Earliest evidence of elephant butchery at Olduvai Gorge (Tanzania) reveals the evolutionary impact of early human megafaunal exploitation

Spatially concurrent lithic tools and proboscidean bones have a long history in human evolution. Such concurrency (sometimes in conjunction with other variables) has been frequently interpreted as the result of hominin butchery (Adrien Hannus, 2018; Agam and Barkai, 2018; Agam and Barkai, 2016; Altamura et al., 2020; Barkai, 2019; Ben-Dor and Barkai, 2020; Berthelet and Chavaillon, 2001; Boschian and Saccà, 2015; Chavaillon et al., 1987; Chavaillon and Berthelet, 2004; Delagnes et al., 2006; Gaudzinski et al., 2005; Gaudzinski-Windheuser et al., 2023a; Gaudzinski-Windheuser et al., 2023b; Gomez et al., 1978; Goren-Inbar et al., 1994; Haynes, 2022; Konidaris et al., 2021; Leakey, 1971; Lemorini et al., 2023; Lev and Barkai, 2016; Mussi and Villa, 2008; Reshef and Barkai, 2015; Rocca et al., 2021; Saccà, 2012; Santucci et al., 2016; Solodenko et al., 2015; Villa et al., 2005; Yravedra et al., 2019; Yravedra et al., 2010; Rabinovich et al., 2012). Spatial concurrency is insufficient to establish a causal relationship between tools and megafaunal remains. Given the time-averaged nature of a large part of the Paleolithic record, the deposition of tools and elephants could have occurred sequentially and unrelatedly in several sites; especially taking into account that most of these sites were in paleo-floodplains where elephants and hippos were intensively active and formed natural taphocoenoses. Examples of this abound in the paleobiological record. For example, at BK (upper Bed II, Olduvai Gorge, Tanzania), some Pelorovis carcasses were naturally deposited on the same alluvial bar and among the fauna processed by hominins, thereby creating an artificial spatial concurrence between independent depositional events and agencies (Organista et al., 2016). In the Middle Pleistocene deposits of Torralba and Ambrona (Spain), artifact-littered areas occur in spatial association with naturally deposited elephant carcasses, only a few of which were also exploited to a limited degree by hominins, as demonstrated by a few proboscidean remains bearing cut marks (Villa, 1990; Villa et al., 2005). Post-depositional/sedimentation processes also might have artificially created associations of faunal remains and unrelated lithic artifacts (Villa, 1990; Villa et al., 2005).

Direct taphonomic evidence of hominin exploitation of megafauna (In this study, megafauna refers to animals with a body mass exceeding 800 kg) (in the form of cutmarked or dynamically-percussed bone) has been so far the main argument to establish a causal link between hominin agency and megafaunal remains (Tappen et al., 2022). If restricted to the Early Pleistocene, most sites containing partial carcasses of megafauna (e.g. elephant, hippopotamus) either show poor preservation, do not exhibit any of these anthropogenic traces, or these modifications cannot be securely identified (Delagnes et al., 2006; Domínguez-Rodrigo et al., 2007; Haynes, 2022; Haynes, 1991; Isaac and Isaac, 2023; Leakey, 1971). In contrast, for the Middle and Upper Pleistocene, there is more abundant evidence of hominin butchery of megafaunal remains, especially of elephants (Gaudzinski-Windheuser et al., 2023a; Gaudzinski-Windheuser et al., 2023b; Haynes, 2022; Yravedra et al., 2010; Rabinovich et al., 2012); see review of taphonomic evidence in Agam and Barkai, 2018, Haynes, 2022 and Konidaris et al., 2021. The question that we address here is if such a behavior has its roots in the early Pleistocene.

