Singular or plural Oldowan tool-makers during the Lower Pleistocene in eastern Africa?

palevol comptes rendus 2025  24  27 Comptes Rendus Palevol est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris et l’Académie des sciences, Paris Comptes Rendus Palevol is a fast track journal published by the Museum Science Press, Paris and the Académie des sciences, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie. L’Académie des sciences publie aussi / The Académie des sciences also publishes: Comptes Rendus Mathématique, Comptes Rendus Physique, Comptes Rendus Mécanique, Comptes Rendus Chimie, Comptes Rendus Géoscience, Comptes Rendus Biologies. 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Christopher Beard (University of Kansas, Lawrence) Paléoanthropologie/Palaeoanthropology Aurélien Mounier (CNRS/Muséum national d’Histoire naturelle, Paris) Archéologie préhistorique (Paléolithique et Mésolithique)/Prehistoric archaeology (Palaeolithic and Mesolithic) Nicolas Teyssandier* (CNRS/Université de Toulouse, Toulouse) Archéologie préhistorique (Néolithique et âge du bronze)/Prehistoric archaeology (Neolithic and Bronze Age) Marc Vander Linden (Bournemouth University, Bournemouth) référés / reviewers : https://sciencepress.mnhn.fr/fr/periodiques/comptes-rendus-palevol/referes-du-journal couverture / cover : Silhouettes of Homo habilis and Paranthropus boisei by T. Michael Keesey (https://www.phylopic.org/), lanscape of Turkana and stone artefact. Credits: S. Prat/MPK-WTAP. Comptes Rendus Palevol est indexé dans / Comptes Rendus Palevol is indexed by: – Cambridge Scientific Abstracts – Current Contents® Physical – Chemical, and Earth Sciences® – ISI Alerting Services® – Geoabstracts, Geobase, Georef, Inspec, Pascal – Science Citation Index®, Science Citation Index Expanded® – Scopus®. Les articles ainsi que les nouveautés nomenclaturales publiés dans Comptes Rendus Palevol sont référencés par / Articles and nomenclatural novelties published in Comptes Rendus Palevol are registered on: – ZooBank® (http://zoobank.org) 545 COMPTES RENDUS PALEVOL • 2025 • 24 (27) © Publications scientifiques du Muséum et/and Académie des sciences, Paris. www.cr-palevol.fr Sandrine PRAT HNHP/CNRS/MNHN/UPVD, Musée de l’Homme, 17 Place du Trocadéro, F-75016 Paris (France) [email protected] Submitted on 15 February 2024 | Accepted on 3 July 2025 | Published on 5 December 2025 Singular or plural Oldowan tool-makers during the Lower Pleistocene in eastern Africa? urn:lsid:zoobank.org:pub:AC3F6968-1912-4C55-BF76-5D980241A4ED Prat S. 2025. — Singular or plural Oldowan tool-makers during the Lower Pleistocene in eastern Africa? in Hublin J.-J., Mounier A. & Teyssandier N. (eds), Lucy’s Heirs – Tribute to Yves Coppens. Comptes Rendus Palevol 24 (27): 545-561. https://doi.org/10.5852/cr-palevol2025v24a27 ABSTRACT Over the last twenty-five years, the study of hominin evolution and archaeological assemblages has become increasingly complex, due to new archaeological and paleoanthropological discoveries, and the advent of innovative analytical approaches. Findings from paleontological and archaeological sites and localities along the East African Rift Valley reveal substantial cultural and biological diversity during the Lower Pleistocene. From this context emerge critical questions about the relationships between the different material cultures and hominins. To explore these questions, I propose a crossdisciplinary approach that examines both the skeletal anatomy of hominins and the associations between hominins and archaeological assemblages in Lower Pleistocene sites and localities in East Africa. Anatomical evidence suggests that the dexterity required to produce Oldowan assemblages was likely not exclusive to the genus Homo. Moreover, multiple hominin species appear to have been involved in tool-making, particularly within Oldowan