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Human dispersal and plant processing in the Pacific 55 000–50 000 years ago

Published online by Cambridge University Press:  13 August 2024

Dylan Gaffney*
Affiliation:
School of Archaeology, University of Oxford, UK Archaeology Programme, University of Otago, Dunedin, New Zealand
Daud A. Tanudirjo*
Affiliation:
Department of Archaeology, Universitas Gadjah Mada, Yogyakarta, Indonesia
Erlin Novita Idje Djami
Affiliation:
Pusat Riset Arkeologi Prasejarah dan Sejarah, Badan Riset dan Inovasi Nasional, Jayapura, Indonesia
Zubair Mas'ud
Affiliation:
Pusat Riset Arkeologi Prasejarah dan Sejarah, Badan Riset dan Inovasi Nasional, Makassar, Indonesia
Abdul Razak Macap
Affiliation:
Balai Pelestarian Kebudayaan Wilayah XXIII, Manokwari, Indonesia
Tristan Russell
Affiliation:
Archaeology Programme, University of Otago, Dunedin, New Zealand Southern Pacific Archaeological Research, University of Otago, Dunedin, New Zealand
Moses Dailom
Affiliation:
Kampung Warsambin, Warsambin, Indonesia
Yulio Ray
Affiliation:
Department of Archaeology, Universitas Gadjah Mada, Yogyakarta, Indonesia
Thomas Higham
Affiliation:
Department of Evolutionary Anthropology, University of Vienna, Austria
Fiona Bradshaw
Affiliation:
School of Archaeology, University of Oxford, UK
Fiona Petchey
Affiliation:
Waikato Radiocarbon Dating Laboratory, University of Waikato, New Zealand
S. Anna Florin
Affiliation:
School of Archaeology and Anthropology, Australian National University, Canberra, Australia
Patrick Roberts
Affiliation:
Department of Archaeology, Max Planck Institute of Geoanthropology, Jena, Germany isoTROPIC Research Group, Max Planck Institute for Geoanthropology, Jena, Germany
Mary Lucas
Affiliation:
Department of Archaeology, Max Planck Institute of Geoanthropology, Jena, Germany
Monica Tromp
Affiliation:
Archaeology Programme, University of Otago, Dunedin, New Zealand Southern Pacific Archaeological Research, University of Otago, Dunedin, New Zealand
Karen Greig
Affiliation:
Archaeology Programme, University of Otago, Dunedin, New Zealand Southern Pacific Archaeological Research, University of Otago, Dunedin, New Zealand
Hermine Xhauflair
Affiliation:
Archaeological Studies Program, University of the Philippines, Quezon City, Philippines
Alvaro Montenegro
Affiliation:
Department of Geography, The Ohio State University, Columbus, USA
Robert Hall
Affiliation:
Department of Earth Sciences, Royal Holloway University, London, UK
Clara Boulanger
Affiliation:
Department of Modern Society and Civilization, Minpaku Museum of Ethnology, Osaka, Japan Japan Society for the Promotion of Science, Tokyo, Japan Histoire Naturelle de l'Homme Préhistorique, Muséum National d'Histoire Naturelle, Paris, France
Rintaro Ono
Affiliation:
Department of Modern Society and Civilization, Minpaku Museum of Ethnology, Osaka, Japan
Annette Oertle
Affiliation:
Human Evolution and Archaeological Sciences, University of Vienna, Austria
Denis Scholz
Affiliation:
Institute for Geosciences, Johannes Gutenberg University of Mainz, Germany
Megan Spitzer
Affiliation:
Department of Vertebrate Zoology, Smithsonian Institute, Washington D.C., USA
Katherine Szabo
Affiliation:
Pre-Construct Archaeology, Cambridge, UK
Irene Bertelli
Affiliation:
Department of Chemistry and Industrial Chemistry, University of Pisa, Italy
Erika Ribechini
Affiliation:
Department of Chemistry and Industrial Chemistry, University of Pisa, Italy
Simon Haberle
Affiliation:
School of Culture, History and Language, Australian National University, Canberra, Australia
*
*Authors for correspondence ✉ [email protected] & [email protected]
*Authors for correspondence ✉ [email protected] & [email protected]
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Abstract

The dynamics of our species’ dispersal into the Pacific remains intensely debated. The authors present archaeological investigations in the Raja Ampat Islands, north-west of New Guinea, that provide the earliest known evidence for humans arriving in the Pacific more than 55 000–50 000 years ago. Seafaring simulations demonstrate that a northern equatorial route into New Guinea via the Raja Ampat Islands was a viable dispersal corridor to Sahul at this time. Analysis of faunal remains and a resin artefact further indicates that exploitation of both rainforest and marine resources, rather than a purely maritime specialisation, was important for the adaptive success of Pacific peoples.

Type
Research Article
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Copyright
Copyright © The Author(s), 2024. Published by Cambridge University Press on behalf of Antiquity Publications Ltd

Introduction

When Homo sapiens dispersed from Eurasia to the Pacific they moved through the islands of Wallacea to reach Sahul (the Pleistocene continent connecting New Guinea with Australia) (Figure 1). To facilitate these journeys, humans devised seafaring technologies and crossed several biogeographic thresholds, pushing beyond the Afro-Eurasian landmass for the first time into increasingly small and faunally depauperate islands (O'Connell & Allen Reference O'Connell and Allen2015; Shipton et al. Reference Shipton, O'Connor and Kealy2021). These dispersals encouraged major behavioural adaptations—not seen among other hominin species such as Homo erectus, Homo floresiensis and Homo luzonensis—as our species learned to live among, and transform, these tropical islands (O'Connor et al. Reference O'Connor, Louys, Kealy and Carro2017; Gaffney Reference Gaffney2022). The timing and route of these movements remain unresolved and highly contested (Kealy et al. Reference Kealy, Louys and O'Connor2018; Norman et al. Reference Norman, Inglis, Clarkson, Faith, Shulmeister and Harris2018; Bird et al. Reference Bird2019; Bradshaw et al. Reference Bradshaw2019; Allen & O'Connell Reference Allen and O'Connell2020).

Figure 1. Northern and southern routes through Wallacea, showing Mololo Cave on Waitanta. Inset A) seafaring simulations and chance of successful landing on Waitanta from nearby landmasses, based on averages taken from 1.5kt paddling at -30m and -50m sea levels (Table S6); Inset B) Waitanta shoreline and Mololo Cave at 50 000 years ago and at the Last Glacial Maximum (20 000 years ago) based on multibeam and GEBCO bathymetry data (figure by authors).

