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Home / Equine Research Bank / Sci Rep / Neanderthal coasteering and the first Portuguese hominin tra...
Scientific reports2025; 15(1); 23785; doi: 10.1038/s41598-025-06089-4

Neanderthal coasteering and the first Portuguese hominin tracksites.

Abstract: Multiple sources of evidence for the systematic use of coastal ecosystems and resources by Neanderthals are known. Fossil hominin footprints offer direct portraits of individual or social group presence and locomotor behavior, and interspecific interactions, in the coastal ecospace. Here we describe the first two hominin tracksites found in the southwestern most region of Europe. At Monte Clérigo, dated to 78 ± 5 ka, trackways of three individuals demonstrate how Neanderthals navigated dune landscapes. These behaviors suggest route planning, with dune systems serving as advantageous settings for ambush hunting or stalking prey. A single footprint at Praia do Telheiro site, dated to 82 ± 5 ka, sustains the presence of Neanderthals in the dune ecosystem during Marine Isotope Stage (MIS) 5a. Network analysis provided dietary preferences and ecological interactions of Neanderthals in coastal areas. A review of the Neanderthal coastal sites associated with faunal evidence shows that their diet was primarily centered on cervids, horses and hares. The consistent presence of these mammal taxa highlights their role as reliable food sources, irrespective of the varying environments inhabited by Neanderthals. In addition, the Neanderthal diet also incorporated animals from neighboring littoral habitats, indicating a broad foraging strategy that capitalized on local biodiversity.
© 2025. The Author(s).
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Publication Date: 2025-07-03 PubMed ID: 40610499PubMed Central: PMC12229643DOI: 10.1038/s41598-025-06089-4Google Scholar: Lookup
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Summary

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The article unveils prehistoric Neanderthal activity along the coastlines of Europe, their navigational habits, food preferences and ecosystem interactions based on fossil footprints found at two sites in Portugal. Also proposed are notions of these Neanderthals’ strategic route planning and diverse dietary habits.

Discovery of Hominin Tracksites

  • The focus of this study revolves around the first two hominin tracksites discovered in the southwestern most region of Europe, specifically at Monte Clérigo and Praia do Telheiro in Portugal.
  • The Monte Clérigo site, dated to 78 ± 5 ka (thousand years ago), reveals trackways of three individuals, demonstrating how these Neanderthals navigated the landscape primarily made up of sand dunes.
  • The footprints at Praia do Telheiro, dated to 82 ± 5 ka, further corroborates the presence of Neanderthals within the dune ecosystem during the Marine Isotope Stage 5a (a period of ancient earth’s climate history).

Implications of Behaviour and Interaction

  • The researchers infer that the way in which these trackways were positioned suggest conscious route planning by Neanderthals. Dune systems may have been used as strategic sites for hunting or stalking prey, possibly indicating an understanding of the utility of landscape in hunting techniques.
  • Network analysis gives a clearer picture about the dietary preferences and ecological interactions of Neanderthals in coastal areas. A broad range of animals including cervids, horses and hares feature predominantly in their diets. This information, derived from faunal evidence related to Neanderthal coastal sites, shows that these specific animal species were frequently relied upon as food resources regardless of the different environments that the Neanderthals lived in.
  • Additionally, the study found that Neanderthals also included animals from neighbouring coastal habitats in their diets, suggesting a diversified foraging strategy that fully utilized the local biodiversity.

Cite This Article

APA
de Carvalho CN, Cunha PP, Belo J, Muñiz F, Baucon A, Cachão M, Figueiredo S, Buylaert JP, Galán JM, Belaústegui Z, Cáceres LM, Zhang Y, Ferreira C, Rodríguez-Vidal J, Finlayson S, Finlayson G, Finlayson C. (2025). Neanderthal coasteering and the first Portuguese hominin tracksites. Sci Rep, 15(1), 23785. https://doi.org/10.1038/s41598-025-06089-4

Publication

Scientific reports
ISSN: 2045-2322
NlmUniqueID: 101563288
Country: England
Language: English
Volume: 15
Issue: 1
Pages: 23785
PII: 23785