assemblages. These assemblages have been associated with Homo sp. and Homo habilis (n = 10), Homo erectus/ergaster (n = 6), and with a significant number, of Paranthropus remains (n = 8). Given this evidence, Paranthropus should now be recognized as a stone tool-maker alongside early Homo. RÉSUMÉ Singularité ou pluralité des artisans de l’Oldowayen au Pléistocène inférieur en Afrique de l’Est ? Au cours des vingt-cinq dernières années, de nouvelles découvertes archéologiques et paléoanthropologiques, ainsi que des approches analytiques innovantes, ont considérablement enrichi notre compréhension de l’évolution des hominines et des assemblages archéologiques. Les découvertes de sites et de localités paléontologiques et archéologiques le long de la vallée du Rift Est Africain révèlent une diversité culturelle et biologique substantielle durant le Pléistocène inférieur. Cela soulève des questions fondamentales sur les relations entre les hominines et les différentes cultures matérielles. Pour explorer ces questions, je propose une approche interdisciplinaire qui examine à la fois l’anatomie de la main des hominines et les associations entre hominines et assemblages archéologiques dans les sites et localités du Pléistocène inférieur en Afrique de l’Est. Les preuves anatomiques suggèrent que la dextérité requise pour produire des assemblages oldowayens n’est probablement pas exclusive au genre Homo. En outre, plusieurs espèces d’hominines semblent avoir été les artisans d’artefacts lithiques, en particulier pour les assemblages oldowayens. Ceux-ci sont associés à Homo sp. et Homo habilis (n = 10), Homo erectus/ergaster (n = 6) et, à un nombre significatif de restes de Paranthropus (n = 8). Compte tenu de ces éléments, Paranthropus devrait donc être reconnu comme un artisan d’artefacts lithiques au même titre que les premiers représentants du genre Homo. KEY WORDS Material culture, hominins, Africa, Lower Pleistocene, Oldowan, Acheulean. MOTS CLÉS Culture matérielle, hominines, Afrique, Pleistocène inférieur, Oldowayen, Acheuléen. 546 COMPTES RENDUS PALEVOL • 2025 • 24 (27) Prat S. INTRODUCTION Hominin evolutionary history depicts a bushy rather than a linear evolutionary history with multiple contemporaneous species or genera in Africa during the Pliocene and Pleistocene and sympatric relationship, for example between Homo erectus and Paranthropus boisei, as illustrated by footprint evidence (Hatala etal. 2024). This indicates significant morphological diversity (Fig. 1) with a reticulated biological evolution and ghost lineages, especially for the genus Homo (e.g. Foley 2002; Wood & Lonergan 2008; Wood & Boyle 2016; LorenteGaldos etal. 2019; Caparros & Prat 2021). Likewise, the archaeological record reveals a diversity of assemblages and a non-linear pattern of cultural evolution, exemplified by some Acheulean sites actually older than Oldowan sites (e.g. Kokiselei 4 [Lepre etal. 2011]; Konso-Gardula [Asfaw etal. 1992; Beyene etal. 2023]). Moreover, the oldest Homo fossils from Ledi Geraru in Ethiopia (Villmoare etal. 2015, 2025) as well as the specimens older than 2.3 Ma for example from the lower Omo Valley (e.g. Hlusko etal. 2024) and from the eastern and western parts of Lake Turkana (e.g. KNM-ER 5431 (Suwa etal. 1996; Villmoare etal. 2015) and KNM-WT 42718 (Prat etal. 2005)) postdate stone tools discovered at the LOM3 archeological site in Kenya by about 500 000 years-1 million years (Ma) (3.3 Ma, Harmand etal. 2015, but see also for discussion DomínguezRodrigo & Alcalá 2016, 2019; Harmand etal. 2019; Archer etal. 2020; Key & Proffitt 2024) and cut-marked bones dated to 3.4 Ma from Dikika locality in Ethiopia (McPherron etal. 2010, but see also for discussion Domínguez-Rodrigo etal. 2011; Thompson etal. 2015; Domínguez-Rodrigo & Alcalá 2016). The Dikika area is also known for the discovery of a juvenile