Archaeological evidence from Madjedbebe, a rockshelter in Arnhem Land, northern Australia, suggests that humans initially arrived in Sahul via a southern route that included the Lesser Sunda Islands, around 65–60ka (thousand years ago) (Clarkson et al. Reference Clarkson2017). Along the southern route, evidence of H. sapiens at Liang Bua (Flores Island), Makpan (Alor) and Laili, Asitau Kuru and Lene Hara (Timor), however, stands at maximally 44ka (Hawkins et al. Reference Hawkins2017; Sutikna et al. Reference Sutikna2018; Shipton et al. Reference Shipton, O'Connor, Jankowski, O'Connor-Veth, Maloney, Kealy and Boulanger2019; Kealy et al. Reference Kealy2020). This disparity has led some archaeologists to argue that the northern Australian dates are erroneous and that a later arrival to Sahul after 50ka, also involving a northern equatorial route, is more convincing (O'Connell et al. Reference O'Connell2018). Although cave art from the Maros Karst sites (Sulawesi) (Brumm et al. Reference Brumm2021) and Lubang Jeriji Saléh (Borneo) (Aubert et al. Reference Aubert2018) include figurative hunting scenes that date to at least 45ka, the earliest sites along the small islands of the northern island chain—Golo (Gebe Island), Liang Lemdubu (Aru Islands) and Liang Sarru (Talaud Islands)—date to less than 40ka (Tanudirjo Reference Tanudirjo2001; O'Connor et al. Reference O'Connor, Aplin, Szabó, Pasveer, Veth, Spriggs, O'Connor, Spriggs and Veth2005; Bellwood et al. Reference Bellwood, Irwin, Tanudirjo, Nitihaminoto, Siswanto, Bowdery and Bellwood2019a). Determining the nature of H. sapiens migrations to the Pacific is crucial for understanding how our species diversified outside of Africa, adapted to the challenges and opportunities of these novel environments, including insularity and tropical forest cover, and interacted with other hominin species present in the region. Northern dispersals may have brought H. sapiens into close contact with Denisovan hominins, perhaps present around Sulawesi (Carlhoff et al. Reference Carlhoff2021), or Homo luzonensis in the Philippines (Détroit et al. Reference Détroit2019), while southern movements may have seen our species cohabit islands, however briefly, with Homo floresiensis (Sutikna et al. Reference Sutikna2016).

Here, we report the first excavations from the Raja Ampat Islands, off the north-west coast of New Guinea, a key island group along the ‘northern route’ to Sahul. We first provide bathymetric (ocean floor depth) reconstructions of island size and voyage simulations for the northern entry to Sahul, and then detail excavations, radiometric dating and multidisciplinary archaeological and palaeoecological analyses at the site of Mololo Cave on Waigeo Island to clarify when and how humans moved along the northern route into Sahul, prior to 50ka.

Coastline reconstruction and seafaring modelling

From 65–50ka, eustatic sea level fluctuated between approximately 30–50m below present levels (Kealy et al. Reference Kealy, Louys and O'Connor2017). Multibeam imagery demonstrates that Waigeo and Batanta islands were, at times, connected by a shallow platform (Figures 1B & S1, see also online supplementary material (OSM)). We call this palaeo-island Waitanta (wai = water, tanta = that extends in front of your eyes). This island was separated from Sahul by the deep Sagewin Strait throughout the Pleistocene, even during the Last Glacial Maximum (LGM). As such, humans would have needed watercraft to move between Waitanta and Sahul. At 50ka, the Sagewin Strait was 5–6km wide on average and 2.5km at its minimal distance, and at the LGM the average width was about 5km with a minimal crossing distance of 1.5km.

Our novel seafaring simulations show that eastward-moving crossings were most easily achieved between Obi–Kofiau–Waitanta, even with minimal propulsion (Figures 1A & S2, Tables S1–S9, see OSM for simulation methods and results). Successful arrivals on Waitanta by paddling at 1.5kt (0.77m/s) were nearly guaranteed when moving from Kofiau, Gebe and Halmahera in the west or Sahul in the east. Trips from Sahul would be quickest, followed by Gebe, Kofiau and Halmahera, lasting 2–3 days. Crossings from Talaud and Morotai to the north were highly unlikely. Even in suboptimal weather, movements between Waitanta and Sahul across the Sagewin Strait were safe for rafts paddled at 1.5kt but were also possible with slower paddling during ideal conditions (Table S10).

Mololo Cave excavations and chronostratigraphy

Mololo Cave (0°18′21.0″S, 130°55′01.4″E; Raja Ampat Archaeological Project site code: WAI-1) lies within the Rabia Strait, leading into Waigeo Island's Mayalibit Bay (Figures 2A & S3, see also OSM). A transect across Rabia Strait indicates it is maximally 46m deep and would have been a valley system during the Pleistocene, becoming inundated with seawater after c. 9.5ka; at 50ka, Mololo was located about 15km inland. Formed from Miocene limestone, Mololo includes an outer chamber, exposed to daylight owing to roof collapse, and a dark inner chamber, home to several bat colonies (Figure 2). A 1 × 1m test pit had been excavated by the Centre for Papuan Archaeology near the cave entrance to 0.8m deep in 2012. Excavations in 2018–2019, reported here, targeted three parts of the cave system: Area 1 near the cave entrance; Area 2 on a flat, elevated space in the outer chamber; and Area 3 at the edge of the inner chamber.

Figure 2. Mololo Cave system showing excavation units. Inset A) location of Mololo Cave at the entrance to Mayalibit Bay. Inset B) transect of entrance to Mololo Cave, relative to mean sea level (MSL) (figure by authors).

Trench 1 was excavated to a depth of 2.57m (Figure 3A, Table S11). The upper stratigraphy consists of interleaved clays, midden material and fire ash, indicating recurring frequentation between c. 15 and 2.1ka (Figure 4, Table S12). In three instances, charcoal with a Holocene age underlies terminal Pleistocene strata, possibly associated with bioturbation or disturbance during the production of hearths and large bone middens. Below this is undisturbed, indurated guano, likely deposited by small bats roosting on the roof above the area where the trench is located. Charcoal incorporated within one indurated context dates to c. 44–43ka, but it is unclear if it is anthropogenic. Owing to its small size, the charcoal was prepared with acid/base/acid rather than acid-base-oxidation, a pre-treatment that can produce younger-than-actual dates for material over about 20ka (Higham et al. Reference Higham, Barton, Turney, Barker, Bronk-Ramsey and Brock2009). As such, 43ka is likely a minimum possible age for these indurated contexts. They overlie looser guano associated with numerous small bat bones that could not be radiocarbon dated owing to poor collagen preservation (Table S13). At the base of the excavation, uranium-series dating indicates that sedimentary calcite encrusted on limestone bedrock dates to 51ka, and coral gravel (Porites sp.) overlying the bedrock dates to 125ka (Marine Isotope Stage 5e, the last interglacial) and 200ka (Marine Isotope Stage 7). This indicates Area 1 was originally submerged and has been subsequently uplifted to its present location (Table S14).