Researcher Affiliations

de Carvalho, Carlos Neto
  • Geology Office of the Municipality of Idanha-a-Nova, Naturtejo UNESCO Global Geopark, Avenida Joaquim Morão, 6060-101, Idanha-a-Nova, Portugal. carlos.praedichnia@gmail.com.
  • Instituto D. Luiz, Faculty of Sciences, University of Lisbon, Lisboa, Portugal. carlos.praedichnia@gmail.com.
Cunha, Pedro Proença
  • MARE/ARNET, Earth Sciences Department-FCTUC, University of Coimbra, Coimbra, Portugal.
Belo, João
  • CGEO-Geosciences Center-University of Coimbra, University of Coimbra-Pólo II, Rua Sílvio Lima, 3030-790, Coimbra, Portugal.
  • FlyGIS-UAV Surveys, Coimbra, Portugal.
Muñiz, Fernando
  • Department of Crystallography, Mineralogy and Agricultural Chemistry, Faculty of Chemistry, University of Seville, C/Prof. García González, s/n, 41012, Seville, Spain.
Baucon, Andrea
  • DISTAV, University of Genova, Corso Europa, 26, 16132, Genova, GE, Italy.
Cachão, Mário
  • Instituto D. Luiz, Faculty of Sciences, University of Lisbon, Lisboa, Portugal.
  • Faculty of Sciences, Department of Geology, University of Lisbon, Campo Grande, 1749-016, Lisboa, Portugal.
Figueiredo, Silvério
  • CGEO-Geosciences Center-University of Coimbra, University of Coimbra-Pólo II, Rua Sílvio Lima, 3030-790, Coimbra, Portugal.
  • Department of Archaeology, Conservation and Heritage, Polytechnical Institute of Tomar, Quinta do Contador-Estrada da Serra, 2300-313, Tomar, Portugal.
  • Centro Português de Geo-História e Pré-História, Praceta ao Campo das Amoreiras, Lote: 1-2º O, 1750-021, Lisboa, Portugal.
Buylaert, Jan-Pieter
  • Department of Physics, Technical University of Denmark, DTU Risø Campus, 4000, Roskilde, Denmark.
Galán, José María
  • Administrative Center of El Acebuche, Doñana National Park, Matalascañas, Huelva, Spain.
Belaústegui, Zain
  • Departament of Earth and Ocean Dynamics, Faculty of Earth Sciences, University of Barcelona (UB), Martí i Franquès s/n, 08028, Barcelona, Spain.
  • Institut of Research in Biodiversity (IRBio), University of Barcelona, Barcelona, Spain.
Cáceres, Luis Miguel
  • Department of Earth Sciences, Faculty of Experimental Sciences, University of Huelva, Campus El Carmen Avda. de las Fuerzas Armadas, s/n, 21007, Huelva, Spain.
Zhang, Yilu
  • Henan Academy of Land Spatial Survey and Planning, N. 41 Huanghe Road., Jinshui District, Zhengzhou, Henan, People's Republic of China.
Ferreira, Cristiana
  • CGEO-Geosciences Center-University of Coimbra, University of Coimbra-Pólo II, Rua Sílvio Lima, 3030-790, Coimbra, Portugal.
Rodríguez-Vidal, Joaquín
  • The National Gibraltar Museum, 18-20 Bomba House Lane, Gibraltar, Gibraltar.
Finlayson, Stewart
  • The National Gibraltar Museum, 18-20 Bomba House Lane, Gibraltar, Gibraltar.
Finlayson, Geraldine
  • The National Gibraltar Museum, 18-20 Bomba House Lane, Gibraltar, Gibraltar.
Finlayson, Clive
  • The National Gibraltar Museum, 18-20 Bomba House Lane, Gibraltar, Gibraltar.

MeSH Terms

  • Animals
  • Neanderthals / physiology
  • Fossils
  • Portugal
  • Ecosystem
  • Hominidae / physiology
  • Diet
  • Humans

Grant Funding

  • (PIDDAC):UID/50019/2025 and LA/P/0068/2020 https://doi.org/10.54499/LA/P/0068/2020 / This work is supported by the Portuguese Fundau00e7u00e3o para a Ciu00eancia e Tecnologia, FCT, I.P./MCTES
  • (PIDDAC):UID/50019/2025 and LA/P/0068/2020 https://doi.org/10.54499/LA/P/0068/2020 / This work is supported by the Portuguese Fundau00e7u00e3o para a Ciu00eancia e Tecnologia, FCT, I.P./MCTES
  • MARE (BASE) (https://doi.org/10.54499/UIDB/04292/2020), / MARE (BASE)
  • MARE (Programu00e1tico) (https://doi.org/10.54499/UIDP/04292/2020) / MARE (Programu00e1tico)
  • IDL and ARNET (https://doi.org/10.54499/LA/P/0069/2020) / IDL and ARNET
  • (https://doi.org/10.54499/UIDP/00073/2020). / Centro de Geociu00eancias
  • (https://doi.org/10.54499/UIDP/00073/2020). / Centro de Geociu00eancias
  • (https://doi.org/10.54499/UIDP/00073/2020). / Centro de Geociu00eancias
  • (https://doi.org/10.54499/UIDP/00073/2020). / Centro de Geociu00eancias

Conflict of Interest Statement

Declarations. Competing interests: The authors declare no competing interests.

