-UNEMAT, Cáceres, MT, 78217-900, BrazilJuliano A. BogoniMamirauá Institute for Sustainable Development (MISD), Estrada do Bexiga, nº 2584, Tefé, AM, BrazilLouise MaranhãoAssociação Onçafari, São Paulo, SP, BrazilMarcos Roberto Monteiro de BritoSan Diego Zoo Wildlife Alliance, Conservation Science and Wildlife Health, 15600 San Pasqual Valley Road, Escondido, CA, 92027, USAMathias W. ToblerNatural History Museum, University of Oslo, POB 1172 Blindern, 0318, Oslo, NorwayØystein WiigCentro Nacional de Pesquisa e Conservação de Mamíferos Carnívoros (CENAP-ICMBio), Estrada Municipal Hisaichi Takebayashi, 8600, Atibaia, SP, 12952-011, BrazilRicardo SampaioAlianza Jaguar AC, Lab. Vida Silvestre, Fac. Biol. Universidad Michoacana, Morelia, Michoacán, MexicoRodrigo NuñezWorld Wide Fund for Nature (WWF) UK, The Living Planet Centre, Brewery Road, Woking, GU214LL, UKValeria BoronEnvironmental Change Institute, School of Geography and the Environment, University of Oxford, South Parks Road, Oxford, UKYadvinder MalhiLeverhulme Centre for Nature Recovery, University of Oxford, South Parks Road, Oxford, UKYadvinder MalhiAuthorsGuilherme Costa AlvarengaView author publicationsSearch author on:PubMed Google ScholarCaroline C. SartorView author publicationsSearch author on:PubMed Google ScholarSamuel A. CushmanView author publicationsSearch author on:PubMed Google ScholarAlexandra ZimmermannView author publicationsSearch author on:PubMed Google ScholarAna Carolina Srbek-AraujoView author publicationsSearch author on:PubMed Google ScholarAna Cristina Mendes-OliveiraView author publicationsSearch author on:PubMed Google ScholarBart HarmsenView author publicationsSearch author on:PubMed Google ScholarCarlos De AngeloView author publicationsSearch author on:PubMed Google ScholarCarolina Franco EstevesView author publicationsSearch author on:PubMed Google ScholarClaudia B. de CamposView author publicationsSearch author on:PubMed Google ScholarDaiana Jeronimo PolliView author publicationsSearch author on:PubMed Google ScholarDiego F. Passos VianaView author publicationsSearch author on:PubMed Google ScholarDiogo Maia GräbinView author publicationsSearch author on:PubMed Google ScholarEmiliano DonadioView author publicationsSearch author on:PubMed Google ScholarEmiliano E. RamalhoView author publicationsSearch author on:PubMed Google ScholarEsteban PayánView author publicationsSearch author on:PubMed Google ScholarFernando C. C. AzevedoView author publicationsSearch author on:PubMed Google ScholarFrancisco PalomaresView author publicationsSearch author on:PubMed Google ScholarGeorge V. N. PowellView author publicationsSearch author on:PubMed Google ScholarGerardo CeballosView author publicationsSearch author on:PubMed Google ScholarGrasiela PorfirioView author publicationsSearch author on:PubMed Google ScholarHeliot ZarzaView author publicationsSearch author on:PubMed Google ScholarIvonne CassaigneView author publicationsSearch author on:PubMed Google ScholarJuliano A. BogoniView author publicationsSearch author on:PubMed Google ScholarLeonardo SenaView author publicationsSearch author on:PubMed Google ScholarLouise MaranhãoView author publicationsSearch author on:PubMed Google ScholarMarcos Roberto Monteiro de BritoView author publicationsSearch author on:PubMed Google ScholarMathias W. ToblerView author publicationsSearch author on:PubMed Google ScholarØystein WiigView author publicationsSearch author on:PubMed Google ScholarRebecca J. FosterView author publicationsSearch author on:PubMed Google ScholarRicardo SampaioView author publicationsSearch author on:PubMed Google ScholarRodrigo NuñezView author publicationsSearch author on:PubMed Google ScholarRonaldo G. MoratoView author publicationsSearch author on:PubMed Google ScholarValeria BoronView author publicationsSearch author on:PubMed Google ScholarWener Hugo Arruda MorenoView author publicationsSearch author on:PubMed Google ScholarYadvinder MalhiView author publicationsSearch author on:PubMed Google ScholarDavid W. MacdonaldView author publicationsSearch author on:PubMed Google ScholarŻaneta KasztaView author publicationsSearch author on:PubMed Google ScholarContributionsGuilherme Costa Alvarenga: Conceptualization, Investigation, Data curation, Methodology, Formal analysis, Validation, Visualization, Writing – original draft, Writing – review & editing. Caroline C. Sartor: Methodology, Formal analysis, Writing – review & editing. Samuel (A) Cushman: Conceptualization, Methodology, Formal analysis, Writing – review & editing, Supervision, Project administration. Alexandra Zimmermann: Writing – review & editing. Ana Carolina Srbek-Araujo: Investigation, Resources, Writing – review & editing. Ana Cristina Mendes-Oliveira: Investigation, Resources, Writing – review & editing. Bart Harmsen: Investigation, Resources, Writing – review & editing. Carlos De Angelo: Investigation, Resources, Writing – review & editing. Carolina Franco Esteves: Investigation, Resources, Writing – review & editing. Claudia (B) de Campos: Investigation, Resources, Writing – review & editing. Daiana Jeronimo Polli: Investigation, Resources, Writing – review & editing. Diego F. Passos Viana: Investigation, Resources, Writing – review & editing. Diogo Maia Gräbin: Investigation, Resources, Writing – review & editing. Emiliano Donadio: Investigation, Resources, Writing – review & editing. Emiliano E. Ramalho: Investigation, Resources, Writing – review & editing. Esteban Payán: Investigation, Resources, Writing – review & editing. Fernando (C) C. Azevedo: Investigation, Resources, Writing – review & editing. Francisco Palomares: Investigation, Resources, Writing – review & editing. George V. N. Powell: Investigation, Resources, Writing – review & editing. Gerardo Ceballos: Investigation, Resources, Writing – review & editing. Grasiela Porfirio: Investigation, Resources, Writing – review & editing. Heliot Zarza: Investigation, Resources, Writing – review & editing. Ivonne Cassaigne: Investigation, Resources, Writing – review & editing. Juliano A. Bogoni: Investigation, Resources, Writing – review & editing. Leonardo Sena: Investigation, Resources, Writing – review & editing. Louise Maranhão: Investigation, Resources, Writing – review & editing. Marcos Roberto Monteiro de Brito: Investigation, Resources, Writing – review & editing. Mathias W. Tobler: Investigation, Resources, Writing – review & editing. Øystein Wiig: Investigation, Resources, Writing – review & editing. Rebecca J. Foster: Investigation, Resources, Writing – review & editing. Ricardo Sampaio: Investigation, Resources, Writing – review & editing. Rodrigo Nuñez: Investigation, Resources, Writing – review & editing. Ronaldo G. Morato: Investigation, Resources, Writing – review & editing. Valeria Boron: Investigation, Resources, Writing – review & editing. Wener Hugo Arruda Moreno: Investigation, Resources, Writing – review & editing. Yadvinder Malhi: Writing – review & editing. David W. Macdonald: Resources, Writing – review & editing, Funding acquisition. Zaneta Kaszta: Conceptualization, Methodology, Formal analysis, Writing – review & editing, Supervision, Project administration.Corresponding authorCorrespondence to
    Guilherme Costa Alvarenga.Ethics declarations