Figure 3. Mololo Cave stratigraphy described by context number and layer. A) Trench 1, south and west walls; B) Trench 2, south and west walls (figure by authors).

Figure 4. Calibrated terrestrial (charcoal and resin) radiocarbon dates from Mololo Cave, organised by stratigraphic location. Red outlines indicate the calibration threshold at 55 000 years ago and periods of inversion in the radiocarbon dates. The lower image shows the global oxygen isotope data reflecting changing temperatures (low values are higher temperatures) (Siddall et al. Reference Siddall, Rohling, Almogi-Labin, Hemleben, Meischner, Schmelzer and Smeed2003). LGM = Last Glacial Maximum; YD = Younger Dryas; H = Heinrich event; MIS = Marine Isotope Stage (figure by authors).

Trench 2 extended to 1.68m deep, including clays and two Late Holocene middens in the upper deposit (Figure 3B, Table S15). Area 2 was repeatedly used by humans between c. 9.1 and 2.1ka (Figure 4, Table S12). There is inversion in layer 6 with Late Holocene charcoal underlying Early Holocene strata, possibly owing to bioturbation. The lower part of the excavation transitions from highly indurated clay into flowstone and limestone bedrock. In layer 7 (context 088; a highly indurated, brown clay), a tree resin artefact was directly radiocarbon dated to greater than 55–49.6ka. Many plant resins begin to polymerise upon burial and are highly resistant to post-depositional contamination (Kocsis et al. Reference Kocsis2020); the carbon in the Mololo resin is exceptionally well-preserved and provided a pretreatment yield of 94.5% (%C = 65.7). The date pushes the upper limit of the IntCal20 calibration curve (Reimer et al. Reference Reimer2020), and the maximum range lies sometime before 55ka. Fruit bat bones from layer 7 could not be radiocarbon dated owing to a lack of collagen (Table S13).

Test pit 3 was excavated to 1.4m and comprised several clay layers overlying dark clay mixed with guano (Table S16). Below this a soft brown clay overlies limestone bedrock. Sparse charcoal and archaeological material in the upper clay layers indicate frequentation of the dark inner chamber by at least 8.6ka (Table S12).

Tree resin processing

The tree resin artefact provides evidence for complex plant processing during early human dispersal (Figure 5A–D). The artefact is rectilinear, totally unlike globular, naturally formed resins, indicating that it was collected by humans directly from the tree. Scanning electron microscope (SEM) analysis suggests it was produced in a multi-step sequence. The resin is 13.5mm wide and 5mm thick, with planilateral dorsal, ventral and lateral surfaces, which suggests it pooled in a small rectilinear channel, possibly cut into the trunk of the tree itself. Curved parallel striations are present on the ventral surface and SEM analysis of the surface microtopography shows internal micro-striations reflective not of ripple marks but scraping on the resin itself. Minor scraping marks are also present on the dorsal surface, along with chipping along the edge, which occurred after hardening, perhaps when extracting the resin from the channel in the tree. Ethnographic observation on Waigeo indicates that people cut the bark of trees and allow the resin to accumulate before they return to extract the hardened resin one or more days later (see OSM). Concave and convex snap fractures at each end imply that, following removal from the channel, the resin was manually snapped into a 19mm long piece (similar to snaps on stone tools such as truncated burins (Hilbert et al. Reference Hilbert2018)) or broken after discard owing to footfall in the cave (as seen on silicious stone artefacts (Hiscock Reference Hiscock1985)).

Figure 5. Tree resin artefact (WAI-1-1024) from trench 2, directly dated to >55 000–49 620 cal BP: A) lateral side with rough surface texture; B) dorsal side with flat surface, chipping and minor scraping marks; C) ventral side showing parallel marks, possibly from scraping or the imprint of a scraped surface; D) lateral side with rough surface texture showing snaps. E illustrates a provisional hypothesis for the sequence of resin artefact production (figure by authors).

Gas chromatography/mass spectrometry (GC/MS) and pyrolysis gas chromatography/mass spectrometry (Py-GC/MS) identify the artefact as a triterpenoid resin from a flowering tree or shrub (Angiosperm), ruling out a conifer source such as Agathis labillardierei, known to have been used in the more recent past on Waigeo (Figures 6 & S4). A triterpenoid resin also used on Waigeo, Vatica rassak or a similar species from the Dipterocarpaceae family, shares chemical similarities but is not a direct match for the Mololo artefact (Figure 6, Table S17, see also OSM). Although there are numerous ethnographic uses of resins in Island Southeast Asia (Bradshaw et al. Reference Bradshaw, Stern, Thompson, Barker and Farr2013), the Mololo artefact does not preserve evidence for its intended function.

Figure 6. Mass spectra of Mololo archaeological resin compared with reference specimens. A–E are total ion chromatograms based on GC/MS: A) Mololo archaeological resin (WAI-1-1024); B) ethnographic ‘damar merah’ from Waigeo (Ref_D2); C) Vatica rassak (Ref_R3_K57); D) ethnographic ‘damar putih’ from Waigeo (Ref_D3); E) Agathis labillardierei (Ref_R1_K08). F–I are pyrograms based on Py-GC/MS: F) Mololo archaeological resin (WAI-1-1024 GCMS1); G) Mololo archaeological resin (WAI-1-1024 GCMS2); H) Canarium salomonense (Ref_R4_K21); I) Prumnopitys ladei (Ref_R5_S19) (figure by authors).

No lithic pieces from Late Pleistocene contexts are unequivocally manually produced or used, but one siliceous limestone object from trench 2, layer 7 may be a core (Figure 7, Table S18, see also OSM). One anthropic limestone flake and two possible flakes are present in trench 2, layer 6, but their dating is imprecise. Moreover, there is no evidence for resin being used to haft any of the stones in the cave. Even in the Late Holocene layers, where cultural material was abundant, only six small radiolarian chert and five limestone artefacts were recovered. The sparseness of stone artefacts is characteristic of north-eastern Wallacea and north-west New Guinea, where many tools were likely made from organic rather than lithic materials (Bellwood et al. Reference Bellwood, Irwin, Tanudirjo and Bellwood2019b); alternatively, lithics may have been discarded offsite or in another part of the cave.

