Bergström, A., Stringer, C., Hajdinjak, M., Scerri, E. M. L. & Skoglund, P. Origins of modern human ancestry. Nature 590, 229–237 (2021).
Fu, Q. et al. The genetic history of Ice Age Europe. Nature 534, 200–205 (2016).
Seguin-Orlando, A. et al. Genomic structure in Europeans dating back at least 36,200 years. Science 346, 1113–1118 (2014).
Fu, Q. et al. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature 514, 445–449 (2014).
Sikora, M. et al. Ancient genomes show social and reproductive behavior of early Upper Paleolithic foragers. Science 358, 659–662 (2017).
Sikora, M. et al. The population history of northeastern Siberia since the Pleistocene. Nature 570, 182–188 (2019).
Yang, M. A. et al. 40,000-year-old individual from Asia provides insight into early population structure in Eurasia. Curr. Biol. 27, 3202–3208 (2017).
Sümer, A. P. et al. Earliest modern human genomes constrain timing of Neanderthal admixture. Nature 638, 711–717 (2025).
Iasi, L. N. M. et al. Neanderthal ancestry through time: insights from genomes of ancient and present-day humans. Science 386, eadq3010 (2024).
Groucutt, H. S. et al. Rethinking the dispersal of Homo sapiens out of Africa. Evol. Anthropol. 24, 149–164 (2015).
Groucutt, H. S. et al. Homo sapiens in Arabia by 85,000 years ago. Nat. Ecol. Evol. 2, 800–809 (2018).
Grün, R. et al. U-series and ESR ***yses of bones and teeth relating to the human burials from Skhul. J. Hum. Evol. 49, 316–334 (2005).
Valladas, H. Merrier, N., Joron, J.-L. & Reyss, J.-L. in Neandertals and Modern Humans in Western Asia (eds Akazawa, T., Aoki, K. & Bar-Yosef, O.) 69–75 (Springer, 1998).
Hershkovitz, I. et al. The earliest modern humans outside Africa. Science 359, 456–459 (2018).
Harvati, K. et al. Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia. Nature 571, 500–504 (2019).
Freidline, S. E. et al. Early presence of Homo sapiens in Southeast Asia by 86–68 kyr at Tam Pà Ling, Northern Laos. Nat. Commun. 14, 3193 (2023).
Breeze, P. et al. Palaeohydrological corridors for hominin dispersals in the Middle East ~250–70,000 years ago. Quat. Sci. Rev. 144, 155–185 (2016).
Carto, S. L., Weaver, A. J., Hetherington, R., Lam, Y. & Wiebe, E. C. Out of Africa and into an ice age: on the role of global climate change in the late Pleistocene migration of early modern humans out of Africa. J. Hum. Evol. 56, 139–151 (2009).
Mellars, P. Why did modern human populations disperse from Africa ca. 60,000 years ago? A new model. Proc. Natl Acad. Sci. 103, 9381–9386 (2006).
Klein, R. G. Out of Africa and the evolution of human behavior. Evol. Anthropol. 17, 267–281 (2008).
Mellars, P., Gori, K. C., Carr, M., Soares, P. A. & Richards, M. B. Genetic and archaeological perspectives on the initial modern human colonization of southern Asia. Proc. Natl Acad. Sci. 110, 10699–10704 (2013).
d’Errico, F. et al. Identifying early modern human ecological niche expansions and ***ociated cultural dynamics in the South African Middle Stone Age. Proc. Natl Acad. Sci. 114, 7869–7876 (2017).
d’Errico, F. & Banks, W. E. Identifying mechanisms behind Middle Paleolithic and Middle Stone Age cultural trajectories. Curr. Anthropol. 54, S371–S387 (2013).
Wilkins, J., Schoville Benjamin, J., Brown Kyle, S. & Chazan, M. Evidence for early hafted hunting technology. Science 338, 942–946 (2012).
Backwell, L. et al. The antiquity of bow-and-arrow technology: evidence from Middle Stone Age layers at Sibudu Cave. Antiquity 92, 289–303 (2018).
Bouzouggar, A. et al. 82,000-year-old shell beads from North Africa and implications for the origins of modern human behavior. Proc. Natl Acad. Sci. 104, 9964–9969 (2007).