References

This article includes 192 references
  1. Langejans GHJ, van Niekerk KL, Dusseldorp GL, Thackeray JF. Middle stone age shellfish exploitation: potential indications for mass collecting and resource intensification at Blombos cave and Klasies river, South Africa.. 80–94 (2012).
  2. Marean CW. The transition to foraging for dense and predictable resources and its impact on the evolution of modern humans.. 20150239 (2016).
    pmc: PMC4920296pubmed: 27298470
  3. Zilhão J et al. Last interglacial Iberian neandertals as fisher-hunter-gatherers.. eaaz7943 (2020).
    pubmed: 32217702
  4. Bicho N, Esteves EP. Pleistocene hunter-gatherer coastal adaptations in Atlantic Iberia.. 957214 (2022).
  5. Marean CW. The origins and significance of coastal resource use in Africa and Western Eurasia.. 17–40 (2014).
    pubmed: 25498601
  6. Finlayson C. On the importance of coastal areas in the survival of neanderthal populations during the late pleistocene.. (23), 2246–2252 (2008).
  7. Stringer CB et al. Neanderthal exploitation of marine mammals in Gibraltar.. 14319–14324 (2008).
    pmc: PMC2567146pubmed: 18809913
  8. Hardy BL, Moncel MH. Neanderthal use of fish, mammals, birds, starchy plants and wood 125–250,000 years ago.. 0–9 (2011).
    pmc: PMC3161061pubmed: 21887315
  9. Cortés-Sanchéz M et al. Earliest known use of marine resources by neanderthals.. (9), e24026 (2011).
    pmc: PMC3173367pubmed: 21935371
  10. Cortés-Sánchez M et al. Shellfish collection on the Westernmost mediterranean, Bajondillo cave (∼160 – 35 cal Kyr BP): A case of behavioral convergence?. 284–296 (2019).
  11. Fa DA et al. Marine mollusc exploitation as evidenced by the gorham’s cave (Gibraltar) excavations 1998–2005: the Middle-Upper palaeolithic transition.. 16–28 (2016).
  12. Rios-Garaizar J. Microlithic lithic technology of neandertal shellfishers from El Cuco rockshelter (Cantabrian region, Northern Spain).. 102201 (2020).
  13. Villa P et al. Neandertals on the beach: use of marine resources at Grotta dei moscerini (Latium, Italy).. (1), e0226690 (2020).
    pmc: PMC6961883pubmed: 31940356
  14. Bicho N, Haws J. At the land’s end: marine resources and the importance of fluctuations in the coastline in the prehistoric hunter–gatherer economy of Portugal.. 2166–2175 (2008).
  15. Allen JRL. Subfossil mammalian tracks (Flandrian) in the Severn estuary, S.W. Britain: mechanics of formation, preservation and distribution.. 481–518 (1997).
  16. Sedrati M et al. A late pleistocene hominin footprint site on the North African Coast of Morocco.. 1962 (2024).
    pmc: PMC10806055pubmed: 38263453
  17. Carvalho M et al. Neanderthal palaeoecology in the late middle palaeolithic of Western iberia: a stable isotope analysis of ungulate teeth from Lapa do Picareiro (Portugal).. (2), 300–319 (2021).
  18. Roach NT et al. Pleistocene animal communities of a 1.5 million–year–old lake margin grassland and their relationship to paleoecology.. 70–83 (2018).
    pubmed: 29970233
  19. Altamura F et al. Ichnological and archaeological evidence from Gombore II OAM, Melka Kunture, Ethiopia: an integrated approach to reconstruct local environments and biological presences between 1.2 and 0.85 Ma.. 106506 (2020).
  20. D’Août K, Meert L, Van Gheluwe B, De Clercq D, Aerts P. Experimentally generated footprints in sand: analysis and consequences for the interpretation of fossil and forensic footprints.. 515–525 (2010).
    pubmed: 19927372
  21. Hatala KG et al. Snapshots of human anatomy, locomotion, and behavior from late pleistocene footprints at Engare sero, Tanzania.. 7740 (2021).
    pmc: PMC7224389pubmed: 32409726
  22. Bennett MR et al. Early hominin foot morphology based on 1.5-million-year-old footprints from ileret, Kenya.. 1197–1201 (2009).
    pubmed: 19251625
  23. Bennett, M. R. & Morse, S. (Springer International Publishing, 2014).
  24. Ashton N et al. Hominin footprints from early pleistocene deposits at happisburgh, UK.. (2), e88329 (2014).
    pmc: PMC3917592pubmed: 24516637
  25. Dingwall HL, Hatala KG, Wunderlich RE, Richmond BG. Hominin stature, body mass, and walking speed estimates based on 1.5 million-year-old fossil footprints at ileret, Kenya.. (6), 556–568 (2013).
    pubmed: 23522822
  26. Duveau J, Berillon G, Verna C, Laisné G, Cliquet D. The composition of a Neandertal social group revealed by the hominin footprints at Le Rozel (Normandy, France).. 1901789116 (2019).
    pmc: PMC6765299pubmed: 31501334
  27. Baucon A et al. Principles of ichnoarchaeology: new frontiers for studying past times.. 43–72 (2008).
  28. Neto de Carvalho C et al. Walk the world and beyond: the hominin footprints.. .
  29. Finlayson C et al. Late survival of neanderthals at the southernmost extreme of Europe.. 850–853 (2006).
    pubmed: 16971951