skeleton of A.afarensis (Alemseged etal. 2006). The Lomekwi archaeological site LOM3 (3.3 Ma) is in the same chronological range as the species Australopithecus afarensis (3.8-3.0 Ma) and was discovered in the Lomekwi area, where hominin remains attributed to Kenyanthropus platyops were also found (Leakey etal. 2001). However, due to the high level of distorsion of KNM-WT 40000 (K. platyops holotype which is the only cranium of this species), it is difficult to establish whether it is a valid taxon or a variant of A. afarensis (White 2003), a species considered by some authors to be the ancestor of the genera Homo and Paranthropus (see Kimbel & Delezene 2009 for a review). Therefore, these findings from Lomekwi and Dikika challenge the exclusive association of lithic artefacts (stone-tool use and stone-tool making) with the genus Homo. They question specific attribution of lithic artefact manufacture to the genus Homo and the validity of this criterion for defining our genus. Indeed, the diagnosis of the genus Homo and the criteria for this classification are still hotly debated, and take into account anatomical, morphological, behavioural, and philosophical considerations (see for a review Prat 2022). Among the criteria proposed by Leakey and colleagues (Leakey etal. 1964) power and precision grip manual abilities were central. However, tool-making was mentioned in the section untitled “Cultural Association” but not in the Homo habilis diagnosis. Consequently, the role of archaeological remains in defining the genus Homo highlights the potential relationships in East Africa between Oldowan, Acheulean and Lower Pleistocene hominins. The discoveries made in the lower Omo Valley, in the extreme south-west of Ethiopia, are of particular interest in this context. Research in this region initially began with work by Camille Arambourg in the early 1930s, following surveys carried out by Dr Brumpt as part of the explorations led by Viscount Robert du Bour de Bozas in 1902. New expeditions were carried out from 1966 onwards, as part of the International Omo Research Expedition (IORE) initiated by Francis Clark Howell, Louis Leakey, Camille Arambourg and Yves Coppens (who led the French side of the expedition with Camille Arambourg, and then alone from 1970 to 1976, and to whom this issue is dedicated). The aim of this international, multidisciplinary expedition was to prospect and excavate the rich fossil deposits of the lower Omo Valley. Research in this region was resumed in 2006 as part of the OGRE (Omo Group Research Expedition) led by Jean-Renaud Boisserie (e.g. Boisserie etal. 2008). This region, with a sedimentary sequence almost 800 metres thick, is of considerable importance for the study of hominins, palaeoenvironmental reconstructions and relationships between hominin evolution, environment and material culture. The lower Omo Valley comprises several Oldowan sites (e.g. Chavaillon 1970; Delagnes etal. 2011; Delagnes etal. 2023) and a very large number of hominin remains (e.g. Coppens 1977; Hlusko etal. 2024). In addition, the Usno and Shungura formations are among one the longest sedimentary sequences bear hominin remains and also provide data for tracing climatic changes between 3 and 1 Ma (e.g. Coppens 1978; Alemseged etal. 2007; Boisserie etal. 2008; Bibi etal. 2013; Bedane etal. 2025). This overview is an updated of a paper published in 2023 (Prat 2023). It focuses on the morphological attributes underlying hominin remains manual skills, and the associations between eastern african hominin remains (more than 950 in the study corpus) and Oldowan assemblages as evidenced in palaeontological and archaeological records from 3.0 to 1.2 Ma, with an update of some datations. A site description is included where hominin remains were discovered in an archaeological site, with a particular focus on the sites