Abstract
Alpine Upper Palaeolithic contexts exhibit specialised subsistence strategies, heavily dependent on Capra ibex. Among them, the rock shelter Riparo Dalmeri stands out, with C. ibex dominating faunal remains across all occupation phases, spanning the Pleistocene-Holocene transition. This evidence positions Riparo Dalmeri as a key site for exploring the interaction between human groups and C. ibex during this period of significant climatic and cultural shifts in human evolution. Here, we present the first multidisciplinary study on Late Palaeolithic C. ibex teeth from Riparo Dalmeri, integrating direct radiocarbon dating, stable isotope (δ13C, δ18O) and 87Sr/86Sr analyses, proteomic, and aDNA data. We generated the earliest aDNA sequences for C. ibex and contextual evidence on mobility, seasonality, and sex ratios. We found that most C. ibex were local to the area despite consistent human presence. They reveal significant dietary differences between sexes as well as increased seasonality at the Pleistocene-Holocene transition. Our results identify Riparo Dalmeri as belonging to an extinct branch of the ibex mtDNA phylogeny, offering unprecedented insights into ibex ecology and evolution that resonate with present-day issues on the conservation of this species in the face of climate change.
Data availability
Raw sequencing data are publicly available in the European Nucleotide Archive under project number PRJEB87623 (data is private pending evaluation). All the proteomic raw data, the MaxQuant evidence file, and the deamidation R script were uploaded to Zenodo (https://doi.org/10.5281/zenodo.15024003). Isotope data are available within the manuscript and its supplementary material. The OxCal code used for modelling the radiocarbon dates of Riparo Dalmeri was uploaded to Zenodo (https://doi.org/10.5281/zenodo.15063853 ).
References
Parrini, F., Cain, J. W. & Krausman, P. R. Capra Ibex (Artiodactyla: Bovidae). Mammalian Species. 830, 1–12 (2009).
Stüwe, M. & Nievergelt, B. Recovery of Alpine ibex from near extinction: The result of effective protection, captive breeding, and reintroductions. Appl. Anim. Behav. Sci. 29, 379–387 (1991).
Toïgo, C., Brambilla, A., Grignolio, S. & Pedrotti, L. The IUCN Red List of Threatened Species 2020: e.T42397A161916377. https://doi.org/10.2305/IUCN.UK.2020- 2.RLTS.T42397A161916377.en.
Semenzato, P. et al. Behavioural heat-stress compensation in a cold‐adapted ungulate: Forage‐mediated responses to warming alpine summers. Ecol. Lett. 24, 1556–1568 (2021).
Brivio, F. et al. Forecasting the response to global warming in a heat-sensitive species. Sci. Rep. 9, 3048 (2019).
Robin, M. et al. Ancient mitochondrial and modern whole genomes unravel massive genetic diversity loss during near extinction of Alpine ibex. Mol. Ecol. 31, 3548–3565 (2022).
Grossen, C., Guillaume, F., Keller, L. F. & Croll, D. Purging of highly deleterious mutations through severe bottlenecks in Alpine ibex. Nat. Commun. 11, 1001 (2020).
Fiore, I. & Tagliacozzo, A. Lo sfruttamento delle risorse animali nei siti di altura e di fondovalle nel Tardiglaciale dell’Italia nord-orientale. in Preistoria Alpina (eds Malerba, G. & Visintini, P.) 97–109 (2005).
Bertola, S. et al. L’Epigravettiano recente nell’area prealpina e alpina orientale. in L’Italia Tra 15.000 E 10.000 anni fa: Cosmopolitismo e regionalità nel Tardoglaciale (ed Martini, F.). Museo fiorentino di preistoria Paolo Graziosi, Firenze (2007).
Tagliacozzo, A. & Fiore, I. Hunting strategies in a mountain environment during the late glacial in North Eastern Italy. Preistoria Alpina. 44, 79–93 (2009).
Dalmeri, G. et al. Riparo Dalmeri: Le pietre dipinte dell’area rituale. Preistoria Alpina. 45, 67–117 (2011).
