Javanese Homo erectus on the move in SE Asia circa 1.8 Ma - Nature
erectus fossils remains, skull is Sangiran-17. ... & Granger, D. E. Age of Zhoukoudian Homo erectus determined with 26Al/10Be burial dating.
Javanese Homo erectus on the move in SE Asia circa 1.8 Ma \| Scientific Reports

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Javanese _Homo erectus_ on the move in SE Asia _circa_ 1.8 Ma


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# Javanese _Homo erectus_ on the move in SE Asia _circa_ 1.8 Ma

- [Laurent Husson](#auth-Laurent-Husson-Aff1) [1](#Aff1),
- [Tristan Salles](#auth-Tristan-Salles-Aff2) [2](#Aff2),
- [Anne-Elisabeth Lebatard](#auth-Anne_Elisabeth-Lebatard-Aff3) [3](#Aff3),
- [Swann Zerathe](#auth-Swann-Zerathe-Aff1) [1](#Aff1),
- [Régis Braucher](#auth-R_gis-Braucher-Aff3) [3](#Aff3),
- [Sofwan Noerwidi](#auth-Sofwan-Noerwidi-Aff4) [4](#Aff4),
- [Sonny Aribowo](#auth-Sonny-Aribowo-Aff1-Aff5) [1](#Aff1), [5](#Aff5),
- [Claire Mallard](#auth-Claire-Mallard-Aff2) [2](#Aff2),
- [Julien Carcaillet](#auth-Julien-Carcaillet-Aff1) [1](#Aff1),
- [Danny H. Natawidjaja](#auth-Danny_H_-Natawidjaja-Aff5) [5](#Aff5),
- [Didier Bourlès](#auth-Didier-Bourl_s-Aff3) [3](#Aff3) &
- [ASTER team](#group-1)

Show authors

[_Scientific Reports_](/srep) **volume 12**, Article number: 19012 (2022)
[Cite this article](#citeas)

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### Subjects

- [Animal migration](/subjects/animal-migration)
- [Archaeology](/subjects/archaeology)
- [Geomorphology](/subjects/geomorphology)

## Abstract

The migration of _Homo erectus_ in Southeast Asia during Early Pleistocene is cardinal to our comprehension of the evolution of the genus _Homo_. However, the limited consideration of the rapidly changing physical environment, together with controversial datings of hominin bearing sites, make it challenging to secure the robust timeline needed to unveil the behavior of early humans. Here, we reappraise the first appearance datum of Javanese _H. erectus_ by adding the most reliable age constraints based on cosmogenic nuclides \\(^{10}\\)Be and \\(^{26}\\)Al produced _in situ_ to a compilation of earlier estimates. We find that _H. erectus_ reached Java and dwelled at Sangiran, Java, _ca._ 1.8 Ma. Using this age as a baseline, we develop a probabilistic approach to reconstruct their dispersal routes, coupling ecological movement simulations to landscape evolution models forced by reconstructed geodynamic and climatic histories. We demonstrate that the hospitable _terra firma_ conditions of Sundaland facilitated the prior dispersal of hominins to the edge of Java, where they conversely could not settle until the Javanese archipelago emerged from the sea and connected to Sundaland. The dispersal of _H. erectus_ across Sundaland occurred over at least tens to hundreds kyr, a time scale over which changes in their physical environment, whether climatic or physiographic, may have become primary forcings on their behavior. Our comprehensive reconstruction method to unravel the peopling timeline of SE Asia provides a novel framework to evaluate the evolution of early humans.

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ArticleOpen access05 October 2021

## Introduction

_When and how did_ _Homo erectus_ _disperse in SE Asia?_ The first part of the question, _when_, has received great attention since the early 20\\(^{th}\\) century discovery of fossil remains in Java[1](/articles/s41598-022-23206-9#ref-CR1 "Dubois, E. On Pithecanthropus Erectus: a transitional form between man and the apes. J. Anthropol. Inst. G. B. Irel. 25, 240–255 (1896)."), [2](/articles/s41598-022-23206-9#ref-CR2 "von Koenigswald, G. H. R. Neue Pithecanthropus-funde, 1936-1938 : ein beitrag zur Kenntnis der Praehominiden Wetenschappelijke Mededeelingen ; no. 28 (Landsdrukkerij, Batavia, 1940)."). The second part, however, has mostly been overlooked even though understanding _how_ _H. erectus_ dispersed holds clues not only on the climatic[3](/articles/s41598-022-23206-9#ref-CR3 "Janssen, R. et al. Tooth enamel stable isotopes of Holocene and Pleistocene fossil fauna reveal glacial and interglacial paleoenvironments of hominins in Indonesia. Quatern. Sci. Rev. 144, 145–154 (2016)."), ecological[4](/articles/s41598-022-23206-9#ref-CR4 "Bettis, E. A. et al. Way out of Africa: Early Pleistocene paleoenvironments inhabited by Homo erectus in Sangiran, Java. J. Hum. Evol. 56(1), 11–24 (2009)."), geological[5](/articles/s41598-022-23206-9#ref-CR5 "Huffman, O. Geologic context and age of the Perning/Mojokerto Homo erectus, East Java. J. Hum. Evol. 40(4), 353–362 (2001)."), or physiographic[6](/articles/s41598-022-23206-9#ref-CR6 "Sarr, A.-C. et al. Subsiding Sundaland. Geology (Boulder) 47(2), 119–122 (2019)."), [7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021).") environment suitable for the dispersal of hominins, but also on some of the physical and behavioral characteristics of early humans. This question is all the more relevant given the singular place that Javanese _H. erectus_ occupy, not only as a historical landmark[1](/articles/s41598-022-23206-9#ref-CR1 "Dubois, E. On Pithecanthropus Erectus: a transitional form between man and the apes. J. Anthropol. Inst. G. B. Irel. 25, 240–255 (1896)."), but also owing to their cardinal position at the southeastern end of the realm of Early Pleistocene hominins. Here, we jointly tackle both parts of the question: first, we reappraise the age at which _H. erectus_ colonized the region and second, we evaluate their migration pathways using paleo-environmental and ecological modeling techniques.

In the very dynamic landscapes of Java and Sundaland (Fig. [1](/articles/s41598-022-23206-9#Fig1)), constantly reshaped by the joint action of geodynamics and climate during the Quaternary[7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."), [8](/articles/s41598-022-23206-9#ref-CR8 "Husson, L., Boucher, F. C., Sarr, A., Sepulchre, P. & Cahyarini, S. Y. Evidence of Sundaland’s subsidence requires revisiting its biogeography. J. Biogeogr. 47(4), 843–853 (2020)."), a precise knowledge of the chronological framework is requisite to reconstruct the physical environment. We therefore directly dated the earliest arrival of _H. erectus_ in Java, which defines the baseline that we then use to assess the past physiography and dispersal routes from mainland Asia to Java. Javanese site Sangiran (Fig. [1](/articles/s41598-022-23206-9#Fig1)) is pivotal in that respect because it yields the largest number of hominin finds, and also because it counts amongst the earliest _H. erectus_ bearing sites in SE Asia. Reconstructing the pathways and drivers of hominin dispersal across Eurasia often relies on the purported age of first appearance of _H. erectus_ in Sangiran[9](/articles/s41598-022-23206-9#ref-CR9 "Winder, I. C. et al. Evolution and dispersal of the genus Homo: A landscape approach. J. Hum. Evol. 87, 48–65 (2015)."), [10](/articles/s41598-022-23206-9#ref-CR10 "Carotenuto, F. et al. Venturing out safely: The biogeography of Homo erectus dispersal out of Africa. J. Hum. Evol. 95, 1–12 (2016)."), which was commonly bracketed between 1.5 Ma and 1.7 Ma based on \\(^{40}\\)Ar/\\(^{39}\\)Ar and paleomagnetic dating[11](#ref-CR11 "Larick, R. et al. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. Proc. Natl. Acad. Sci. PNAS 98(9), 4866–4871 (2001)."), [12](#ref-CR12 "Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G. & Suprijo, A. Age of the earliest known hominids in Java, Indonesia. Science 263(5150), 1118–1121 (1994)."), [13](#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000)."), [14](/articles/s41598-022-23206-9#ref-CR14 "Bettis, E. et al. Landscape development preceding Homo erectus immigration into Central Java, Indonesia: The Sangiran Formation Lower Lahar. Palaeogeogr. Palaeoclimatol. Palaeoecol. 206(1), 115–131 (2004)."). However, a recent study opposed a younger age of \\(\\sim\\)1.3 Ma, based on U-Pb radiometric dating and zircon fission tracks[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020).").

Dating in Java is notoriously difficult owing to the scarcity of datable material in the volcano-clastic sediments that would unambiguously determine the age of arrival of _H. erectus_ with an unbiased method, but also because of the geological setting of the Sangiran dome (Fig. [1](/articles/s41598-022-23206-9#Fig1) c). Despite its tectonic deformation providing good exposure of fossiliferous layers, sedimentary deposits are often inconveniently reworked. Yet, within the available dataset, only the latest dating[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020).") is in fact difficult to reconcile with the others. While the large uncertainties and the overdispersion of the data revealed by the mean square weighted deviation (Fig. 3 in[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020).")) may explain this outlier, their study indisputably revives the controversy. In addition, most previous dating methods suffer from large uncertainties arising from the poor constraints on the pre-burial history. Instead, Terrestrial Cosmogenic Nuclide dating (TCN) is, to our knowledge, the only available method to focus on the burial time and thus to minimize uncertainties associated to the pre-burial history. For that reason, this technique is increasingly applied in archaeology and paleoanthropology, and sheds new lights on hominin dispersal[16](#ref-CR16 "Granger, D. E. & Muzikar, P. F. Dating sediment burial with in situ-produced cosmogenic nuclides: Theory, techniques, and limitations. Earth Planet. Sci. Lett. 188(1), 269–281 (2001)."), [17](#ref-CR17 "Shen, G., Gao, X., Gao, B. & Granger, D. E. Age of Zhoukoudian Homo erectus determined with 26Al/10Be burial dating. Nature 458(7235), 198–200 (2009)."), [18](#ref-CR18 "Pappu, S. et al. Early Pleistocene presence of Acheulian Hominins in South India. Science 331(6024), 1596–1599 (2011)."), [19](#ref-CR19 "Lebatard, A.-E. et al. Dating the Homo erectus bearing travertine from Kocabaş (Denizli, Turkey) at at least 1.1 Ma. Earth Planet. Sci. Lett.390, 8–18 (2014)."), [20](#ref-CR20 "Lebatard, A.-E., Bourlès, D. L. & Braucher, R. Absolute dating of an Early Paleolithic site in Western Africa based on the radioactive decay of in situ-produced 10Be and 26Al. Nucl. Instrum. Methods Phys. Res. Sect. B 456, 169–179 (2019)."), [21](/articles/s41598-022-23206-9#ref-CR21 "Braucher, R., Oslisly, R., Mesfin, I., Ntoutoume Mba, P. P. & Team, A. In situ-produced 10 Be and 26 Al indirect dating of Elarmékora Earlier Stone Age artifacts: First attempt in a savannah forest mosaic in the middle Ogooué valley, Gabon. Philos. Trans. Biol. Sci. (2021) ."). Here, we date the first appearance of Javanese _H. erectus_ using TCN relying on the concentration of two _in-situ_ produced cosmogenic nuclides (\\(^{10}\\)Be and \\(^{26}\\)Al) in quartz grains from the sedimentary layers of interest. These isotopes accumulate in the grains when they are exposed to cosmic radiations during exhumation and transport phases, and conversely decay when the grains are shielded during burial periods. The burial duration in terrigenous sediments, and exhumation history until sampling, is thus encoded in the sediments by the amount of each isotope they contain.

**Figure 1**

[![figure 1](//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41598-022-23206-9/MediaObjects/41598_2022_23206_Fig1_HTML.png)](/articles/s41598-022-23206-9/figures/1)

Geology of Sangiran. ( **a**) Regional view. Black line delinates continental Sundaland (120 m isodepth). Stars indicate possible northern entry points (thereafter set to Myanmar, Thailand and Vietnam) for _H. erectus_. ( **b**) Java island and main biostratigraphic sites (hominin-bearing sites: Sangiran (s), Mojokerto (m), Trinil (t), Ngandong (n); and faunal site Bumiayu ( **b**). ( **c**) 3D view of the Sangiran dome. Geological map redrawn from[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."), [22](/articles/s41598-022-23206-9#ref-CR22 "Grimaud-Hervé, D. et al. Position of the posterior skullcap fragment from Sendang Klampok (Sangiran Dome, Java, Indonesia) among the Javanese Homo erectus record. Quatern. Int. 416, 193–209 (2016)."). Red star locates Bukuran creek outcrop; skull is Sangiran-17[23](/articles/s41598-022-23206-9#ref-CR23 "Sartono, S. Observations on a new skull of Pithecanthropus erectus (Pithecanthropus VIII), from Sangiran, Central Java. Koninklijke Akademie Wetenschappen te Amsterdam 74, 185–194 (1971)."). Maps on panels ( **a**, **b**) were created using GMT 5 ( [www.generic-mapping-tools.org](http://www.generic-mapping-tools.org))[24](/articles/s41598-022-23206-9#ref-CR24 "Wessel, P., Smith, W. H. F., Scharroo, R., Luis, J. & Wobbe, F. Generic mapping tools: Improved version released. EOS Trans. Am. Geophys. Union 94(45), 409–410.                    https://doi.org/10.1002/2013EO450001                                     (2013)."); on panel ( **c**). using ArcGIS ( [www.arcgis.com](http://www.arcgis.com)).

[Full size image](/articles/s41598-022-23206-9/figures/1)

A refined knowledge of this time period is crucial to decipher the dispersal and behavior of Javanese _H. erectus_ for at least two reasons. First, depending on this age, it is unclear if Javanese _H. erectus_ directly arrived along the coastlines in a single _out-of-Africa_ episode, or discontinuously stemmed from smaller size groups that expanded earlier in China[25](/articles/s41598-022-23206-9#ref-CR25 "Antón, S., Potts, R. & Aiello, L. Evolution of Early Homo: An integrated biological perspective. Science (New York, N.Y.)345 (2014).                    https://doi.org/10.1126/science.1236828                                    ."), [26](/articles/s41598-022-23206-9#ref-CR26 "Luo, L. et al. The first radiometric age by isochron 26Al/10Be burial dating for the Early Pleistocene Yuanmou hominin site, southern China. Quat. Geochronol. 55, 101022.                    https://doi.org/10.1016/j.quageo.2019.101022                                     (2019)."); it implies that early humans could have entered Sundaland from different points in mainland Asia (Fig. [1](/articles/s41598-022-23206-9#Fig1) a). Second, the subsequent trajectories of Javanese _H. erectus_ across Sundaland were conditioned by the extremely transient paleoenvironmental conditions. While it is at present-day largely flooded, Sundaland was permanently continental during Early Pleistocene[6](/articles/s41598-022-23206-9#ref-CR6 "Sarr, A.-C. et al. Subsiding Sundaland. Geology (Boulder) 47(2), 119–122 (2019)."); conversely, while Sundaland was slowly drowning, Java was an uplifting chain of volcanoes emerging from a shallow sea. The settlement of terrestrial faunas and hominins in Java occurs soon after the transition from marine to more terrestrial environments[2](/articles/s41598-022-23206-9#ref-CR2 "von Koenigswald, G. H. R. Neue Pithecanthropus-funde, 1936-1938 : ein beitrag zur Kenntnis der Praehominiden Wetenschappelijke Mededeelingen ; no. 28 (Landsdrukkerij, Batavia, 1940)."), [4](/articles/s41598-022-23206-9#ref-CR4 "Bettis, E. A. et al. Way out of Africa: Early Pleistocene paleoenvironments inhabited by Homo erectus in Sangiran, Java. J. Hum. Evol. 56(1), 11–24 (2009)."), [11](/articles/s41598-022-23206-9#ref-CR11 "Larick, R. et al. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. Proc. Natl. Acad. Sci. PNAS 98(9), 4866–4871 (2001)."), [13](/articles/s41598-022-23206-9#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000)."), [27](/articles/s41598-022-23206-9#ref-CR27 "Zaim, Y. et al. New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J. Hum. Evol.61(4), 363–376 (2011)."), [28](/articles/s41598-022-23206-9#ref-CR28 "Rizal, Y. et al. Last appearance of Homo erectus at Ngandong, Java, 117,000–108,000 years ago. Nature 577(7790), 381–385 (2020)."). Within this broad geographical framework, the direction and pace of movements of _H. erectus_ were driven by the contingencies of the local physical environment at the time of their migration, as defined by the river network, relief, and vegetation cover of Sundaland and Java.