where Pr. Yves Coppens worked. MATERIAL AND METHODS The discussion on hominin remains (anatomical parts, taxonomy), archaeological sites, spatial context, and other relevant data (i.e., as primary data, without entering into discussions) is based on data from the literature, used here at face value. However, finding detailed contextual information on hominin remains, and depositional processes, whether in situ or on the surface, and their proximity to archaeological artefacts can be challenging. Despite thorough research, it would appear that certain specific associations may be largely overlooked. Regarding the taxonomy of hominin remains, some 547 Singular or plural Oldowan tool-makers COMPTES RENDUS PALEVOL • 2025 • 24 (27) specimens are referred to Homo ergaster or Homo erectus by different authors, according to diverse nomenclatures (Reed etal. 2023). To avoid confusion, in this context, I refer to them as Homo ergaster/erectus. According to Key and Proffitt, the earliest Oldowan assemblages may date from 3.020 or 3.181 Ma and, according to an optimal linear estimation (OLE), they fall within the range of 2.622-3.436 Ma (Key & Proffitt 2024). In archaeological terminology, the terms Oldowan and Acheulean are used without distinguishing the early Oldowan and Developed Oldowan, despite differences between early (c.2.6-2.3 Ma) and later Oldowan sites (c.2.0-1.7 Ma) (e.g. Harmand 2009; Roche etal. 2009; de la Torre 2011; GoldmanNeuman & Hovers 2012; Potts 2012; Delagnes etal. 2023). The Oldowan can be considered as multiple specialized adaptations (Clark & Linares-Matás 2024). Furthermore, in many publications, particularly the earliest ones, the only terminology used is Oldowan or Acheulean (e.g. Leakey 1971; Chavaillon 1976; Harris & Herbich 1978; Kimbel etal. 1996; Plummer etal. 2023). Determining whether a hominin found at an archaeological site or on the surface was the stone tool-makers is always challenging. The place of death may not correspond to the living area, and sites can sometimes be palimpsests and/or have undergone complex geological and sedimentation processes over time. However, applying Occam’s razor, the most parsimonious hypothesis is that the hominins present at a site are the ones responsible for the material culture found there. fig. 1 . — Chronological framework of hominin taxa (above) and material cultures (below). Australopithecus Paranthropus Homo H. ergaster H. rudolfensis H. habilis Homo sp. Kenyanthropus platyops Au. garhi Au. deyiremeda P. boisei Au. bahrelghazali Paranthropus aethiopicus Au. afarensis P. robustus A ustralopithecus anamensis Au. prometheus/Au. africanus Au. sediba Lomekwian Oldowan Acheulean 4321million years 548 COMPTES RENDUS PALEVOL • 2025 • 24 (27) Prat S. Specifically, it is more reasonable to assume that stone toolmakers correspond to the species identified in the site’s sedimentary layers rather than to species absent from them. This is generally the case when specimens of the genus Homo are found, but is less systematic for other genera. Moreover, this continues to be one of the most widely accepted hypotheses, especially for Middle and Upper Paleolithic sites. The reliability of the association between hominins and archaeological sites is categorized following the framework used by Grine etal. (2022) for species attribution of postcranial remains. Two categories are proposed: 1) High Confidence 1 (HC1), where hominin remains were found in archaeological site (in archaeological layers and on the surface of the excavation due to erosion processes); and 2) High Confidence 2 (HC2), categorizing hominin remains found less than 400 metres from the archaeological site, within the same chrono-stratigraphic layer. RESULTS Overview Of anatOmical cOnstraints Concerning the anatomical prerequisites, the morphological evidence indicates that enhanced dexterity