Angelucci, D. E. et al. La successione esterna del Riparo Dalmeri (Trento, Italia). Prime informazioni geoarcheologiche. Preistoria Alpina. 45, 127–146 (2011).
Ravazzi, C. An overview of the Quaternary continental stratigraphic units based on biological and climatic events in Italy. J. Quat Sci. 16, 11–18 (2003).
Dalmeri, G. et al. The ochre painted stones from the Riparo dalmeri (Trento). Development of the research on the art and rituality of the Epigravettian site. Preistoria Alpina. 44, 95–119 (2009).
Phoca-Cosmetatou, N. Specialisation & diversification: A Tale of two subsistence strategies from late glacial Italy. Before Farming. 3, 1–29 (2009).
Lugli, F. et al. A strontium isoscape of Italy for provenance studies. Chem. Geol. 587, 120624 (2022).
Lehner, B. & Grill, G. Global river hydrography and network routing: Baseline data and new approaches to study the world’s large river systems. Hydrol. Process. 27, 2171–2186 (2013).
Hartmann, J. & Moosdorf, N. The new global lithological map database glim: A representation of rock properties at the Earth surface. Geochem. Geophys. Geosyst. 13, 2012GC004370 (2012).
Kohn, M. J. Carbon isotope compositions of terrestrial C3 plants as indicators of (paleo)ecology and (paleo)climate. Proc. Natl. Acad. Sci. U.S.A. 107, 19691–19695 (2010).
Still, C. J., Berry, J. A., Collatz, G. J. & DeFries, R. S. Global distribution of C3 and C4 vegetation: Carbon cycle implications. Global Biogeochem. Cycles 17, (2003).
Pederzani, S. & Britton, K. Oxygen isotopes in bioarchaeology: Principles and applications, challenges and opportunities. Earth-Sci. Rev. 188, 77–107 (2019).
Nelson, D. B., Basler, D. & Kahmen, A. Precipitation isotope time series predictions from machine learning applied in Europe. Proc. Natl. Acad. Sci. U.S.A. 118, e2024107118 (2021).
Couturier, M. A. Le bouquetin des Alpes: Capra Aegagrus Ibex Ibex L. Grenoble (1962).
Fiore, I., Tagliacozzo, A. & Cassoli, P. F. Ibex exploitation at Dalmeri rockshelter (TN) and specialized hunting in the sites of the Eastern Alps during the Tardiglacial and the Early Holocene. Preistoria Alpina. 34, 173–183 (2001).
Villa, G. & Giacobini, G. Nuovi denti umani dai livelli epigravettiani di Riparo Dalmeri (TN). Preistoria Alpina. 41, 245–250 (2005).
Alciati, G. et al. Human deciduous dental crowns from the epigravettian layers at Riparo Dalmeri (Trento). A preliminary descriptive note. Preistoria Alpina. 34, 197–200 (2001).
Fiore, I. & Tagliacozzo, A. Lo Sfruttamento Dello Stambecco nel tardiglaciale Di Riparo dalmeri (TN): Il Livello 26c. in Archaeozoological Studies in Honour of Alfredo Riedel (Bolzano, 2006).
Seguinot, J. et al. Modelling last glacial cycle ice dynamics in the Alps. Cryosphere 12, 3265–3285 (2018).
Boxleitner, M. et al. Late Pleistocene – Holocene surface processes and landscape evolution in the central Swiss Alps. Geomorphology 295, 306–322 (2017).
Brambilla, A., Von Hardenberg, A., Nelli, L. & Bassano, B. Distribution, status, and recent population dynamics of Alpine ibex Capra Ibex in Europe. Mamm. Rev. 50, 267–277 (2020).
Curci, A. & Tagliacozzo, A. Determinazione dell’età di morte e della stagione di cattura attraverso lo studio dei livelli di accrescimento di cemento e dentina nei denti di mammiferi: L’esempio Di Riparo Dalmeri (TN). in Atti Del 2° Convegno Nazionale Di Archeozoologia (Asti, 1997), ABACO Edizioni, Forlì, (2000).