  30. Zilhão J et al. Symbolic use of marine shells and mineral pigments by Iberian neanderthals.. (3), 1023–1028 (2010).
    pmc: PMC2824307pubmed: 20080653
  31. Higham T, Douka K, Wood R. The timing and Spatiotemporal patterning of neanderthal disappearance.. 306–309 (2014).
    pubmed: 25143113
  32. Alcaraz-Castaño M et al. First modern human settlement recorded in the Iberian hinterland occurred during Heinrich stadial 2 within harsh environmental conditions.. 15161 (2021).
    pmc: PMC8313528pubmed: 34312431
  33. de Neto C et al. Neanderthal footprints in the Matalascañas trampled surface (SW Spain): new OSL dating and Mousterian lithic industry.. 108200 (2023).
  34. Muñiz F et al. Following the last neanderthals: mammal tracks in late pleistocene coastal dunes of Gibraltar (S Iberian Peninsula).. 297–309 (2019).
  35. de Neto C et al. First tracks of newborn straight-tusked elephants ().. 17311 (2021).
    pmc: PMC8445925pubmed: 34531420
  36. Ashton N. The Happisburgh footprints and their connections with the past.. 153–158 (2021).
  37. De Lumley H. Les Fouilles de Terra Amata à nice. Premiers résultats.. 29–51 (1966).
  38. De Lumley H. Découverte d’habitats de l’achéuleen ancien Dans des dépôts mindéliens Sur Le site de Terra Amata.. 801–804 (1967).
  39. De Lumley MA et al. Une empreinte de pied Humain acheuleen Dans La Dune Littorale du site de Terra Amata.. 483–507 (2011).
  40. Meldrum DJ. Ichnotaxonomy of the earliest hominid footprints.. 225–231 (2007).
  41. Scaillet S, Vita-Scaillet G, Guillou H. Oldest human footprints dated by ar/ar.. 320–325 (2008).
  42. Panarello A, Santello L, Belvedere M, Mietto P. Is it human? Discriminating between real tracks and track-like structures.. (1), 66–75 (2018).
  43. Helm CW et al. Dating the pleistocene hominin ichnosites on South africa’s cape South Coast.. (1), 49–68 (2023).
  44. Roberts DL. Last interglacial hominid and associated vertebrate fossil trackways in coastal eolianites, South Africa.. 190–207 (2008).
  45. Helm CW, Lockley MG, Cole K, Noakes TD, McCrea RTH. Hominin tracks in Southern africa: A review and an approach to identification.. 81–96 (2018).
  46. Helm CW et al. Newly identified hominin trackways from the cape South Coast of South Africa.. (9/10), 8156 (2020).
  47. Park KH et al. Geologic age dating of fossil hominid footprints from coast of Namjeju.. 136 (2005).
  48. Kim JY, Kim KS, Kim SH, Lee CZ, Lim JD. Preliminary report on hominid and other vertebrate footprints from the late quaternary strata of Jeju Island.. 1–11 (2009).
  49. Kim CB, Kim JY, Kim KS, Lim HS. New age constraints for hominid footprints found on Jeju island, South Korea.. 3338–3343 (2010).
  50. McLaren D et al. Terminal pleistocene epoch human footprints from the Pacific Coast of Canada.. e0193522 (2018).
    pmc: PMC5873988pubmed: 29590165
  51. Aramayo SA. Late Quaternary paleoichnological sites at Southern Atlantic coast of Buenos Aires Province, Argentina.. 112–130 (2004).
  52. Mountain ED. Footprints in calcareous sandstone of Nahoon point.. 103–111 (1966).
  53. Jacobs Z, Roberts DL. Last interglacial age for aeolian and marine deposits and the Nahoon fossil human footprints, Southeast Coast of South Africa.. 160–169 (2009).
  54. Roberts D, Berger LR. Last interglacial (c. 117 kyr) human footprints from South Africa.. (8), 349–350 (1997).
  55. Helm CW, Cawthra HC, De Vynck JC, Hattingh R, Lockley MG. Possible pleistocene hominin tracks from South africa’s West Coast.. (1/2), Art11842 (2022).
  56. McLaren D, Mackie Q, Fedje D. Experimental re-creation of the depositional context in which late pleistocene tracks were found on the Pacific Coast of Canada.. 91–100 (2021).
  57. Lucas SG. Human footprints and the peopling of the Americas.. (2), 97–100 (2017).
  58. Bayón C, Manera T, Politis G, Aramayo S. Following the tracks of the first South Americans.. 205–217 (2011).
  59. Aramayo SA, De Bianco TM. Late quaternary palaeoichnological sites from the Southern Atlantic Coast of Buenos Aires province, argentina: mammal, bird and hominid evidence.. 25–32 (2009).
  60. Aramayo SA, Manera de Bianco T, Bastianelli NV, Melchor RN. Pehuén Co: Updated taxonomic review of a late Pleistocene ichnological site in Argentina.. 144–165 (2015).
  61. Mayoral E et al. Tracking late pleistocene neandertals on the Iberian Coast.. 4103 (2021).
    pmc: PMC7952904pubmed: 33707474
  62. Mayoral E et al. New dating of the Matalascañas footprints provides new evidence of the middle pleistocene (MIS 9 8) hominin paleoecology in Southern Europe.. 17505 (2022).
    pmc: PMC9581921pubmed: 36261474
  63. Onac BP, Viehmann I, Lauritzen LJ, Stringer SE, Popitã C. U-Th ages constraining the neanderthal footprint at vârtop cave, Romania.. 1151–1157 (2005).
  64. Onac BP, Veres DS, Stringer C. Hominin footprints in caves from Romanian Carpathians.. 200–210 (2021).