and anatomical capabilities were required for using or making stone tools. This infers in particular “pad-to-pad” precision grasping, as defined by Napier (1956) and Almécija & Alba (2014), involving the “perfect” opposition of the thumb and finger’s proximal pulp areas, which can be facilitated by a longer thumb relative to the remaining fingers. This also includes forceful precision gripping and precision handling as described by Marzke (1997), deduced from the carpometacarpal joint of the thumb and the metacarpophalangeal joint of the thumb and the fingers involved. Such morphological evidence and muscular insertions are observed on two additional hominin genera to Homo (for a review see also Kivell 2015): Australopithecus (specifically A. afarensis [e.g. Marzke 1983, 1997, 2013; Tocheri etal. 2003; Kivell etal. 2011, 2020; Almécija & Alba 2014; Kunze etal. 2024], A. africanus [e.g. Green & Gordon 2008; Skinner etal. 2015; Richmond etal. 2016], A. sediba [e.g. Kivell etal. 2011; Bardo etal. 2018; Dunmore etal. 2020]) and Paranthropus (P. robustus and P. boisei see below), and therefore extends beyond the genus Homo. Experimental biomechanics and musculoskeletal modelling also offer valuable insights into the functional anatomy of the upper limb, particularly in stone tool production (Tocheri etal. 2003; Rolian etal. 2011; Williams etal. 2012; Williams-Hatala etal. 2018; Key etal. 2019; Macchi etal. 2021; Manafzadeh & Gatesy 2021; Williams-Hatala etal. 2021; Kivell etal. 2022) and can complement anatomical and archaeological field data. Concerning the Pleistocene fossil record after 2 Ma, the anatomy of hand remains from Olduvai Gorge (misspelling of the Maa word Oldupai, Tanzania, i.e., OH 7 (1.84 Ma) and OH 86 (1.86 Ma), and from Ileret (Kenya, i.e., KNMER 47000, 1.5 Ma) was conducive to stone tool use and stone tool-making. In particular, the OH 7 late juvenile hand (1.84 Ma) from Oldupai Gorge shows a fully opposable thumb and capabilities for power and precision gripping (Leakey etal. 1964). Leakey, Tobias and Napier in 1964, pointed out the tool-making abilities of this specimen, whereas Napier (1962) was more cautious, highlighthing that “There is less certainty about toolmaking which involves not only a peripheral but also a central intellectual factor as Oakley has long insisted” (Napier 1962: 411)”. The mixed attributes of OH 7 (precision grip abilities and climbing adaptation) were emphasized by Susman & Creel (1979). Furthermore, although OH 7 has been allocated to Homo habilis (Leakey etal. 1964), some authors consider that the phalangeal morphologies are closer to those of Paranthropus (Moyà-Solà etal. 2008). Moreover, the manual proximal phalanx (OH 86, 1.84 Ma) allocated to the genus Homo, is considered to represent the earliest modern human-like hand anatomy, showing reduced postural and locomotor pressures and a decrease in arboreal adaptations (DomínguezRodrigo etal. 2015). OH 86 is distinct in size and shape from both Australopithecus/Paranthropus and the OH 7 hand bones. Its taxonomic classification is challenging, but it is generally considered as Homo, different from OH 7 (H.habilis holotype) and attributed rather to H.erectus type (Diez-Martín etal. 2015). Moreover, the third metacarpal styloid process found in Kaitio, West Turkana, Kenya (with a morphology typical of the modern humans/Neandertal hand but absent in Pliocene and Lower Pleistocene hominins) suggests that this distinctive complex of radial carpometacarpal joints emerged at early as 1.42 Ma for the genus Homo (Ward etal. 2014). Concerning Paranthropus, the hand anatomy of P. boisei, illustrated by the KNM-ER 47000 specimen (Ileret, East Turkana, Kenya), dated to 1.5 Ma, indicates that the fine precision gripping capacity and manual dexterity of this species is compatible with stone tool use or stone tool-making as