To quantitatively unravel their migration pathways across this complex environment, we build upon two recent advances in paleo-environmental and ecological modeling techniques. First, we reconstruct past physiographies over geological time, accounting for the joint effects of geodynamic deformations and climatic forcings, by applying a Landscape Evolution Model (LEM). Concurrently, an array of mechanistic models have emerged from conservationist studies[29](#ref-CR29 "McRae, B. & Beier, P. Circuit theory predicts gene flow in plant and animal populations. Proc. Natl. Acad. Sci. USA 104, 19885–90.                    https://doi.org/10.1073/pnas.0706568104                                     (2008)."), [30](#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019)."), [31](/articles/s41598-022-23206-9#ref-CR31 "Landau, V. A., Shah, V. B., Anantharaman, R. & Hall, K. R. Omniscape.jl: Software to compute omnidirectional landscape connectivity. J. Open Source Softw.6(57), 2829 (2021).                    https://doi.org/10.21105/joss.02829                                    .") to simulate contemporary species displacements across a given landscape, based on a series of morphometric biases that prompt or hinder the motion of species, like relief and drainage, but also climate or vegetation cover. Here, we reconstruct the past physiography using the LEM _goSPL_ [32](/articles/s41598-022-23206-9#ref-CR32 "Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. J. Open Source Softw.5(56), 2804 (2020).                    https://doi.org/10.21105/joss.02804                                    .") that simulates the joint effects of erosion, sediment transport and deposition on the relief and drainage network with an adequate resolution (\\(\\sim\\)500 m) to address biogeographical purposes[7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."). The modelled landscape is sculpted at the time of hominin migration by the interplay between geodynamics -which deforms the surface of the Sunda shelf[6](/articles/s41598-022-23206-9#ref-CR6 "Sarr, A.-C. et al. Subsiding Sundaland. Geology (Boulder) 47(2), 119–122 (2019)."), [7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."), [33](/articles/s41598-022-23206-9#ref-CR33 "Husson, L. et al. Slow geodynamics and fast morphotectonics in the far East Tethys. Geochem. Geophys. Geosyst. 23(1), n/a (2022).")\- and climate evolution[34](/articles/s41598-022-23206-9#ref-CR34 "Valdes, P., Scotese, C. & Lunt, D. Deep ocean temperatures through time. Climate Past 17, 1483–1506.                    https://doi.org/10.5194/cp-17-1483-2021                                     (2021).") -which sets the amount of precipitation. The reconstructed landscape then defines the environmental conditions for the mechanistic model of ecological displacement _SiMRiv_ [30](/articles/s41598-022-23206-9#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019)."), in which the random trajectories of species are conditioned by landscape complexity, perceptual range, and partial memory of past displacement (semi-correlated Lévy-flight)[30](/articles/s41598-022-23206-9#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019)."). Applying this approach to the longterm dispersal of _H. erectus_ across Sundaland, we opt for a probabilistic assessment of a large set of simulations (3000 realizations with 5 millions iterations each), which we use to evaluate the travel distances and times, and the likelihood of the presence of hominins at any given location within the region. For reference, we additionally compute the least-cost path end-member solution (upon the incongruous hypothesis that hominins had a destination and a roadmap).

## Results

### Dating Sangiran

We dated the earliest episode of continentalization of the Sangiran Dome (Fig. [2](/articles/s41598-022-23206-9#Fig2) a) in Java island, that was colonized by hominins shortly after[13](/articles/s41598-022-23206-9#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000)."). The general lithostratigraphy spans more than 2 Ma[36](/articles/s41598-022-23206-9#ref-CR36 "Brasseur, B., Sémah, F., Sémah, A.-M. & Djubiantono, T. Pedo-sedimentary dynamics of the Sangiran dome hominid bearing layers (Early to Middle Pleistocene, central Java, Indonesia): A palaeopedological approach for reconstructing ‘Pithecanthropus’ (Javanese Homo erectus) palaeoenvironment. Quatern. Int. 376, 84–100 (2015)."). The marine Puren _fm._ forms the earliest layers. It is overlain by the Sangiran _fm._, wherein the earliest terrestrial faunas are found in the _Lower Lahar_ unit[2](/articles/s41598-022-23206-9#ref-CR2 "von Koenigswald, G. H. R. Neue Pithecanthropus-funde, 1936-1938 : ein beitrag zur Kenntnis der Praehominiden Wetenschappelijke Mededeelingen ; no. 28 (Landsdrukkerij, Batavia, 1940)."). Hominin fossils appear in the uppermost part of the Sangiran _fm._ where shallow marine environments gradually give space to more terrestrial settings. The conglomerates of the _Grenzbank_ unit, at the base of the Bapang _fm._ mark the onset of more permanent and hospitable conditions. The overlying Bapang _fm._, predominantly fluviatile, hosts the vast majority of _H. erectus_ finds until the _Upper Tuff_ unit. It is unconformably overlain by the Pohjajar _fm._, and ultimately by recent alluvium from the Solo river; to date, no hominin fossil has been retrieved from these uppermost formations. Following their deposition, the Bapang and Pohjajar formations have been eroded during the deformation of the Sangiran dome, exposing the deeper fossil-bearing formations. In order to most comprehensively evaluate the chronology of the events, we benefit from available ages for the entire stratigraphic series that we complement by TCN dating of the critical _Grenzbank_ unit.

**Figure 2**

[![figure 2](//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41598-022-23206-9/MediaObjects/41598_2022_23206_Fig2_HTML.png)](/articles/s41598-022-23206-9/figures/2)

Bukuran sampling site, stratigraphic setting, and analytical results. ( **a**) General lithostratigraphy of the Sangiran Dome[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."), [22](/articles/s41598-022-23206-9#ref-CR22 "Grimaud-Hervé, D. et al. Position of the posterior skullcap fragment from Sendang Klampok (Sangiran Dome, Java, Indonesia) among the Javanese Homo erectus record. Quatern. Int. 416, 193–209 (2016)."), [35](/articles/s41598-022-23206-9#ref-CR35 "Hyodo, M. et al. High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia. Proc. Natl. Acad. Sci. PNAS 108(49), 19563–19568 (2011)."), depositional environment, and available datings\\(\\dagger\\). Blue bars indicate proven occurrence of _H. erectus_ fossils remains, skull is Sangiran-17[23](/articles/s41598-022-23206-9#ref-CR23 "Sartono, S. Observations on a new skull of Pithecanthropus erectus (Pithecanthropus VIII), from Sangiran, Central Java. Koninklijke Akademie Wetenschappen te Amsterdam 74, 185–194 (1971)."). ( **b**) Stratigraphic log of the sampling site. ( **c**) Sampling site, lithologies, and sample location (labeled by their sample name suffix: _1_ is SAN18-1). The Bukuran stream flows at the bottom of the outcrop. ( **d,e**): \\(^{10}\\)Be and \\(^{26}\\)Al concentrations and \\(^{26}\\)Al/\\(^{10}\\)Be ratios over depth. Red and blue dots are \\(^{26}\\)Al and \\(^{10}\\)Be concentrations (S.I. Table 1). _L. Lahar_: _Lower Lahar_; _U. Tuff_: _Upper Tuff_; _al._: alluvium; _Gb._: _Grenzbank_ unit. \\(\\dagger\\) Earlier datings: _H11_, paleomag[35](/articles/s41598-022-23206-9#ref-CR35 "Hyodo, M. et al. High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia. Proc. Natl. Acad. Sci. PNAS 108(49), 19563–19568 (2011)."). _L01_, \\(^{40}\\)Ar/\\(^{39}\\)Ar[11](/articles/s41598-022-23206-9#ref-CR11 "Larick, R. et al. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. Proc. Natl. Acad. Sci. PNAS 98(9), 4866–4871 (2001).") ; _M20a_, U-Pb and ZFT[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."); _M20b_, U-Pb[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."); _M20c_, U-Pb and ZFT[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."); _S94_, \\(^{40}\\)Ar/\\(^{39}\\)Ar[12](/articles/s41598-022-23206-9#ref-CR12 "Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G. & Suprijo, A. Age of the earliest known hominids in Java, Indonesia. Science 263(5150), 1118–1121 (1994)."); _S00_, \\(^{40}\\)Ar/\\(^{39}\\)Ar, paleomag.[13](/articles/s41598-022-23206-9#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000)."); _B04_, \\(^{40}\\)Ar/\\(^{39}\\)Ar[14](/articles/s41598-022-23206-9#ref-CR14 "Bettis, E. et al. Landscape development preceding Homo erectus immigration into Central Java, Indonesia: The Sangiran Formation Lower Lahar. Palaeogeogr. Palaeoclimatol. Palaeoecol. 206(1), 115–131 (2004).").

[Full size image](/articles/s41598-022-23206-9/figures/2)

Samples were extracted in the Bukuran creek in the SE of the Sangiran dome (Fig. [1](/articles/s41598-022-23206-9#Fig1) c), less than 30 m north of the find spot of a maxilla[27](/articles/s41598-022-23206-9#ref-CR27 "Zaim, Y. et al. New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J. Hum. Evol.61(4), 363–376 (2011)."), at the exact location of a section that was previously dated with discordant ages at \\(\\sim\\)1.5 Ma[11](/articles/s41598-022-23206-9#ref-CR11 "Larick, R. et al. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. Proc. Natl. Acad. Sci. PNAS 98(9), 4866–4871 (2001).") and \\(\\sim\\)0.9 Ma[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."). The local stratigraphy (Fig. [2](/articles/s41598-022-23206-9#Fig2) b,c) has been documented in great details[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."), [27](/articles/s41598-022-23206-9#ref-CR27 "Zaim, Y. et al. New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J. Hum. Evol.61(4), 363–376 (2011)."), and corresponds to the base of the Bapang _fm._, the basal conglomerate being the _Grenzbank_ unit, and the dark clay the uppermost unit of Sangiran _fm_.

\\(^{10}\\)Be and \\(^{26}\\)Al concentrations and ratios do not show significant depth dependence (Fig. [2](/articles/s41598-022-23206-9#Fig2) d,e), while an exponential depth decrease of cosmogenic isotope concentration could instead be expected _a priori_ due to the depth attenuation of the incoming cosmic ray particles (Methods). This uniformity results from the long-lasting denudation of the \\(\\sim\\)100 m thick layers of the Bapang and Pohjajar formations: This episode, which necessarily postdates the deposition of Pohjajar _fm._ at 0.15 to 0.25 Ma[36](/articles/s41598-022-23206-9#ref-CR36 "Brasseur, B., Sémah, F., Sémah, A.-M. & Djubiantono, T. Pedo-sedimentary dynamics of the Sangiran dome hominid bearing layers (Early to Middle Pleistocene, central Java, Indonesia): A palaeopedological approach for reconstructing ‘Pithecanthropus’ (Javanese Homo erectus) palaeoenvironment. Quatern. Int. 376, 84–100 (2015)."), corresponds to mean denudation rates of 400 to 600 m/Ma. As it proceeds, this sustained yet recent exhumation advects and constantly trims the uppermost, \\(^{10}\\)Be and \\(^{26}\\)Al enriched layers while they accumulate cosmogenic isotopes. Radioactive decay is modulated by post-burial production during the final stage of exhumation, which then has to be accounted for[18](/articles/s41598-022-23206-9#ref-CR18 "Pappu, S. et al. Early Pleistocene presence of Acheulian Hominins in South India. Science 331(6024), 1596–1599 (2011)."), [20](/articles/s41598-022-23206-9#ref-CR20 "Lebatard, A.-E., Bourlès, D. L. & Braucher, R. Absolute dating of an Early Paleolithic site in Western Africa based on the radioactive decay of in situ-produced 10Be and 26Al. Nucl. Instrum. Methods Phys. Res. Sect. B 456, 169–179 (2019)."). Inverting \\(^{10}\\)Be and \\(^{26}\\)Al concentrations yields a mean burial age of 1.78±0.20 Ma, pre-burial denudation rates ranging from 21 to 80 m/Ma, and post-deposition denudation rates from 334 to 608 m/Ma (Methods). These mean denudation rates are consistent with geological estimates of erosion rates following the deposition of the Pohjajar _fm._, and the tenfold increase of pre-burial to post-burial rates is consistent with the respective lithologies of the source rocks (volcanic rocks) and of the unconsolidated Pleistocene sediments. For reference, we also computed the minimal burial age, given by the end-member case considering the (quasi-)instantaneous removal of tens of meters of sediments over the outcrop, while ignoring the longer term erosion of the Bapang and Pohjajar formations. In that case, only the time span since deposition sets the present-day ratios; this hypothesis yields a minimum weighted mean burial duration of 1.45±0.10 Ma (Methods).

This TCN age (1.78±0.35 Ma) of the _Grenzbank_ unit arguably belongs to the older end of the range of earlier estimates (Fig. [2](/articles/s41598-022-23206-9#Fig2) a). The dated unit is bracketed by layers of compatible ages, from the base of the Sangiran _fm._ (\\(^{40}\\)Ar/\\(^{39}\\)Ar and paleomagnetic ages of \\(\\sim\\)1.9[14](/articles/s41598-022-23206-9#ref-CR14 "Bettis, E. et al. Landscape development preceding Homo erectus immigration into Central Java, Indonesia: The Sangiran Formation Lower Lahar. Palaeogeogr. Palaeoclimatol. Palaeoecol. 206(1), 115–131 (2004).") and \\(\\sim\\)1.7 Ma[12](/articles/s41598-022-23206-9#ref-CR12 "Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G. & Suprijo, A. Age of the earliest known hominids in Java, Indonesia. Science 263(5150), 1118–1121 (1994)."), [13](/articles/s41598-022-23206-9#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000).")) and from its core (U-Pb ages of \\(\\sim\\)1.7 Ma[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020).")), to the _Upper Tuff_ unit in the upper part of the Bapang _fm._ (paleomagnetic ages, \\(\\sim\\)0.8 Ma[35](/articles/s41598-022-23206-9#ref-CR35 "Hyodo, M. et al. High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia. Proc. Natl. Acad. Sci. PNAS 108(49), 19563–19568 (2011).")). Although the new TCN dating advocates for an older age, it remains compatible, within uncertainty, with the commonly used \\(\\sim\\)1.5 Ma \\(^{40}\\)Ar/\\(^{39}\\)Ar age obtained at the same site and stratigraphic position at the base of the Bapang _fm._ [11](/articles/s41598-022-23206-9#ref-CR11 "Larick, R. et al. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. Proc. Natl. Acad. Sci. PNAS 98(9), 4866–4871 (2001)."). Conversely, this TCN age contradicts recently estimated ages of \\(\\sim\\)0.9 to 1.0 Ma for the bottom part of the Bapang _fm._ and _Grenzbank_ unit (U-Pb and ZFT[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020).")) and earlier reported age estimates for the Bapang _fm._ of \\(\\sim\\)0.8 from the northern Sangiran Ngebung site[37](/articles/s41598-022-23206-9#ref-CR37 "Falguéres, C. et al. Geochronology of early human settlements in Java: What is at stake?. Quatern. Int. 416, 5–11 (2016)."). Besides these outliers, the overall contemporaneity of most ages below the lower Bapang _fm._ can be explained by the rapid sedimentation in the shallow marine to swampy environments of the Sangiran _fm._, which likely laid the sedimentary pile in less than 100 kyr. These layers thus only represent a snapshot in the history of Javanese _H. erectus_, for which the oldest age provides a conservative estimate of the first appearance datum, _ca._ 1.8 Ma. Conversely during the deposition of the more lacustrine and fluviatile Bapang _fm._, which may span up to 1 Ma until the last appearance of _H. erectus_ in Sangiran _ca._ 0.78 Ma[35](/articles/s41598-022-23206-9#ref-CR35 "Hyodo, M. et al. High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia. Proc. Natl. Acad. Sci. PNAS 108(49), 19563–19568 (2011)."), sedimentation rates are much lower. It implies that, as a simple effect of sedimentation rates, hominin remains could be more diluted in the fast sedimenting Sangiran _fm._ (likely above 1 mm/a) than in the slow-sedimenting Bapang _fm._ (possibly as low as \\(\\sim\\)0.05 mm/a), consistent with the distribution and frequency of remains in these in the two formations. More generally, the highly variable sedimentation rates of the available archive might bias the interpretation of the taphonomic, paleo-anthropological, paleo-environmental and paleo-ecological conditions at the time of first appearance of _H. erectus_.

### Peopling Sundaland to Java

The slow drowning of the shallow Sunda shelf implies that the entire Sundaland was subaerial during most of the Pleistocene[6](/articles/s41598-022-23206-9#ref-CR6 "Sarr, A.-C. et al. Subsiding Sundaland. Geology (Boulder) 47(2), 119–122 (2019)."). In order to reconstruct the landscape at the time when _H. erectus_ reached Java, we compiled and interpolated estimates of uplift and subsidence rates[7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."), [33](/articles/s41598-022-23206-9#ref-CR33 "Husson, L. et al. Slow geodynamics and fast morphotectonics in the far East Tethys. Geochem. Geophys. Geosyst. 23(1), n/a (2022).") over the entire region (Methods). The backward integration of these rates over time allows to restore the paleo-elevation of Sundaland at large spatial scales. In addition, in order to comprehensively retrodict the physiography at the local scale and determine the migratory behavior of _H. erectus_, we model the surface processes overprint on this long-wavelength relief using LEM _goSPL_ [32](/articles/s41598-022-23206-9#ref-CR32 "Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. J. Open Source Softw.5(56), 2804 (2020).                    https://doi.org/10.21105/joss.02804                                    ."). We performed these reconstructions opting for a baseline age of 1.8 Ma, which, based on the TCN age and existing earlier dating, is most representative of the period of dispersal, but uncertainties in the ages and vertical land motions of course propagate into our physiographic reconstructions.