  65. Kiparissi-Apostolika N, Manolis SK. Reconsideration of the antiquity of the middle palaeolithic footprints from Theopetra cave (Thessaly, Greece).. 169–182 (2021).
  66. Duveau J, Berillou G, Verna C. On the tracks of neandertals: the Ichnological assemblages from Le Rozel (Normandy, France).. 183–200 (2021).
  67. Duveau J. The Le Rozel footprints: snapshots of neandertal groups in the late pleistocene. A combined morphometric and experimental approach.. (2), 53–64 (2021).
  68. Mercier N, Martin L, Kreutzer S, Moineau V, Cliquet D. Dating the palaeolithic footprints of Le Rozel (Normandy, France).. 271–277 (2019).
  69. Ruff CB et al. Body mass Estimation from footprint size in hominins.. 102997 (2021).
    pubmed: 33993031
  70. de Neto C et al. First vertebrate tracks and palaeoenvironment in a MIS-5 context in the Doñana National park (Huelva, SW Spain).. 106508 (2020).
  71. de Neto C et al. Aurochs roamed along the SW Coast of Andalusia (Spain) during late pleistocene.. 9911 (2022).
    pmc: PMC9198092pubmed: 35701579
  72. Zazo C et al. Palaeoenvironmental evolution of the Barbate-Trafalgar Coast (Cadiz) during the last ~ 140 ka: climate, sea-level interactions and tectonics.. 212–222 (2008).
  73. Manolis SK, Aiello LC, Henessy R, Kyparissi-Apostolika N. The Middle Palaeolithic footprints from Theopetra cave (Thessaly, Greece).. 87–93 (2000).
  74. Valladas H et al. TL age-estimates for the middle palaeolithic layers of Theopetra cave (Greece).. 303–308 (2007).
  75. Lockley M, Meldrum J, Kim JY. Major events in hominin evolution.. 411–448 (2016).
  76. Bennett MR, Reynolds SC. Inferences from footprints: archaeological best practice.. 15–39 (2021).
  77. Holowka N, O’Neill MC, Thompson NA, Demes B. Chimpanzee and human midfoot motion during bipedal walking and the evolution of the longitudinal arch of the foot.. 23–31 (2017).
    pubmed: 28317554
  78. Tuttle RH. Kinesiological inferences and evolutionary implications from Laetoli bipedal trails G-1, G-2/3 and A.. 503–523 (1987).
  79. Belvedere M, Farlow JO. A numerical scale for quantifying the quality of preservation of vertebrate tracks.. 92–99 (2016).
  80. Falkingham PL et al. A standard protocol for documenting modern and fossil Ichnological data.. (4), 469–480 (2018).
  81. Bucchi A, Luengo J, Fuentes R, Arellano-Villalón M, Lorenzo C. Recommendations for improving photo quality in close range photogrammetry, exemplified in hand bones of chimpanzees and gorillas.. (2), 348–355 (2020).
  82. Romilio et al. A multidisciplinary approach to digital mapping of dinosaurian tracksites in the lower cretaceous (Valanginian-Barremian) Broome sandstone of the dampier peninsula, Western Australia.. e3013 (2017).
    pmc: PMC5363262pubmed: 28344899
  83. M Petti F et al. The use of aerial and close-range photogrammetry in the study of dinosaur tracksites: lower cretaceous (upper aptian/lower Albian) Molfetta ichnosite (Apulia, Southern Italy).. 1–18 (2018).
  84. Lkebir N et al. Anza palaeoichnological site, late cretaceous, morocco. Part III: comparison between traditional and photogrammetric records.. 103985 (2020).
  85. Belvedere M et al. Stat-tracks and mediotypes: powerful tools for modern ichnology based on 3D models.. e4247 (2018).
    pmc: PMC5767334pubmed: 29340246
  86. Belvedere M et al. Late jurassic globetrotters compared: A closer look at large and giant theropod tracks of North Africa and Europe.. 103547 (2019).
  87. Lallensack J. Automatic generation of objective footprint outlines.. e7203 (2019).
    pmc: PMC6599673pubmed: 31293834
  88. AliceVision. Meshroom: V2020.1.1. GNU-GPL.. .
  89. Cignoni P et al. MeshLab: an Open-Source Mesh Processing Tool.. 129–136 (2008).
  90. CloudCompare. V2.11.0. GNU-GPL.. .
  91. Inkscape. Draw freely. V0.92.3 GNU-GPL.. .
  92. Baucon A, Ronchi A, Felletti F, Neto de Carvalho C. Evolution of crustaceans at the edge of the end-Permian crisis: ichnonetwork analysis of the fluvial succession of Nurra (Permian-Triassic, sardinia, Italy).. (1), 74–103 (2014).
  93. Baucon A, Neto de Carvalho C, Felletti F, Tosadori G, Antonelli A. Small-world dynamics drove phanerozoic divergence of burrowing behaviors.. (6), 748–752 (2021).
  94. Baucon A, Felletti F. The ichnogis method: network science and geostatistics in ichnology. Theory and application (Grado lagoon, Italy).. 83–111 (2013).
  95. Jaccard P. Étude comparative de La distribution Florale Dans Une portion des Alpes et des Jura.. 547–579 (1901).
  96. Baucon A, Neto de Carvalho CN. Can AI get a degree in geoscience?? Performance analysis of a GPT-Based artificial intelligence system trained for Earth science (GeologyOracle).. (4), 121 (2024).
  97. Huntley DJ, Baril MR. The K content of the K-feldspars being measured in optical dating or in thermoluminescence dating.. 11–13 (1997).