also seen in earlier australopithecine species such as A. africanus (e.g. Kivell etal. 2011; Richmond etal. 2016), but not a robust thumb as observed in H. erectus/H. ergaster. This reflects adaptations to the intensification of precision gripping and tool use (Richmond etal. 2020). Paranthropus individuals present increased wrist mobility, which could improve the accuracy of force production during stone tool-making and stone tool use (e.g. Williams etal. 2010; Williams etal. 2014 ), or even during throwing, as shown in H. erectus (Roach etal. 2013; Roach & Lieberman 2014). However, Moyà-Solà etal. (2008) proposed that the Paranthropus hand structure which lacks the enlarged first metacarpal typical of Homo, might also reflect adaptations for feeding activities as seen in extant hominids (e.g. Marzke etal. 2015; Neufuss etal. 2018; Gérard etal. 2022). The anatomy of Paranthropus, particularly elements like OH 80 (e.g. radial morphology compatible with grasping abilities, used for power and precision grips) aligns with the abilities required for stone tool production (DomínguezRodrigo etal. 2013). However, as noted by Grine etal. 2022, it is very difficult to associate isolated post-cranial elements more specifically with the genus Homo or Paranthropus, given that these two genera are stratigraphically associated. 549 Singular or plural Oldowan tool-makers COMPTES RENDUS PALEVOL • 2025 • 24 (27) Overview Of the archaeOlOgical data In Eastern Africa, between 3 and 1.2 Ma, 25 documented instances (which correspond to 20 archaeological sites) of a high-confidence association between lithic and palaeoanthropological remains have been reported (Table 1). Association between Oldowan assemblages and Homo habilis sensu lato The association between the Oldowan and early Homo specimens can be observed in six occurrences (in Ethiopia, at Hadar, A.L. 666 (HC1), and Fejej, Fj-1a (HC2); in Kenya, in the western part of Lake Turkana, Lokalalei site complex (HC2); in Tanzania, in Oldupai Gorge, MNK Skull site (HC1), DK site (HC2) and Ewass Oldupa site (HC2) dated between 2.3 and 1.67 Ma (Table 1; Fig. 2). I describe here the two archaeological sites where hominin remains were discovered in archaeological layers or at the surface of the excavation (HC1). The first is the archaeological site named A.L. 666, in the Hadar region in Ethiopia. The lithic assemblage (34 artefacts including 14 found in situ) is typical of the Oldowan (Kimbel etal. 1996; Plummer 2004; Goldman-Neuman & Hovers 2012). These assemblages are dated to c.2.35 Ma by the Bouroukie Tuff 3 (Campisano & Feibel 2008) and between 2.3 and 1.9 Ma based on the fauna (Rowan etal. 2022). A maxilla (A.L. 666-1) was found on the surface of the hill (at the base). This latter shows clear early Homo traits and was assigned to Homo aff. H. habilis (Kimbel etal. 1996; Kimbel etal. 1997). However, Spoor etal. (2015) have considered that its morphology is not compatible with OH7, and therefore with H. habilis, and furthermore does not present the facial morphology observed in H. rudolfensis. For the abovementioned authors, the morphology of A.L. 666-1 is more derived toward early H. erectus/ergaster. The second high-confidence site (HC1) is MNK Skull site from Middle Bed II at Oldupai (Leakey 1971; de la Torre etal. 2021), which yielded an Oldowan assemblage and two in situ Homo habilis specimens (OH 13, paratype of Homo habilis), as well as dental remains catalogued as OH 15 (Leakey etal. 1964; Tobias 1991) dated to c. 1.67 Ma (de la Torre etal. 2021). Association between Oldowan assemblages and Paranthropus (P. boisei) The association between Paranthropus boisei specimens and Oldowan assemblages can be observed in four sites (in Kenya at Homa Peninsula, Nyayanga site (HC1); in the western part of Lake Turkana, Kokiselei 1 (HC1) and at Koobi Fora, fig. 2 . — Location of