Henshilwood, C., d’Errico, F., Vanhaeren, M., van Niekerk, K. & Jacobs, Z. Middle Stone Age shell beads from South Africa. Science 304, 404 (2004).
d’Errico, F., Henshilwood, C., Vanhaeren, M. & van Niekerk, K. N***arius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age. J. Hum. Evol. 48, 3–24 (2005).
Vanhaeren, M. et al. Middle Paleolithic shell beads in Israel and Algeria. Science 312, 1785–1788 (2006).
Steele, T., Álvarez-Fernández, E. & Hallett-Desguez, E. Special Issue: Early personal ornaments: a review of shells as personal ornamentation during the African Middle Stone Age. PaleoAnthropology 2019, 24–51 (2019).
Shea, J. J. The Middle Paleolithic of the East Mediterranean Levant. J. World Prehist. 17, 313–394 (2003).
Scerri, E. M. L. The North African Middle Stone Age and its place in recent human evolution. Evol. Anthropol. 26, 119–135 (2017).
Will, M., Conard, N. J. & Tryon, C. A. in Modern Human Origins and Dispersal (eds Sahle, Y., Reyes-Centeno, H. & Bentz, C.) 25–72 (Kerns Verlag, 2019).
Scerri, E. M. L. & Will, M. The revolution that still isn’t: the origins of behavioral complexity in Homo sapiens. J. Hum. Evol. 179, 103358 (2023).
Brown, K. S. et al. An early and enduring advanced technology originating 71,000 years ago in South Africa. Nature 491, 590–593 (2012).
Brown, K. S. et al. Fire as an engineering tool of early modern humans. Science 325, 859–862 (2009).
Henshilwood, C. S. et al. A 100,000-year-old ochre-processing workshop at Blombos Cave, South Africa. Science 334, 219–222 (2011).
Clark, J. D. The Middle Stone Age of East Africa and the beginnings of regional identity. J. World Prehist. 2, 235–305 (1988).
Tryon, C. A. & Faith, J. T. Variability in the Middle Stone Age of eastern Africa. Curr. Anthropol. 54, S234–S254 (2013).
Dibble, H. L. et al. On the industrial attributions of the Aterian and Mousterian of the Maghreb. J. Hum. Evol. 64, 194–210 (2013).
Jacobs, Z., Roberts, R. G., Nespoulet, R., El Hajraoui, M. A. & Debénath, A. Single-grain OSL chronologies for Middle Palaeolithic deposits at El Mnasra and El Harhoura 2, Morocco: implications for Late Pleistocene human–environment interactions along the Atlantic coast of northwest Africa. J. Hum. Evol. 62, 377–394 (2012).
Scerri, E. M. L. et al. Did our species evolve in subdivided populations across Africa, and why does it matter? Trends Ecol. Evol. 33, 582–594 (2018).
Powell, A., Shennan, S. & Thomas, M. G. Late Pleistocene demography and the appearance of modern human behavior. Science 324, 1298–1301 (2009).
Skoglund, P. et al. Reconstructing prehistoric African population structure. Cell 171, 59–71 (2017).
Lipson, M. et al. Ancient West African foragers in the context of African population history. Nature 577, 665–670 (2020).
Poznik, G. D. et al. Punctuated bursts in human male demography inferred from 1,244 worldwide Y-chromosome sequences. Nat. Genet. 48, 593–599 (2016).
Grove, M. Population density, mobility, and cultural transmission. J. Archaeol. Sci. 74, 75–84 (2016).
Brooks, A. S. et al. Long-distance stone transport and pigment use in the earliest Middle Stone Age. Science 360, 90–94 (2018).
Zeller, E. & Timmermann, A. The evolving three-dimensional landscape of human adaptation. Sci. Adv. 10, eadq3613 (2024).
Roberts, P. & Stewart, B. A. Defining the ‘generalist specialist’ niche for Pleistocene Homo sapiens. Nat. Hum. Behav. 2, 542–550 (2018).
Houlahan, J. E., McKinney, S. T., Anderson, T. M. & McGill, B. J. The priority of prediction in ecological understanding. Oikos 126, 1–7 (2017).
Faith, J. T. et al. Rethinking the ecological drivers of hominin evolution. Trends Ecol. Evol. 36, 797–807 (2021).
Fretwell, S. D. & Lucas, H. L. On territorial behavior and other factors influencing habitat distribution in birds. I. Theoretical development. Acta Biotheor. 19, 16–36 (1969).