Once hominin agency is empirically demonstrated at taphonomically-supported butchery sites, a second inferential step is attributing carcass butchery to hunting or scavenging. Occasionally, the occurrence of embedded flint in proboscidean remains has been taken as direct evidence of predatory behaviors (Agam and Barkai, 2018). This evidence is marginal and chronologically restricted to the late Upper Pleistocene with composite hafted tools. Recently, a supportive argument used to detect proboscidean hunting lies on the intensity of cut marks on carcasses and mortality profiles dominated by male prime adults (Gaudzinski-Windheuser et al., 2023a; Gaudzinski-Windheuser et al., 2023b). Prime adult mortality is documented also in other natural palimpsests (Haynes, 1991), in some cases probably caused by fights leading also to broken tusks (Villa and d’Errico, 2001). Ethnoarchaeological observation of modern foragers butchering elephants and hippos shows that they leave very few cut marks after bulk defleshing (Crader, 1983). Modern elephant butchery after culling events (based on a sample of >500 carcasses) also shows that bulk defleshing virtually leaves no traces on bones (Haynes and Krasinski, 2021). Only butchery aiming at the extraction of meat scraps after bulk defleshing imparts most of the cut marks documented experimentally (Haynes and Krasinski, 2021). This casts doubts on whether cutmarking intensity relates to hunting, scavenging, or a maximization strategy feasible in all scenarios. To complicate things more, there is a clear mismatch between the butchery intensity documented at several sites and the resulting tool kit associated with it. Some Middle Pleistocene elephant carcasses display abundant cut marks, but there are very few or no stone tools associated with them (Gaudzinski-Windheuser et al., 2023a), whereas at other sites elephant carcasses exhibit not a single cut mark, but they occur with hundreds or thousands of associated tools (Chavaillon et al., 1987; Delagnes et al., 2006; Yravedra et al., 2010; Table 1). In the latter case, it is probable that in some cases the bulk of tools may have been used to process other non-megafaunal fauna. In the particular case of the Nadung´a elephant (Kenya, 1–0.7 Ma), that explanation for the presence of lithics is improbable since the only other mammal carcass remains found are from one bovid and one suid, most likely a background scatter associated with the bird, fish, and reptile remains with which they are spatially associated. The question remains as to why such a divergent range of tool sets (both in quantity and typological composition) accompany similar partial megafaunal skeletal remains if they are truly functionally related to them.

To make interpretations even more challenging, many butchery sites exhibit fairly complete and (semi)articulated carcasses with limb bones intact. In some of these instances, elements were dismembered but not demarrowed or degreased (Gaudzinski-Windheuser et al., 2023a). Defleshing or demarrowing/degreasing does not require disarticulating bones, and the energy costs involved in cutting through tendons and ligaments of a proboscidean carcass are better explained as a maximizing strategy aiming at reducing the costs of element transportation (neither defleshing nor demarrowing/degreasing requires disarticulation), and/or further exploitation of within-bone fat resources. None of this is clearly documented in those sites. Fat was a crucial resource during human evolution (Ben-Dor and Barkai, 2024; Ben-Dor et al., 2011; Reshef and Barkai, 2015; Solodenko et al., 2015), and the large fat deposits stored in proboscidean carcasses rationally would have been a target of prehistoric butchery, especially in strategies of carcass maximization (as documented in the abundant cut mark record at some megafaunal butchery sites). The question of why our ancestors chose to maximize flesh extraction from proboscideans, and yet frequently abandoned their fat deposits untouched remains unanswered. Modern Bisa people (Zambia) butcher elephants and do not exploit their fat, but they follow a ‘satisficing’ strategy (Haynes and Krasinski, 2021), through bulk defleshing (barely impacting bone surfaces; Crader, 1983), and not a ‘maximizing’ strategy, as suggested for some highly anthropogenically impacted archaeological proboscidean sites (Gaudzinski-Windheuser et al., 2023a). It could be argued that the meat of healthy elephants contains already a high amount of fat and that other anatomical areas (like the podal pads) contain extremely nutritious fat, or that the relevance of fat exploitation varies according to latitude and climate, but if the strategy was to maximize food resources, long bone medullary contents should also have been consumed (Ben-Dor et al., 2011; Haynes et al., 2021), as documented in sites like Castel di Guido (Italy), Preresa (Spain; Boschian and Saccà, 2015; Yravedra et al., 2012), or BK (see below). Megafaunal long bone green breakage (linked to continuous spiral fractures on thick cortical bone accompanied by modification [in some cases, intensive] of cortical and medullary surfaces of breakage planes) is probably a less ambiguous trace of butchery than ‘cut marks’, since many of the latter could be equifinal and harder to identify in contexts of moderate to poor preservation, especially in contexts of high abrasion and trampling (Haynes et al., 2021; Haynes et al., 2020). For example, the purported cut-marked pelvis from Fuentenueva 3 (Yravedra et al., 2024) exhibits intense biostratinomic—in addition to diagenetic—modifications impacted by abrasive processes, which renders the pristine experimental reference collection inadequate to confidently classify it as human-made. This is further supported by the fact that the experimental trampling collection used depicts trampling marks as extremely shallow and with wide divergent walls. This does not include the large diversity of morphologies generated by trampling marks, whose cross-sections frequently are indistinguishable from stone-tool cut marks (Domínguez-Rodrigo et al., 2010b). Given the abrasive sedimentary context, the abundance of abrasion marks on the affected specimens, and the lack of description of microscopic features that could unambiguously be used to identify those traces as human-made, we remain skeptical of their anthropogenic nature.