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    The authors declare no competing interests.

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    Reprints and permissionsAbout this articleCite this articleAlvarenga, G.C., Sartor, C.C., Cushman, S.A. et al. Range-wide assessment of habitat suitability for jaguars using multiscale species distribution modelling.
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    National human footprint maps for Peru and Ecuador

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    Data availability

    The full dataset is available at https://doi.org/10.6084/m9.figshare.30226402.
    Code availability

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    Download referencesAcknowledgementsThe NASA Biodiversity and Ecological Forecasting Program funded the work under the 2016 A.8 Sustaining Living Systems in a Time of Climate Variability and Change solicitation under Grant number 80NSSC19K0186. We want to thank everyone at the Life on Land project for all their work. We thank the following individuals for their valuable support in this study: Mauricio Trujillo, Maria Olga Borja, Rodrigo Torres, Cícero Augusto, Gonzalo Morales, and everyone at the Conservation Solutions Lab and the GIS Lab at UNBC. ChatGPT was used for language suggestions.Author informationAuthors and AffiliationsNatural Resources and Environmental Studies Institute, University of Northern British Columbia, Prince George, BC, CanadaJose Aragon-Osejo & Oscar VenterMinisterio del Ambiente, Agua y Transición Ecológica de Ecuador (MAATE), Quito, EcuadorLenin Beltrán, Juan Iglesias, Hólger Zambrano, Daniel Borja, Carlos Oñate, Francisco Simbaña, Freddy Valencia, Karen Rodríguez De la Vera, Luis Poveda, Fernando Proaño, Lorena Parra & María ChacónMinisterio del Ambiente del Perú (MINAM), Lima, PerúWilliam Llactayo, Tatiana Pequeño Saco, Walter Huamani, German Marchand, Raúl Tinoco & Luis QuispeUniversidad Nacional Federico Villarreal, Lima, PerúWilliam LlactayoRainforest Alliance (RA), Lima, PerúTatiana Pequeño SacoServicio Nacional Forestal y de Fauna Silvestre (SERFOR), Lima, PerúAlexs AranaUnited Nations Development Programme, New York, NY, USAAnne Lucy Stilger VirnigPrograma de las Naciones Unidas del Desarrollo – Perú (PNUD), Lima, PerúPatricia HuertaIndependent Contributor, Quito, EcuadorKarla Jiménez & Pedro Tipula TipulaIndependent Contributor, Lima, PerúPedro Tipula TipulaInstituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, ColombiaSusana RodríguezGeoIS, Austin, TX, USARodrigo SierraProfessor Emeritus of Ecology, Montana State University, Bozeman, MT, USAAndrew J. HansenAuthorsJose Aragon-OsejoView author publicationsSearch author on:PubMed Google ScholarLenin BeltránView author publicationsSearch author on:PubMed Google ScholarJuan IglesiasView author publicationsSearch author on:PubMed Google ScholarHólger ZambranoView author publicationsSearch author on:PubMed Google ScholarDaniel BorjaView author publicationsSearch author on:PubMed Google ScholarCarlos OñateView author publicationsSearch author on:PubMed Google ScholarFrancisco SimbañaView author publicationsSearch author on:PubMed Google ScholarFreddy ValenciaView author publicationsSearch author on:PubMed Google ScholarKaren Rodríguez De la VeraView author publicationsSearch author on:PubMed Google ScholarLuis PovedaView author publicationsSearch author on:PubMed Google ScholarFernando ProañoView author publicationsSearch author on:PubMed Google ScholarLorena ParraView author publicationsSearch author on:PubMed Google ScholarMaría ChacónView author publicationsSearch author on:PubMed Google ScholarWilliam LlactayoView author publicationsSearch author on:PubMed Google ScholarTatiana Pequeño SacoView author publicationsSearch author on:PubMed Google ScholarWalter HuamaniView author publicationsSearch author on:PubMed Google ScholarGerman MarchandView author publicationsSearch author on:PubMed Google ScholarRaúl TinocoView author publicationsSearch author on:PubMed Google ScholarLuis QuispeView author publicationsSearch author on:PubMed Google ScholarAlexs AranaView author publicationsSearch author on:PubMed Google ScholarAnne Lucy Stilger VirnigView author publicationsSearch author on:PubMed Google ScholarPatricia HuertaView author publicationsSearch author on:PubMed Google ScholarKarla JiménezView author publicationsSearch author on:PubMed Google ScholarPedro Tipula TipulaView author publicationsSearch author on:PubMed Google ScholarSusana RodríguezView author publicationsSearch author on:PubMed Google ScholarRodrigo SierraView author publicationsSearch author on:PubMed Google ScholarAndrew J. HansenView author publicationsSearch author on:PubMed Google ScholarOscar VenterView author publicationsSearch author on:PubMed Google ScholarContributionsJ.A.-O. conducted the analysis and wrote the manuscript. R.S. provided inputs. All co-authors conceived the study and reviewed and edited the manuscript.Corresponding authorCorrespondence to
    Jose Aragon-Osejo.Ethics declarations