**Figure 3**

[![figure 3](//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41598-022-23206-9/MediaObjects/41598_2022_23206_Fig3_HTML.png)](/articles/s41598-022-23206-9/figures/3)

Migration pathways of _H. erectus_ across Sundaland during Early Pleistocene. ( **a**) Reconstructed landscape _ca._ 1.8 Ma (model _goSPL_ [32](/articles/s41598-022-23206-9#ref-CR32 "Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. J. Open Source Softw.5(56), 2804 (2020).                    https://doi.org/10.21105/joss.02804                                    .")). The underlying physiography, with overprinted modeled rivers, converts into a map of resistance to displacement that serves to compute the migration pathways, either using a least-cost path algorithm shown for the Myanmar entry point (purple), or a mechanistic movement model ( _SiMRiv_ [30](/articles/s41598-022-23206-9#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019).")) exemplified for the Vietnam entry point (orange). Blue star locates the Thailand entry point. ( **b**) Statistical analysis of the pathways of _H. erectus_ migration to Sangiran, from 3000 simulations (1000 per entry point). The kernel density estimation (kde, normalized) indicates the likelihood that hominins came across a given region (combined for the three entry points). Map on panel ( **a**, **b**). were respectively made using Paraview ( [www.paraview.org](http://www.paraview.org)) and GMT 5 ( [www.generic-mapping-tools.org](http://www.generic-mapping-tools.org))[24](/articles/s41598-022-23206-9#ref-CR24 "Wessel, P., Smith, W. H. F., Scharroo, R., Luis, J. & Wobbe, F. Generic mapping tools: Improved version released. EOS Trans. Am. Geophys. Union 94(45), 409–410.                    https://doi.org/10.1002/2013EO450001                                     (2013).").

[Full size image](/articles/s41598-022-23206-9/figures/3)

These reconstructions primarily show that Sundaland was permanently continental _ca._ 1.8 Ma (Fig. [3](/articles/s41598-022-23206-9#Fig3) a), regardless of glacio-eustatic sea level oscillations, while tectonic uplift in the Indonesian arc completed the land bridge from mainland Asia to the emerging relief of the Javanese archipelago. The elevation gently decreased from more than 1000 m in the East towards the lowlands of what can be referred to as the “Mekong isthmus” that separated the internally drained Thailand sea (now Gulf of Thailand) from the South China Sea to the North, and towards the marginal seas and alluvial plains that border the inner margins of Sumatra and Java to the Southwest. The drainage network finely dissected the surface of the continent, with a set of major rivers, radially diverging from the core of Sundaland outwards, that incised the shelf; in the South in particular, N-S trending rivers profoundly imprinted the landscape (Fig. [3](/articles/s41598-022-23206-9#Fig3) a and S.I. Fig. [1B](/articles/s41598-022-23206-9#MOESM1)). Because of their sizes (the largest computed discharge compares to that of the present-day Yangtze), these rivers might have acted as barrier to migration and could have channelized terrestrial faunas and early humans[38](/articles/s41598-022-23206-9#ref-CR38 "Roach, N. et al. Pleistocene footprints show intensive use of lake margin habitats by Homo erectus groups. Sci. Rep. 121 (2016).                    https://doi.org/10.1038/srep26374                                    .") towards the shallow marginal seas that separated the shelf from the emerging landmasses of Java, where marine conditions prevailed until Early Pleistocene (Fig. [3](/articles/s41598-022-23206-9#Fig3)). The protracted subsidence of Sundaland, stimulated by the slow deformation of the Earth interior[33](/articles/s41598-022-23206-9#ref-CR33 "Husson, L. et al. Slow geodynamics and fast morphotectonics in the far East Tethys. Geochem. Geophys. Geosyst. 23(1), n/a (2022).") changed the landscape during mid-Pleistocene[7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."). Lowlands gradually invaded most of the shelf, with only a limited set of rivers draining towards the South China, Flores, and Andaman seas, followed by ever more pervasive marine incursions over the latest interglacial stages.

Once the land bridges connecting Java to South Sundaland were opened during Early Pleistocene, _H. erectus_ reached Java, neither as seafarers nor islanders, but as continental walkers. The stratigraphic record in Java ubiquitously indicates that terrestrial faunas and _H. erectus_ dwelled as soon as the environment became continental enough to support them[13](/articles/s41598-022-23206-9#ref-CR13 "Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. J. Archaeol. Sci. 27(9), 763–769 (2000)."). Emersion was triggered by Plio-Pleistocene tectonic shortening and uplift. It started in East-Central Java -near Sangiran- and gradually propagated westward[39](/articles/s41598-022-23206-9#ref-CR39 "Simandjuntak, T. O. & Barber, A. J. Contrasting tectonic styles in the Neogene orogenic belts of Indonesia. Geol. Soc. Spec. Pub. 106(1), 185–201 (1996)."), [40](/articles/s41598-022-23206-9#ref-CR40 "Clements, B., Hall, R., Smyth, H. R. & Cottam, M. A. Thrusting of a volcanic arc; a new structural model for Java. Pet. Geosci. 15(2), 159–174 (2009).") and northward, suggesting that the Sangiran dome emerged and hosted hominins and terrestrial faunas earlier than the northern and western counterparts, for instance near the more recently occupied site of Trinil[41](/articles/s41598-022-23206-9#ref-CR41 "Joordens, J., Wesselingh, F., de Vos, J., Vonhof, H. & Kroon, D. Relevance of aquatic environments for hominins: A case study from Trinil (Java, Indonesia). J. Hum. Evol. 57(6), 656–671 (2009)."), [42](/articles/s41598-022-23206-9#ref-CR42 "Berghuis, H. et al. Hominin homelands of East Java: Revised stratigraphy and landscape reconstructions for Plio-Pleistocene Trinil. Quatern. Sci. Rev. 260, 106912 (2021).") (Fig. [1](/articles/s41598-022-23206-9#Fig1) b).

In order to develop a quantitative approach of the peopling of Sundaland and Java, we converted this contrasted physiography into a cost map of landscape complexity, that quantifies the resistance of the physiography to the displacement of species, integrating the effects of slope, distance to water bodies (rivers and shores), and river size (Methods). Both least-cost paths and random (Markovian) walks eventually lead hominins from mainland Asia to Java, but with extremely variable routes and travelled distances (Fig. [3](/articles/s41598-022-23206-9#Fig3) a). Depending on the entry point (Myanmar, Thailand or Vietnam), least-cost paths show two preferential routes, either following the western coastline of the epicontinental Thailand sea or opting for an eastern path through the “Mekong isthmus”.

Markovian walks instead are more exploratory (see animated examples in [Supplementary Information](/articles/s41598-022-23206-9#MOESM1)), and show that the entire region was made available to hominins, albeit with a variable probability. The likelihood that the species came across a given location (revealed by the kernel density estimation of all 3000 modeled random walks, Fig. [3](/articles/s41598-022-23206-9#Fig3) b) unsurprisingly reaches the highest values in the northern part of the region, near their entry points. En route to Sundaland, while the leasts cost paths includes a western trajectory through the Malay peninsula, the probabilistic approach instead indicates that this western option is unlikely due to the rugged topography of the Malay peninsula. Instead, the “Mekong isthmus” channelized them between the Thailand and South China seas. In these lowlands, landscape biases are low (absence of relief, favorable drainage network, see [Methods](/articles/s41598-022-23206-9#Sec6)), which further reinforces the likelihood of hominin occurence. Likewise, the likelihood is slightly higher in the lowlands of eastern Sumatra than in more mountainous regions of Sundaland, and in the North, in the flat Chao Praya basin and Khorat plateau of Thailand (Fig. [3](/articles/s41598-022-23206-9#Fig3) b). While these results indicate that peopling Sundaland was ubiquitously possible, the likelihood that hominins joined the rims of Sundaland is however relatively low. This result, seemingly at odds with the Javanese yield of fossil remains, could reveal that our reconstructions and parametrizations are inadequate to the point of being misleading, and that better accounting for habitat suitability, using species distribution modeling in particular, would yield alternative and more compatible solutions. Alternatively, this incongruence can be simply explained by the inconvenient concealment of fossil bearing sedimentary layers in subsiding, fast sedimenting basins (as in Thailand) or below sea level (as in the “Mekong isthmus”).

How long did the journey take? While least-cost paths are \\(\\sim\\)6,000 km long on average from mainland Asia to Java (Fig. [4](/articles/s41598-022-23206-9#Fig4) a), the mean travelled distances are \\(\\sim\\)50 times longer for Markovian walks (327\\(\\times 10^3\\) km), regardless of the north entry point, and almost always in excess of 100\\(\\times 10^3\\) km. The corresponding time span depends on the migration speed. Estimates are available for more recent palaeolithic hominins, ranging from 1 to 10 km/yr[43](#ref-CR43 "Fort, J., Pujol, T. & Cavalli-Sforza, L. Palaeolithic populations and waves of advance. Camb. Archaeol. J. 14, 53–61.                    https://doi.org/10.1017/S0959774304000046                                     (2004)."), [44](#ref-CR44 "Hamilton, M. & Buchanan, B. Spatial gradients in Clovis-age radiocarbon dates across North America suggest rapid colonization from the north. Proc. Natl. Acad. Sci. USA 104, 15625–30.                    https://doi.org/10.1073/pnas.0704215104                                     (2007)."), [45](/articles/s41598-022-23206-9#ref-CR45 "Hazelwood, L. & Steele, J. Spatial dynamics of human dispersals: Constraints on modelling and archaeological validation. J. Archaeol. Sci. 31, 669–679.                    https://doi.org/10.1016/j.jas.2003.11.009                                     (2004)."). At similar speeds, the least-cost paths convert into an optimal travel time of 0.6 to 6 kyr, which is thus the minimal time to cross Sundaland. Markovian walks imply much longer durations before hominins reached Java, between 25-40 kyr at fast moving rates, and 250-400 kyr at slow moving rates (Fig. [4](/articles/s41598-022-23206-9#Fig4) b). These relatively long delays at average or slow moving rates are difficult to reconcile with the early arrival to Java _ca._ 1.8 Ma, following departure from Georgia or China only a few 100 kyr before. We suggest that because these characteristic travel times scale with the characteristic time scale of changes of the physical environment, external forcings impelled hominins to migrate and boosted their displacement. Climatic variations notoriously impact the habitats of hominins[46](/articles/s41598-022-23206-9#ref-CR46 "Bae, C., Li, F., Liuling, C., Wang, W. & Hanlie, H. Hominin distribution and density patterns in pleistocene China: Climatic influences. Palaeogeogr. Palaeoclimatol. Palaeoecol. 512 (2018).                    https://doi.org/10.1016/j.palaeo.2018.03.015                                    ."), [47](/articles/s41598-022-23206-9#ref-CR47 "Timmermann, A. et al. Climate effects on archaic human habitats and species successions. Nature 604, 1–7.                    https://doi.org/10.1038/s41586-022-04600-9                                     (2022)."), but here we suggest that the fast physiographic changes of geodynamic origin and environmental stochasticity could have also prompted the dispersal of _H. erectus_.

**Figure 4**

[![figure 4](//media.springernature.com/lw685/springer-static/image/art%3A10.1038%2Fs41598-022-23206-9/MediaObjects/41598_2022_23206_Fig4_HTML.png)](/articles/s41598-022-23206-9/figures/4)

Travelled distance and time of _H. erectus_ migration to Sangiran, statistical analysis (3000 simulations, 1000 per entry point - Myanmar, Thailand, and Vietnam). ( **a**) Migration distance from the least-cost path (LCP) analysis and successful mechanistic movement realizations (Markovian random walk) for the three entry points, and all combined. ( **b**) Travel times, for three dispersal speeds (slow: 1 km/yr[43](/articles/s41598-022-23206-9#ref-CR43 "Fort, J., Pujol, T. & Cavalli-Sforza, L. Palaeolithic populations and waves of advance. Camb. Archaeol. J. 14, 53–61.                    https://doi.org/10.1017/S0959774304000046                                     (2004)."), medium: 5 km/year[44](/articles/s41598-022-23206-9#ref-CR44 "Hamilton, M. & Buchanan, B. Spatial gradients in Clovis-age radiocarbon dates across North America suggest rapid colonization from the north. Proc. Natl. Acad. Sci. USA 104, 15625–30.                    https://doi.org/10.1073/pnas.0704215104                                     (2007)."), and fast: 10 km/yr[45](/articles/s41598-022-23206-9#ref-CR45 "Hazelwood, L. & Steele, J. Spatial dynamics of human dispersals: Constraints on modelling and archaeological validation. J. Archaeol. Sci. 31, 669–679.                    https://doi.org/10.1016/j.jas.2003.11.009                                     (2004).")).

[Full size image](/articles/s41598-022-23206-9/figures/4)

## Discussion

Eugène Dubois[1](/articles/s41598-022-23206-9#ref-CR1 "Dubois, E. On Pithecanthropus Erectus: a transitional form between man and the apes. J. Anthropol. Inst. G. B. Irel. 25, 240–255 (1896).") wondered early on whether his revolutionary findings in Java, as opposed to his unfruitful attempts in Sumatra, were due to the habitats of early humans or to their preservation and outcrop. This question is in fact a general one, on which our joint analysis of the chronology, physical environment, and dispersal trajectories brings a new light on: Hominin finds in Java mark the onset of continental conditions there _ca._ 1.8 Ma rather than the timing of their migration across SE Asia. The high probability that earlier groups of hominins thrived in the nearby Sundaland and were ready to disperse in Java, only waiting for more continental conditions in the Javanese archipelago, suggests that the immediate ancestors of Javanese _H. erectus_ could be abundantly buried in the sediments of the nearby Sunda shelf more than exposed in the Indonesian arc. During Early Pleistocene, _H. erectus_ likely clustered in lowlands during the peopling of Sundaland, in particular in the “Mekong isthmus” that connected mainland Asia to the Sunda shelf, or in a few hotspots in mainland Asia. These sites are currently buried underneath sediments or seawater, or both, which explains that fossil finds are few elsewhere than Java.

This analysis is based on a large number of set of random walks across the region. While it advantageously indicates the probability distribution of the trajectories, distances and tempos, a corollary limitation arises from the unaccounted possibility that the peopling of SE Asia could have been the case of a limited number of groups only. It would imply that a limited number of stochastic events, including the random selection of low cost paths, could have been influential on the final peopling of the region.

While the impact of physiography on hominin evolution has been identified[48](/articles/s41598-022-23206-9#ref-CR48 "Bailey, G. N., Reynolds, S. C. & King, G. C. Landscapes of human evolution: Models and methods of tectonic geomorphology and the reconstruction of hominin landscapes. J. Hum. Evol. 60(3), 257–280 (2011)."), this case study of the iconic Javanese _H. erectus_ emphasizes the need to tie a quantified reconstruction of the past physiography to a robust time frame before attempting to unravel the migration trajectories. This novel approach sheds new lights on the dispersal of the genus _Homo_, by recasting hominins within their regional environmental framework. First, our results emphasize the role of the physiography -lowlands, rivers, and reliefs in particular- in controlling hominin habitats. Second, and perhaps more importantly, they highlight the impact of the transience of the environmental factors, by demonstrating that hominin dispersal operates at the same characteristic time scale than that of physiographic changes, which could have spurred their displacement.

These results open new perspectives to untangle the physical and behavioral capacities of _H. erectus_ within their changing physical environments. We find that _H. erectus_ migrated across Sundaland at remarkably fast rates (>10 km/yr) with respect to their prior journey across Eurasia, and to the displacement rates of other lineages of the genus _Homo_. We suggest that the quickly changing physiography of Sundaland could have prompted their fast migration, which implies that extrinsic (environmental) forcings prevailed over intrinsic factors (either cultural or demographic)[10](/articles/s41598-022-23206-9#ref-CR10 "Carotenuto, F. et al. Venturing out safely: The biogeography of Homo erectus dispersal out of Africa. J. Hum. Evol. 95, 1–12 (2016)."). This proposition invites to go beyond the current study and comprehensively investigate the joint evolution of the entire ecological and physical -climatic and physiographic- environments. Physiographic changes in SE Asia triggered a chain of modifications that impacted the behavior of _H. erectus_, including the regional climate and its feedback relationships with the vegetation cover[49](/articles/s41598-022-23206-9#ref-CR49 "Sarr, A., Sepulchre, P. & Husson, L. Impact of the Sunda Shelf on the Climate of the Maritime Continent. J. Geophys. Res. Atmos. 124(5), 2574–2588 (2019)."), and ultimately on the foodchain on which hunter-gatherers relied[50](/articles/s41598-022-23206-9#ref-CR50 "Louys, J. & Roberts, P. Environmental drivers of megafauna and hominin extinction in Southeast Asia. Nature 586(7829), 402–406 (2020)."). In order to more thoroughly address the impact of this changes on habitat suitability, species distribution modeling would be a natural avenue to more precisely reconstruct the evolution of early humans[47](/articles/s41598-022-23206-9#ref-CR47 "Timmermann, A. et al. Climate effects on archaic human habitats and species successions. Nature 604, 1–7.                    https://doi.org/10.1038/s41586-022-04600-9                                     (2022)."), [51](/articles/s41598-022-23206-9#ref-CR51 "Raia, P. et al. Past extinctions of homo species coincided with increased vulnerability to climatic change. One Earth 3(4), 480–490 (2020).").