  98. Cunha PP et al. The lowermost Tejo river terrace at Foz do enxarrique, portugal: A palaeoenvironmental archive from c. 60–35 ka and its implications for the last neanderthals in Westernmost Iberia.. (1), 3, 31–60 (2019).
  99. Murray AS, Helsted L, Autzen M, Jain M, Buylaert JP. Measurement of natural radioactivity: calibration and performance of a high-resolution gamma spectrometry facility.. 215–220 (2018).
  100. Cresswell AJ, Carter J, Sanderson DCW. Dose rate conversion parameters: assessment of nuclear data.. 195–201 (2018).
  101. Prescott JR, Hutton JT. Cosmic ray contributions to dose rates for luminescence and ESR dating: large depths and long-term variations.. 497–500 (1994).
  102. Murray A et al. Optically stimulated luminescence dating using quartz.. (1), art. no. 72 (2021).
  103. Murray AS, Wintle AG. The single aliquot regenerative dose protocol: potential for improvements in reliability.. 377–381 (2003).
  104. Buylaert JP et al. A robust feldspar luminescence dating method for middle and late pleistocene sediments.. 435–451 (2012).
  105. Monge Soares AM, Pereira AMM, Matos JM, Portela PJ. Radiocarbon dating of aeolianite formations.. 27–41 (2012).
  106. Martins J. A Plataforma Continental Algarvia como arquivo de paleoambientes e paleoclimas holocénicos: o papel do 14 C no seu estudo.. 278 (2014).
  107. Heaton T et al. Marine20—The marine radiocarbon age calibration curve (0–55,000 cal BP).. (4), 779–820 (2020).
  108. Neto de Carvalho C. Vertebrate tracksites from the Mid-Late pleistocene eolianites of portugal: the first record of elephant tracks in Europe.. (4), 407–414 (2009).
  109. Gouveia MP et al. Electron spin resonance dating of the culminant allostratigraphic unit of the Mondego and lower Tejo cenozoic basins (W Iberia), which predates fluvial incision into the basin-fill sediments.. 103081 (2020).
  110. Pereira AR. Geomorphology of the littoral platform: ilha do Pessegueiro.. 59–67 (2007).
  111. Cabral J. Neotectonics of Mainland portugal: state of the Art and future perspectives.. (1), 71–84 (2012).
  112. Pereira AR, Ramos C. The Southwest coast of Portugal.. 8, 109–115 (2020).
  113. Goy JL et al. Paleolandscape evolution along the Coasts of the Baixo Alentejo (Portugal) during the quaternary.. 60–75 (2024).
  114. Ressurreição R, Cabral J, Dias RP, Cunha PP. Tectonic vertical displacements in the Sines-Vila Nova de Milfontes coastal sector (Alentejo, SW Portugal).. 255–256 (2018).
  115. Figueiredo PM. Neotectonics of southwest Portugal mainland: implications on the regional seismic hazard.. 263 (2015).
  116. Figueiredo PM, Cabral J, Rockwell TK. Recognition of pleistocene marine terraces in the Southwest of Portugal (Iberian Peninsula): evidences of regional quaternary uplift.. (6), S0672 (2013).
  117. Figueiredo PM, Rockwell TK, Cabral J, Ponte C. Morphotectonics in a low tectonic rate area: analysis of the Southern Portuguese Atlantic coastal region.. 132–151 (2019).
  118. de Neto C et al. Coastal raptors and raiders: new bird tracks in the pleistocene of SW Iberian Peninsula.. 108185 (2023).
  119. Sánchez-Goñi MF et al. High resolution palynological record off the Iberian margin: direct land-sea correlation for the last interglacial complex.. 123–137 (1999).
  120. Sánchez-Goñi MF et al. Increasing vegetation and climate gradient in Western Europe over the last glacial inception (122–110 ka): data-model comparison.. 111–130 (2005).
  121. Sánchez-Goñi MF et al. Interactions vegetation-climat Au cours des derniers 425 000 Ans En Europe occidentale. Le message du Pollendes archives Marines.. 3–25 (2006).
  122. Banks WE et al. Neanderthal extinction by competitive exclusion.. (12), e3972 (2008).
    pmc: PMC2600607pubmed: 19107186
  123. Minckley T, Haws J, Benedetti M, Brewer S, Forman S. Last interglacial vegetation and climate history from the Portuguese Coast.. 59–69 (2015).
  124. Daura J et al. Cova Del rinoceront (Castelldefels, Barcelona): a terrestrial record for the Last Interglacial period (MIS 5) in the Mediterranean coast of the Iberian Peninsula.. 203–227 (2015).
  125. Rasmussen SO et al. A stratigraphic framework for abrupt Climatic changes during the last glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphy.. 14–28 (2014).
  126. Staubwasser M et al. Impact of climate change on the transition of Neanderthals to modern humans in Europe.. (37), 9116–9121 (2018).
    pmc: PMC6140518pubmed: 30150388
  127. Morse SA et al. Holocene footprints in namibia: the influence of substrate on footprint variability.. 265–279 (2013).
    pubmed: 23640691
  128. Bustos D et al. Footprints preserve terminal pleistocene hunt? Human-sloth interactions in North America.. (4), eaar7621 (2018).
    pmc: PMC5916513pubmed: 29707640