Figure 7. Selection of Mololo lithic artefacts and possible artefacts. A) WAI-1-183 (trench 2, layer 3)—flake with impact point indicated by converging hackles (marked by arrows), clear platform, ventral face with bulb, and dorsal face with a rougher texture. Edges are post-depositionally altered preventing the detection of possible use-wear; B) WAI-1-2162 (trench 1, layer 3) – flake with platform, partly cortical dorsal face and one removal. No distal end or trace of utilisation; C) WAI-1-701-B-ii (trench 2, layer 6)—smooth ventral face (iv) and platform, and dorsal face with smooth surface within two previous removals compared with rougher dorsal face (i). The edge of fracture on ventral face (pink) presents rounding and micro-polish (ii) likely resulting from scraping motion. The sharp edge (blue) does not show utilisation (iii); D) WAI-1-701-B-i (trench 2, layer 6)—flake with cortical convex face and fresher flatter face possibly indicating knapping, potential platform, and long irregular sharp edge with no clear evidence of utilisation; E) WAI-1-700 (trench 2, layer 6)—possible small flake with platform, smooth face with small bulb, and long possible flake removal on opposite side. Micro-scars on distal part of right ventral edge, possibly use-related or taphonomic; F) WAI-1-1005-C (trench 2, layer 7)—fragment with two possible concave flake removals, no evidence of use-wear (figure by authors).

Pleistocene environments and subsistence

The Pleistocene fauna from trench 1 reflect natural deposition and are characteristic of cave sequences in the tropics; small insectivorous bats (including Hipposideros maggietaylorae and Miniopterus sp.), medium-sized fruit bats (Dobsonia sp.), frogs (Anura) and rodents (Muridae) (Table S19 & OSM). The remains of insectivorous bats include Hipposideros, a taxon that forages in closed-canopy forests, but Miniopterus, a genus that forages on forest edges, is also present, indicating some clearings around Mololo. Post-depositional alteration such as polish, brown discolouration and minor pitting on these bones suggest dead animals were incorporated with acidic guano, with some perhaps accumulated by avian or marsupial predators.

The Late Pleistocene fauna from trench 2, closely associated with or underlying the resin artefact of layer 7, includes forest species, some likely hunted by humans. Most specimens are pteropodids (number of identified specimens (NISP) = 35), probably all deriving from New Guinea bare-backed fruit bats (Dobsonia magna), a species commonly hunted around northern Sahul (see Hawkins et al. Reference Hawkins, O'Connor and Kealy2016; Tables S20 & S21, OSM); only a single insectivorous bat bone was identified. Of the four Dobsonia magna teeth identified from layers 7 and 8, none display tooth wear, indicating the animals were young adults at death and unlikely to have died naturally. Other specimens suggestive of human predation include those from a ground-dwelling bird (Phasianidae) and mid-sized marsupials, while the New Guinea spiny rat (Rattus praetor) could have been deposited by human or non-human predators. Attempted species identification of non-diagnostic bones from the Pleistocene deposits based on mass spectrometry failed owing to a lack of collagen. Bleached colouration and a lack of pitting on most specimens indicate the animals were deposited onto clays exposed to light rather than into guano. Although no cutmarks are found on the Late Pleistocene fauna, neither is there evidence of consumption by non-human predators (large fruit bats and medium-sized terrestrial birds and marsupials are not known to be common targets of raptors around New Guinea).

One sea urchin (Echinodermata) spine was recovered from trench 1, layer 10, suggestive of manual importation from the coast by humans before 43ka (Figure 8). Another spine with asymmetrical abrasion along one side was found in trench 2, layer 6, dating to between >55 and 7.5ka (Figure 8). It is possible that this has been worked by humans, but recrystallisation has removed the original surface features. A tooth from a fast-swimming, carnivorous fish (Scombroidei sp. possibly Sphyraena cf. barracuda or Gymnosarda unicolor) was also found in trench 2, layer 6. The black polished surface of the fish tooth closely matches discolouration of terrestrial long bones from trench 2, layers 7 and 8, and suggests it dates to the Late Pleistocene. Unfortunately, diagenesis means these specimens are not suitable for radiocarbon dating.

Figure 8. Aquatic resources at Mololo. A) Hyridella cf. misoolensis (WAI-1-1022) from Mololo Cave, trench 2, layer 5. Note delamination and recrystallisation of aragonite shown in inset SEM micrograph. B) Sea urchin specimens from Mololo Cave: WAI-1-2086-A-xiii, small Echinoderm spine from trench 1, layer 5; WAI-1-1169-E, large Echinoderm spine from trench 1, layer 10; WAI-1-1551-A-ii, large Echinoderm spine with abrasion from trench 1, layer 5A. C) Sea urchin spine (WAI-1-696-B) with an edge alteration from Mololo Cave, trench 2, layer 6. D) Scombroidei sp. teeth from Mololo Cave: WAI-1-773-B, subfossilised tooth from trench 2, layer 6; WAI-1-2058, tooth from trench 1, layer 5A. Both specimens preserve diagnostic lateral margins and concave shape at the apical end (figure by authors).

One hand-collected fragment of charred monocotyledonous stem tissue from trench 1, layer 6, likely deriving from an underground storage organ, was directly dated to 13–12.8ka (Figure 9, Table S22, OSM). Owing to fine grain physical erosion, no pollen or phytoliths were identified in Late Pleistocene sediments, but contexts older than 8.2ka preserve fern phytoliths. In contrast, those from Late Holocene contexts contain pine pollen, fern, grass, palm and possible breadfruit phytoliths, and sponge spicules (Figure 9, Table S23, OSM), the latter incorporated into the site at the same time as marine shell.

Figure 9. Archaeobotanical remains from Mololo. A–D are SEM images of charred monocotyledonous stem tissue from trench 1, layer 5–6 (WAI-1-1161) and layer 6 (WAI-1-1236), vb = vascular bundle. A & B) longitudinal section of WAI-1-1161, with a vascular bundle exposed longitudinally within a parenchyma matrix, B shows a close-up of the vascular bundle; C & D) transverse section of WAI-1-1236, with several vitrified vascular bundles, within a parenchyma matrix, D shows a close up of vitrified vascular bundles. E–J are biogenic silica recovered from Mololo Cave sediment samples: E) sponge spicule from trench 1, layer 2 (WAI-1-1612-C); F) sponge spicule from trench 1, context 011 (WAI-1-1379-B); G) sponge asterone microsclere from trench 1, layer 3 (WAI-1-1422-B); H) sponge spicule from trench 1, layer 3 (WAI-1-1422-B); I) fern spore from trench 2, context 098 (WAI-1-424-B); J) an elongate entire and a blocky phytolith, likely from grasses, from trench 2, layer 2 (WAI-1-606-B) (figure by authors).