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    Reprints and permissionsAbout this articleCite this articleAragon-Osejo, J., Beltrán, L., Iglesias, J. et al. National human footprint maps for Peru and Ecuador.
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    Respiratory bacteriome and its predicted functional profiles in blue whales (Balaenoptera musculus)

    AbstractThe respiratory microbiome plays a critical role in the health of organisms and studying it in natural populations can reveal interactions between hosts and their environment, as well as help predict responses to environmental stressors. We characterized the core respiratory bacteriome and functional profiles of Eastern North Pacific blue whales (Balaenoptera musculus) sampled in the Gulf of California using next-generation sequencing. Our compositional analysis identified 15 dominant bacterial phyla in the respiratory tract, with Proteobacteria (34.44%), Firmicutes (26.98%), Bacteroidota (20.26%), Fusobacteriota (7.61%), and Actinobacteria (5.55%) as the most abundant. Nineteen ASVs, representing 12 bacterial genera (primarily Corynebacterium, Oceanivirga, Tenacibaculum, and Psychrobacter), were shared by over 60% of whales, with a relative abundance greater than 0.02%. These bacteria, proposed to be the core respiratory bacteriome of blue whales, contributed to functional pathways associated with metabolism, environmental information processing, and cellular processes. Notably, two whales with high relative abundance of Mycoplasma spp. and of Streptococcus spp., exhibited overrepresented pathways related to nucleotide metabolism and translation, suggesting a suboptimal immune status or dysbiosis. To our knowledge, this is the first functional profiling of the bacteriome in any cetacean. Future studies are needed to explore how the blue whale respiratory bacteriome may vary over time, seasonally or across geographical locations. This study establishes a baseline for future research on the plasticity of the bacteriome, its associations with other microbiome components, the impact of environmental changes on its diversity, and its relevance for health. Our novel approach underscores the ecological and physiological importance of the bacteriome and its potential for long-term monitoring of a sentinel marine species in a rapidly changing ocean.