  129. Alexander RM. Stride length and speed for adults, children, and fossil hominids.. 23–27 (1984).
    pubmed: 6422766
  130. Avanzini M, Mietto P, Panarello A, De Angelis M, Rolandi G. The devil’s trails: middle pleistocene human footprints preserved in a volcanoclastic deposit of Southern Italy.. 179–189 (2008).
  131. Cohen A et al. Modern vertebrate track taphonomy at lake Manyara.. 371–389 (1991).
  132. Cohen A et al. Modern vertebrate tracks from lake manyara, Tanzania and their Paleobiological implications.. 433–458 (1993).
  133. Sorrentino R et al. Unique foot posture in neanderthals reflects their body mass and high mechanical stress.. 103093 (2021).
    pubmed: 34749003
  134. Trinkaus, R. (Academic, 1983).
  135. Trinkaus R et al. Robusticity versus shape: the functional interpretation of neandertal appendicular morphology.. (3), 257–278 (1991).
  136. DeSilva J et al. One small step: A review of Plio-Pleistocene hominin foot evolution.. (567), 63–140 (2019).
    pubmed: 30575015
  137. Pablos A et al. Metric and morphological analysis of the foot in the Middle Pleistocene sample of Sima de los Huesos (Sierra de Atapuerca, Burgos, Spain).. (Part A), 103–113 (2017).
  138. Pearson OM, Pablos A, Rak Y, Hovers E. A partial neandertal foot from the late middle paleolithic of Amud cave, Israel.. 98–125 (2020).
  139. Bicho N. The middle paleolithic occupation of Southern Portugal.. 513–531 (2004).
  140. Haws JA et al. Coastal wetlands and the neanderthal settlement of Portuguese estremadura.. 709–744 (2010).
  141. Pereira T, Hawks J, Bicho N. O paleolítico médio no território Português.. 11–30 (2012).
  142. Zilhão J et al. Precise dating of the Middle-to-Upper paleolithic transition in Murcia (Spain) supports late neandertal persistence in Iberia.. (11), e00435 (2017).
    pmc: PMC5696381pubmed: 29188235
  143. Wiseman ALA et al. The morphological affinity of the early pleistocene footprints from happisburgh, england, with other footprints of pliocene, pleistocene, and holocene age.. 102776 (2020).
    pubmed: 32505032
  144. Pérez-Díaz S, López‐Sáez JA. Late pleistocene environmental dynamics and human occupation in Southwestern Europe.. 39–53 (2021).
  145. Trájer AJ. Ecological evaluation of the development of neanderthal niche exploitation.. 108127 (2023).
  146. Jiménez-Espejo FJ et al. Climate forcing and neanderthal extinction in Southern iberia: insights from a multiproxy marine record.. (7/8), 836–852 (2007).
  147. Vidal-Cordasco M et al. Ecosystem productivity affected the Spatiotemporal disappearance of neanderthals in Iberia.. 1644–1657 (2022).
    pmc: PMC9630105pubmed: 36175541
  148. Finlayson C et al. Close encounters vs. missed connections? A critical review of the evidence for late pleistocene hominin interactions in Western Eurasia.. 108307 (2023).
  149. Hatala KG, Roach NT, Behrensmeyer AK. Fossil footprints and what they mean for hominin paleobiology.. 39–53 (2022).
    pubmed: 36223539
  150. Sévêque, N. Variabilité des comportements alimentaires au Paléolithique moyen en France septentrionale: apports des études archéozoologiques. PhD. Thesis, Université de Lille3—Charles de Gaulle, 730 (2017).
  151. Cliquet D et al. Le site paléolithique Moyen du Pou Au Rozel (Manche): des Aires des travaux spécialisés et des habitats Vieux d’environ 80000 Ans.. 13–35 (2018).
  152. Helm CW et al. A new pleistocene hominin tracksite from the cape South coast, South Africa.. 3772 (2018).
    pmc: PMC5830700pubmed: 29491482
  153. Burns A. The mesolithic footprints retained in one bed of the former saltmarshes at Formby point, Sefton coast, North West England.. 295–316 (2021).
  154. Wiseman ALA, Vicari D, Belvedere M, de Groote I. Neolithic track sites from Formby point, england: new data and insights.. 103546 (2022).
  155. Roberts G, Gonzalez S, Huddart D. Intertidal holocene footprints and their archaeological significance.. 647–651 (1996).
  156. Roberts G. Ephemeral, subfossil mammalian, avian and hominid footprints within Flandrian sediment exposures at Formby point, Sefton coast, North West England.. 33–48 (2009).
  157. Burns A, Woodward J, Conneller C, Reimer P. Footprint beds record holocene decline in large mammal diversity on the Irish sea Coast of Britain.. 1553–1563 (2022).
    pubmed: 36163258
  158. Muller S, Carlsohn A, Muller J, Baur H, Mayer F. Static and dynamic foot characteristics in children aged 1–13 years: a cross-sectional study.. 389–394 (2012).
    pubmed: 22118730
  159. Rosas A, Ríos L, Estalrrich A. The growth pattern of neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain).. (6357), 1282–1287 (2017).
    pubmed: 28935804
  160. Altamura F et al. Archaeology and ichnology at Gombore II-2, Melka kunture, ethiopia: everyday life of a mixed-age hominin group 700,000 years ago.. 2815 (2018).