The wide variability in carbon stable isotope (δ13C) values of D. magna enamel during the Late Pleistocene indicates their selection of diverse forest patches for foraging, with open forests, woodlands and closed-canopy forests all available (Figure S5, Table S24, OSM). Isotope data indicate the expansion of dense rainforests after the Last Glacial Maximum, c. 14–11ka (Figure 10). Enamel δ13C values for rodents (Muridae) provide a more localised signal of Late Pleistocene forest cover and indicate there were closed-canopy forests near Mololo (Figure S5, Table S24, OSM). Patchy Late Pleistocene forest mosaics were probably produced by minimally compressed temperatures, but clearances by humans would have further encouraged forest gaps.

Figure 10. Reconstruction of Mayalibit Valley ecology from the Late Pleistocene to the Late Holocene alongside fruitbat and rodent δ13C and δ18O isotope results. The Late Pleistocene was characterised by patches of open and closed tropical forest, followed by expansion of closed canopy forests in the Terminal Pleistocene. The infilling of Mayalibit Bay in the Early Holocene created open areas near Mololo, possibly further encouraged by clearances. Diversification of forest patches in the Late Holocene may be caused by burning (figure by authors).

Discussion

The Mololo investigations provide critical, albeit sparse, evidence for occupation along the northern equatorial route to Sahul before 50ka (possibly >55ka). H. sapiens probably produced this archaeological record, moving by raft/boat through the humid tropics. However, it should be noted that based on modern population genetic evidence (Jacobs et al. Reference Jacobs2019) it is not currently possible to exclude the possibility that Denisovans or people who carried both H. sapiens and Denisovan ancestry—having admixed in the islands of Wallacea or continental Eurasia—migrated along this route.

Based on our evidence, the northern route to what is today New Guinea through Waitanta is a viable corridor for the early colonisation of Sahul. Primarily owing to chronometric imprecision at Raja Ampat, Timorese and Australian sites, the timing of people's movements through the northern route, and how those relate to dispersals along the southern route, needs clarification. At Madjedbebe in northern Australia, the minimum estimate for initial occupation is 59.3ka, although it may begin several millennia earlier (Clarkson et al. Reference Clarkson2017). Occupation of other Australian sites probably post-dates 50ka but may begin as early as 52ka (Norman et al. Reference Norman2022). Similarly, the earliest known sites from the mainland of New Guinea, in the Ivane Valley and Huon Peninsula, and on offshore islands such as New Ireland and New Britain, all post-date 50ka and most post-date 45ka (Summerhayes et al. Reference Summerhayes, Field, Shaw and Gaffney2017). It is likely that Waitanta was first frequented by those living further west in Wallacea, but it remains possible that humans initially entered Sahul via Australia and rapidly moved north-west, arriving at Waitanta from what is today the Bird's Head Peninsula of New Guinea. The dynamics of these movements therefore need explication, especially how humans dispersed along these different chains, whether either of these island groups hosted ‘back-migrations’ from Sahul after initial colonisation, and whether there might have been latitudinal movement between these chains. Ongoing excavations at Mololo, alongside luminescence and radiocarbon dating of the lowest deposits, aim to provide clarity about the earliest human occupation of the site.

The earliest humans on Waitanta adapted to insular tropical forests by using native vegetation and animals. The presence of multi-stage resin processing at Mololo supports evidence from Madjedbebe (Australia) and Niah (Borneo) that indicates H. sapiens were engaging with diverse and complex plant processing techniques (Barker et al. Reference Barker2007; Bradshaw et al. Reference Bradshaw, Stern, Thompson, Barker and Farr2013; Florin et al. Reference Florin2020). Although resins have been used since the late Middle Pleistocene (Mazza et al. Reference Mazza2006; Wadley et al. Reference Wadley, Hodgskiss and Grant2009), the Mololo resin is the earliest known use of tree exudates by our species outside of Africa. Alongside its possible use as a fuel source, resin could be used as an adhesive to haft stone and bamboo onto composite organic tools, and in the construction of boats (Allen & O'Connell Reference Allen and O'Connell2020). Watercraft were necessary to transport scores of people between Wallacea and Sahul, creating biologically viable founder populations of around 1000 individuals over several centuries, as indicated by demographic modelling (Bradshaw et al. Reference Bradshaw2019).

Despite Waitanta being faunally depauperate, hunters targeted small native animals such as terrestrial birds, marsupials and perhaps fruit bats in the rainforest and may have transported marine resources from the coast. This study therefore contributes to debate about H. sapiens dietary breadth (O'Connell & Allen Reference O'Connell and Allen2012; Florin Reference Florin2022), showing that humans along the northern route were not exclusively marine specialists as argued for those along the southern route 42ka (O'Connor et al. Reference O'Connor, Ono and Clarkson2011; Roberts et al. Reference Roberts2020), but moved between the coast and the interior to access the array of resources offered by island rainforests.

Conclusion

Multi-proxy archaeological and palaeoecological analyses of the Mololo Cave sequence on the palaeo-island of Waitanta provide evidence for the earliest known peopling of the Pacific region >55 000–50 000 years ago. These humans practised complex plant processing and engaged with both coastal and tropical forest ecologies. The Raja Ampat archaeological record provides some of the earliest global evidence for humans exploring rainforests outside Africa and the earliest evidence of our species using small islands. Capacities for adaptive flexibility and environmental transformation likely stimulated human movements into insular rainforests, previously beyond the range of other hominin species. These settings help us to understand the process of cultural and biological diversification generated as our species dispersed around the planet and began to push the boundaries of novel habitats, and how humans have become enmeshed in these ecologies for tens of millennia.

Acknowledgements

Warsambin and Lopintol communities permitted excavation at Mololo Cave and supported fieldwork, Herman Soor (Raja Ampat regency government) and Wolter Gaman (Intelektual Suka Maya) provided assistance during survey. Excavation assistants included Melkion Amberbaken, Olos, Nathanial, Otniel, Otto, Robert, Isak, Demianus, Demersis, Septinus, Peter, Yan, Fernandes and Paul. We thank Gilar, Lazuardi and Yaqin, research assistants in Yogkayarta, Antoine Lourdeau, Pierre Voinchet, Xavier Gallet, Camille Daujeard, Sylvie Beyries and Gilles Durenmath for advice and assistance in lithic analysis, M. Großkopf and M. Weber for U-series laboratory assistance, Cam Webb for botanical advice, Christoph Parsch for zoological advice and Laura Arnold for linguistic advice. We thank two anonymous reviewers for their supportive comments. Fieldwork was carried out under Indonesian research permit 359/SIP/FRP/E5/Dit.KI/X/2018.