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    IntroductionThe advent of modern technologies that allow for the identification of bacteria in environmental or clinical samples1 has led to a surge in studies examining the abundance, diversity, and structure of microbiomes across species2. Increasing our understanding of the microbiome is crucial because microbial communities associated with specific organs or tissues can significantly impact host physiology3 and health4. For instance, respiratory infections may arise when opportunistic microorganisms—normally part of a healthy respiratory tract—proliferate under certain conditions1, disrupting the diversity and composition of the microbial community in a phenomenon known as dysbiosis5, which can contribute to disease. Additionally, respiratory disease can result from exposure to non-commensal microorganisms with pathogenic potential. This underscores the importance of microbiome composition as a potential predictor of health and disease progression, often more so than the mere presence of specific microorganisms commonly associated with disease. Understanding how microbiomes differ between individuals could, therefore, become a valuable tool for assessing health6.When using the microbiome to assess health status, it is important to distinguish between commensal, opportunistic, and transient bacteria7. This distinction is complex, as the symbiotic relationships of bacteria can vary both across species and among individuals8 To help differentiate potential commensal and mutualistic bacteria, it is necessary to identify the core microbiome—the microbial taxa that predominate within a community and are common in apparently healthy individuals9. Defining the core bacteriome (the bacterial community of the microbiome) involves setting the detection threshold (relative abundance) and determining the minimum occurrence percentage (prevalence) of bacterial taxa to include10. However, because biological justifications for these prevalence and threshold values are often lacking11, it is important to exercise caution when interpreting results10. Despite varying definitions, the core bacteriome tends to be relatively stable, particularly when samples from closely related individuals are analyzed11,12.Microbial taxonomic composition provides a basic understanding of the microbiome, but it does not fully capture the intricate microbial contributions to host health13. Bacteria within the mammalian microbiome exist in composite communities1, whose diversity and abundance result from complex interactions between species14. The metabolic contributions of these communities are important to the host and depend on their composition15. This is where functional profiling becomes important, as it reveals the metabolic and ecological roles of microbial communities16,17. Combined taxonomic and functional analyses offer a deeper understanding of the microbiome’s dual nature, as both a diverse community and a functional unit that essential for the holobiont’s processes18. Functional predictions are based on genetic data derived from sequencing the 16 S rRNA gene18,19, which is mapped to reference databases, correlating specific taxa with known functional abilities20. These functions are organized hierarchically, from broad functional categories to specific metabolic pathways21. This approach allows researchers to infer the ecological and metabolic roles of the microbiome without the need for whole-genome sequencing, making it a powerful and accessible tool for microbiome research19. By integrating taxonomic and functional analyses, we can gain deeper insights into a host’s microbiome and its role in holobiont resilience, particularly in the context of health22.In cetaceans, characterizing the microbiome offers a unique opportunity to link microbial community structure and function with host ecology, physiology, and responses to environmental change, providing valuable insights for conservation and health monitoring23,24. Whales, as long-lived animals that play a critical role in the ocean’s carbon movement and storage, are vital to marine ecosystems25, and are often considered sentinels of ocean health26. The study of the cetacean microbiome is still in its early stages. Microbial diversity has been assessed for a few species1,27,28,29,30,31, and some opportunistic pathogens in the respiratory tracts of free-ranging cetaceans have also been described32,33. However, to our knowledge, no study has yet combined taxonomic and functional profiling of the microbiome in any cetacean species. Blue whales, among the world’s largest and most iconic animals, play an essential role in marine ecosystems34. Their long migrations and diverse habitats make them valuable indicators of ocean health35. Despite this, only one published study has examined the respiratory microbiome of blue whales in the wild36. Given the growing importance of understanding blue whale health in their natural environment, studying their respiratory microbiome is both timely and relevant. Not only would it provide insights into their exposure with potential pathogens32, but it would also establish a baseline of core bacteria and functional profiles in apparently healthy individuals. This baseline could facilitate the identification of dysbiosis, help predict potential diseases and ultimately inform conservation strategies and management plans for the species37. This is a pressing need, especially in light of the global and local environmental changes currently affecting oceans38. Here, we characterized the common core and functional profiles of the respiratory bacteriome in Eastern North Pacific blue whales from the Gulf of California using next-generation sequencing on blow samples collected from 17 adult blue whales via a non-invasive drone-based technique39.ResultsA total of 19 samples were analysed, including 17 photo-identified blue whales, one technical control, and one seawater sample. Exhaled breath was collected from the whales using a drone-based method previously described39, with no adverse behavior observed before, during, or after sampling. After filtering, denoising, merging, and chimera elimination (2.38% of reads), we obtained 68,922 sequences (mean per sample: 3514.8 [SD = 1998.3]), which corresponded to 1304 amplicon sequence variants (ASVs). We removed 51 ASVs classified as Archaea (n = 2), chloroplasts (n = 27), or mitochondria (n = 7), as well as those not classified at the phylum level (n = 15), and 22 ASVs identified as contaminants using the Decontam algorithm based on the LabControl sample reads. This left 1231 ASVs remained, with 500 ASVs classified as “Others” (representing less than 0.02% relative abundance).Species richness (S) in the blow samples ranged from 62 to 404 (mean = 189.63.06 [SD = 113.71]), and Simpson’s diversity index (D) ranged from 0.49 to 0.98 (mean = 0.94 [SD = 0.11]). The compositional analysis identified 15 bacterial genera (Fig. 1, Table S1) with Psychrobacter spp. (mean = 12.07% [SD = 6.09%]), Oceanovirga spp. (mean = 10.93% [SD = 4.42%]), Tenacibaculum spp. (8.87% [SD = 6.39%]), and Streptococcus spp. (6.79% [SD = 20.13%]) being the most abundant. Notably, two blow samples Bm057 and Bm044) exhibited a high relative abundance of the opportunistic pathogens Mycoplasma spp. (27.22%), and Streptococcus spp. (74.37%). In addition, we identified Bacteroides sp. in sample Bm042 at a high relative abundance of 13.96% compared with the other samples (mean = 0.02% [SD = 0.04%]). For this whale, mucus was also retrieved during blow sampling, which had a noticeable bad smell and a yellowish coloration; features that were not observed in any other samples.Fig. 1Stacked bar plot depicting relative abundance of the top 15 bacterial genera. Each vertical bar depicts the relative abundance of adjusted sequence variants (ASVs) and associated taxa that were recovered per sample. Plot shows the top fifteen identified bacterial genera, unclassified, and “others” (sum of bacteria that did not reach the detection threshold of 0.02%).Full size imageFunctional profiling, at 97% similarity, was possible for 32.53% of the ASVs. At KEGG Level 1, the most predominant pathways were associated with metabolism (mean = 74.88% [SD = 4.36%]), followed by environmental information processing (mean = 9.98% [SD = 1.82%]), cellular processes (mean = 5.54% [SD = 1.67%]), and genetic information processing (mean = 5.40% [SD = 1.57%]). At KEGG Level 2, the top subcategories included global and overview maps (mean = 38.37% [SD = 2.96%]), carbohydrate metabolism (mean = 9.65% [SD = 1.14%]), amino acid metabolism (mean = 7.23% [SD = 1.45%]), and membrane transport (mean = 6.55% [SD = 1.32%]). At KEGG Level 3, the most abundant pathways were metabolic pathways, biosynthesis of secondary metabolites, ABC transporters, and microbial metabolism in diverse environments (Fig. 2).Fig. 2Alluvial diagram of the top 20 predicted functional pathways (at different KEEG levels) associated with the bacteriome in the respiratory tract of blue whales.Full size imageIn two blow samples (Bm057 and Bm044) with the highest relative abundance of opportunistic pathogens, functional profiling revealed overrepresentation of pathways such as nucleotide metabolism, membrane transport, translation, folding, sorting and degradation, and carbohydrate metabolism; while pathways related to amino acid metabolism, cofactor and vitamin metabolism, lipid metabolism, and biosynthesis of secondary metabolites were underrepresented (Fig. 3). Among the bacterial genera identified, Psychrobacter (26.83%) contributed most to the functional pathways predicted in the blue whale respiratory tract, followed by Tenacibaculum (17.48%) and Porphyromonas (13.01%). Genera such as Suttonella and Streptococcus contributed less (4.07% and 3.25%, respectively; Fig. S1). Despite variation in taxonomic composition, functional profiles across individuals were consistent (Fig. S2).Fig. 3Boxplot of the relative abundance of functional pathways (at KEGG Level 2) across all blow samples. Red dots represent blow sample Bm057 (the whale that had a high relative abundance of Mycoplasma sp.), while yellow dots correspond to blow sample Bm042 (the whale that had a high relative abundance of Streptococcus sp.). Functional pathways that were overrepresented in