Stephen Cantrell, R., Cosner, C., Deangelis, D. L. & Padron, V. The ideal free distribution as an evolutionarily stable strategy. J. Biol. Dyn. 1, 249–271 (2007).
Avgar, T., Betini, G. S. & Fryxell, J. M. Habitat selection patterns are density dependent under the ideal free distribution. J. Anim. Ecol. 89, 2777–2787 (2020).
Hutchinson, G. E. Concluding remarks. Cold Spring Harbor Symp. Quant. Biol. 22, 415–427 (1957).
Banks, W. E. et al. Eco-cultural niche modeling: new tools for reconstructing the geography and ecology of past human populations. PaleoAnthropology 2006, 68–83 (2006).
Banks, W. E. et al. Human ecological niches and ranges during the LGM in Europe derived from an application of eco-cultural niche modeling. J. Archaeol. Sci. 35, 481–491 (2008).
Timmermann, A. et al. Climate effects on archaic human habitats and species successions. Nature 604, 495–501 (2022).
Raia, P. et al. Past extinctions of Homo species coincided with increased vulnerability to climatic change. One Earth 3, 480–490 (2020).
Elith, J. & Leathwick, J. R. Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697 (2009).
Beyer, R. M., Krapp, M. & Manica, A. High-resolution terrestrial climate, bioclimate and vegetation for the last 120,000 years. Sci. Data 7, 236 (2020).
Yun, K. S. et al. A transient coupled general circulation model (CGCM) simulation of the past 3 million years. Clim. Past 19, 1951–1974 (2023).
Zeller, E. et al. Human adaptation to diverse biomes over the past 3 million years. Science 380, 604–608 (2023).
Leonardi, M., Boschin, F., Boscato, P. & Manica, A. Following the niche: the differential impact of the last glacial maximum on four European ungulates. Commun. Biol. 5, 1038 (2022).
Araújo, M. B. & New, M. Ensemble forecasting of species distributions. Trends Ecol. Evol. 22, 42–47 (2007).
Hirzel, A. H., Le Lay, G., Helfer, V., Randin, C. & Guisan, A. Evaluating the ability of habitat suitability models to predict species presences. Ecol. Model. 199, 142–152 (2006).
Humphrey, L., Bello, S. M., Turner, E., Bouzouggar, A. & Barton, N. Iberomaurusian funerary behaviour: evidence from Grotte des Pigeons, Taforalt, Morocco. J. Hum. Evol. 62, 261–273 (2012).
Stewart, B. A. et al. Ostrich eggshell bead strontium isotopes reveal persistent macroscale social networking across late Quaternary southern Africa. Proc. Natl Acad. Sci. 117, 6453–6462 (2020).
Crevecoeur, I., Dias-Meirinho, M.-H., Zazzo, A., Antoine, D. & Bon, F. New insights on interpersonal violence in the Late Pleistocene based on the Nile valley cemetery of Jebel Sahaba. Sci. Rep. 11, 9991 (2021).
Thompson, J. C. et al. Early human impacts and ecosystem reorganization in southern-central Africa. Sci. Adv. 7, eabf9776 (2021).
Texier, P.-J. et al. A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa. Proc. Natl Acad. Sci. 107, 6180–6185 (2010).
Texier, P.-J. et al. The context, form and significance of the MSA engraved ostrich eggshell collection from Diepkloof Rock Shelter, Western Cape, South Africa. J. Archaeol. Sci. 40, 3412–3431 (2013).
Clark, J. L. & Kandel, A. W. The evolutionary implications of variation in human hunting strategies and diet breadth during the Middle Stone Age of Southern Africa. Curr. Anthropol. 54, S269–S287 (2013).
Smith, M. The Archaeology of Australia’s Deserts (Cambridge Univ. Press, 2013).
Böhm, E. et al. Strong and deep Atlantic meridional overturning circulation during the last glacial cycle. Nature 517, 73–76 (2015).
Schaebitz, F. et al. Hydroclimate changes in eastern Africa over the past 200,000 years may have influenced early human dispersal. Commun. Earth Environ. 2, 123 (2021).
Nutz, A. et al. A 60–50 ka African Humid Period modulated by stadial Heinrich events HE6 and HE5a in northwestern Africa. Palaeogeogr. Palaeoclimatol. Palaeoecol. 635, 111952 (2024).