Chauveau, V. et al. Identifying the environmental drivers of corridors and predicting connectivity between seasonal ranges in multiple populations of Alpine ibex (Capra Ibex) as tools for conserving migration. Divers. Distrib. 30, e13894 (2024).
Parrini, F., Grignolio, S., Luccarini, S., Bassano, B. & Apollonio, M. Spatial behaviour of adult male Alpine ibex Capra Ibex Ibex in the Gran Paradiso National Park, Italy. Acta Theriol. 48, 411–423 (2003).
Herfindal, I., Anderwald, P., Filli, F., Campell Andri, S. & Rempfler, T. Climate, competition and weather conditions drive vertical displacement and habitat use of an alpine ungulate in a highly topographic landscape. Landsc. Ecol. 34, 2523–2539 (2019).
Grignolio, S., Rossi, I., Bassano, B. & Apollonio, M. Predation risk as a factor affecting sexual segregation in Alpine ibex. J. Mammal. 88, 1488–1497 (2007).
Grignolio, S., Rossi, I., Bassano, B., Parrini, F. & Apollonio, M. Seasonal variations of spatial behaviour in female alpine Ibex (Capra Ibex Ibex) in relation to Climatic conditions and age. Ethol. Ecol. Evol. 16, 255–264 (2004).
Cerling, T. E. & Harris, J. M. Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia 120, 347–363 (1999).
Manzella, G. et al. Paleoecology of mid-mountain Alps (Trentino, Italy) between Greenland interstadial 1 and Early Holocene. Carbon and nitrogen stable isotope analysis of ibex and red deer. Q. Sci. Rev. 328, 108549 (2024).
Lazzerini, N. et al. The isotope record (δ13C, δ18O) of vertical mobility in incremental tissues (tooth enamel, hair) of modern livestock: A reference set from the Mongolian Altai. Quat Int. 595, 128–144 (2021).
Heaton, T. H. E. Spatial, species, and Temporal variations in the 13C/12C ratios of C3plants: Implications for Palaeodiet studies. J. Archaeol. Sci. 26, 637–649 (1999).
Körner, C., Farquhar, G. D. & Roksandic, Z. A global survey of carbon isotope discrimination in plants from high altitude. Oecologia 74, 623–632 (1988).
Makarewicz, C. A., Arbuckle, B. S. & Öztan, A. Vertical transhumance of sheep and goats identified by intra-tooth sequential carbon (δ13C) and oxygen (δ18O) isotopic analyses: Evidence from chalcolithic Köşk Höyük, central Turkey. J. Archaeol. Sci. 86, 68–80 (2017).
Tornero, C. et al. The altitudinal mobility of wild sheep at the epigravettian site of Kalavan 1 (Lesser Caucasus, Armenia): Evidence from a sequential isotopic analysis in tooth enamel. J. Hum. Evol. 97, 27–36 (2016).
Brambilla, A. et al. Dietary differences in males and females of a strongly sexually dimorphic ungulate. Eur. J. Wildl. Res. 70, 98 (2024).
Tichon, J. et al. Population dynamics and the effect of drought in the threatened Nubian ibex. Wildl. Soc. Bull. 47, e1397 (2023).
Makarewicz, C. A. & Pederzani, S. Oxygen (δ18O) and carbon (δ13C) isotopic distinction in sequentially sampled tooth enamel of co-localized wild and domesticated caprines: Complications to Establishing seasonality and mobility in herbivores. Palaeogeogr Palaeoclimatol Palaeoecol. 485, 1–15 (2017).
Schenk, F. et al. Warm summers during the Younger Dryas cold reversal. Nat. Commun. 9, 1634 (2018).
Bromley, G. R. M. et al. Younger Dryas deglaciation of Scotland driven by warming summers. Proc. Natl. Acad. Sci. U.S.A. 111, 6215–6219 (2014).
Mussi, M. & Peresani, M. Human settlement of Italy during the Younger Dryas. Quat Int. 242, 360–370 (2011).
Brivio, F. et al. Seeking temporal refugia to heat stress: Increasing nocturnal activity despite predation risk. Proc. R Soc. B. 291, 20231587 (2024).