both Bm057 and Bm044 compared to all other samples (grey) are highlighted as light green columns, while underrepresented pathways are shown as light blue columns. The functional pathway that was underrepresented in the bacteriome of Bm057 but over represented in Bm042 is highlighted as light purple columns, and the functional pathways were over represented only in Bm057 are shown in as light orange columns.Full size imageThe core bacteriome analysis identified 19 ASVs from 12 bacterial families (Fig. 4, Table S2), with Tenacibaculum (ASV3) and Oceavivirga (ASV7) the being the most abundant genera (30.01% [SD = 15.94] and 28.78% [SD = 9.76], respectively]). The core functional profile derived from these core ASVs was composed mainly by metabolic pathways (24.99%), biosynthesis of secondary metabolites (11.09%), biosynthesis of antibiotics (8.73%), and microbial metabolism in diverse environments (8.34%) (KEEG level 1; Fig. S3).Fig. 4Relative abundances of bacterial genera that constitute the core respiratory bacteriome of the blue whale. The figure includes the seven ASVs that were present in more than 60% of the samples and that had a relative abundance of over 0.02%. The relative abundance of each ASV shown in this plot is confined to the core microbiome members and not the entire microbiome of each sample.Full size imageThe Bayesian approach used to estimate the contribution of seawater diversity to blow samples indicated that seawater contributed on average of 1.68% (SD = 0.81). Herbaspirillum sp., the most common genus in seawater (20.23% relative abundance; Fig. 1), was also detected in blow samples, albeit at a lower average abundance (3.39%; SD = 5.97). Interestingly, three whale blows (from individuals Bm023, Bm043, and Bm059) exhibited notably higher levels of Herbaspirillum sp. (9.44%, 15.65%, and 16.82%, respectively).DiscussionA healthy microbiome is generally characterized by high diversity, which helps both the microbiome and the host cope with external challenges30. In our study, the respiratory bacteriome of the blue whale exhibited considerable diversity, with significant variation in bacterial richness and abundance across samples. These fluctuations may arise from several factors, including bacterial immigration from the environment during inhalation, mucociliary clearance, and community growth rates40, all of which can vary among healthy individuals27. However, variations could also stem from sampling techniques, such as differences in the number of blows, volume of sample collected, whale size and behavior (e.g. dive depth and duration) 39,41. Notably, the bacterial diversity observed in blue whale blow samples was similar to that reported for humpback whales and bottlenose dolphins41,42, although the blue whale blow showed greater taxonomic richness. This may be attributed to differences in methods used to resolve taxonomy43,44,45 or the identification of rare bacterial species44,45, which play an important role in microbiome resilience, given their contribution as a seed bank of genetic resources that can lead to the restoration of the core microbiome46.The presence of a complex respiratory bacteriome is beneficial for a host, as higher microbial diversity supports vital ecosystem functions47. Functional analysis of the blue whale bacteriome revealed overrepresentation of pathways related to macromolecular metabolism and environmental information processing and signal transduction, indicating a potential role in adapting to environmental changes48,49. This result reinforces the idea that bacteriome diversity serves a protective role for the host18, as these pathways are critical for maintaining host health and epithelial immune function50,51 by enabling microbial communication with host immune cells via molecular signals that activate pattern recognition receptors, triggering cytokine production and immune cell recruitment52, including dendritic cells53.Our findings indicate that the respiratory bacteriome of blue whales is dominated by members of Proteobacteria, Firmicutes, Bacteroidota, Actinobacteria, and Fusobacteriota, which are common bacterial phyla in the respiratory microbiome of other mammals1. Particularly noteworthy is the consistent presence of Psychrobacter sp. and Tenacibaculum sp., which are known commensal bacteria54,55 that contribute to respiratory and skin health4,27,29,30,56, although they can also be implicated in pathological conditions in other organs57,58. Additionally, the respiratory core bacteriome included Oceanivirga sp., a bacterium common to the respiratory tract of various marine mammals from different geographical locations59, and identified as part of the core respiratory bacteriome of humpback whales41. Given that Oceanivirga sp., was present in most of the blue whales sampled, it is reasonable to consider it a key member of their respiratory bacteriome, reflecting a healthy respiratory epithelium.It is important to recognize that while the bacterial taxa in the blue whale’s respiratory bacteriome share similarities with those found in the oropharynx and nasopharynx of terrestrial mammals60, cetaceans lack anatomical connections between the mouth and nasopharynx41. Thus, the bacteria identified in this study are more likely associated with the respiratory tract rather than the oral cavity of the blue whales. In addition, it is important to note that this composition may vary over time and space, and could be influenced by factors such as fasting, reproductive stage6,61, or other physiological variables1,62,63.Interestingly, four bacterial genera (Psychrobacter, Tenacibaculum, Staphylococcus, and Corynebacterium) identified in the blow samples are typically found in the skin of humans and other terrestrial mammals64,65. These genera were also identified in the skin microbiota of both captive and free-ranging cetaceans1,6,28,55,56. Given that strict protocols were followed to minimize contamination during sampling, processing or sequencing, their presence in whale blow suggests that they colonize the epithelial lining of the blowhole and are forcefully expelled during exhalation41. Moreover, Psychrobacter and Tenacibaculum, contributed significantly to metabolic and environmental processing pathways, suggesting their role in maintaining microbial and host homeostasis. We hypothesize that these bacteria establish a commensal or mutualistic associations with the blue whale, potentially offering a protective role against dysbiosis and environmental stressors. Furthermore, it is possible that these taxa play a crucial role in maintaining respiratory health in this species, and more detailed functional analyses will be necessary in the future to clarify their ecological and physiological roles.Our study also found that approximately 2% of the microbial diversity in blow samples overlapped that of seawater, indicating some influence of the marine environment on the respiratory bacteriome, possibly as carryover during diving immersions. However, this overlap should be interpreted with caution, as seawater sampling was limited in number and not conducted for every breath sample. The absence of more water samples restricts our ability to fully assess the extent to which environmental microorganisms contribute to the respiratory bacteriome composition. Regardless, the detection of Psychrobacter, Oceanivirga, Tenacibaculum, Helcococcus, Porphyromonas, Mycoplasma, Dielma, Synechococcus, and Suttonella, in blue whale blow, but not in seawater, adds support to the notion that these taxa are intrinsic to the blue whale’s respiratory microbiome. Variations in the relative abundance of Herbaspirillum sp. in certain samples suggest that whale diving behavior, environmental factors and technical sampling conditions may also influence bacterial detection.We identified Bacteroides spp. in blow Bm042 at a relative abundance of 13.96%. Bacteroides spp. can influence airway immune responses by inducing regulatory T cells and associated cytokines and has been shown to promote transient PD-L1 expression and modulate general aeroallergen responses66. This genus has also been reported in increased abundance during tracheobronchitis, suggesting potential roles in modulating respiratory immune function66,67. Interestingly, whale Bm042 also presented mucus with a yellowish coloration, which may indicate a high concentration of airway mucin, which is associated with various pulmonary diseases68. The excessive synthesis of mucin can result from increased neutrophil recruitment, reflecting an acute inflammatory response to bacterial infection in the airways69,70. Given its immunomodulatory capacity, the elevated abundance of Bacteroides spp. in the blow of whale Bm042 may reflect a role in host immune regulation during localized airway infection or inflammation. Its presence alongside signs of mucus suggests a potential microbial shift and underscores the need to consider both protective and pathogenic roles of Bacteroides spp. in the respiratory tract.Two unidentified species from Mycoplasma and Streptococcus were found in the blow of two whales. As 16 S rRNA gene sequencing does not allow reliable species-level resolution, our assignments were limited to the genus level, and we acknowledge that the detected Streptococcus and Mycoplasma taxa may include both commensal and opportunistic members. This taxonomic uncertainty underscores the importance of continued monitoring, since shifts at the genus level can still provide meaningful indicators of host health. Various species within these bacterial genera are known respiratory tract opportunists in mammals71,72 and have been detected in the lungs of stranded marine mammals73,74, although their presence does not necessarily indicate disease since they can also occur in healthy hosts72. This is essential to consider when studying the bacteriome of individuals, as the type of relationship between host and bacteria can depend on different factors, including the status of the immune system8,71. The low prevalence of these pathogens in our study likely suggests that they are not common members of the respiratory bacterial community and highlights the natural diversity of the blue whale respiratory microbiome. As the blue whales migrate through coastal areas, they could become exposed to transient bacteria which do not normally