At a more general scale, our TCN ages obtained for the first appearance datum of _H. erectus_ in Sangiran advocate for an early peopling of Sundaland by _H. erectus_, contemporary with their Chinese[26](/articles/s41598-022-23206-9#ref-CR26 "Luo, L. et al. The first radiometric age by isochron 26Al/10Be burial dating for the Early Pleistocene Yuanmou hominin site, southern China. Quat. Geochronol. 55, 101022.                    https://doi.org/10.1016/j.quageo.2019.101022                                     (2019)."), [52](/articles/s41598-022-23206-9#ref-CR52 "Zhu, Z. et al. Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago. Nature 559(7715), 608–612 (2018).") and Georgian[53](/articles/s41598-022-23206-9#ref-CR53 "Gabunia, L. et al. Earliest Pleistocene hominid cranial remains from Dmanisi, Republic of Georgia: Taxonomy, geological setting, and age. Science 288, 1019–1025.                    https://doi.org/10.1126/science.288.5468.1019                                     (2000)."), [54](/articles/s41598-022-23206-9#ref-CR54 "Lordkipanidze, D. et al. A complete skull from Dmanisi, Georgia, and the evolutionary biology of early homo. Science 342(6156), 326–331 (2013).") counterparts, whose endocranial capacity was much smaller[54](/articles/s41598-022-23206-9#ref-CR54 "Lordkipanidze, D. et al. A complete skull from Dmanisi, Georgia, and the evolutionary biology of early homo. Science 342(6156), 326–331 (2013).") than specimens from Sangiran[55](/articles/s41598-022-23206-9#ref-CR55 "Baba, H. et al. Homo erectus calvarium from the pleistocene of java. Sci. (Am. Assoc. Adv. Sci.) 299 (5611), 1384–1388 (2003) ."). These results reignite the long-standing controversy over the origin and dispersal pathways of archaic humans[27](/articles/s41598-022-23206-9#ref-CR27 "Zaim, Y. et al. New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J. Hum. Evol.61(4), 363–376 (2011)."), [52](/articles/s41598-022-23206-9#ref-CR52 "Zhu, Z. et al. Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago. Nature 559(7715), 608–612 (2018)."), [56](/articles/s41598-022-23206-9#ref-CR56 "Ciochon, R. L. & Bettis, E. A. III. Asian Homo erectus converges in time. Nature 458(7235), 153–154 (2009).") and invites a reexamination of the out-of-Africa paradigm, which provides a global roadmap for the dispersal of the genus _Homo_, but which one-way direction may be questioned[12](/articles/s41598-022-23206-9#ref-CR12 "Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G. & Suprijo, A. Age of the earliest known hominids in Java, Indonesia. Science 263(5150), 1118–1121 (1994)."), [57](#ref-CR57 "Dennell, R. & Roebroeks, W. An Asian perspective on early human dispersal from Africa. Nature 438(7071), 1099–1104 (2005)."), [58](#ref-CR58 "Martinon-Torres, M. et al. Dental evidence on the hominin dispersals during the Pleistocene. Proc. Natl. Acad. Sci. PNAS 104(33), 13279–13282 (2007)."), [59](#ref-CR59 "Wood, B. Did early Homo migrate “out of’’ or “in to’’ Africa?. Proc. Natl. Acad. Sci. PNAS 108(26), 10375–10376 (2011)."), [60](/articles/s41598-022-23206-9#ref-CR60 "Shen, G. et al. Isochron 26Al/10Be burial dating of Xihoudu: Evidence for the earliest human settlement in northern China. Anthropologie 124, 102790.                    https://doi.org/10.1016/j.anthro.2020.102790                                     (2020).").

## Methods

### Terrestrial Cosmogenic Nuclide dating (TCN)

The Bukuran creek outcrop (Fig. [1](/articles/s41598-022-23206-9#Fig1), red star, \\(110^\\circ 51'18'' E, 7^\\circ 27'58.3'' S, 99 m.a.s.l.\\)) was sampled in July 2018. The 6 m high vertical section (Fig. [2](/articles/s41598-022-23206-9#Fig2)) is continuously rejuvenated by lateral erosion by the Bukuran stream, thus offering ideal conditions for TCN application. \\(^{26}\\)Al/\\(^{10}\\)Be burial dating[16](/articles/s41598-022-23206-9#ref-CR16 "Granger, D. E. & Muzikar, P. F. Dating sediment burial with in situ-produced cosmogenic nuclides: Theory, techniques, and limitations. Earth Planet. Sci. Lett. 188(1), 269–281 (2001)."), [18](#ref-CR18 "Pappu, S. et al. Early Pleistocene presence of Acheulian Hominins in South India. Science 331(6024), 1596–1599 (2011)."), [19](#ref-CR19 "Lebatard, A.-E. et al. Dating the Homo erectus bearing travertine from Kocabaş (Denizli, Turkey) at at least 1.1 Ma. Earth Planet. Sci. Lett.390, 8–18 (2014)."), [20](/articles/s41598-022-23206-9#ref-CR20 "Lebatard, A.-E., Bourlès, D. L. & Braucher, R. Absolute dating of an Early Paleolithic site in Western Africa based on the radioactive decay of in situ-produced 10Be and 26Al. Nucl. Instrum. Methods Phys. Res. Sect. B 456, 169–179 (2019).") is based on the relative decay of \\(^{26}\\)Al and \\(^{10}\\)Be cosmogenic nuclides produced _in situ_ in quartz (SiO\\(\_2\\)) minerals upon exposure to cosmic rays. Given the half-lives of \\(^{10}\\)Be (1.387±0.012 Ma[61](/articles/s41598-022-23206-9#ref-CR61 "Chmeleff, J., von Blanckenburg, F., Kossert, K. & Jakob, D. Determination of the 10Be half-life by multicollector ICP-MS and liquid scintillation counting. Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms268(2), 192–199 (2010)."), [62](/articles/s41598-022-23206-9#ref-CR62 "Korschinek, G. et al. A new value for the half-life of 10Be by Heavy-Ion Elastic Recoil Detection and liquid scintillation counting. Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms 268(2), 187–191 (2010) .")) and \\(^{26}\\)Al (0.705±0.024 Ma[63](/articles/s41598-022-23206-9#ref-CR63 "Nishiizumi, K. Preparation of 26Al AMS standards. Nucl. Inst. and Meth. in Phys. Res. 223-224, 388–392 (2004)."), [64](/articles/s41598-022-23206-9#ref-CR64 "Norris, T. L., Gancarz, A. J., Rokop, D. J. & Thomas, K. W. Half-life of 26Al. J. Geophys. Res. Solid Earth 88(S01), B331–B333 (1983).")), it is applicable over a time frame of 200 ka to \\(\\sim\\)6 Ma[16](/articles/s41598-022-23206-9#ref-CR16 "Granger, D. E. & Muzikar, P. F. Dating sediment burial with in situ-produced cosmogenic nuclides: Theory, techniques, and limitations. Earth Planet. Sci. Lett. 188(1), 269–281 (2001)."). During burial, the attenuation of secondary cosmic ray particles with depth is marked by an exponential depth decay of cosmogenic isotope concentration, while posterior erosion flattens out the depth dependence.

The top of the sequence, down to \\(\\sim\\)1.5 m depth, is composed of a yellowish tuff and quartz poor siltstone. A light pink and yellowish tuffaceous sand, also quartz poor, follows down to 3 m below. Between 3 m and 4.6 m, a bedded sand and gravel layer yields a higher concentration of quartz grains. From 4.6 m to 5.6 m, an indurated conglomerate with a higher concentration of quartz is observed, followed by a layer of dark clays, down to the stream bed. 18 samples have been collected at 15 different depths from the surface (SAN18-1) down to the stream bed (SAN18-18), the deepest samples (SAN-15 to SAN-18) being precisely extracted from the _Grenzbank_ unit, as identified in earlier studies[15](/articles/s41598-022-23206-9#ref-CR15 "Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367(6474), 210–214 (2020)."), [27](/articles/s41598-022-23206-9#ref-CR27 "Zaim, Y. et al. New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). J. Hum. Evol.61(4), 363–376 (2011).").

Measured \\(^{10}\\)Be and \\(^{26}\\)Al concentrations and ratios are depth independent and in agreement (see raw and statistical values in S.I. Table 1). This is explained by low and uniform inherited concentrations, and by long-lasting denudation at a fast enough rate to dampen the depth dependent exposure of quartz grains to cosmic rays. By modeling radioactive decay, we estimate the burial age, pre-burial and post-burial denudation rates[18](/articles/s41598-022-23206-9#ref-CR18 "Pappu, S. et al. Early Pleistocene presence of Acheulian Hominins in South India. Science 331(6024), 1596–1599 (2011)."), [65](/articles/s41598-022-23206-9#ref-CR65 "Braucher, R., Merchel, S., Borgomano, J. & Bourlès, D. Production of cosmogenic radionuclides at great depth: A multi element approach. Earth Planet. Sci. Lett. 309(1), 1–9 (2011)."). We assume that all samples were buried deep enough to be fully shielded from cosmic rays, but account for post-burial production of nuclides since deposition (burial _with_ post-production). We also model pre-burial \\(^{26}\\)Al/\\(^{10}\\)Be ratios (assuming that samples were at steady state before deposition). As all samples underwent the same burial-exhumation history, we inverted for a common burial time and post-production for all samples at once, while leaving individual inheritance independent for each sample. This yields a burial age of 1.78±0.20 Ma ( S.I. Table 1). Denudation rates of the source rocks range from 21 to 80 m/Ma, while post-deposition denudation rates range from 334 to 608 m/Ma with a best fit value of 436 m/Ma. Within the topmost meter of the sequence, the fraction of post-burial \\(^{10}\\)Be and \\(^{26}\\)Al respectively ranges from 14 to 28% and from 26 to 47%, but then decreases at greater depths to less than 10% for \\(^{10}\\)Be and less than 20%, mostly, for \\(^{26}\\)Al. Given the inferred high denudation rates, the corresponding time span is on the order of 4000 years.

For reference, we also consider the end-member scenario, wherein the upper part of the profile has been either quasi-instantaneously truncated by several tens of meters (burial _without_ post-production), in order to determine the minimal age. Individual minimum burial durations ( _i.e._ treating each sample independently) range from 0.58±0.20 Ma to 2.25±1.04 Ma, and pre-burial denudation rates range from 23 to 101 m/Ma (S.I. Table 2). \\(\\chi ^2\\) test isolated three outliers (SAN18-2, SAN18-4 and SAN18-16). The remaining 13 samples yield a minimum weighted mean burial duration of 1.45±0.10 Ma.

Chemical sample preparation and \\(^{10}\\)Be and \\(^{26}\\)Al measurements were carried out at the French Accelerator Mass Spectrometry national facility, ASTER (CEREGE, Aix-en-Provence) following the standard procedure[19](/articles/s41598-022-23206-9#ref-CR19 "Lebatard, A.-E. et al. Dating the Homo erectus bearing travertine from Kocabaş (Denizli, Turkey) at at least 1.1 Ma. Earth Planet. Sci. Lett.390, 8–18 (2014)."). A challenging aspect for TCN application at the Bukuran creek site is the low quartz content of the fluvio-volcanic sediments. As the amount of quartz mineral is notoriously poor, a mass of 0.5 to 2 kg of sediment per sample was collected, depending on the lithology (S.I. Table 3). Acid attacks were repeated until the maximum amount of pure SiO\\(\_2\\) was retrieved, within the availability limit of materials. All samples (besides SAN18-7 to SAN18-9) yielded enough quartz to perform suitable measurements. After purification, the quartz content was dissolved in HF 48% after addition of 150 \\(\\mu l\\) of a 3025±9 ppm \\(^9\\)Be solution. Natural Al content was determined by ICP-OES using an ICAP6500 from Thermo. BeO and Al\\(\_2\\)O\\(\_3\\) were respectively mixed with niobium and silver powders prior to measurements. Beryllium data were directly calibrated against the STD11 standard[66](/articles/s41598-022-23206-9#ref-CR66 "Braucher, R. et al. Preparation of ASTER in-house 10Be/9Be standard solutions. Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms361, 335–340 (2015) .") with a \\(^{10}\\)Be/\\(^9\\)Be ratio of (1.191±0.013) x 10\\(^{-11}\\). Aluminum measurements were performed against in-house standard SM-Al-11, with a \\(^{26}\\)Al/\\(^{27}\\)Al ratio of (7.401±0.064) x 10\\(^{-12}\\), previously cross-calibrated against the primary standards certified by a round-robin exercise[67](/articles/s41598-022-23206-9#ref-CR67 "Merchel, S. & Bremser, W. First international 26Al interlaboratory comparison—Part I. Nucl. Instrum. Methods Phys. Res. 223–224, 393–400 (2004)."), [68](/articles/s41598-022-23206-9#ref-CR68 "Arnold, M. et al. The French accelerator mass spectrometry facility ASTER: Improved performance and developments. Nucl. Instrum. Methods Phys. Res. 268(11), 1954–1959 (2010)."). The successive measurement batches are characterized by stable, high level \\(^9\\)Be and \\(^{27}\\)Al currents during AMS measurements (S.I. Table 3) that attest for their high reliability (despite low \\(^{10}\\)Be and \\(^{26}\\)Al counts at the AMS receptors). Analytical uncertainties (reported as 1\\(\\sigma\\)) include uncertainties associated with AMS counting statistics, AMS external error (0.5% for \\(^{10}\\)Be), chemical blank measurement, and \\(^{26}\\)Al, \\(^{27}\\)Al measurements. Measurements of chemically processed blank yield ratios on the order of (2.0±0.75) x 10\\(^{-15}\\) for \\(^{10}\\)Be and (2.0±2.0) x 10\\(^{-15}\\) for \\(^{26}\\)Al. A sea level high latitude spallation production rate of 4.02±0.32 \\(at.g^{-1}.a^{-1}\\)[69](/articles/s41598-022-23206-9#ref-CR69 "Borchers, B. et al. Geological calibration of spallation production rates in the CRONUS-Earth project. Quat. Geochronol. 31, 188–198 (2016).") was used and scaled using Stone’s polynomials[70](/articles/s41598-022-23206-9#ref-CR70 "Stone, J. O. Air pressure and cosmogenic isotope production. J. Geophys. Res. Solid Earth 105(B10), 23753–23759 (2000)."). The \\(^{26}\\)Al/\\(^{10}\\)Be production ratio induced by the standardization used at ASTER is 6.61±0.50.

### Sundaland physical landscape reconstruction _ca._ 1.8 Ma

In order to reconstruct the landscape at 1.8 Ma, we first compiled rates of vertical land motions inferred from geomorphological indicators, stratigraphic, and seismic data (S.I. Fig. [1a](/articles/s41598-022-23206-9#MOESM1) adapted from[6](/articles/s41598-022-23206-9#ref-CR6 "Sarr, A.-C. et al. Subsiding Sundaland. Geology (Boulder) 47(2), 119–122 (2019)."), [7](/articles/s41598-022-23206-9#ref-CR7 "Salles, T. et al. Quaternary landscape dynamics boosted species dispersal across Southeast Asia. Commun. Earth Environ. 2(1), 1–12 (2021)."), [33](/articles/s41598-022-23206-9#ref-CR33 "Husson, L. et al. Slow geodynamics and fast morphotectonics in the far East Tethys. Geochem. Geophys. Geosyst. 23(1), n/a (2022).")). This dataset provides discrete informations that we interpolated in order to obtain a continuous map of uplift rates over the region, which backward integration in time provides a paleo-elevation map. To reconstruct a realistic landscape at 1.8 Ma, we ran the Landscape Evolution Model (LEM) _goSPL_ [32](/articles/s41598-022-23206-9#ref-CR32 "Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. J. Open Source Softw.5(56), 2804 (2020).                    https://doi.org/10.21105/joss.02804                                    .") over a 10,000 years period starting from an adjusted paleo-elevation surface (1 km resolution) derived from the uplift map and past absolute sea level, set to -30 m during Early Pleistocene sea level highstands[71](/articles/s41598-022-23206-9#ref-CR71 "Bintanja, R. & van de Wal, R. S. W. North American ice-sheet dynamics and the onset of 100,000-year glacial cycles. Nature 454, 869–872.                    https://doi.org/10.1038/nature07158                                     (2008)."). Continuity of mass is the main equation of the model, and is expressed by:

$$\\begin{aligned} \\frac{\\partial z}{\\partial t} = U + \\kappa \\nabla ^2 z + \\epsilon P^d (PA)^m \\nabla z^n \\end{aligned}$$

(1)


where _z_ is surface elevation (m), _t_ is time (yr), _U_ is vertical land motion (\\(\\text {m}/\\text {yr}\\)) and \\(\\kappa\\) the diffusion coefficient for soil creep (set to \\(5\\times 10^{-3}\\) m\\(^2\\)/yr). The last term in the right-hand side of Eq. ( [1](/articles/s41598-022-23206-9#Equ1)) represents fluvial processes based on a stream power law with erodibility \\(\\epsilon\\) (set to \\(4\\times 10^{-6}\\) m\\(^{-1}\\)yr\\(^{-1}\\)), dimensionless constants _m_ and _n_ (empirically set to 0.5 and 1) and water flux _PA_ combining upstream watershed area _A_ and precipitation _P_. The stream power law here incorporates the effect of local mean annual precipitation rate (obtained from paleoclimate model _HadCM3BL-M2.1aD_ [34](/articles/s41598-022-23206-9#ref-CR34 "Valdes, P., Scotese, C. & Lunt, D. Deep ocean temperatures through time. Climate Past 17, 1483–1506.                    https://doi.org/10.5194/cp-17-1483-2021                                     (2021).")), _d_ being a positive exponent (set to 0.42), fingerprinted in the paleo-landscape and paleo-drainage maps (see resulting landscape after 10 kyr of concurrent action of riverine and hillslope processes in S.I. Fig. [1b](/articles/s41598-022-23206-9#MOESM1)). Both processes erode, transport and deposit sediments with maximum erosion (>500 m) along the Barisan Mountains in Sumatra and deposition (up to 400 m) in several endhoreic lakes and basins across Borneo and East Malay Peninsula (S.I. Fig. [2a](/articles/s41598-022-23206-9#MOESM1)).