Homo habilis sensu lato remains in association with Oldowan sites. Credits: background map NASA/Internet Archive. AL 666 AL 666-1 Homo aff. H. habilis 2.34 +/- 0.07 Ma Lokalalei 1 KNM-WT 42718 early Homo 2.34 +/- 0.04 Ma MNK Skull, Middle Bed II OH 13, OH 15 Homo habilis c. 1.67 Ma Ewass Oldupa site, Bed I OH 65 Homo habilis c. 1.82 Ma DK site, Bed I OH 24 Homo habilis c. 1.87-1.84 Ma early Homo Fj-1a Fj-1-Hd1, -Hd3, -Hd4 Homo aff. H. habilis 1.951.9 Ma Oldowan 550 COMPTES RENDUS PALEVOL • 2025 • 24 (27) Prat S. table 1. — Association between hominin remains and archaeological assemblage with a High Confidence level (HC). Abbreviations: HC1, hominin remains have been found in an archaeological site, which includes both in archaeological layers and remains found on the surface of the excavation due to erosion processes; HC2, hominin remains have been found less than 400 meters from the archaeological site, within the same chrono-stratigraphic layer. Locality Region Age (Ma) Hominins Industry Category of reliability Association between Oldowan assemblages and Homo (H. habilis sensu lato and Homo sp.) A.L. 666 Hadar (Ethiopia) c. 2.35 (Campisano & Feibel 2008) 2.3 and 1.9 (Rowan et al. 2022) A.L. 666-1 Homo aff. H. habilis (Kimbel et al. 1997) Oldowan (Kimbel et al. 1996) HC 1 Lokalalei 1, Gajh5 West Turkana (Kenya) 2.34 +/- 0.04 (Prat et al. 2005) KNM-WT 42718 Early Homo (Prat et al. 2005) Oldowan (Prat et al. 2005) HC 2 FJ-1a Fejej (Ethiopia) 1.95-1.90 (Chapon et al. 2005) Fj-1-Hd1, Hd3, Hd4 Homo aff. Homo habilis (de Lumley & Marchal 2004) Oldowan (e.g. Asfaw et al. 1991; de Lumley & Beyene 2004; Barsky et al. 2011) HC 2 DK Oldupai Gorge (Tanzania) 1.87-1.84 (Deino 2012) OH 24 Homo habilis (Leakey et al. 1971) Oldowan (Leakey 1971; de la Torre & Mora 2005) HC 2 Ewass Oldupa Oldupai Gorge (Tanzania) c. 1.82 (Blumenschine et al. 2003; Deino 2012; McHenry 2012) OH 65 Homo habilis (Blumenschine et al. 2012; Clarke 2012) Oldowan (Mercader et al. 2021) HC 2 MNK Skull Oldupai Gorge (Tanzania) c. 1.67 (de la Torre et al. 2021) OH 13 Paratype Homo habilis OH 15 Homo habilis (Leakey et al. 1964; Tobias 1991) Oldowan (Leakey 1971; de la Torre et al. 2021) HC 1 Association between Oldowan assemblages and Paranthropus Nyayanga, excavation 3 Lake Victoria (Kenya) 3.03-2.58 (Plummer et al. 2023) KNM-NG 77316 Paranthropus (Plummer et al. 2023) Oldowan (Plummer et al. 2023) HC 1 Kokiselei 1 FxJh6 West Turkana (Kenya) 1.79 (Boës et al. 2024) KNM-WT 37744, 37747, 37748, 37100 Paranthropus boisei (Prat et al. 2003) Oldowan (Roche et al. 2003; Arroyo et al. 2020) HC 1 FxJj 20 Koobi Fora (Kenya) 1.65 +/- 0.05 (Feibel et al. 1989) KNM-ER 3230 Paranthropus boisei (Wood 1991) Oldowan (Harris & Herbich 1978) HC 1 BK site level 4, Bed II Olduvai Gorge (Tanzania) 1.34 (Domínguez-Rodrigo et al. 2013) OH 80 Paranthropus boisei (Domínguez-Rodrigo et al. 2013) Oldowan (Domínguez-Rodrigo et al. 2009) HC 1 Association between Oldowan assemblages, Homo and Paranthropus OMO 57/5 Lower Omo Valley (Ethiopia) 2.32-2.23 (McDougall & Brown 2008) OMO 57/5-1972-371 Paranthropus aff. aethiopicus (Coppens 1977; Suwa 1990; Wood & Leakey 2011) OMO 57/5-1972-319 Indet. (White 1988) Non robust indet. hominin (Suwa 1990) Homo aff. Homo sp. (Wood & Leakey 2011) Oldowan (Coppens et al. 1973; Chavaillon 1976) HC 1 FLK NN level 3 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 7 cranial remains Homo habilis (Holotype) (Leakey et al. 1964) Oldowan (Leakey et al. 1964; Leakey 1971) HC 1 FLK NN level 3 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 7 hand bones Homo habilis (Leakey et al. 1964) or Paranthropus (Moyà-Solà et al. 2008) Oldowan (Leakey et al. 1964; Leakey 1971) HC 1 FLK NN level 3 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 8 foot bones Homo habilis (paratype) (Day & Napier 1964; Leakey et al. 1964) Oldowan (Leakey et al. 