manage to colonize the respiratory epithelium. However, the intense maritime traffic and potential human interactions75 could act as stressors that affect immune regulation of bacterial communities in susceptible hosts and favor the growth of transient or opportunistic bacteria68,69,70,76,77. Therefore, it is plausible that the detection of these bacteria could indicate underlying health conditions, a suboptimal immune status, or chronic stress in these individuals78. We have some support for this argument as the respiratory bacteriome of the two whales that harbored Mycoplasma spp. and Streptococcus spp. exhibited distinct functional pathway patterns than the other whales, whose bacteriome functional profiles remained largely stable across individuals. Namely the bacteriome of these two whales showed overexpression of nucleotide metabolism, translation, and replication and repair pathways, which have been associated with various diseases in humans79. In contrast, pathways involved in lipid metabolism and biosynthesis of other secondary metabolites were underrepresented in these whales, suggesting possible vulnerabilities in their immune responses, as has been shown for humans50,80. As these functional profiles were inferred from 16 S rRNA gene data, incorporating functional analyses based on transcriptomics or other omics approaches in future studies would provide a more comprehensive understanding of the microbiome’s functional potential. The identification of these bacterial genera and the distinct functional profile of the bacteriome of the whales that harbored them, highlights the need for ongoing monitoring specific microbial taxa, regardless of their perceived roles as commensal, mutualistic or opportunistic in other mammals, and underscores the importance of considering natural fluctuations in the respiratory bacteriome when assessing the health of blue whales.Given the current threats facing marine ecosystems, that include habitat degradation, pollution, and other anthropogenic stressors26, the taxonomic and functional study of the blue whale respiratory bacteriome offers valuable insights into their health and resilience. Respiratory microbiome data can serve as an early warning system by detecting shifts associated with environmental change, disease, or human activities41. Monitoring such changes in bacterial composition and functionality over time can help inform conservation efforts and management strategies23 to protect these iconic species and the ecosystems they inhabit. While our study is based on a modest number of individuals, it represents a meaningful fraction of the population migrating through the Loreto area. Future studies incorporating multiple blow samples per individual could capture temporal variability more effectively, reduce potential sampling bias, and further strengthen the value of microbiome monitoring for conservation and health assessments.Methods Sample collection Using a small Phantom 3® quadrocopter drone (DJI Innovations, China) with floaters and sterile Petri dishes, we collected 17 blows samples from 17 different individual blue whales sampled between February and March 2016 and 2017 in Loreto Bay National Park (25° 51′ 51″ N, 111° 07′ 18″ O) within the Gulf of California, Mexico. The number of sampled whales represents 17% of the estimated 100 blue whales that reside during winter/spring in the southwestern Gulf of California (mark-recapture data from 1994 to 200681. Each whale was photo-identified prior to sample collection81. The approach of the drone to the whale was done from the caudal fin towards the head to minimize disturbance, and sampling was conducted at a height between 3 and 4 m above the blowhole39. We observed the whale body condition (see Supplementary Material) for each individual and recorded characteristics of their blow, such as color and odor when we were sufficiently close to the whale during sampling.For each sample, blow droplets were swabbed directly from the Petri dish using one sterile cotton-tipped swab per individual. These were then transferred to a sterile 1.5 mL cryogenic microtube containing 500 µL of 96% molecular grade ethanol and kept frozen in a liquid nitrogen container until processing. To address potential contamination, all necessary precautions were taken, always including the use of sterile gloves and face masks during sample processing. In addition, we included a technical control, termed “LabControl” (a template-free DNA negative extraction control), to identify any contaminants during sample processing. Furthermore, we included a seawater sample, termed “seawater” (a DNA sample extracted from 1mL water collected at a depth of 0.10 m in the same area where we sampled the whale blows), to consider potential sources of bacterial diversity for the blow samples. DNA extraction, PCR amplification and sequencing Total DNA was isolated from the whale blow, seawater, and LabControl samples in one batch using a QIAamp ® DNA Mini Kit (QIAGEN, Germany). The primers used for sequencing the 16S rRNA V3 and V4 regions were 341F (5′-CCTACGGGNGGCWGCAG) and 785R (5′-GACTACHVGGGTATCTAATCC), which amplified a single product of 444 bp82. The PCR program used an initial denaturation step at 95 °C for 3 min; 25 cycles of 95 °C for 30 s, 55 °C for 30 s, and 72 °C for 30 s; and a final extension step at 72 °C for 5 min. Each 25 µL-reaction contained 12.5 ng of extracted DNA, 5 µM of barcoded primers and 2x KAPA HiFi HotStart Ready Mix (KAPABIOSYSTEM, Cape Town, South Africa). 1 µl of each sample was run on a 2100 Bioanalyzer (Agilent Technologies, CA, USA) with an Agilent DNA 1000 chip (Agilent Technologies, CA, USA) to verify amplicon size. AMPure XP beads (New England BioLabs, USA) were used to remove unused primers and primer dimers. Amplicons were sequenced over 2- by 250-bp MiSeq at the Unit of Sequencing and Identification of Polymorphisms of the National Institute of Genomic Medicine (Instituto Nacional de Medicina Genómica, Unidad de Secuenciación e Identificación de Polimorfismos, INMEGEN) in Mexico. Dual index barcodes were used to avoid index hopping83. The protocol used by INMEGEN can be seen in: https://support.illumina.com/documents/documentation/chemistry_documentation/16s/16s-metagenomic-library-prep-guide-15044223-b.pdf. 16 S rRNA sequence data processing A quality control overview was performed using FASTQC84. This allowed us to obtain a quick impression of the data and avoid downstream problems. The raw sequences were then imported into R v.4.2.185, where all subsequent analyses were carried out. We used the Divisive Amplicon Denoising Algorithm 2 (dada2) v.1.26.044 to infer exact ASVs. This approach is preferable over the rough and less precise 16 S rRNA OTU clustering approach86 that groups the sequences with a 97% identity87. First, we filtered by quality (trunQ = 25) and discarded the sequences that presented more than two Ns (maxN = 0) or more than two expected errors (maxEE = 2). Next, the forward and reverse reads for each sample were combined into a single merged contig sequence, and we grouped all identical reads into unique sequences to determine their abundance. After building the ASVs table and removing chimeras (detected using self-referencing), sequences were classified and identified with Decipher v.2.26.088, using the SILVA rRNA sequence database v.138.1 as the taxa reference89. We used phyloseq v.1.42.090 to classify and remove any sequence not classified at the kingdom and Phylum level or belonging to Archaea, Eukarya, chloroplasts, or mitochondria. Contamination assessmentAt present, there is no standard approach for minimizing or controlling potential contaminants in 16 S rRNA gene sequencing experiments91. In our study, we employed two methods to limit and eliminate contaminant sequences from downstream analyses. First, we used metagMisc v.0.5.092 to eliminate ASVs with less than ten reads (minabund = 1093). Next, we used Decontam version 1.18.094 to identify sequences that had a negative relationship with DNA concentration. We classified ASVs found in the LabControl sample as potential contaminants if they were identified as true contaminants by the Decontam algorithm. To ensure result accuracy, we then removed the identified contaminant sequences from the analysis. Respiratory bacteriome analysis and identification of functional pathways To get a sense of the bacterial community composition of the samples, we used phyloseq to identify the distribution of read counts from all the samples and to plot the relative abundance stacked bar plot at genus level. In addition, we used SourceTracker95, a Bayesian approach that allowed us to estimate the proportion of the bacterial community in the blue whale blows samples that are also detected in the seawater sample. Using microbiome v.1.2096, we identified the common core bacteriome (threshold detection set at ≥ 0.02%, prevalence set at ≥ 60%). We selected these values because we wanted a more conservative approach. Finally, we calculated alpha diversity indices: richness (S) and Simpson’s diversity index (D) using vegan v.2.6.497. Bacterial functional profiles and pathways were inferred from 16 S rRNA gene sequencing data and annotated at a 97% similarity threshold using the ref99NR database as a reference, employing the Tax4Fun2 package19, which is based on the Kyoto Encyclopedia of Genes and Genomes (KEGG; 20). All graphs were rendered using Tableau v.2024.398 and RAWGraphs v 2.099.Use of animals in research All methods were performed in accordance with the relevant international guidelines and regulations of Mexican authorities. See ethical approval.We confirm that our manuscript complies with the ARRIVE Essential 10 guidelines. The study design, experimental groups, and units are clearly described, with exact sample sizes reported. All outcome measures, including the primary outcome, are clearly defined. Statistical methods and assumptions are detailed, along with the software used. Comprehensive information on the animals, including species and probable health status, is provided. Experimental procedures are described with sufficient detail to allow replication, including what was done, how, when, where, and why. Results are presented with descriptive statistics and measures of variability, along with confidence intervals where appropriate.