    pmc: PMC5809588pubmed: 29434269
  161. Altamura F, Mussi M. Archeologia e impronte fossili nel Sito Acheuleano Di Gombore II (0,85 Ma), Melka kunture, etiopia.. (1), 21–35 (2017).
  162. Salazar-García DC et al. Neanderthal diets in central and southeastern mediterranean Iberia.. 3–18 (2013).
  163. Hockett B, Haws JA. Continuity in animal resource diversity in the Late Pleistocene human diet of central Portugal.. Before Farming 2009/2 Article 2, 1–15 (2009).
  164. Haws J et al. Paleolithic seascapes along the West Coast of Portugal.. 203–246 (2011).
  165. Demay L, Péan S, Patou-Mathis M. Mammoths used as food and Building resources by neanderthals: Zooarchaeological study applied to layer 4, Molodova I (Ukraine).. 212–226 (2012).
  166. Yravedra J et al. Neanderthal and interactions at EDAR culebro 1 (Madrid, Spain).. 500–508 (2014).
  167. Germonpré M, Udrescu M, Fiers E. Possible evidence of mammoth hunting at the neanderthal site of spy (Belgium).. 28–42 (2014).
  168. Romandini M, Nannini N, Tagliacozzo A, Peresani M. The ungulate assemblage from layer A9 at Grotta Di fumane, italy: a Zooarchaeological contribution to the reconstruction of neanderthal ecology.. 11–27 (2014).
  169. Smith GM. Neanderthal megafaunal exploitation in Western Europe and its dietary implications. A context assessment of La Cotte de st. Brelade (Jersey).. 181–201 (2015).
    pubmed: 25454779
  170. Marín J, Saladié P, Rodríguez-Hidalgo A, Carbonell E. Neanderthal hunting strategies inferred from mortality profiles within the abric Romaní sequence.. (11), e0186970 (2017).
    pmc: PMC5699840pubmed: 29166384
  171. Marín J, Saladié P, Rodríguez-Hidalgo A, Carbonell E. Ungulate carcass transport strategies at the middle palaeolithic site of abric Romaní (Capellades, Spain).. 103–121 (2017).
  172. Agam A, Barkai M. Elephant and mammoth hunting during the paleolithic: A review of the relevant archeological, ethnographic and ethno-historical records.. (3), 10.3390 (2018).
  173. Gaudzinski-Windheuser S et al. Evidence for close-range-hunting by last interglacial neanderthals.. 1087–1092 (2018).
    pubmed: 29942012
  174. Aranguren B et al. Poggetti Vecchi (Tuscany, Italy): A late Middle Pleistocene case of human–elephant interaction.. 32–60 (2019).
    pubmed: 31358183
  175. Richards MP et al. Neanderthal diet at Vindija and Neanderthal predation: the evidence from stable isotopes.. (13), 7663–7666 (2000).
    pmc: PMC16602pubmed: 10852955
  176. Bocherens H, Drucker DG, Billiou D, Patou-Mathis M, Vandermeersch B. Isotopic evidence for diet and subsistence pattern of the Saint-Césaire I neanderthal: review and use of a multi-source mixing model.. 71–87 (2005).
    pubmed: 15869783
  177. Richards MP, Trinkaus E. Isotopic evidence for the diets of European neanderthals and early modern humans.. (38), 16034–16039 (2009).
    pmc: PMC2752538pubmed: 19706482
  178. Bocherens H. Diet and ecology of neanderthals: implications from C and N isotopes: insights from bone and tooth biogeochemistry.. 73–85 (2011).
  179. Jaouen K et al. Exceptionally high δ15N values in collagen single amino acids confirm neandertals as high-trophic level carnivores.. 4928–4933 (2019).
    pmc: PMC6421459pubmed: 30782806
  180. Brown K, Fa DA, Finlayson G, Finalyson C. Small game and marine resource exploitation by neanderthals: the evidence from Gibraltar.. 247–272 (2011).
  181. Figueiredo S, Cunha PP, Carvalho IS. Marine mammals fossil remains and synthesis of the sedimentary and paleontological record of the Furninha cave pleistocene (Peniche, Portugal).. (1–2), 11–26 (2022).
  182. Sanz M, Daura J, Égüez N, Cabanes D. On the track of anthropogenic activity in carnivore dens: altered combustion structures in Cova Del Gegant (NE Iberian Peninsula).. 102–114 (2017).
  183. Mellars, P. (Princeton University Press, 1996).
  184. Réka A, Barabási AL. Statistical mechanics of complex networks.. (1), 48–97 (2002).
  185. Everett M, Borgatti SP. Ego network betweenness.. (1), 31–38 (2005).
  186. Wassermann, S. & Faust, K. (Cambridge University Press, 1994).
  187. Fortunato S. Community detection in graphs.. (3–5), 75–174 (2010).
  188. Newman MEJ. Communities, modules and large-scale structure in networks.. (1), 25–31 (2011).
  189. Blondel VD, Guillaume JL, Lambiotte R, Lefebvre E. Fast unfolding of communities in large networks.. (10), P10008 (2008).
  190. Finlayson C et al. Gorham’s cave, Gibraltar—The persistence of a neanderthal population.. 64–71 (2008).
  191. Finlayson C, Carrión J. Rapid ecological turnover and its impact on neanderthal and other human populations.. (4), 213–222 (2007).
    pubmed: 17300854
  192. Braza F, Álvarez F. Habitat use by red deer and fallow deer in Doñana National park.. 363–367 (1987).

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