### Resistance to species displacement from regional physiography

To compute the dispersal of _H. erectus_ in Sundaland and its migration to Sangiran, we extracted from the landscape evolution model three physiographic features known to influence early human dispersal[72](#ref-CR72 "Field, J. & Mirazon Lahr, M. Assessment of the Southern Dispersal: GIS-Based Analyses of Potential Routes at Oxygen Isotopic Stage 4. J. World Prehist. 19, 1–45 (2005).                    https://doi.org/10.1007/s10963-005-9000-6                                    ."), [73](#ref-CR73 "Howey, M. Multiple pathways across past landscapes: Circuit theory as a complementary geospatial method to least cost path for modeling past movement. J. Archaeol. Sci. 38, 2523–2535.                    https://doi.org/10.1016/j.jas.2011.03.024                                     (2011)."), [74](#ref-CR74 "Tassi, F. et al. Early modern human dispersal from Africa: Genomic evidence for multiple waves of migration. Investig. Genet. 6, 13.                    https://doi.org/10.1186/s13323-015-0030-2                                     (2015)."), [75](/articles/s41598-022-23206-9#ref-CR75 "Kealy, S., Louys, J. & O’Connor, S. Least-cost pathway models indicate northern human dispersal from Sunda to Sahul. J. Hum. Evol. 125, 59–70.                    https://doi.org/10.1016/j.jhevol.2018.10.003                                     (2018)."). These features (S.I. Fig. [2](/articles/s41598-022-23206-9#MOESM1)) are then converted into cost values at a \\(\\sim\\)1 km\\(^2\\) resolution, with high costs corresponding to resistant regions impeding movement. First, we consider endorheic lakes and oceans as absolute dispersal barriers (normalized cost set to 1.0) and assume that major rivers (> 5.5\\(\\times\\)10\\(^3\\) m\\(^3\\)/s - approximately the Niger river flow rates) represent strong barriers to crossing[72](/articles/s41598-022-23206-9#ref-CR72 "Field, J. & Mirazon Lahr, M. Assessment of the Southern Dispersal: GIS-Based Analyses of Potential Routes at Oxygen Isotopic Stage 4. J. World Prehist. 19, 1–45 (2005).                    https://doi.org/10.1007/s10963-005-9000-6                                    ."). From our simulation, the largest river flow rate reaches 30.5\\(\\times\\)10\\(^3\\) m\\(^3\\)/s, one sixth of the current discharge of the Amazon river, comparable to the discharge of the Yangtze river. As opposed to earlier studies[72](/articles/s41598-022-23206-9#ref-CR72 "Field, J. & Mirazon Lahr, M. Assessment of the Southern Dispersal: GIS-Based Analyses of Potential Routes at Oxygen Isotopic Stage 4. J. World Prehist. 19, 1–45 (2005).                    https://doi.org/10.1007/s10963-005-9000-6                                    ."), we do not deem these major rivers as fully impermeable in the cost calculation, and let their normalized values to vary linearly with flow rates from 0.7 to 0.95. Second, we estimate a riparian buffer zone by computing the distance to rivers over the entire region. In addition to rivers, we also set maritime and lacustrine coastlines as preferential pathways[38](/articles/s41598-022-23206-9#ref-CR38 "Roach, N. et al. Pleistocene footprints show intensive use of lake margin habitats by Homo erectus groups. Sci. Rep. 121 (2016).                    https://doi.org/10.1038/srep26374                                    .") and add these distances in our cost calculation. Computed distances are then converted into an exponential series[75](/articles/s41598-022-23206-9#ref-CR75 "Kealy, S., Louys, J. & O’Connor, S. Least-cost pathway models indicate northern human dispersal from Sunda to Sahul. J. Hum. Evol. 125, 59–70.                    https://doi.org/10.1016/j.jhevol.2018.10.003                                     (2018)."), to account for the exponentially increasing difficulty of travel as _H. erectus_ moves away from freshwaters[72](/articles/s41598-022-23206-9#ref-CR72 "Field, J. & Mirazon Lahr, M. Assessment of the Southern Dispersal: GIS-Based Analyses of Potential Routes at Oxygen Isotopic Stage 4. J. World Prehist. 19, 1–45 (2005).                    https://doi.org/10.1007/s10963-005-9000-6                                    ."). Finally, we account for topographic complexity by calculating the local slope Slopes below 0.5\\(^{\\circ }\\) are considered costless, and the normalized cost increases linearly between 0.1 and 0.6 from 0.5\\(^{\\circ }\\) to 5\\(^{\\circ }\\) and set to 0.8 above 5\\(^{\\circ }\\). Combining individual cost from each variable, we obtain a final normalized cost map (S.I. Fig. [1c](/articles/s41598-022-23206-9#MOESM1)) that favour routes along riparian areas and coastal plains and avoid hilly and mountainous regions. Assuming foraging time and energy expenditure minimization, this cost map is then used to compute the dispersal of _H. erectus_ across South East Asia and colonization of Sundaland.

We then defined three entry points, corresponding to western, central and eastern routes, at places with low resistance to displacement (riparian areas with low topographic complexity). First, from its migration out of Africa via the Levantine corridor, traces of a western route can be found in Riwat, Pakistan[76](/articles/s41598-022-23206-9#ref-CR76 "Dennell, R. W., Rendell, H. M. & Hailwood, E. Late pliocene artefacts from northern Pakistan. Curr. Anthropol. 29(3), 495–498 (1988)."), which could lead to the Himalayan foreland, Ganges-Brahmaputra rivers, down to the North of the Irrawaddy delta in Myanmar. Second, hominin fossil remains in south China near Yuanmou[77](/articles/s41598-022-23206-9#ref-CR77 "Zhu, R. et al. Early evidence of the genus homo in east asia. J. Hum. Evol. 55(6), 1075–1085 (2008)."), that we found contemporaneous to _H. erectus_ from Sangiran, set an eastern route, hypothesizing that the Mekong and Red rivers acted as barriers and channelized early humans to the coast south of the Red river in Vietnam. Last, an intermediate northern route upstream of the Chao Phraya plain in Thailand, hypothesizes that _H. erectus_ either crossed the Shan plateau in Myanmar or the Yunnan-Guizhou plateau, bounded by the Salween and Mekong rivers on its western and eastern sides.

### Assuming _H. erectus_ omniscience: least-cost paths

A convenient theoretical approach to predict movement of species through a landscape relies on a least-cost path (LCP) analysis[75](/articles/s41598-022-23206-9#ref-CR75 "Kealy, S., Louys, J. & O’Connor, S. Least-cost pathway models indicate northern human dispersal from Sunda to Sahul. J. Hum. Evol. 125, 59–70.                    https://doi.org/10.1016/j.jhevol.2018.10.003                                     (2018)."), [78](/articles/s41598-022-23206-9#ref-CR78 "Gowen, K. M. & de Smet, T. S. Testing least cost path (LCP) models for travel time and kilocalorie expenditure: Implications for landscape genomics. PLoS ONE 15(9), 1–20.                    https://doi.org/10.1371/journal.pone.0239387                                     (2020)."). Here, LCP are calculated using the Dijkstra’s algorithm accounting for diagonal connectivity between cells and implemented using the scikit-image library[79](/articles/s41598-022-23206-9#ref-CR79 "Walt, S. et al. scikit-image: Image processing in Python. PeerJ 2, e453.                    https://doi.org/10.7287/peerj.preprints.336v2                                     (2014)."). LCPs not only require the positions of the entry points but also the location of the ending point (here set to Sangiran). Both the traveled distance and landscape resistance (S.I. Fig. [1c](/articles/s41598-022-23206-9#MOESM1)) determine the most parsimonious route connecting these endpoints. While stochasticity imposes to consider this route as a possible option, LCP otherwise implicitly rests on the unrealistic assumption that the species had a destination, a compass and a roadmap. LCPs thus give a reference for the minimum travelled distance across Sundaland. Estimated walked distances for all 3 cases are comparable (5545, 5620, and 7050 km, respectively for the Myanmar, Thailand, and Vietnam entry points). The western and central routes merge after \\(\\sim\\)1500 km downstream of their entry points (S.I. Fig. [3](/articles/s41598-022-23206-9#MOESM1)), while the eastern route merges with them South of the endorheic Thailand sea, between the Malay peninsula and the “Mekong isthmus” that connected South Vietnam to the Sunda shelf. As expected from the cost surface, LCP follows preferentially alluvial plains and coastal regions. LCP occasionally significant detours from the shortest distance path, specifically where large rivers hamper dispersal and force migration across upstream smaller tributaries of lower discharge.

### Mechanistic movements conditioned by landscape heterogeneity: Markovian walks

Migration models assuming a destination ( _e.g._, LCP) would rarely reflect the actual movements of species across a heterogeneous landscape, which impairs their predictive capacity. In reality, most species progress without a planned destination during their lifespan[80](/articles/s41598-022-23206-9#ref-CR80 "Mueller, T. & Fagan, W. Search and navigation in dynamic environments—From individual behaviors to population distributions. Oikos 117, 654–664.                    https://doi.org/10.1111/j.0030-1299.2008.16291.x                                     (2008).") and have no memory of longterm prior displacements: their movements are only determined from local information[81](/articles/s41598-022-23206-9#ref-CR81 "Bastille-Rousseau, G., Douglas-Hamilton, I., Blake, S., Northrup, J. & Wittemyer, G. Applying network theory to animal movements to identify properties of landscape space use. Ecol. Appl. 28 (2018).                    https://doi.org/10.1002/eap.1697                                    .") and immediate prior displacement. To lift the aforementioned LCP limitations, we adapted the recently developed mechanistic model SiMRiv[30](/articles/s41598-022-23206-9#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019).") that simulates spatially explicit stochastic movements (multi-state Markov model[82](/articles/s41598-022-23206-9#ref-CR82 "Michelot, T., Langrock, R. & Patterson, T. moveHMM: An R package for the statistical modelling of animal movement data using hidden Markov models. Methods Ecol. Evol. 7 (2016).                    https://doi.org/10.1111/2041-210X.12578                                     .")) accounting for landscape heterogeneity and perceptual range ( _i.e._, the radius up to which an individual perceives its surroundings) (S.I. Fig. [4a](/articles/s41598-022-23206-9#MOESM1)). Here again, the normalized resistance map (S.I. Fig [1c](/articles/s41598-022-23206-9#MOESM1)) defines the underlying environmental complexity of the region and _H. erectus_ movement is simulated with a two-state movement (Lévy-like walker[83](/articles/s41598-022-23206-9#ref-CR83 "Benhamou, S. How many animals really do the Lévy Walk. Ecology 88, 1962–9.                    https://doi.org/10.1890/06-1769.1                                     (2007).")) that alternates between random and correlated random walks[84](/articles/s41598-022-23206-9#ref-CR84 "Turchin, P. Quantitative Analysis of Movement: Measuring and Modeling Population Redistribution of Plants and Animals (Sinauer Associates, Sunderland, 1998)."), where the incremental azimuthal direction is correlated with the prior direction (turning angle concentration parameter of the wrapped normal distribution is set to 0.99 in our simulations) (S.I. Fig. [4b](/articles/s41598-022-23206-9#MOESM1)). To inform the mechanistic model on the probabilities of changing between the two states, SiMRiv provides a transition matrix (which values are here set to 0.01 and 0.002). We also set the step length (unit of movement) for both states to 100 m in order to avoid unrealistic jumps over high cost cells. Finally, we set the perceptual range to 5 km for both states: the movement decisions at each step accounts for the resistance that the walker _sees_ in a 5 km radius around its current location ( _i.e._, it attempts to bypass high-resistance areas within this range, see S.I. Fig. [4b](/articles/s41598-022-23206-9#MOESM1), which illustrates the two-state mechanistic movement for a given realization with multiple phases of acceleration and deceleration as _H. erectus_ moves over the heterogeneous landscape). To have statistically significant results, we run 1000 realizations over 5 million steps each for the three routes (Myanmar, Thailand, and Vietnam entry points). One third of the 3000 realizations reaches Sangiran (respectively 29%, 34% and 38%). As expected, Markovian walks to Sangiran are much longer than LCP (LCP lengths amount to 7-9% of the shortest mechanistic realizations - Myanmar: 83,000 km, Thailand: 66,000 km, and Vietnam: 76,000 km - S.I. Fig. [5a](/articles/s41598-022-23206-9#MOESM1)). We then compute the likelihood of _H. erectus_ occupation at any given location, when combining all realizations successfully passing through Sangiran. We use a kernel density estimate analysis (KDE, S.I. Fig. [5b](/articles/s41598-022-23206-9#MOESM1)) weighted by the inverse normalized cost surface, which not only allows to illuminate the most favourable physiographic settings ( _e.g._, riparian areas, lowlands and coastal plains) but also to account for the slower progression (and proportional number of steps) in complex landscapes (S.I. Fig. [4b](/articles/s41598-022-23206-9#MOESM1)).

### _H. erectus_ migration speed range

Estimating the dispersal velocity across Sundaland remains challenging for _H. erectus_, and we rely on available estimates for better documented, more recent hunter-gatherers hominins. In light of the limited available data on their diet and anatomy, _H. erectus_ are also commonly thought to be hunter-gatherers[85](/articles/s41598-022-23206-9#ref-CR85 "Lieberman, D. E. The Story of the Human Body: Evolution, Health, and Disease (Pantheon Books, New York, 2013)."), [86](/articles/s41598-022-23206-9#ref-CR86 "Braun, D. et al. Early hominin diet included diverse terrestrial and aquatic animals 1.95 Ma in East Turkana, Kenya. Proc. Natl. Acad. Sci. USA 107, 10002–7 (2010).                    https://doi.org/10.1073/pnas.1002181107                                    ."), although controversy persists[87](/articles/s41598-022-23206-9#ref-CR87 "O’Connor, S., Louys, J., Kealy, S. & Samper Carro, S. C. Hominin dispersal and settlement east of huxley’s line: The role of sea level changes, island size, and subsistence behavior. Curr. Anthropol. 58(S17), S567–S582 (2017)."). Recolonization speed estimates for late glacial hunter-gatherers of northern Europe, deduced from reproduction and mobility parameters, occurred at rates of 0.7 to 1.4 km/yr[43](/articles/s41598-022-23206-9#ref-CR43 "Fort, J., Pujol, T. & Cavalli-Sforza, L. Palaeolithic populations and waves of advance. Camb. Archaeol. J. 14, 53–61.                    https://doi.org/10.1017/S0959774304000046                                     (2004)."). The distribution of the Clovis-age occupations across North America suggests a speed of 5-8 km/yr[44](/articles/s41598-022-23206-9#ref-CR44 "Hamilton, M. & Buchanan, B. Spatial gradients in Clovis-age radiocarbon dates across North America suggest rapid colonization from the north. Proc. Natl. Acad. Sci. USA 104, 15625–30.                    https://doi.org/10.1073/pnas.0704215104                                     (2007)."). The first peopling of the Americas, and the Neolithic transition in Europe, occurred at rates between 6 and 10 km/yr, based on archaeological boundary conditions[45](/articles/s41598-022-23206-9#ref-CR45 "Hazelwood, L. & Steele, J. Spatial dynamics of human dispersals: Constraints on modelling and archaeological validation. J. Archaeol. Sci. 31, 669–679.                    https://doi.org/10.1016/j.jas.2003.11.009                                     (2004)."). mtDNA variation in isolated populations in southeast Asia shows a migration speed of 4 km/yr for the dispersal of modern humans from Eurasia to Australasia[88](/articles/s41598-022-23206-9#ref-CR88 "Macaulay, V. et al. Single, rapid coastal settlement of asia revealed by analysis of complete mitochondrial genomes. Science (New York, N.Y.)308, 1034–6 (2005).                    https://doi.org/10.1126/science.1109792                                    ."). Based on these estimates, we conservatively assume plausible migration speeds ranging from 1 to 10 km/yr for Southeast Asian _H. erectus_, and use this range to compute the travel time for each of the routes obtained from both LCP and Markovian walks.

## Data availability

All necessary data are available in the text and tables.