Bökönyi, S. A new method for the determination of the number of individuals in animal bone material. Am. J. Archaeol. 74, 291–292 (1970).
Holt, E., Evans, J. A. & Madgwick, R. Strontium (87Sr/86Sr) mapping: A critical review of methods and approaches. Earth Sci. Rev. 216, 103593 (2021).
Longin, R. New method of collagen extraction for radiocarbon dating. Nature 230, 241–242 (1971).
D’Elia, M. et al. Sample Preparation and blank values at the AMS radiocarbon facility of the university of Lecce. Nucl. Instrum. Methods Phys. Res. B. 223–224, 278–283 (2004).
Calcagnile, L., Quarta, G. & D’Elia, M. High-resolution accelerator-based mass spectrometry: Precision, accuracy and background. Appl. Radiat. Isot. 62, 623–629 (2005).
Stuiver, M. & Polach, H. A. Discussion reporting of 14C data. Radiocarbon 19, 355–363 (1977).
Reimer, P. J. et al. The IntCal20 Northern hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 62, 725–757 (2020).
Bronk Ramsey, C. Bayesian analysis of radiocarbon dates. Radiocarbon 51, 337–360 (2009).
Bronk Ramsey, C. Dealing with outliers and offsets in radiocarbon dating. Radiocarbon 51, 1023–1045 (2009).
Ramsey, B. OxCal v.4.4.4. (2021).
Argentino, C., Lugli, F., Cipriani, A. & Panieri, G. Testing miniaturized extraction chromatography protocols for combined 87Sr /86Sr and δ88/86Sr analyses of pore water by MC-ICP‐MS. Limnol. Oceanography: Methods. 19, 431–440 (2021).
Lugli, F., Cipriani, A., Tavaglione, V., Traversari, M. & Benazzi, S. Transhumance pastoralism of Roccapelago (Modena, Italy) early-modern individuals: Inferences from Sr isotopes of hair strands. Am. J. Phys. Anthropol. 167, 470–483 (2018).
Lugli, F., Cipriani, A., Peretto, C., Mazzucchelli, M. & Brunelli, D. In situ high Spatial resolution 87Sr/86Sr ratio determination of two Middle Pleistocene (c.a. 580 ka) Stephanorhinus hundsheimensis teeth by LA–MC–ICP–MS. Int. J. Mass Spectrom. 412, 38–48 (2017).
Berglund, M. & Wieser, M. E. Isotopic compositions of the elements 2009 (IUPAC Technical Report). Pure Appl. Chem. 83, 397–410 (2011).
McArthur, J. M., Howarth, R. J. & Bailey, T. R. Strontium isotope stratigraphy: LOWESS version 3: Best fit to the marine Sr-isotope curve for 0–509 Ma and accompanying look‐up table for deriving numerical age. J. Geol. 109, 155–170 (2001).
Vonhof, H. B. et al. High-precision stable isotope analysis of < 5 µg CaCO3 samples by continuous‐flow mass spectrometry. Rapid Comm. Mass. Spectrom. 34, e8878 (2020).
Passey, B. H. et al. Inverse methods for estimating primary input signals from time-averaged isotope profiles. Geochim. Cosmochim. Act. 69, 4101–4116 (2005).
Passey, B. H. & Cerling, T. E. Tooth enamel mineralization in ungulates: Implications for recovering a primary isotopic time-series. Geochim. Cosmochim. Act. 66, 3225–3234 (2002).
Kohn, M. J., Comment, B. H., Passey & Cerling, T. E. Geochim. Cosmochim. Act. 68, 403–405 (2004). (2002).
Pederzani, S. et al. Stable isotopes show homo sapiens dispersed into cold steppes ~ 45,000 years ago at Ilsenhöhle in Ranis, Germany. Nat. Ecol. Evol. 8, 578–588 (2024).
Pederzani, S. et al. Subarctic climate for the earliest Homo sapiens in Europe. Sci. Adv. 7, eabi4642 (2021).