Funding statement

Fieldwork: National Geographic Explorer Grant (HJ-058ER-17), Evans Fund Fellowship, University of Cambridge School of the Humanities and Social Sciences Research Grant, Magdalene College Anthropology and Archaeology Fund, Leakey Foundation Research Grant. Radiocarbon dating: Royal Anthropological Institute Horniman and Sutasoma Awards, the Evans Fund, NRCF (NERC) Radiocarbon Fund. Py-GC/MS analysis: Quaternary Research Association Award. Isotope analysis: Max Planck Society. U-series dating: Deutsche Forschungsgemeinschaft (INST 247/889-1 FUGG).

Supplementary material

To view supplementary material for this article, please visit https://doi.org/10.15184/aqy.2024.83.

References

Allen, J. & O'Connell, J.F.. 2020. A different paradigm for the colonisation of Sahul. Archaeology in Oceania 55: 114. https://doi.org/10.1002/arco.5207CrossRefGoogle Scholar
Aubert, M., et al. 2018. Palaeolithic cave art in Borneo. Nature 564: 254–57. https://doi.org/10.1038/s41586-018-0679-9CrossRefGoogle ScholarPubMed
Barker, G., et al. 2007. The ‘human revolution’ in lowland tropical Southeast Asia: the antiquity and behavior of anatomically modern humans at Niah Cave (Sarawak, Borneo). Journal of Human Evolution 52: 243–61. https://doi.org/10.1016/j.jhevol.2006.08.011CrossRefGoogle Scholar
Bellwood, P., Irwin, G., Tanudirjo, D., Nitihaminoto, G., Siswanto, J. & Bowdery, D.. 2019a. Investigations on Gebe Island, in Bellwood, P. (ed.) The Spice Islands in prehistory: archaeology in the Northern Moluccas, Indonesia: 1544. Canberra: Australia National University Press.Google Scholar
Bellwood, P., Irwin, G. & Tanudirjo, D.. 2019b. Lithic and other non-ceramic artefacts, in Bellwood, P. (ed.) The Spice Islands in prehistory: archaeology in the Northern Moluccas, Indonesia: 107119. Canberra: Australia National University Press.Google Scholar
Bird, M.I., et al. 2019. Early human settlement of Sahul was not an accident. Scientific Reports 9. https://doi.org/10.1038/s41598-019-42946-9CrossRefGoogle Scholar
Bradshaw, C.J.A., et al. 2019. Minimum founding populations for the first peopling of Sahul. Nature Ecology and Evolution 3: 1057–63. https://doi.org/10.1038/s41559-019-0902-6CrossRefGoogle ScholarPubMed
Bradshaw, F., Stern, B. & Thompson, G.. 2013. Archaeological ‘dammar’ resins from the West Mouth, in Barker, G. & Farr, L. (ed.) Archaeological investigations in the Niah Caves, Sarawak: 363–72. Cambridge: McDonald Institute of Archaeological Research.Google Scholar
Brumm, A., et al. 2021. Oldest cave art found in Sulawesi. Science Advances 7. https://doi.org/10.1126/sciadv.abd4648CrossRefGoogle ScholarPubMed
Carlhoff, S., et al. 2021. Genome of a middle Holocene hunter-gatherer from Wallacea. Nature 596: 543–47. https://doi.org/10.1038/s41586-021-03823-6CrossRefGoogle ScholarPubMed
Clarkson, C., et al. 2017. Human occupation of northern Australia by 65,000 years ago. Nature 547: 306–10. https://doi.org/10.1038/nature22968CrossRefGoogle ScholarPubMed
Détroit, F., et al. 2019. A new species of Homo from the Late Pleistocene of the Philippines. Nature 568: 181–86. https://doi.org/10.1038/s41586-019-1067-9CrossRefGoogle ScholarPubMed
Florin, S.A. 2022. The role of plant foods in the evolution and dispersal of early humans: a perspective from across the Wallace Line. Mitteilungen der Gesellschaft für Urgeschichte 30: 1339. https://doi.org/10.51315/mgfu.2021.30002CrossRefGoogle Scholar
Florin, S.A., et al. 2020. The first Australian plant foods at Madjedbebe, 65,000–53,000 years ago. Nature Communications 11. https://doi.org/10.1038/s41467-020-14723-0CrossRefGoogle ScholarPubMed
Gaffney, D. 2022. Peopling island rainforests: global trends from the Early Pleistocene to the Late Holocene. World Archaeology 54: 338–62. https://doi.org/10.1080/00438243.2022.2121316CrossRefGoogle Scholar
Hawkins, S., O'Connor, S. & Kealy, S.. 2016. Late Quaternary hominin-bat (Chiroptera) interactions in the Asia-Pacific. Archaeology in Oceania 51: 717. https://doi.org/10.1002/arco.5084CrossRefGoogle Scholar
Hawkins, S., et al. 2017. Oldest human occupation of Wallacea at Laili Cave, Timor-Leste, shows broad-spectrum foraging responses to Late Pleistocene environments. Quaternary Science Reviews 171: 5872. https://doi.org/10.1016/j.quascirev.2017.07.008CrossRefGoogle Scholar
Higham, T.F.G., Barton, H., Turney, C.S.M., Barker, G., Bronk-Ramsey, C. & Brock, F.. 2009. Radiocarbon dating of charcoal from tropical sequences: results from the Niah Great Cave, Sarawak, and their broader implications. Journal of Quaternary Science 24: 189–97. https://doi.org/10.1002/jqs.1197Google Scholar
Hilbert, Y.H., et al. 2018. Woodworking sites from the Late Paleolithic of South Arabia: functional and technological analysis of burins from Dhofar, Oman. Journal of Archaeological Science: Reports 20: 115–34. https://doi.org/10.1016/j.jasrep.2018.04.010Google Scholar
Hiscock, P. 1985. The need for a taphonomic perspective in stone artefact analysis. Queensland Archaeological Research 2: 8297.CrossRefGoogle Scholar
Jacobs, G.S., et al. 2019. Multiple deeply divergent Denisovan ancestries in Papuans. Cell 177: 1010–21. https://doi.org/10.1016/j.cell.2019.02.035CrossRefGoogle ScholarPubMed
Kealy, S., Louys, J. & O'Connor, S.. 2017. Reconstructing palaeogeography and inter-island visibility in the Wallacean Archipelago during the likely period of Sahul colonization, 65–45 000 years ago. Archaeological Prospection 24: 259–72. https://doi.org/10.1002/arp.1570CrossRefGoogle Scholar
Kealy, S., Louys, J. & O'Connor, S.. 2018. Least-cost pathway models indicate northern human dispersal from Sunda to Sahul. Journal of Human Evolution 125: 5970. https://doi.org/10.1016/j.jhevol.2018.10.003CrossRefGoogle ScholarPubMed