## Code availability

goSPL[32](/articles/s41598-022-23206-9#ref-CR32 "Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. J. Open Source Softw.5(56), 2804 (2020).                    https://doi.org/10.21105/joss.02804                                    ."), is available from [https://github.com/Geodels/goSPL](https://github.com/Geodels/goSPL) and the software documentation can be found at [https://goSPL.readthedocs.io](https://goSPL.readthedocs.io). SiMRiv[30](/articles/s41598-022-23206-9#ref-CR30 "Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. Mov. Ecol.                    https://doi.org/10.1186/s40462-019-0154-8                                     (2019).") is available at [https://github.com/miguel-porto/SiMRiv](https://github.com/miguel-porto/SiMRiv).

## References

01. Dubois, E. On Pithecanthropus Erectus: a transitional form between man and the apes. _J. Anthropol. Inst. G. B. Irel._ **25**, 240–255 (1896).

    [Google Scholar](http://scholar.google.com/scholar_lookup?&title=On%20Pithecanthropus%20Erectus%3A%20a%20transitional%20form%20between%20man%20and%20the%20apes&journal=J.%20Anthropol.%20Inst.%20G.%20B.%20Irel.&volume=25&pages=240-255&publication_year=1896&author=Dubois%2CE)

02. von Koenigswald, G. H. R. _Neue Pithecanthropus-funde, 1936-1938 : ein beitrag zur Kenntnis der Praehominiden_ Wetenschappelijke Mededeelingen ; no. 28 (Landsdrukkerij, Batavia, 1940).

03. Janssen, R. _et al._ Tooth enamel stable isotopes of Holocene and Pleistocene fossil fauna reveal glacial and interglacial paleoenvironments of hominins in Indonesia. _Quatern. Sci. Rev._ **144**, 145–154 (2016).

    [Article](https://doi.org/10.1016%2Fj.quascirev.2016.02.028) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2016QSRv..144..145J) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Tooth%20enamel%20stable%20isotopes%20of%20Holocene%20and%20Pleistocene%20fossil%20fauna%20reveal%20glacial%20and%20interglacial%20paleoenvironments%20of%20hominins%20in%20Indonesia&journal=Quatern.%20Sci.%20Rev.&doi=10.1016%2Fj.quascirev.2016.02.028&volume=144&pages=145-154&publication_year=2016&author=Janssen%2CR)

04. Bettis, E. A. _et al._ Way out of Africa: Early Pleistocene paleoenvironments inhabited by Homo erectus in Sangiran, Java. _J. Hum. Evol._ **56**(1), 11–24 (2009).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2008.09.003) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19007966) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Way%20out%20of%20Africa%3A%20Early%20Pleistocene%20paleoenvironments%20inhabited%20by%20Homo%20erectus%20in%20Sangiran%2C%20Java&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2008.09.003&volume=56&issue=1&pages=11-24&publication_year=2009&author=Bettis%2CEA)

05. Huffman, O. Geologic context and age of the Perning/Mojokerto Homo erectus, East Java. _J. Hum. Evol._ **40**(4), 353–362 (2001).

    [Article](https://doi.org/10.1006%2Fjhev.2001.0464) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=11312587) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Geologic%20context%20and%20age%20of%20the%20Perning%2FMojokerto%20Homo%20erectus%2C%20East%20Java&journal=J.%20Hum.%20Evol.&doi=10.1006%2Fjhev.2001.0464&volume=40&issue=4&pages=353-362&publication_year=2001&author=Huffman%2CO)

06. Sarr, A.-C. _et al._ Subsiding Sundaland. _Geology (Boulder)_ **47**(2), 119–122 (2019).

    [Article](https://doi.org/10.1130%2FG45629.1) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2019Geo....47..119S) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Subsiding%20Sundaland&journal=Geology%20%28Boulder%29&doi=10.1130%2FG45629.1&volume=47&issue=2&pages=119-122&publication_year=2019&author=Sarr%2CA-C)

07. Salles, T. _et al._ Quaternary landscape dynamics boosted species dispersal across Southeast Asia. _Commun. Earth Environ._ **2**(1), 1–12 (2021).

    [Article](https://doi.org/10.1038%2Fs43247-021-00311-7) [MathSciNet](http://www.ams.org/mathscinet-getitem?mr=3628105) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Quaternary%20landscape%20dynamics%20boosted%20species%20dispersal%20across%20Southeast%20Asia&journal=Commun.%20Earth%20Environ.&doi=10.1038%2Fs43247-021-00311-7&volume=2&issue=1&pages=1-12&publication_year=2021&author=Salles%2CT)

08. Husson, L., Boucher, F. C., Sarr, A., Sepulchre, P. & Cahyarini, S. Y. Evidence of Sundaland’s subsidence requires revisiting its biogeography. _J. Biogeogr._ **47**(4), 843–853 (2020).

09. Winder, I. C. _et al._ Evolution and dispersal of the genus Homo: A landscape approach. _J. Hum. Evol._ **87**, 48–65 (2015).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2015.07.002) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=26235482) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Evolution%20and%20dispersal%20of%20the%20genus%20Homo%3A%20A%20landscape%20approach&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2015.07.002&volume=87&pages=48-65&publication_year=2015&author=Winder%2CIC)

10. Carotenuto, F. _et al._ Venturing out safely: The biogeography of Homo erectus dispersal out of Africa. _J. Hum. Evol._ **95**, 1–12 (2016).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2016.02.005) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=27260171) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Venturing%20out%20safely%3A%20The%20biogeography%20of%20Homo%20erectus%20dispersal%20out%20of%20Africa&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2016.02.005&volume=95&pages=1-12&publication_year=2016&author=Carotenuto%2CF)

11. Larick, R. _et al._ Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central Jawa, Indonesia. _Proc. Natl. Acad. Sci. PNAS_ **98**(9), 4866–4871 (2001).

    [Article](https://doi.org/10.1073%2Fpnas.081077298) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2001PNAS...98.4866L) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=11309488) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Early%20Pleistocene%2040Ar%2F39Ar%20ages%20for%20Bapang%20Formation%20hominins%2C%20Central%20Jawa%2C%20Indonesia&journal=Proc.%20Natl.%20Acad.%20Sci.%20PNAS&doi=10.1073%2Fpnas.081077298&volume=98&issue=9&pages=4866-4871&publication_year=2001&author=Larick%2CR)

12. Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G. & Suprijo, A. Age of the earliest known hominids in Java, Indonesia. _Science_ **263**(5150), 1118–1121 (1994).

    [Article](https://doi.org/10.1126%2Fscience.8108729) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=1994Sci...263.1118S) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=8108729) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Age%20of%20the%20earliest%20known%20hominids%20in%20Java%2C%20Indonesia&journal=Science&doi=10.1126%2Fscience.8108729&volume=263&issue=5150&pages=1118-1121&publication_year=1994&author=Swisher%2CCC&author=Curtis%2CGH&author=Jacob%2CT&author=Getty%2CAG&author=Suprijo%2CA)

13. Sémah, F., Saleki, H., Falguŕes, C., Féraud, G. & Djubiantono, T. Did early man reach Java during the Late Pliocene?. _J. Archaeol. Sci._ **27**(9), 763–769 (2000).

    [Article](https://doi.org/10.1006%2Fjasc.1999.0482) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Did%20early%20man%20reach%20Java%20during%20the%20Late%20Pliocene%3F&journal=J.%20Archaeol.%20Sci.&doi=10.1006%2Fjasc.1999.0482&volume=27&issue=9&pages=763-769&publication_year=2000&author=S%C3%A9mah%2CF&author=Saleki%2CH&author=Falgu%C5%95es%2CC&author=F%C3%A9raud%2CG&author=Djubiantono%2CT)

14. Bettis, E. _et al._ Landscape development preceding Homo erectus immigration into Central Java, Indonesia: The Sangiran Formation Lower Lahar. _Palaeogeogr. Palaeoclimatol. Palaeoecol._ **206**(1), 115–131 (2004).

    [Article](https://doi.org/10.1016%2Fj.palaeo.2004.01.016) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Landscape%20development%20preceding%20Homo%20erectus%20immigration%20into%20Central%20Java%2C%20Indonesia%3A%20The%20Sangiran%20Formation%20Lower%20Lahar&journal=Palaeogeogr.%20Palaeoclimatol.%20Palaeoecol.&doi=10.1016%2Fj.palaeo.2004.01.016&volume=206&issue=1&pages=115-131&publication_year=2004&author=Bettis%2CE)

15. Matsu’ura, S. _et al._ Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. _Science_ **367**(6474), 210–214 (2020).

    [Article](https://doi.org/10.1126%2Fscience.aau8556) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2020Sci...367..210M) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=31919224) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Age%20control%20of%20the%20first%20appearance%20datum%20for%20Javanese%20Homo%20erectus%20in%20the%20Sangiran%20area&journal=Science&doi=10.1126%2Fscience.aau8556&volume=367&issue=6474&pages=210-214&publication_year=2020&author=Matsu%E2%80%99ura%2CS)

16. Granger, D. E. & Muzikar, P. F. Dating sediment burial with in situ-produced cosmogenic nuclides: Theory, techniques, and limitations. _Earth Planet. Sci. Lett._ **188**(1), 269–281 (2001).

    [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2001E%26PSL.188..269G) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Dating%20sediment%20burial%20with%20in%20situ-produced%20cosmogenic%20nuclides%3A%20Theory%2C%20techniques%2C%20and%20limitations&journal=Earth%20Planet.%20Sci.%20Lett.&volume=188&issue=1&pages=269-281&publication_year=2001&author=Granger%2CDE&author=Muzikar%2CPF)

17. Shen, G., Gao, X., Gao, B. & Granger, D. E. Age of Zhoukoudian Homo erectus determined with 26Al/10Be burial dating. _Nature_ **458**(7235), 198–200 (2009).

    [Article](https://doi.org/10.1038%2Fnature07741) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2009Natur.458..198S) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19279636) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Age%20of%20Zhoukoudian%20Homo%20erectus%20determined%20with%2026Al%2F10Be%20burial%20dating&journal=Nature&doi=10.1038%2Fnature07741&volume=458&issue=7235&pages=198-200&publication_year=2009&author=Shen%2CG&author=Gao%2CX&author=Gao%2CB&author=Granger%2CDE)

18. Pappu, S. _et al._ Early Pleistocene presence of Acheulian Hominins in South India. _Science_ **331**(6024), 1596–1599 (2011).

    [Article](https://doi.org/10.1126%2Fscience.1200183) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011Sci...331.1596P) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=21436450) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Early%20Pleistocene%20presence%20of%20Acheulian%20Hominins%20in%20South%20India&journal=Science&doi=10.1126%2Fscience.1200183&volume=331&issue=6024&pages=1596-1599&publication_year=2011&author=Pappu%2CS)

19. Lebatard, A.-E. _et al._ Dating the Homo erectus bearing travertine from Kocabaş (Denizli, Turkey) at at least 1.1 Ma. _Earth Planet. Sci. Lett._ **390**, 8–18 (2014).

20. Lebatard, A.-E., Bourlès, D. L. & Braucher, R. Absolute dating of an Early Paleolithic site in Western Africa based on the radioactive decay of in situ-produced 10Be and 26Al. _Nucl. Instrum. Methods Phys. Res. Sect. B_ **456**, 169–179 (2019).

    [Article](https://doi.org/10.1016%2Fj.nimb.2019.05.052) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2019NIMPB.456..169L) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Absolute%20dating%20of%20an%20Early%20Paleolithic%20site%20in%20Western%20Africa%20based%20on%20the%20radioactive%20decay%20of%20in%20situ-produced%2010Be%20and%2026Al&journal=Nucl.%20Instrum.%20Methods%20Phys.%20Res.%20Sect.%20B&doi=10.1016%2Fj.nimb.2019.05.052&volume=456&pages=169-179&publication_year=2019&author=Lebatard%2CA-E&author=Bourl%C3%A8s%2CDL&author=Braucher%2CR)

21. Braucher, R., Oslisly, R., Mesfin, I., Ntoutoume Mba, P. P. & Team, A. In situ-produced 10 Be and 26 Al indirect dating of Elarmékora Earlier Stone Age artifacts: First attempt in a savannah forest mosaic in the middle Ogooué valley, Gabon. _Philos. Trans. Biol. Sci._ (2021) .

22. Grimaud-Hervé, D. _et al._ Position of the posterior skullcap fragment from Sendang Klampok (Sangiran Dome, Java, Indonesia) among the Javanese Homo erectus record. _Quatern. Int._ **416**, 193–209 (2016).

    [Article](https://doi.org/10.1016%2Fj.quaint.2015.10.003) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Position%20of%20the%20posterior%20skullcap%20fragment%20from%20Sendang%20Klampok%20%28Sangiran%20Dome%2C%20Java%2C%20Indonesia%29%20among%20the%20Javanese%20Homo%20erectus%20record&journal=Quatern.%20Int.&doi=10.1016%2Fj.quaint.2015.10.003&volume=416&pages=193-209&publication_year=2016&author=Grimaud-Herv%C3%A9%2CD)

23. Sartono, S. Observations on a new skull of Pithecanthropus erectus (Pithecanthropus VIII), from Sangiran, Central Java. _Koninklijke Akademie Wetenschappen te Amsterdam_ **74**, 185–194 (1971).

    [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Observations%20on%20a%20new%20skull%20of%20Pithecanthropus%20erectus%20%28Pithecanthropus%20VIII%29%2C%20from%20Sangiran%2C%20Central%20Java&journal=Koninklijke%20Akademie%20Wetenschappen%20te%20Amsterdam&volume=74&pages=185-194&publication_year=1971&author=Sartono%2CS)

24. Wessel, P., Smith, W. H. F., Scharroo, R., Luis, J. & Wobbe, F. Generic mapping tools: Improved version released. _EOS Trans. Am. Geophys. Union_ **94**(45), 409–410. [https://doi.org/10.1002/2013EO450001](https://doi.org/10.1002/2013EO450001) (2013).

    [Article](https://doi.org/10.1002%2F2013EO450001) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2013EOSTr..94..409W) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Generic%20mapping%20tools%3A%20Improved%20version%20released&journal=EOS%20Trans.%20Am.%20Geophys.%20Union&doi=10.1002%2F2013EO450001&volume=94&issue=45&pages=409-410&publication_year=2013&author=Wessel%2CP&author=Smith%2CWHF&author=Scharroo%2CR&author=Luis%2CJ&author=Wobbe%2CF)

25. Antón, S., Potts, R. & Aiello, L. Evolution of Early Homo: An integrated biological perspective. _Science (New York, N.Y.)_ **345** (2014). [https://doi.org/10.1126/science.1236828](https://doi.org/10.1126/science.1236828).

26. Luo, L. _et al._ The first radiometric age by isochron 26Al/10Be burial dating for the Early Pleistocene Yuanmou hominin site, southern China. _Quat. Geochronol._ **55**, 101022\. [https://doi.org/10.1016/j.quageo.2019.101022](https://doi.org/10.1016/j.quageo.2019.101022) (2019).

    [Article](https://doi.org/10.1016%2Fj.quageo.2019.101022) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=The%20first%20radiometric%20age%20by%20isochron%2026Al%2F10Be%20burial%20dating%20for%20the%20Early%20Pleistocene%20Yuanmou%20hominin%20site%2C%20southern%20China&journal=Quat.%20Geochronol.&doi=10.1016%2Fj.quageo.2019.101022&volume=55&publication_year=2019&author=Luo%2CL)

27. Zaim, Y. _et al._ New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia). _J. Hum. Evol._ **61**(4), 363–376 (2011).

28. Rizal, Y. _et al._ Last appearance of Homo erectus at Ngandong, Java, 117,000–108,000 years ago. _Nature_ **577**(7790), 381–385 (2020).

    [Article](https://doi.org/10.1038%2Fs41586-019-1863-2) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=31853068) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Last%20appearance%20of%20Homo%20erectus%20at%20Ngandong%2C%20Java%2C%20117%2C000%E2%80%93108%2C000%20years%20ago&journal=Nature&doi=10.1038%2Fs41586-019-1863-2&volume=577&issue=7790&pages=381-385&publication_year=2020&author=Rizal%2CY)

29. McRae, B. & Beier, P. Circuit theory predicts gene flow in plant and animal populations. _Proc. Natl. Acad. Sci. USA_ **104**, 19885–90. [https://doi.org/10.1073/pnas.0706568104](https://doi.org/10.1073/pnas.0706568104) (2008).

    [Article](https://doi.org/10.1073%2Fpnas.0706568104) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2007PNAS..10419885M) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Circuit%20theory%20predicts%20gene%20flow%20in%20plant%20and%20animal%20populations&journal=Proc.%20Natl.%20Acad.%20Sci.%20USA&doi=10.1073%2Fpnas.0706568104&volume=104&pages=19885-90&publication_year=2008&author=McRae%2CB&author=Beier%2CP)

30. Quaglietta, L. & Porto, M. SiMRiv: An R package for mechanistic simulation of individual, spatially-explicit multistate movements in rivers, heterogeneous and homogeneous spaces incorporating landscape bias. _Mov. Ecol._ [https://doi.org/10.1186/s40462-019-0154-8](https://doi.org/10.1186/s40462-019-0154-8) (2019).