Ideally, cut marks should complement green breakage patterns to identify butchery episodes, but the virtual lack of cut marks in ‘satisfying’ proboscidean exploitation renders their presence/absence non-indicative of the type of anthropic action. Despite the occurrence of green fractures on naturally broken bones, such as those trampled by elephants (Haynes et al., 2020), and those occurring through traumatic fracturing or gnawed by carnivores (Haynes and Hutson, 2020), these fail to reproduce the elongated, extensive, or helicoidal spiral fractures (uninterrupted by stepped sections), accompanied by the overlapping conchoidal scars (both cortical and medullary), the reflected scarring, the inflection points, or the impact hackled break surfaces and flakes typical of dynamic percussive breakage. Evidence of this type of green breakage had not been documented earlier for the Early Pleistocene proboscidean or hippopotamid carcasses, beyond the documentation of flaked bone with the purpose of elaboration of bone tools (Backwell and d’Errico, 2004; Pante et al., 2020; Sano et al., 2020). Such type of dynamic breaking, in contrast, has been documented in Pelorovis and Sivatherium carcasses at the 1.3 Ma site of BK (Olduvai Gorge; Domínguez-Rodrigo et al., 2014a; Labrado, 2017; Organista et al., 2016). One reviewer suggested that such variables have also been documented in non-anthropogenic contexts, including helicoidal spiral fractures attributed to trampling or carnivore activity (Haynes, 1983), adjacent or flake-like scars produced by carnivore gnawing (Villa and Bartram, 1996), hackled fracture surfaces allegedly resulting from heavy passive breakage such as trampling or sediment pressure (Haynes, 1983), and ‘impact-like’ bone flakes documented in carnivore-broken bones. However, this interpretation is epistemologically problematic because it does not satisfy the fundamental criteria for valid analogy as outlined by Bunge, 1981, namely substantial, structural, and environmental affinity. Specifically, the cited examples involve agents, materials, and contexts that differ markedly in composition, mechanical properties, and loading regimes from those considered here. Experimental and actualistic studies demonstrate that carnivores—rather than trampling—are also capable of producing spiral fractures and overlapping bone scarring, but these observations are restricted to faunal remains of substantially smaller body size than elephants, which carnivores can gnaw (Haynes, 1983; see also Appendix 1—figures 30–36). To date, no carnivore has been documented as producing comparable fracture morphologies or surface damage on late juvenile and adult elephant bones. Consequently, the proposed analogy is not supported. Moreover, Haynes, 1983 provides no empirical evidence that sediment pressure or trampling can generate hackled fracture surfaces. Such features are instead associated with dynamic loading conditions, whereas passive breakage processes have not been shown to produce these types of modifications (Lyman, 2006). This reasoning also applies to impact flakes on elephant limb bones, which can only be produced by the sole modern agent documented to dynamically fracture non-infantile green proboscidean long bones: humans.