1964; Leakey 1971) HC 1 551 Singular or plural Oldowan tool-makers COMPTES RENDUS PALEVOL • 2025 • 24 (27) FXJj 20 (HC1); in Tanzania, at Oldupai Gorge, BK site (HC1)), dated between 3.03-2.58 and 1.34 Ma (Table 1; Fig. 3). Here I describe the oldest and youngest sites with hominin remains discovered in archaeological layers or at the surface of the excavation (HC1). The oldest site with an association between Oldowan and Paranthropus remains is Nyayanga, which in the Homa Peninsula near Lake Victoria’s Winam Gulf in Kenya. This site revealed one of the oldest Oldowan assemblages, dated between 3.032 Ma and 2.581 Ma (Plummer etal. 2023), with 135 in situ artefacts (excavations 3 and 5) and faunal remains, some of which bear cutmarks. The lithic artefacts show evidence of unifacial, bifacial, and multifacial reduction, and are comparable to lithics from other Oldowan sites. Use wear analyses indicate the processing of plant and animal tissues. From excavation 3, KNM-NG 77316 (a left lower molar fragment) was found in situ, associated with Oldowan artefacts (HC1) and KNM-NG 77315 (a left upper molar) from the surface. Both teeth are assigned to Paranthropus based on their morphology and metrics. The youngest site illustrating an association between Paranthropus specimens and Oldowan artefacts is BK site in Oldupai Gorge (Bed II) in Tanzania. In this site dated to 1.34 Ma, a partial Paranthropus boisei skeleton (OH 80) was discovered in situ in level 4 (HC1) (Domínguez-Rodrigo etal. 2013). This partial skeleton includes four postcranial bones and nine upper teeth, allocated to P. boisei on the basis of their morphology, dimensions, and postcranial bone section properties. This find represents the first partial skeleton attributed to P.boisei. A large Oldowan assemblage and numerous vertebrate fossils, including some with cut marks, were also unearthed in level 4 (Domínguez-Rodrigo etal. 2009). Table 1. — Continuation. Locality Region Age (Ma) Hominins Industry Category of reliability FLK level 22 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 5 Holotype of Paranthropus boisei (Leakey 1959) Oldowan (Leakey 1971) HC 1 FLK level 22 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 6 Homo habilis (Tobias 1991) Oldowan (Leakey 1971) HC 1 FLK level 22 Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 35 Paranthropus boisei or Homo habilis (Leakey et al. 1964) Oldowan (Leakey 1971) HC 1 FxJj 38 Koobi Fora (Kenya) 1.78-1.52 (Mana et al. 2019) 1.85 (Feibel et al. 1989) KNM-ER 1805 Homo sp. but taxonomical allocation under discussion (Leakey 1974; Prat 2002) Oldowan (Harris & Herbich 1978) HC 1 FxJj 38 Koobi Fora (Kenya) 1.78-1.52 (Mana et al. 2019) 1.85 (Feibel et al. 1989) KNM-ER 1806 Paranthropus boisei (Leakey 1974; Wood 1991) Oldowan (Harris & Herbich 1978) HC 1 Association between Oldowan assemblages and Homo erectus/Homo ergaster Garba IV, level E Melka Kunture (Ethiopia) Slightly older than 1.7 (Morgan et al. 2012; Tamrat et al. 2014; Gallotti & Mussi 2015) 2 Ma (Mussi et al. 2023) MK 81 GAR IVE 0043 early Homo erectus/Homo ergaster (Condemi 2004) Oldowan (Gallotti & Mussi 2015; Mussi et al. 2023) HC 1 PTK site Oldupai Gorge (Tanzania) 1.84 (Deino 2012) OH 86 Homo erectus-like (Diez-Martín et al. 2015; Domínguez-Rodrigo et al. 2015) Oldowan (Domínguez-Rodrigo et al. 2015) HC 1 Naiyena Engol 1 FxJh5 West Turkana (Kenya) 1.75 +/- 0.05 (Roche et al. 2003) KNM-WT 37745 Homo sp. (aff. H. ergaster) (Prat et al. 2003) Oldowan (Roche et al. 2003) HC 1 Gombore I Melka Kunture (Ethiopia) 1.6-1.5 (Morgan et al. 2012; Tamrat et al. 2014) Gombore IB-7594 early Homo erectus Homo sp. 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American Journal of Physical Anthropology 162 (3): 533-549. https://doi. org/10.1002/ajpa.23135 Submitted on 15 February 2024; accepted on 3 July 2025; published on 5 December 2025.