Kealy, S., et al. 2020. Forty-thousand years of maritime subsistence near a changing shoreline on Alor Island (Indonesia). Quaternary Science Reviews 249. https://doi.org/10.1016/j.quascirev.2020.106599CrossRefGoogle Scholar
Kocsis, L., et al. 2020. The Bruneian record of “Borneo Amber”: a regional review of fossil tree resins in the Indo-Australian Archipelago. Earth-Science Reviews 201. https://doi.org/10.1016/j.earscirev.2019.103005CrossRefGoogle Scholar
Mazza, P.P.A., et al. 2006. A new Palaeolithic discovery: tar-hafted stone tools in a European Mid-Pleistocene bone-bearing bed. Journal of Archaeological Science 33: 1310–18. https://doi.org/10.1016/j.jas.2006.01.006CrossRefGoogle Scholar
Norman, K., Inglis, J., Clarkson, C., Faith, J.T., Shulmeister, J. & Harris, D.. 2018. An early colonisation pathway into northwest Australia 70–60,000 years ago. Quaternary Science Reviews 180: 229–39. https://doi.org/10.1016/j.quascirev.2017.11.023CrossRefGoogle Scholar
Norman, K., et al. 2022. Human occupation of the Kimberley coast of northwest Australia 50,000 years ago. Quaternary Science Reviews 288. https://doi.org/10.1016/j.quascirev.2022.107577CrossRefGoogle Scholar
O'Connell, J.F. & Allen, J.. 2012. The restaurant at the end of the universe: modelling the colonisation of Sahul. Australian Archaeology 74: 517. https://doi.org/10.1080/03122417.2012.11681932CrossRefGoogle Scholar
O'Connell, J.F. & Allen, J.. 2015. The process, biotic impact, and global implications of the human colonization of Sahul about 47,000 years ago. Journal of Archaeological Science 56: 7384. https://doi.org/10.1016/j.jas.2015.02.020CrossRefGoogle Scholar
O'Connell, J.F., et al. 2018. When did Homo sapiens first reach Southeast Asia and Sahul? Proceedings of the National Academy of Sciences USA 115: 8482–90. https://doi.org/10.1073/pnas.1808385115CrossRefGoogle ScholarPubMed
O'Connor, S., Aplin, K., Szabó, K., Pasveer, J., Veth, P. & Spriggs, M.. 2005. Liang Lemdubu: a Pleistocene cave site in the Aru Islands, in O'Connor, S., Spriggs, M. & Veth, P. (ed.) The archaeology of the Aru Islands, Eastern Indonesia: 171204. Canberra: Australia National University Press.Google Scholar
O'Connor, S., Ono, R. & Clarkson, C.. 2011. Pelagic fishing at 42,000 years before the present and the maritime skills of modern humans. Science 334: 1117–21. https://doi.org/10.1126/science.1207703CrossRefGoogle ScholarPubMed
O'Connor, S., Louys, J., Kealy, S. & Carro, S.C.S.. 2017. Hominin dispersal and settlement east of Huxley's Line the role of sea level changes, island size, and subsistence behavior. Current Anthropology 58: S567–82. https://doi.org/10.1086/694252CrossRefGoogle Scholar
Reimer, P.J., et al. 2020. The IntCal20 Northern Hemisphere radiocarbon age calibration curve (0–55 cal kBP). Radiocarbon 62: 725–57. https://doi.org/10.1017/RDC.2020.41CrossRefGoogle Scholar
Roberts, P., et al. 2020. Isotopic evidence for initial coastal colonization and subsequent diversification in the human occupation of Wallacea. Nature Communications 11. https://doi.org/10.1038/s41467-020-15969-4CrossRefGoogle ScholarPubMed
Shipton, C., O'Connor, S., Jankowski, N., O'Connor-Veth, J., Maloney, T., Kealy, S. & Boulanger, C.. 2019. A new 44,000-year sequence from Asitau Kuru (Jerimalai), Timor-Leste, indicates long-term continuity in human behaviour. Archaeological and Anthropological Sciences 11: 5717–41. https://doi.org/10.1007/s12520-019-00840-5CrossRefGoogle Scholar
Shipton, C., O'Connor, S. & Kealy, S.. 2021. The biogeographic threshold of Wallacea in human evolution. Quaternary International 574: 112. https://doi.org/10.1016/j.quaint.2020.07.028CrossRefGoogle Scholar
Siddall, M., Rohling, E.J., Almogi-Labin, A., Hemleben, C., Meischner, D., Schmelzer, I. & Smeed, D.A.. 2003. Sea-level fluctuations during the last glacial cycle. Nature 423: 853–58. https://doi.org/10.1038/nature01690CrossRefGoogle ScholarPubMed
Summerhayes, G.R., Field, J.H., Shaw, B. & Gaffney, D.. 2017. The archaeology of forest exploitation and change in the tropics during the Pleistocene: the case of Northern Sahul (Pleistocene New Guinea). Quaternary International 448: 1430. https://doi.org/10.1016/j.quaint.2016.04.023CrossRefGoogle Scholar
Sutikna, T., et al. 2016. Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia. Nature 532: 366–69. https://doi.org/10.1038/nature17179CrossRefGoogle Scholar
Sutikna, T., et al. 2018. The spatio-temporal distribution of archaeological and faunal finds at Liang Bua (Flores, Indonesia) in light of the revised chronology for Homo floresiensis. Journal of Human Evolution 124: 5274. https://doi.org/10.1016/j.jhevol.2018.07.001CrossRefGoogle Scholar
Tanudirjo, D.A. 2001. Islands in between: prehistory of the northeastern Indonesian Archipelago. Unpublished PhD dissertation, Australian National University.Google Scholar
Wadley, L., Hodgskiss, T. & Grant, M.. 2009. Implications for complex cognition from the hafting of tools with compound adhesives in the Middle Stone Age, South Africa. Proceedings of the National Academy of Sciences USA 106: 9590–94. https://doi.org/10.1073/pnas.0900957106CrossRefGoogle ScholarPubMed
Figure 0

Figure 1. Northern and southern routes through Wallacea, showing Mololo Cave on Waitanta. Inset A) seafaring simulations and chance of successful landing on Waitanta from nearby landmasses, based on averages taken from 1.5kt paddling at -30m and -50m sea levels (Table S6); Inset B) Waitanta shoreline and Mololo Cave at 50 000 years ago and at the Last Glacial Maximum (20 000 years ago) based on multibeam and GEBCO bathymetry data (figure by authors).