    Data availability

    Data from the Sequence Read Archive (SRA) submission will be released upon publication. Accession ID: PRJNA977688.
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    Karina Acevedo-Whitehouse.Ethics declarations

    Competing interests
    The authors declare no competing interests.

    Ethical approval
    This study complied with the recommendations and methods for approaching blue whales provided by Mexican legislation (NOM-059-SEMARNAT-2010). All procedures were approved by the Bioethics Committee of the Universidad Autónoma de Queretaro (Mexico), and sampling was conducted under permits SGPA/DGVS/00255/16 and SGPA/DGVS/01832/17 issued by the Dirección General de Vida Silvestre to D. Gendron.

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    Reprints and permissionsAbout this articleCite this articleDomínguez-Sánchez, C.A., Gendron, D., Álvarez-Martínez, R.C. et al. Respiratory bacteriome and its predicted functional profiles in blue whales (Balaenoptera musculus).
    Sci Rep 15, 44434 (2025). https://doi.org/10.1038/s41598-025-28025-2Download citationReceived: 03 February 2025Accepted: 07 November 2025Published: 24 December 2025Version of record: 24 December 2025DOI: https://doi.org/10.1038/s41598-025-28025-2Share this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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    Identifying South African marine protected areas at risk from marine heatwaves and cold-spells

    AbstractMarine heatwaves (MHWs) and marine cold-spells (MCSs) can negatively impact biodiversity as species distributions are largely governed by temperature linked to physiological tolerances. These extremes have not been considered in South Africa’s Marine Protected Area (MPA) network design, so understanding frequency and severity of extreme thermal events will be important for assessing their impact. This study characterises MHWs and MCSs in MPAs across the six South African marine ecoregions, using a novel index to compare thermal event severity. Thermal events declined in duration and intensity from west to east, with the least severe events recorded in the Delagoa ecoregion. Walker Bay MPA was identified as most at risk due to the combined impact of MHWs and MCSs. These thermal events may threaten the ability of the MPA to meet its conservation objective as a cetacean sanctuary. If past trends in MHW frequency and cumulative intensity persist, the majority of South African MPAs could experience more severe heatwaves in the future. Our approach will help prioritise sites for in situ monitoring of water temperature and studies of the impact of extreme thermal events, as well as identifying areas for expanding refugia and conservation corridors, supporting adaptive management into the future.

    Data availability

    Data is freely available on Zenodo: Courtaillac et al. (2024) Identifying South African Marine Protected Areas at risk from marine heatwaves and cold spells [Data set]. Zenodo. DOI: 10.5281/zenodo.14900260.
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    Tamara B. Robinson.Ethics declarations

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    Roles of micro/nanoplastics in the spread of antimicrobial resistance through conjugative gene transfer

    AbstractThe role of micro/nanoplastics (M/NPs) in the dissemination of antimicrobial resistance (AMR) remains insufficiently understood. Here, we examine how polystyrene (PS) M/NPs of varying sizes and concentrations affect AMR gene (ARG) transfer in model systems with gram-negative (Escherichia coli) and gram-positive (Enterococcus faecalis) donors. In these systems, the ARG transfer frequency is higher for intrageneric pairs than for intergeneric pairs. The 20- and 120-nm-sized PS broadly facilitate conjugation, whereas the 1-μm-sized PS selectively promotes ARG transfer to E. coli recipients, in addition to altering the expression of conjugation- and pili-associated genes. Notably, an environmentally relevant (0.1 mg/L) concentration of PS M/NPs facilitates AMR transfer in the tested systems, which correlates with increased reactive oxygen species levels, ATP levels, and cell membrane permeability in both donors and recipients. Collectively, our findings underscore the role of M/NPs in facilitating AMR spread in specific bacterial systems, providing valuable insights for understanding their potential ecological risk in water environments.