Armstrong, E., Hopcroft, P. O. & Valdes, P. J. A simulated Northern Hemisphere terrestrial climate dataset for the past 60,000 years. Sci. Data 6, 265 (2019).
Tierney, J. E., deMenocal, P. B. & Zander, P. D. A climatic context for the out-of-Africa migration. Geology 45, 1023–1026 (2017).
Beyer, R. M., Krapp, M., Eriksson, A. & Manica, A. Climatic windows for human migration out of Africa in the past 300,000 years. Nat. Commun. 12, 4889 (2021).
Aitken, M. J. & Stokes, S. in Chronometric Dating in Archaeology (eds Taylor, R. E. & Aitken, M. J.) 1–30 (Springer, 1997).
Taylor, R. E. & Aitken, M. J. (eds). Chronometric Dating in Archaeology (Springer, 1997).
Kandel, A. W. et al. The ROCEEH Out of Africa Database (ROAD): a large-scale research database serves as an indispensable tool for human evolutionary studies. PLoS One 18, e0289513 (2023).
Douka, K. et al. The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya). J. Hum. Evol. 66, 39–63 (2014).
Reimer, P. J. et al. The IntCal20 Northern Hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 62, 725–757 (2020).
Crema, E. R. & Bevan, A. Inference from large sets of radiocarbon dates: software and methods. Radiocarbon 63, 23–39 (2021).
Heaton, T. J. et al. Marine20—the marine radiocarbon age calibration curve (0–55,000 cal BP). Radiocarbon 62, 779–820 (2020).
Reimer, R. W. & Reimer, P. J. An online application for ΔR calculation. Radiocarbon 59, 1623–1627 (2017).
Vogel, J. C., Visser, E. & Fuls, A. Suitability of ostrich eggshell for radiocarbon dating. Radiocarbon 43, 133–137 (2001).
Hogg, A. G. et al. SHCal20 Southern Hemisphere calibration, 0–55,000 years cal BP. Radiocarbon 62, 759–778 (2020).
Cohen, K. M. & Gibbard, P. L. Global chronostratigraphical correlation table for the last 2.7 million years, version 2019 QI-500. Quat. Int. 500, 20–31 (2019).
Valdes, P. J. Earth systems modelling within Bridge: a database of model simulations. BRIDGE (2022).
Leonardi, M., Colucci, M., Pozzi, A. V., Scerri, E. M. L. & Manica, A. tidysdm: leveraging the flexibility of tidymodels for species distribution modelling in R. Methods Ecol. Evol. 15, 1789–1795 (2024).
Leonardi, M., Hallett, E. Y., Beyer, R., Krapp, M. & Manica, A. pastclim 1.2: an R package to easily access and use paleoclimatic reconstructions. Ecography 2023, e06481 (2023).
Will, M., Krapp, M., Stock, J. T. & Manica, A. Different environmental variables predict body and brain size evolution in Homo. Nat. Commun. 12, 4116 (2021).
Wood, S. N. Generalized Additive Models: An Introduction with R, Second Edition (Chapman and Hall/CRC, 2017).
Wood, S. N. Low-rank scale-invariant tensor product smooths for generalized additive mixed models. Biometrics 62, 1025–1036 (2006).
Clay, T. A., Phillips, R. A., Manica, A., Jackson, H. A. & de L. Brooke, M. Escaping the oligotrophic gyre? The year-round movements, foraging behaviour and habitat preferences of Murphy’s petrels. Mar. Ecol. Prog. Ser. 579, 139–155 (2017).
Hartig, F. DHARMa: Residual Diagnostics for Hierarchical (Multi-Level / Mixed) Regression Models. R package version 0.4.4. (2021).
Moran, P. A. P. Notes on continuous stochastic phenomena. Biometrika 37, 17–23 (1950).
Boyce, M. S., Vernier, P. R., Nielsen, S. E. & Schmiegelow, F. K. A. Evaluating resource selection functions. Ecol. Model. 157, 281–300 (2002).
Di Cola, V. et al. ecospat: an R package to support spatial ***yses and modeling of species niches and distributions. Ecography 40, 774–787 (2017).
Simpson, G. L. gratia: Graceful ‘ggplot’-Based Graphics and Other Functions for GAMs Fitted Using ‘mgcv’. R package version 0.6.0. (2021).
Broennimann, O. et al. Measuring ecological niche overlap from occurrence and spatial environmental data. Global Ecol. Biogeogr. 21, 481–497 (2012).