    [Article](https://link.springer.com/doi/10.1186/s40462-019-0154-8) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=30984401) [PubMed Central](http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6444552) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=SiMRiv%3A%20An%20R%20package%20for%20mechanistic%20simulation%20of%20individual%2C%20spatially-explicit%20multistate%20movements%20in%20rivers%2C%20heterogeneous%20and%20homogeneous%20spaces%20incorporating%20landscape%20bias&journal=Mov.%20Ecol.&doi=10.1186%2Fs40462-019-0154-8&publication_year=2019&author=Quaglietta%2CL&author=Porto%2CM)

31. Landau, V. A., Shah, V. B., Anantharaman, R. & Hall, K. R. Omniscape.jl: Software to compute omnidirectional landscape connectivity. _J. Open Source Softw._ **6**(57), 2829 (2021). [https://doi.org/10.21105/joss.02829](https://doi.org/10.21105/joss.02829).

32. Salles, T., Mallard, C. & Zahirovic, S. gospl: Global Scalable Paleo Landscape Evolution. _J. Open Source Softw._ **5**(56), 2804 (2020). [https://doi.org/10.21105/joss.02804](https://doi.org/10.21105/joss.02804).

33. Husson, L. _et al._ Slow geodynamics and fast morphotectonics in the far East Tethys. _Geochem. Geophys. Geosyst._ **23**(1), n/a (2022).

34. Valdes, P., Scotese, C. & Lunt, D. Deep ocean temperatures through time. _Climate Past_ **17**, 1483–1506. [https://doi.org/10.5194/cp-17-1483-2021](https://doi.org/10.5194/cp-17-1483-2021) (2021).

    [Article](https://doi.org/10.5194%2Fcp-17-1483-2021) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2021CliPa..17.1483V) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Deep%20ocean%20temperatures%20through%20time&journal=Climate%20Past&doi=10.5194%2Fcp-17-1483-2021&volume=17&pages=1483-1506&publication_year=2021&author=Valdes%2CP&author=Scotese%2CC&author=Lunt%2CD)

35. Hyodo, M. _et al._ High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia. _Proc. Natl. Acad. Sci. PNAS_ **108**(49), 19563–19568 (2011).

    [Article](https://doi.org/10.1073%2Fpnas.1113106108) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011PNAS..10819563H) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=22106291) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=High-resolution%20record%20of%20the%20Matuyama%E2%80%93Brunhes%20transition%20constrains%20the%20age%20of%20Javanese%20Homo%20erectus%20in%20the%20Sangiran%20dome%2C%20Indonesia&journal=Proc.%20Natl.%20Acad.%20Sci.%20PNAS&doi=10.1073%2Fpnas.1113106108&volume=108&issue=49&pages=19563-19568&publication_year=2011&author=Hyodo%2CM)

36. Brasseur, B., Sémah, F., Sémah, A.-M. & Djubiantono, T. Pedo-sedimentary dynamics of the Sangiran dome hominid bearing layers (Early to Middle Pleistocene, central Java, Indonesia): A palaeopedological approach for reconstructing ‘Pithecanthropus’ (Javanese Homo erectus) palaeoenvironment. _Quatern. Int._ **376**, 84–100 (2015).

    [Article](https://doi.org/10.1016%2Fj.quaint.2014.05.030) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Pedo-sedimentary%20dynamics%20of%20the%20Sangiran%20dome%20hominid%20bearing%20layers%20%28Early%20to%20Middle%20Pleistocene%2C%20central%20Java%2C%20Indonesia%29%3A%20A%20palaeopedological%20approach%20for%20reconstructing%20%E2%80%98Pithecanthropus%E2%80%99%20%28Javanese%20Homo%20erectus%29%20palaeoenvironment&journal=Quatern.%20Int.&doi=10.1016%2Fj.quaint.2014.05.030&volume=376&pages=84-100&publication_year=2015&author=Brasseur%2CB&author=S%C3%A9mah%2CF&author=S%C3%A9mah%2CA-M&author=Djubiantono%2CT)

37. Falguéres, C. _et al._ Geochronology of early human settlements in Java: What is at stake?. _Quatern. Int._ **416**, 5–11 (2016).

    [Article](https://doi.org/10.1016%2Fj.quaint.2015.10.076) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Geochronology%20of%20early%20human%20settlements%20in%20Java%3A%20What%20is%20at%20stake%3F&journal=Quatern.%20Int.&doi=10.1016%2Fj.quaint.2015.10.076&volume=416&pages=5-11&publication_year=2016&author=Falgu%C3%A9res%2CC)

38. Roach, N. _et al._ Pleistocene footprints show intensive use of lake margin habitats by Homo erectus groups. _Sci. Rep._ **121** (2016). [https://doi.org/10.1038/srep26374](https://doi.org/10.1038/srep26374).

39. Simandjuntak, T. O. & Barber, A. J. Contrasting tectonic styles in the Neogene orogenic belts of Indonesia. _Geol. Soc. Spec. Pub._ **106**(1), 185–201 (1996).

    [Article](https://doi.org/10.1144%2FGSL.SP.1996.106.01.12) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Contrasting%20tectonic%20styles%20in%20the%20Neogene%20orogenic%20belts%20of%20Indonesia&journal=Geol.%20Soc.%20Spec.%20Pub.&doi=10.1144%2FGSL.SP.1996.106.01.12&volume=106&issue=1&pages=185-201&publication_year=1996&author=Simandjuntak%2CTO&author=Barber%2CAJ)

40. Clements, B., Hall, R., Smyth, H. R. & Cottam, M. A. Thrusting of a volcanic arc; a new structural model for Java. _Pet. Geosci._ **15**(2), 159–174 (2009).

    [Article](https://doi.org/10.1144%2F1354-079309-831) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Thrusting%20of%20a%20volcanic%20arc%3B%20a%20new%20structural%20model%20for%20Java&journal=Pet.%20Geosci.&doi=10.1144%2F1354-079309-831&volume=15&issue=2&pages=159-174&publication_year=2009&author=Clements%2CB&author=Hall%2CR&author=Smyth%2CHR&author=Cottam%2CMA)

41. Joordens, J., Wesselingh, F., de Vos, J., Vonhof, H. & Kroon, D. Relevance of aquatic environments for hominins: A case study from Trinil (Java, Indonesia). _J. Hum. Evol._ **57**(6), 656–671 (2009).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2009.06.003) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19683789) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Relevance%20of%20aquatic%20environments%20for%20hominins%3A%20A%20case%20study%20from%20Trinil%20%28Java%2C%20Indonesia%29&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2009.06.003&volume=57&issue=6&pages=656-671&publication_year=2009&author=Joordens%2CJ&author=Wesselingh%2CF&author=Vos%2CJ&author=Vonhof%2CH&author=Kroon%2CD)

42. Berghuis, H. _et al._ Hominin homelands of East Java: Revised stratigraphy and landscape reconstructions for Plio-Pleistocene Trinil. _Quatern. Sci. Rev._ **260**, 106912 (2021).

    [Article](https://doi.org/10.1016%2Fj.quascirev.2021.106912) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Hominin%20homelands%20of%20East%20Java%3A%20Revised%20stratigraphy%20and%20landscape%20reconstructions%20for%20Plio-Pleistocene%20Trinil&journal=Quatern.%20Sci.%20Rev.&doi=10.1016%2Fj.quascirev.2021.106912&volume=260&publication_year=2021&author=Berghuis%2CH)

43. Fort, J., Pujol, T. & Cavalli-Sforza, L. Palaeolithic populations and waves of advance. _Camb. Archaeol. J._ **14**, 53–61. [https://doi.org/10.1017/S0959774304000046](https://doi.org/10.1017/S0959774304000046) (2004).

    [Article](https://doi.org/10.1017%2FS0959774304000046) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Palaeolithic%20populations%20and%20waves%20of%20advance&journal=Camb.%20Archaeol.%20J.&doi=10.1017%2FS0959774304000046&volume=14&pages=53-61&publication_year=2004&author=Fort%2CJ&author=Pujol%2CT&author=Cavalli-Sforza%2CL)

44. Hamilton, M. & Buchanan, B. Spatial gradients in Clovis-age radiocarbon dates across North America suggest rapid colonization from the north. _Proc. Natl. Acad. Sci. USA_ **104**, 15625–30. [https://doi.org/10.1073/pnas.0704215104](https://doi.org/10.1073/pnas.0704215104) (2007).

    [Article](https://doi.org/10.1073%2Fpnas.0704215104) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2007PNAS..10415625H) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17898176) [PubMed Central](http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2000382) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Spatial%20gradients%20in%20Clovis-age%20radiocarbon%20dates%20across%20North%20America%20suggest%20rapid%20colonization%20from%20the%20north&journal=Proc.%20Natl.%20Acad.%20Sci.%20USA&doi=10.1073%2Fpnas.0704215104&volume=104&pages=15625-30&publication_year=2007&author=Hamilton%2CM&author=Buchanan%2CB)

45. Hazelwood, L. & Steele, J. Spatial dynamics of human dispersals: Constraints on modelling and archaeological validation. _J. Archaeol. Sci._ **31**, 669–679. [https://doi.org/10.1016/j.jas.2003.11.009](https://doi.org/10.1016/j.jas.2003.11.009) (2004).

    [Article](https://doi.org/10.1016%2Fj.jas.2003.11.009) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Spatial%20dynamics%20of%20human%20dispersals%3A%20Constraints%20on%20modelling%20and%20archaeological%20validation&journal=J.%20Archaeol.%20Sci.&doi=10.1016%2Fj.jas.2003.11.009&volume=31&pages=669-679&publication_year=2004&author=Hazelwood%2CL&author=Steele%2CJ)

46. Bae, C., Li, F., Liuling, C., Wang, W. & Hanlie, H. Hominin distribution and density patterns in pleistocene China: Climatic influences. _Palaeogeogr. Palaeoclimatol. Palaeoecol._ **512** (2018). [https://doi.org/10.1016/j.palaeo.2018.03.015](https://doi.org/10.1016/j.palaeo.2018.03.015).

47. Timmermann, A. _et al._ Climate effects on archaic human habitats and species successions. _Nature_ **604**, 1–7. [https://doi.org/10.1038/s41586-022-04600-9](https://doi.org/10.1038/s41586-022-04600-9) (2022).

    [Article](https://doi.org/10.1038%2Fs41586-022-04600-9) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Climate%20effects%20on%20archaic%20human%20habitats%20and%20species%20successions&journal=Nature&doi=10.1038%2Fs41586-022-04600-9&volume=604&pages=1-7&publication_year=2022&author=Timmermann%2CA)

48. Bailey, G. N., Reynolds, S. C. & King, G. C. Landscapes of human evolution: Models and methods of tectonic geomorphology and the reconstruction of hominin landscapes. _J. Hum. Evol._ **60**(3), 257–280 (2011).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2010.01.004) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=20947132) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Landscapes%20of%20human%20evolution%3A%20Models%20and%20methods%20of%20tectonic%20geomorphology%20and%20the%20reconstruction%20of%20hominin%20landscapes&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2010.01.004&volume=60&issue=3&pages=257-280&publication_year=2011&author=Bailey%2CGN&author=Reynolds%2CSC&author=King%2CGC)

49. Sarr, A., Sepulchre, P. & Husson, L. Impact of the Sunda Shelf on the Climate of the Maritime Continent. _J. Geophys. Res. Atmos._ **124**(5), 2574–2588 (2019).

    [Article](https://doi.org/10.1029%2F2018JD029971) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2019JGRD..124.2574S) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Impact%20of%20the%20Sunda%20Shelf%20on%20the%20Climate%20of%20the%20Maritime%20Continent&journal=J.%20Geophys.%20Res.%20Atmos.&doi=10.1029%2F2018JD029971&volume=124&issue=5&pages=2574-2588&publication_year=2019&author=Sarr%2CA&author=Sepulchre%2CP&author=Husson%2CL)

50. Louys, J. & Roberts, P. Environmental drivers of megafauna and hominin extinction in Southeast Asia. _Nature_ **586**(7829), 402–406 (2020).

    [Article](https://doi.org/10.1038%2Fs41586-020-2810-y) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2020Natur.586..402L) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=33029012) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Environmental%20drivers%20of%20megafauna%20and%20hominin%20extinction%20in%20Southeast%20Asia&journal=Nature&doi=10.1038%2Fs41586-020-2810-y&volume=586&issue=7829&pages=402-406&publication_year=2020&author=Louys%2CJ&author=Roberts%2CP)

51. Raia, P. _et al._ Past extinctions of homo species coincided with increased vulnerability to climatic change. _One Earth_ **3**(4), 480–490 (2020).

    [Article](https://doi.org/10.1016%2Fj.oneear.2020.09.007) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2020OEart...3..480R) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Past%20extinctions%20of%20homo%20species%20coincided%20with%20increased%20vulnerability%20to%20climatic%20change&journal=One%20Earth&doi=10.1016%2Fj.oneear.2020.09.007&volume=3&issue=4&pages=480-490&publication_year=2020&author=Raia%2CP)

52. Zhu, Z. _et al._ Hominin occupation of the Chinese Loess Plateau since about 2.1 million years ago. _Nature_ **559**(7715), 608–612 (2018).

53. Gabunia, L. _et al._ Earliest Pleistocene hominid cranial remains from Dmanisi, Republic of Georgia: Taxonomy, geological setting, and age. _Science_ **288**, 1019–1025. [https://doi.org/10.1126/science.288.5468.1019](https://doi.org/10.1126/science.288.5468.1019) (2000).

    [Article](https://doi.org/10.1126%2Fscience.288.5468.1019) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2000Sci...288.1019G) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=10807567) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Earliest%20Pleistocene%20hominid%20cranial%20remains%20from%20Dmanisi%2C%20Republic%20of%20Georgia%3A%20Taxonomy%2C%20geological%20setting%2C%20and%20age&journal=Science&doi=10.1126%2Fscience.288.5468.1019&volume=288&pages=1019-1025&publication_year=2000&author=Gabunia%2CL)

54. Lordkipanidze, D. _et al._ A complete skull from Dmanisi, Georgia, and the evolutionary biology of early homo. _Science_ **342**(6156), 326–331 (2013).

    [Article](https://doi.org/10.1126%2Fscience.1238484) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2013Sci...342..326L) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=24136960) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=A%20complete%20skull%20from%20Dmanisi%2C%20Georgia%2C%20and%20the%20evolutionary%20biology%20of%20early%20homo&journal=Science&doi=10.1126%2Fscience.1238484&volume=342&issue=6156&pages=326-331&publication_year=2013&author=Lordkipanidze%2CD)

55. Baba, H. _et al._ Homo erectus calvarium from the pleistocene of java. _Sci. (Am. Assoc. Adv. Sci.)_ **299** (5611), 1384–1388 (2003) .

56. Ciochon, R. L. & Bettis, E. A. III. Asian Homo erectus converges in time. _Nature_ **458**(7235), 153–154 (2009).

    [Article](https://doi.org/10.1038%2F458153a) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2009Natur.458..153C) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=19279621) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Asian%20Homo%20erectus%20converges%20in%20time&journal=Nature&doi=10.1038%2F458153a&volume=458&issue=7235&pages=153-154&publication_year=2009&author=Ciochon%2CRL&author=Bettis%2CEA)

57. Dennell, R. & Roebroeks, W. An Asian perspective on early human dispersal from Africa. _Nature_ **438**(7071), 1099–1104 (2005).

    [Article](https://doi.org/10.1038%2Fnature04259) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2005Natur.438.1099D) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=16371999) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=An%20Asian%20perspective%20on%20early%20human%20dispersal%20from%20Africa&journal=Nature&doi=10.1038%2Fnature04259&volume=438&issue=7071&pages=1099-1104&publication_year=2005&author=Dennell%2CR&author=Roebroeks%2CW)

58. Martinon-Torres, M. _et al._ Dental evidence on the hominin dispersals during the Pleistocene. _Proc. Natl. Acad. Sci. PNAS_ **104**(33), 13279–13282 (2007).

    [Article](https://doi.org/10.1073%2Fpnas.0706152104) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2007PNAS..10413279M) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17684093) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Dental%20evidence%20on%20the%20hominin%20dispersals%20during%20the%20Pleistocene&journal=Proc.%20Natl.%20Acad.%20Sci.%20PNAS&doi=10.1073%2Fpnas.0706152104&volume=104&issue=33&pages=13279-13282&publication_year=2007&author=Martinon-Torres%2CM)

59. Wood, B. Did early Homo migrate “out of’’ or “in to’’ Africa?. _Proc. Natl. Acad. Sci. PNAS_ **108**(26), 10375–10376 (2011).

    [Article](https://doi.org/10.1073%2Fpnas.1107724108) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011PNAS..10810375W) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=21677194) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Did%20early%20Homo%20migrate%20%E2%80%9Cout%20of%E2%80%9D%20or%20%E2%80%9Cin%20to%E2%80%9D%20Africa%3F&journal=Proc.%20Natl.%20Acad.%20Sci.%20PNAS&doi=10.1073%2Fpnas.1107724108&volume=108&issue=26&pages=10375-10376&publication_year=2011&author=Wood%2CB)

60. Shen, G. _et al._ Isochron 26Al/10Be burial dating of Xihoudu: Evidence for the earliest human settlement in northern China. _Anthropologie_ **124**, 102790\. [https://doi.org/10.1016/j.anthro.2020.102790](https://doi.org/10.1016/j.anthro.2020.102790) (2020).