    Data availability

    All data generated or analysed during this study are included in this published article, its Supplementary Information and the accompanying Source Data file. All RNA sequencing data have been deposited in the NCBI Gene Expression Omnibus under accession codes GSE248909 and GSE297944. Source data are provided with this paper.
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    Download referencesAcknowledgementsThis work was supported by the National Natural Science Foundation of China (grant nos. 52321005 awarded to A.W., 52293441 awarded to S.-H.G., 52293443 awarded to A.W., and 52070060 awarded to S.-H.G.), the Natural Science Foundation of Guangdong Basic and Applied Basic Research Foundation (Grant No. 2024A1515010085 awarded to S.-H.G.), the Shenzhen Overseas High-level Talents Research Startup Program (No. 20200518750C awarded to S.-H.G.), the Shenzhen Overseas High-Level Talent Innovation and Entrepreneurship Special Fund (No. KQTD20190929172630447 awarded to S.-H.G.) and Shenzhen Science and Technology Program (Nos. GXWD20231127195344001 awarded to A.W. and S.-H.G. and JCYJ20241202123735045 awarded to S.-H.G.), and the State Key Laboratory of Urban-rural Water Resource and Environment (Harbin Institute of Technology) (No.2025TS39 awarded to S.-H.G.). We would like to thank Prof. Zhigang Qiu from the Tianjin Institute of Environmental and Operational Medicine for donating the bacterial strains E. faecalis OG1RF and E. faecalis OG1RS, and Dr. Casey Huang and Dr. Lyman Tze Kin Ngiam from the Australian Centre for Water and Environmental Biotechnology for proofreading the paper. Figure 5 was designed, composed, and edited using BioRender (Kang, Y. (2025) https://BioRender.com/ofu614f), ChemDraw, and Adobe Illustrator.Author informationAuthor notesThese authors contributed equally: Yuanyuan Kang, Shu-Hong Gao.Authors and AffiliationsState Key Laboratory of Urban-Rural Water Resource and Environment, School of Eco-Environment, Harbin Institute of Technology, Shenzhen, ChinaYuanyuan Kang, Shu-Hong Gao, Yusheng Pan, Tianyao Li, Yiyi Su, Wanying Zhang, Bin Liang & Aijie WangState Key Laboratory of Urban-Rural Water Resource and Environment, School of Environment, Harbin Institute of Technology, Harbin, ChinaRui GaoDepartment of Ocean Science and Engineering, Southern University of Science and Technology, Shenzhen, ChinaLu FanAustralian Centre for Water and Environmental Biotechnology, The University of Queensland, Brisbane, QLD, AustraliaZhigang Yu & Jianhua GuoState Key Laboratory for Ecological Security of Regions and Cities, Institute of Urban Environment,, Chinese Academy of Sciences, Xiamen, ChinaJian-Qiang SuState Key Laboratory of Water Pollution Control and Green Resource Recycling, School of the Environment, Nanjing University, Nanjing, ChinaYi LuoSchool of Environmental Science and Engineering, Tiangong University, Tianjin, ChinaYue WangAuthorsYuanyuan KangView author publicationsSearch author on:PubMed Google ScholarShu-Hong GaoView author publicationsSearch author on:PubMed Google ScholarYusheng PanView author publicationsSearch author on:PubMed Google ScholarRui GaoView author publicationsSearch author on:PubMed Google ScholarTianyao LiView author publicationsSearch author on:PubMed Google ScholarLu FanView author publicationsSearch author on:PubMed Google ScholarYiyi SuView author publicationsSearch author on:PubMed Google ScholarWanying ZhangView author publicationsSearch author on:PubMed Google ScholarZhigang YuView author publicationsSearch author on:PubMed Google ScholarBin LiangView author publicationsSearch author on:PubMed Google ScholarJian-Qiang SuView author publicationsSearch author on:PubMed Google ScholarYi LuoView author publicationsSearch author on:PubMed Google ScholarYue WangView author publicationsSearch author on:PubMed Google ScholarJianhua GuoView author publicationsSearch author on:PubMed Google ScholarAijie WangView author publicationsSearch author on:PubMed Google ScholarContributionsY.K. and S.-H.G. designed the overall experiments for this study. Y.K. performed all analyses, conducted the RP4-relevant conjugation experiments within and across genera, measured ROS levels, and detected changes in cell membrane permeability and ATP generation in RP4-relevant strains. S.-H.G. supervised and managed the project and contributed to the writing and revision of the manuscript. Y.K. and S.-H.G. wrote the full manuscript and illustrated all the figures provided. Y.P., T.L. and R.G. performed the pCF10-relevant conjugation experiments and corresponding measurements of ROS production, changes in cell membrane permeability and ATP generation; Y.S. and W.Z. performed the experiments, analysed the data, and revised the manuscript. J.G. and Z.Y. contributed to the initial planning for this study, provided guidance on the research significance of this study, and contributed to revising the manuscript. L.F. and B.L. assisted in analysing the mechanisms, ecological significance, and potential application scenarios of this study. J.S. and Y.L. provided the donor and recipient strains and provided feedback on the conjugation experiments. Y.W. analysed the transcriptomic data and determined the changes in the expression of related genes under different concentrations and particle size PS treatments. A.W. provided guidance on the concentration and particle size of PS in the study and contributed to revising the manuscript. All the authors provided feedback and discussed the manuscript.Corresponding authorsCorrespondence to
    Shu-Hong Gao, Yue Wang or Aijie Wang.Ethics declarations

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    The authors declare no competing interests.

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    Nature Communications thanks Lorenzo Brusetti, Raffaella Sabatino, and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. A peer review file is available.

    Additional informationPublisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.Supplementary informationSupplementary InformationReporting SummaryTransparent Peer Review fileSource dataSource dataRights and permissions
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    Reprints and permissionsAbout this articleCite this articleKang, Y., Gao, SH., Pan, Y. et al. Roles of micro/nanoplastics in the spread of antimicrobial resistance through conjugative gene transfer.
    Nat Commun (2025). https://doi.org/10.1038/s41467-025-67879-yDownload citationReceived: 26 May 2025Accepted: 11 December 2025Published: 24 December 2025DOI: https://doi.org/10.1038/s41467-025-67879-yShare this articleAnyone you share the following link with will be able to read this content:Get shareable linkSorry, a shareable link is not currently available for this article.Copy shareable link to clipboard
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