Zazzo, A. et al. The isotope record of short- and long-term dietary changes in sheep tooth enamel: Implications for quantitative reconstruction of paleodiets. Geochim. Cosmochim. Act. 74, 3571–3586 (2010).
Iacumin, P., Bocherens, H., Mariotti, A. & Longinelli, A. Oxygen isotope analyses of co-existing carbonate and phosphate in biogenic apatite: A way to monitor diagenetic alteration of bone phosphate? EPSL 142, 1–6 (1996).
Delgado-Huertas, A., Iacumin, P., Stenni, B., Sánchez Chillón, B. & Longinelli, A. Oxygen isotope variations of phosphate in mammalian bone and tooth enamel. Geochim. Cosmochim. Acta. 59, 4299–4305 (1995).
Pryor, A. J. E., Stevens, R. E., O’Connell, T. C. & Lister, J. R. Quantification and propagation of errors when converting vertebrate biomineral oxygen isotope data to temperature for palaeoclimate reconstruction. Palaeogeogr Palaeoclimatol Palaeoecol. 412, 99–107 (2014).
Cappellini, E. et al. Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny. Nature 574, 103–107 (2019).
Lugli, F. et al. Enamel peptides reveal the sex of the late antique ‘Lovers of modena’. Sci. Rep. 9, 13130 (2019).
Knapp, M., Clarke, A. C. & Horsburgh, K. A. Matisoo-Smith, E. A. Setting the stage – Building and working in an ancient DNA laboratory. Ann. Anat. 194, 3–6 (2012).
Llamas, B. et al. From the field to the laboratory: Controlling DNA contamination in human ancient DNA research in the high-throughput sequencing era. STAR 3, 1–14 (2017).
Dabney, J. et al. Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proc. Natl. Acad. Sci. U S A. 110, 15758–15763 (2013).
Fontani, F. et al. Bioarchaeological and paleogenomic profiling of the unusual neolithic burial from Grotta Di Pietra Sant’Angelo (Calabria, Italy). Sci. Rep. 13, 11978 (2023).
Kapp, J. D., Green, R. E. & Shapiro, B. A fast and efficient single-stranded genomic library preparation method optimized for ancient DNA. J. Hered. 112, 241–249 (2021).
Fellows Yates, J. A. et al. Reproducible, portable, and efficient ancient genome reconstruction with nf-core/eager. PeerJ 9, e10947 (2021).
Cibulskis, K. et al. Sensitive detection of somatic point mutations in impure and heterogeneous cancer samples. Nat. Biotechnol. 31, 213–219 (2013).
Danecek, P. et al. Twelve years of samtools and BCFtools. GigaScience 10, giab008 (2021).
Alberto, F. J. et al. Convergent genomic signatures of domestication in sheep and goats. Nat. Commun. 9, 813 (2018).
Maixner, F. et al. The Iceman’s last meal consisted of fat, wild meat, and cereals. Curr. Biol. 28, 2348–2355e9 (2018).
Rambaut, A. FigTree v1.4.3. (2012). Available from http://tree.bio.ed.ac.uk/software/figtree/
Bouckaert, R. et al. BEAST 2: A software platform for bayesian evolutionary analysis. PLoS Comput. Biol. 10, e1003537 (2014).
Stadler, T., Kühnert, D., Bonhoeffer, S. & Drummond, A. J. Birth–death skyline plot reveals temporal changes of epidemic spread in HIV and hepatitis C virus (HCV). Proc. Natl. Acad. Sci. U.S.A. 110, 228–233 (2013).
Bouckaert, R. R., Drummond, A. J. & bModelTest Bayesian phylogenetic site model averaging and model comparison. BMC Evol. Biol. 17, 42 (2017).
Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in bayesian phylogenetics using tracer 1.7. Syst. Biol. 67, 901–904 (2018).
Leigh, J. W. & Bryant, D. Popart: Full-feature software for haplotype network construction. Methods Ecol. Evol. 6, 1110–1116 (2015).
Stecher, G., Tamura, K. & Kumar, S. Molecular evolutionary genetics analysis (MEGA) for MacOS. Mol. Biol. Evol. 37, 1237–1239 (2020).