Figure 1

Figure 2. Mololo Cave system showing excavation units. Inset A) location of Mololo Cave at the entrance to Mayalibit Bay. Inset B) transect of entrance to Mololo Cave, relative to mean sea level (MSL) (figure by authors).

Figure 2

Figure 3. Mololo Cave stratigraphy described by context number and layer. A) Trench 1, south and west walls; B) Trench 2, south and west walls (figure by authors).

Figure 3

Figure 4. Calibrated terrestrial (charcoal and resin) radiocarbon dates from Mololo Cave, organised by stratigraphic location. Red outlines indicate the calibration threshold at 55 000 years ago and periods of inversion in the radiocarbon dates. The lower image shows the global oxygen isotope data reflecting changing temperatures (low values are higher temperatures) (Siddall et al. 2003). LGM = Last Glacial Maximum; YD = Younger Dryas; H = Heinrich event; MIS = Marine Isotope Stage (figure by authors).

Figure 4

Figure 5. Tree resin artefact (WAI-1-1024) from trench 2, directly dated to >55 000–49 620 cal BP: A) lateral side with rough surface texture; B) dorsal side with flat surface, chipping and minor scraping marks; C) ventral side showing parallel marks, possibly from scraping or the imprint of a scraped surface; D) lateral side with rough surface texture showing snaps. E illustrates a provisional hypothesis for the sequence of resin artefact production (figure by authors).

Figure 5

Figure 6. Mass spectra of Mololo archaeological resin compared with reference specimens. A–E are total ion chromatograms based on GC/MS: A) Mololo archaeological resin (WAI-1-1024); B) ethnographic ‘damar merah’ from Waigeo (Ref_D2); C) Vatica rassak (Ref_R3_K57); D) ethnographic ‘damar putih’ from Waigeo (Ref_D3); E) Agathis labillardierei (Ref_R1_K08). F–I are pyrograms based on Py-GC/MS: F) Mololo archaeological resin (WAI-1-1024 GCMS1); G) Mololo archaeological resin (WAI-1-1024 GCMS2); H) Canarium salomonense (Ref_R4_K21); I) Prumnopitys ladei (Ref_R5_S19) (figure by authors).

Figure 6

Figure 7. Selection of Mololo lithic artefacts and possible artefacts. A) WAI-1-183 (trench 2, layer 3)—flake with impact point indicated by converging hackles (marked by arrows), clear platform, ventral face with bulb, and dorsal face with a rougher texture. Edges are post-depositionally altered preventing the detection of possible use-wear; B) WAI-1-2162 (trench 1, layer 3) – flake with platform, partly cortical dorsal face and one removal. No distal end or trace of utilisation; C) WAI-1-701-B-ii (trench 2, layer 6)—smooth ventral face (iv) and platform, and dorsal face with smooth surface within two previous removals compared with rougher dorsal face (i). The edge of fracture on ventral face (pink) presents rounding and micro-polish (ii) likely resulting from scraping motion. The sharp edge (blue) does not show utilisation (iii); D) WAI-1-701-B-i (trench 2, layer 6)—flake with cortical convex face and fresher flatter face possibly indicating knapping, potential platform, and long irregular sharp edge with no clear evidence of utilisation; E) WAI-1-700 (trench 2, layer 6)—possible small flake with platform, smooth face with small bulb, and long possible flake removal on opposite side. Micro-scars on distal part of right ventral edge, possibly use-related or taphonomic; F) WAI-1-1005-C (trench 2, layer 7)—fragment with two possible concave flake removals, no evidence of use-wear (figure by authors).

Figure 7

Figure 8. Aquatic resources at Mololo. A) Hyridella cf. misoolensis (WAI-1-1022) from Mololo Cave, trench 2, layer 5. Note delamination and recrystallisation of aragonite shown in inset SEM micrograph. B) Sea urchin specimens from Mololo Cave: WAI-1-2086-A-xiii, small Echinoderm spine from trench 1, layer 5; WAI-1-1169-E, large Echinoderm spine from trench 1, layer 10; WAI-1-1551-A-ii, large Echinoderm spine with abrasion from trench 1, layer 5A. C) Sea urchin spine (WAI-1-696-B) with an edge alteration from Mololo Cave, trench 2, layer 6. D) Scombroidei sp. teeth from Mololo Cave: WAI-1-773-B, subfossilised tooth from trench 2, layer 6; WAI-1-2058, tooth from trench 1, layer 5A. Both specimens preserve diagnostic lateral margins and concave shape at the apical end (figure by authors).

Figure 8

Figure 9. Archaeobotanical remains from Mololo. A–D are SEM images of charred monocotyledonous stem tissue from trench 1, layer 5–6 (WAI-1-1161) and layer 6 (WAI-1-1236), vb = vascular bundle. A & B) longitudinal section of WAI-1-1161, with a vascular bundle exposed longitudinally within a parenchyma matrix, B shows a close-up of the vascular bundle; C & D) transverse section of WAI-1-1236, with several vitrified vascular bundles, within a parenchyma matrix, D shows a close up of vitrified vascular bundles. E–J are biogenic silica recovered from Mololo Cave sediment samples: E) sponge spicule from trench 1, layer 2 (WAI-1-1612-C); F) sponge spicule from trench 1, context 011 (WAI-1-1379-B); G) sponge asterone microsclere from trench 1, layer 3 (WAI-1-1422-B); H) sponge spicule from trench 1, layer 3 (WAI-1-1422-B); I) fern spore from trench 2, context 098 (WAI-1-424-B); J) an elongate entire and a blocky phytolith, likely from grasses, from trench 2, layer 2 (WAI-1-606-B) (figure by authors).

Figure 9

Figure 10. Reconstruction of Mayalibit Valley ecology from the Late Pleistocene to the Late Holocene alongside fruitbat and rodent δ13C and δ18O isotope results. The Late Pleistocene was characterised by patches of open and closed tropical forest, followed by expansion of closed canopy forests in the Terminal Pleistocene. The infilling of Mayalibit Bay in the Early Holocene created open areas near Mololo, possibly further encouraged by clearances. Diversification of forest patches in the Late Holocene may be caused by burning (figure by authors).

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