    [Article](https://doi.org/10.1016%2Fj.anthro.2020.102790) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Isochron%2026Al%2F10Be%20burial%20dating%20of%20Xihoudu%3A%20Evidence%20for%20the%20earliest%20human%20settlement%20in%20northern%20China&journal=Anthropologie&doi=10.1016%2Fj.anthro.2020.102790&volume=124&publication_year=2020&author=Shen%2CG)

61. Chmeleff, J., von Blanckenburg, F., Kossert, K. & Jakob, D. Determination of the 10Be half-life by multicollector ICP-MS and liquid scintillation counting. _Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms_ **268**(2), 192–199 (2010).

62. Korschinek, G. _et al._ A new value for the half-life of 10Be by Heavy-Ion Elastic Recoil Detection and liquid scintillation counting. _Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms_ **268**(2), 187–191 (2010) .

63. Nishiizumi, K. Preparation of 26Al AMS standards. _Nucl. Inst. and Meth. in Phys. Res._ **223-224**, 388–392 (2004).

64. Norris, T. L., Gancarz, A. J., Rokop, D. J. & Thomas, K. W. Half-life of 26Al. _J. Geophys. Res. Solid Earth_ **88**(S01), B331–B333 (1983).

    [Article](https://doi.org/10.1029%2FJB088iS01p0B331) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=1983JGRS...88..331N) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Half-life%20of%2026Al&journal=J.%20Geophys.%20Res.%20Solid%20Earth&doi=10.1029%2FJB088iS01p0B331&volume=88&issue=S01&pages=B331-B333&publication_year=1983&author=Norris%2CTL&author=Gancarz%2CAJ&author=Rokop%2CDJ&author=Thomas%2CKW)

65. Braucher, R., Merchel, S., Borgomano, J. & Bourlès, D. Production of cosmogenic radionuclides at great depth: A multi element approach. _Earth Planet. Sci. Lett._ **309**(1), 1–9 (2011).

    [Article](https://doi.org/10.1016%2Fj.epsl.2011.06.036) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2011E%26PSL.309....1B) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Production%20of%20cosmogenic%20radionuclides%20at%20great%20depth%3A%20A%20multi%20element%20approach&journal=Earth%20Planet.%20Sci.%20Lett.&doi=10.1016%2Fj.epsl.2011.06.036&volume=309&issue=1&pages=1-9&publication_year=2011&author=Braucher%2CR&author=Merchel%2CS&author=Borgomano%2CJ&author=Bourl%C3%A8s%2CD)

66. Braucher, R. _et al._ Preparation of ASTER in-house 10Be/9Be standard solutions. _Nucl. Instrum. Methods Phys. Res. Sect. B Beam Interact. Mater. Atoms_ **361**, 335–340 (2015) .

67. Merchel, S. & Bremser, W. First international 26Al interlaboratory comparison—Part I. _Nucl. Instrum. Methods Phys. Res._ **223–224**, 393–400 (2004).

    [Article](https://doi.org/10.1016%2Fj.nimb.2004.04.076) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2004NIMPB.223..393M) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=First%20international%2026Al%20interlaboratory%20comparison%E2%80%94Part%20I&journal=Nucl.%20Instrum.%20Methods%20Phys.%20Res.&doi=10.1016%2Fj.nimb.2004.04.076&volume=223%E2%80%93224&pages=393-400&publication_year=2004&author=Merchel%2CS&author=Bremser%2CW)

68. Arnold, M. _et al._ The French accelerator mass spectrometry facility ASTER: Improved performance and developments. _Nucl. Instrum. Methods Phys. Res._ **268**(11), 1954–1959 (2010).

    [Article](https://doi.org/10.1016%2Fj.nimb.2010.02.107) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2010NIMPB.268.1954A) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=The%20French%20accelerator%20mass%20spectrometry%20facility%20ASTER%3A%20Improved%20performance%20and%20developments&journal=Nucl.%20Instrum.%20Methods%20Phys.%20Res.&doi=10.1016%2Fj.nimb.2010.02.107&volume=268&issue=11&pages=1954-1959&publication_year=2010&author=Arnold%2CM)

69. Borchers, B. _et al._ Geological calibration of spallation production rates in the CRONUS-Earth project. _Quat. Geochronol._ **31**, 188–198 (2016).

    [Article](https://doi.org/10.1016%2Fj.quageo.2015.01.009) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Geological%20calibration%20of%20spallation%20production%20rates%20in%20the%20CRONUS-Earth%20project&journal=Quat.%20Geochronol.&doi=10.1016%2Fj.quageo.2015.01.009&volume=31&pages=188-198&publication_year=2016&author=Borchers%2CB)

70. Stone, J. O. Air pressure and cosmogenic isotope production. _J. Geophys. Res. Solid Earth_ **105**(B10), 23753–23759 (2000).

    [Article](https://doi.org/10.1029%2F2000JB900181) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Air%20pressure%20and%20cosmogenic%20isotope%20production&journal=J.%20Geophys.%20Res.%20Solid%20Earth&doi=10.1029%2F2000JB900181&volume=105&issue=B10&pages=23753-23759&publication_year=2000&author=Stone%2CJO)

71. Bintanja, R. & van de Wal, R. S. W. North American ice-sheet dynamics and the onset of 100,000-year glacial cycles. _Nature_ **454**, 869–872. [https://doi.org/10.1038/nature07158](https://doi.org/10.1038/nature07158) (2008).

    [Article](https://doi.org/10.1038%2Fnature07158) [ADS](http://adsabs.harvard.edu/cgi-bin/nph-data_query?link_type=ABSTRACT&bibcode=2008Natur.454..869B) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=18704083) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=North%20American%20ice-sheet%20dynamics%20and%20the%20onset%20of%20100%2C000-year%20glacial%20cycles&journal=Nature&doi=10.1038%2Fnature07158&volume=454&pages=869-872&publication_year=2008&author=Bintanja%2CR&author=Wal%2CRSW)

72. Field, J. & Mirazon Lahr, M. Assessment of the Southern Dispersal: GIS-Based Analyses of Potential Routes at Oxygen Isotopic Stage 4. _J. World Prehist._ **19**, 1–45 (2005). [https://doi.org/10.1007/s10963-005-9000-6](https://doi.org/10.1007/s10963-005-9000-6).

73. Howey, M. Multiple pathways across past landscapes: Circuit theory as a complementary geospatial method to least cost path for modeling past movement. _J. Archaeol. Sci._ **38**, 2523–2535. [https://doi.org/10.1016/j.jas.2011.03.024](https://doi.org/10.1016/j.jas.2011.03.024) (2011).

    [Article](https://doi.org/10.1016%2Fj.jas.2011.03.024) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Multiple%20pathways%20across%20past%20landscapes%3A%20Circuit%20theory%20as%20a%20complementary%20geospatial%20method%20to%20least%20cost%20path%20for%20modeling%20past%20movement&journal=J.%20Archaeol.%20Sci.&doi=10.1016%2Fj.jas.2011.03.024&volume=38&pages=2523-2535&publication_year=2011&author=Howey%2CM)

74. Tassi, F. _et al._ Early modern human dispersal from Africa: Genomic evidence for multiple waves of migration. _Investig. Genet._ **6**, 13\. [https://doi.org/10.1186/s13323-015-0030-2](https://doi.org/10.1186/s13323-015-0030-2) (2015).

    [Article](https://link.springer.com/doi/10.1186/s13323-015-0030-2) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=26550467) [PubMed Central](http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4636834) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Early%20modern%20human%20dispersal%20from%20Africa%3A%20Genomic%20evidence%20for%20multiple%20waves%20of%20migration&journal=Investig.%20Genet.&doi=10.1186%2Fs13323-015-0030-2&volume=6&publication_year=2015&author=Tassi%2CF)

75. Kealy, S., Louys, J. & O’Connor, S. Least-cost pathway models indicate northern human dispersal from Sunda to Sahul. _J. Hum. Evol._ **125**, 59–70. [https://doi.org/10.1016/j.jhevol.2018.10.003](https://doi.org/10.1016/j.jhevol.2018.10.003) (2018).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2018.10.003) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=30502898) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Least-cost%20pathway%20models%20indicate%20northern%20human%20dispersal%20from%20Sunda%20to%20Sahul&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2018.10.003&volume=125&pages=59-70&publication_year=2018&author=Kealy%2CS&author=Louys%2CJ&author=O%E2%80%99Connor%2CS)

76. Dennell, R. W., Rendell, H. M. & Hailwood, E. Late pliocene artefacts from northern Pakistan. _Curr. Anthropol._ **29**(3), 495–498 (1988).

    [Article](https://doi.org/10.1086%2F203666) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Late%20pliocene%20artefacts%20from%20northern%20Pakistan&journal=Curr.%20Anthropol.&doi=10.1086%2F203666&volume=29&issue=3&pages=495-498&publication_year=1988&author=Dennell%2CRW&author=Rendell%2CHM&author=Hailwood%2CE)

77. Zhu, R. _et al._ Early evidence of the genus homo in east asia. _J. Hum. Evol._ **55**(6), 1075–1085 (2008).

    [Article](https://doi.org/10.1016%2Fj.jhevol.2008.08.005) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=18842287) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Early%20evidence%20of%20the%20genus%20homo%20in%20east%20asia&journal=J.%20Hum.%20Evol.&doi=10.1016%2Fj.jhevol.2008.08.005&volume=55&issue=6&pages=1075-1085&publication_year=2008&author=Zhu%2CR)

78. Gowen, K. M. & de Smet, T. S. Testing least cost path (LCP) models for travel time and kilocalorie expenditure: Implications for landscape genomics. _PLoS ONE_ **15**(9), 1–20. [https://doi.org/10.1371/journal.pone.0239387](https://doi.org/10.1371/journal.pone.0239387) (2020).

    [Article](https://doi.org/10.1371%2Fjournal.pone.0239387) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Testing%20least%20cost%20path%20%28LCP%29%20models%20for%20travel%20time%20and%20kilocalorie%20expenditure%3A%20Implications%20for%20landscape%20genomics&journal=PLoS%20ONE&doi=10.1371%2Fjournal.pone.0239387&volume=15&issue=9&pages=1-20&publication_year=2020&author=Gowen%2CKM&author=Smet%2CTS)

79. Walt, S. _et al._ scikit-image: Image processing in Python. _PeerJ_ **2**, e453. [https://doi.org/10.7287/peerj.preprints.336v2](https://doi.org/10.7287/peerj.preprints.336v2) (2014).

    [Article](https://doi.org/10.7287%2Fpeerj.preprints.336v2) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=25024921) [PubMed Central](http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4081273) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=scikit-image%3A%20Image%20processing%20in%20Python&journal=PeerJ&doi=10.7287%2Fpeerj.preprints.336v2&volume=2&publication_year=2014&author=Walt%2CS)

80. Mueller, T. & Fagan, W. Search and navigation in dynamic environments—From individual behaviors to population distributions. _Oikos_ **117**, 654–664. [https://doi.org/10.1111/j.0030-1299.2008.16291.x](https://doi.org/10.1111/j.0030-1299.2008.16291.x) (2008).

    [Article](https://doi.org/10.1111%2Fj.0030-1299.2008.16291.x) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Search%20and%20navigation%20in%20dynamic%20environments%E2%80%94From%20individual%20behaviors%20to%20population%20distributions&journal=Oikos&doi=10.1111%2Fj.0030-1299.2008.16291.x&volume=117&pages=654-664&publication_year=2008&author=Mueller%2CT&author=Fagan%2CW)

81. Bastille-Rousseau, G., Douglas-Hamilton, I., Blake, S., Northrup, J. & Wittemyer, G. Applying network theory to animal movements to identify properties of landscape space use. _Ecol. Appl._ **28** (2018). [https://doi.org/10.1002/eap.1697](https://doi.org/10.1002/eap.1697).

82. Michelot, T., Langrock, R. & Patterson, T. moveHMM: An R package for the statistical modelling of animal movement data using hidden Markov models. _Methods Ecol. Evol._ **7** (2016). [https://doi.org/10.1111/2041-210X.12578](https://doi.org/10.1111/2041-210X.12578) .

83. Benhamou, S. How many animals really do the Lévy Walk. _Ecology_ **88**, 1962–9. [https://doi.org/10.1890/06-1769.1](https://doi.org/10.1890/06-1769.1) (2007).

    [Article](https://doi.org/10.1890%2F06-1769.1) [PubMed](http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Abstract&list_uids=17824427) [Google Scholar](http://scholar.google.com/scholar_lookup?&title=How%20many%20animals%20really%20do%20the%20L%C3%A9vy%20Walk&journal=Ecology&doi=10.1890%2F06-1769.1&volume=88&pages=1962-9&publication_year=2007&author=Benhamou%2CS)

84. Turchin, P. _Quantitative Analysis of Movement: Measuring and Modeling Population Redistribution of Plants and Animals_ (Sinauer Associates, Sunderland, 1998).

    [Google Scholar](http://scholar.google.com/scholar_lookup?&title=Quantitative%20Analysis%20of%20Movement%3A%20Measuring%20and%20Modeling%20Population%20Redistribution%20of%20Plants%20and%20Animals&publication_year=1998&author=Turchin%2CP)

85. Lieberman, D. E. _The Story of the Human Body: Evolution, Health, and Disease_ (Pantheon Books, New York, 2013).

    [Google Scholar](http://scholar.google.com/scholar_lookup?&title=The%20Story%20of%20the%20Human%20Body%3A%20Evolution%2C%20Health%2C%20and%20Disease&publication_year=2013&author=Lieberman%2CDE)

86. Braun, D. _et al._ Early hominin diet included diverse terrestrial and aquatic animals 1.95 Ma in East Turkana, Kenya. _Proc. Natl. Acad. Sci. USA_ **107**, 10002–7 (2010). [https://doi.org/10.1073/pnas.1002181107](https://doi.org/10.1073/pnas.1002181107).

87. O’Connor, S., Louys, J., Kealy, S. & Samper Carro, S. C. Hominin dispersal and settlement east of huxley’s line: The role of sea level changes, island size, and subsistence behavior. _Curr. Anthropol._ **58**(S17), S567–S582 (2017).

88. Macaulay, V. _et al._ Single, rapid coastal settlement of asia revealed by analysis of complete mitochondrial genomes. _Science (New York, N.Y.)_ **308**, 1034–6 (2005). [https://doi.org/10.1126/science.1109792](https://doi.org/10.1126/science.1109792).


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## Acknowledgements

Authors are grateful to Nandang from LIPI Geotek for his continued involvement in the field, to the people of Sangiran for their assistance, and to the Museum Situs Manusia Purba Sangiran and Tanto for their guidance. The authors thank J. Arief for his help in the field, F. Semah and C. Falguères for discussions, L. Léanni for her valuable assistance during chemical treatments and ICP-OES measurements, and D. Daddi-Addoun for her support. J. Louys and two anonymous reviewers greatly help improving the manuscript. Our colleague and friend Didier L. Bourlès, who passed away in 2021, instigated this study.

## Funding

ASTER AMS national facility (CEREGE) is supported by INSU/CNRS, ANR “Projets thématiques d’excellence”, and IRD. National Geographic Society and CNRS-INSU provided financial support.

## Author information

Author notes

1. A list of authors and their affiliations appears at the end of the paper.


### Authors and Affiliations

1. ISTerre, CNRS, IRD, Univ. Grenoble Alpes, 38000, Grenoble, France

Laurent Husson, Swann Zerathe, Sonny Aribowo & Julien Carcaillet

2. School of Geosciences, The University of Sydney, Sydney, NSW, 2006, Australia

Tristan Salles & Claire Mallard

3. CEREGE, Aix-Marseille Université CNRS-IRD-Collège de France-INRAE, Technopôle de l’Environnement Arbois-Méditerrannée, 13545, Aix-en-Provence, France

Anne-Elisabeth Lebatard, Régis Braucher, Didier Bourlès, Georges Aumaitre, Didier Bourlès & Karim Keddadouche

4. Research Center for Archaeometry, National Research and Innovation Agency (BRIN), Jakarta, Indonesia

Sofwan Noerwidi

5. Research Center for Geological Disasters, National Research and Innovation Agency (BRIN), Bandung, Indonesia

Sonny Aribowo & Danny H. Natawidjaja


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### Consortia

### ASTER team

- Georges Aumaitre
- , Didier Bourlès
-  & Karim Keddadouche

### Contributions

L.H., T.S., A.-E.L., D.B. designed the study, T.S., A.-E.L., L.H., and ASTER team conducted the experiments, L.H., T.S., A.-E.L., S.Z., R.B., S.N., S.A., C.M., J.C., D.H.N. analyzed the results. All authors reviewed the manuscript.

### Corresponding author

Correspondence to
[Laurent Husson](mailto:laurent.husson@univ-grenoble-alpes.fr).

## Ethics declarations

### Competing interests

The authors declare no competing interests.

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## Supplementary Information

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Supplementary Video 1.

Supplementary Video 2.

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### Cite this article

Husson, L., Salles, T., Lebatard, AE. _et al._ Javanese _Homo erectus_ on the move in SE Asia _circa_ 1.8 Ma.
_Sci Rep_ **12**, 19012 (2022). https://doi.org/10.1038/s41598-022-23206-9

[Download citation](https://citation-needed.springer.com/v2/references/10.1038/s41598-022-23206-9?format=refman&flavour=citation)

- Received: 02 July 2022

- Accepted: 26 October 2022

- Published: 08 November 2022

- DOI: https://doi.org/10.1038/s41598-022-23206-9


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