Paradis, E. Pegas: an R package for population genetics with an integrated–modular approach. Bioinformatics 26, 419–420 (2010).
Nomura, K. et al. Domestication process of the goat revealed by an analysis of the nearly complete mitochondrial protein-encoding genes. PLoS ONE. 8, e67775 (2013).
Tarekegn, G. M. et al. Mitochondrial DNA variation reveals maternal origins and demographic dynamics of Ethiopian indigenous goats. Ecol. Evol. 8, 1543–1553 (2018).
Mittnik, A., Wang, C. C., Svoboda, J. & Krause, J. A molecular approach to the sexing of the triple burial at the upper paleolithic site of Dolní Věstonice. PLoS ONE. 11, e0163019 (2016).
De Flamingh, A., Coutu, A., Roca, A. L. & Malhi, R. S. Accurate sex identification of ancient elephant and other animal remains using low-coverage DNA shotgun sequencing data. G3 (Bethesda). 10, 1427–1432 (2020).
Skoglund, P., Storå, J., Götherström, A. & Jakobsson, M. Accurate sex identification of ancient human remains using DNA shotgun sequencing. J. Archaeol. Sci. 40, 4477–4482 (2013).
Denoyelle, L. et al. VarGoats project: A dataset of 1159 whole-genome sequences to dissect Capra Hircus global diversity. Genet. Sel. Evol. 53, 86 (2021).
Acknowledgements
We thank the Soprintendenza per i Beni Culturali of the Autonomous Province of Trento, and the MUSE – Science Museum of Trento for granting access to the archaeological materials analysed in this study. Dr. Diego Pinetti and Dr. Filippo Genovese of the Centro Interdipartimentale Grandi Strumenti (UNIMORE) are acknowledged for technical assistance during LC-MS/MS analyses. We also thank Prof. Luca Pagani (University of Padua) for his feedback and comments, which have provided helpful insights for this work. Nerobutto sponsored research activities on prehistory carried out at Muse – Science Museum of Trento between 2019 and 2025. F.L. is supported by the European Union’s Horizon Europe Research and Innovation programme under the Marie Skłodowska-Curie Actions PF (grant agreement no. 101104566—AROUSE). M.R. is supported by CHANGES, SPOKE 5 “Science and Technologies for Sustainable Diagnostics of Cultural Heritage”, PE 0000020, CUP B53C22003890006, NRP M4C2, Investment 1.3, funded by the European, Union—NextGenerationEU. S.S. is supported by the ERC FIRSTSTEPS project (Grant Agreement ID: 101019659). Open access publication was funded by the Department of Cultural Heritage through a grant awarded to F.F. and by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement No 724046- SUCCESS, PI: S.B).
Author information
Authors and Affiliations
Contributions
M.R., E.B., S.B., F.L. and A.C. designed the research; M.R., G.D., A.F., N.N. and R.D. conducted fieldwork and sample collection; E.A., F.F., R.I., E.C., A.L., F.L., G.T., S.S., H.V., L.C. and G.Q. performed the experiments and collected data; E.A., F.F., R.I., F.L., A.C., M.R., E.B., L.C. and G.Q. analysed the data and interpreted the results; E.A. and F.F. wrote the manuscript with contributions from all authors; A.C., F.L., M.R. and E.B. revised and edited the manuscript; M.R. supervised the project and provided guidance; M.R., A.C., S.B. and D.L. coordinated research activities and ensured project completion; M.R., S.B., A.C. and F.L. secured funding and managed financial resources.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Supplementary Material 1
Supplementary Material 2
Supplementary Material 3
Supplementary Material 4
Supplementary Material 5
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
Reprints and permissions
About this article
Cite this article
Armaroli, E., Fontani, F., Iacovera, R. et al. Ecology and demographic structure of an extinct ibex population in late Upper Palaeolithic Italian Alps.
Sci Rep (2026). https://doi.org/10.1038/s41598-025-32389-w
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-025-32389-w
Source: Ecology - nature.com