Regardless of the type of butchery evidence—and with the taphonomic caveat that no unambiguous evidence exists to confirm that megafaunal carcasses were hunted or scavenged other than hominins accessed them in different taphonomically defined stages (i.e. early or late)—the principal reasons for exploring megafaunal consumption in early human evolution are its origin, its episodic or temporally patterned occurrence, its impact on hominin adaptation to certain landscapes, and its reflection on hominin group size and site functionality. If hominins actively sought the exploitation of megafauna, especially if targeting early stages of carcass consumption, the recovery of an apparent surplus of resources reflects a substantially different behavior from the small-group/small-site pattern documented at several earlier Oldowan anthropogenic sites (Domínguez-Rodrigo et al., 2019) -or some modern foragers, like the Hadza, who only exploit megafaunal carcasses very sporadically, mostly upon opportunistic encounters (Marlowe, 2010; O’Connell et al., 1992; Wood, 2010; Wood and Marlowe, 2013). Determining when the process of becoming megafaunal commensal started has major implications for human evolution, since it has clear implications for our understanding of past social group sizes and social dynamics.

The multiple taphonomic biases intervening in the palimpsestic nature of most of these butchery sites often prevent the detection of the causal traces linking megafaunal carcasses and hominins. Functional links have commonly been assumed through the spatial concurrence of tools and carcass remains; however, this perception may be utterly unjustified as we argued above. Functional association of both archaeological elements can more securely be detected through objective spatial statistical methods. This has been argued to be foundational for heuristic interpretations of proboscidean butchery sites (Giusti, 2021). Such an approach removes ambiguity and provides a better argument for spatial functional association, as demonstrated at sites like Marathousa 1 (Konidaris et al., 2018) or TK Sivatherium (Panera et al., 2019). This method will play a major role in the present study.

Here, we present the discovery of a new elephant butchery site (Emiliano Aguirre Korongo, EAK), dated to 1.78 Ma, from the base of Bed II at Olduvai Gorge (Tanzania). It is potentially the oldest unambiguous proboscidean butchery site at Olduvai. In combination with its study, we have carried out extensive survey at a landscape scale on one stratigraphic section of Bed I (Oldowan) and three stratigraphic sections of Bed II: lower Bed II-Lower Augitic Sands (1.8.1.69 Ma), upper middle Bed II (from tuff IIB to IID, 1.5 Ma), and upper Bed II (overlying Tuff IID, 1.3 Ma), which are known to contain Acheulian materials. The goal was to find megafaunal fossils bearing traces of hominin modification across the landscape. Starting at 1.78 Ma, Olduvai Gorge records extensive evidence of hominin involvement with megafaunal carcasses (Domínguez-Rodrigo et al., 2014c; Domínguez-Rodrigo et al., 2009b), namely Proboscidea and Hippopotamidae, followed closely by Giraffidae and big Bovidae. The nature of this interaction is elaborated and analyzed in this paper. We have used a spatial approach, combined with a more general taphonomic analysis, and an analysis of hammerstone-broken bone specimens occurring widespread on the three targeted Bed II stratigraphic intervals. Also provided is a technological study of the lithic artifacts.

Evidence shows that the oldest systematic anthropogenic exploitation of proboscidean carcasses is documented (at several paleolandscape scales) in the Late Pleistocene sites of Neumark-Nord (Germany; Gaudzinski-Windheuser et al., 2023a; Gaudzinski-Windheuser et al., 2023b). Evidence of at least episodic access to proboscidean remains goes back in time (see review in Agam and Barkai, 2018; Ben-Dor et al., 2011; Haynes, 2022). Redundant megafaunal exploitation is well documented at some early Pleistocene sites from Olduvai Gorge (Domínguez-Rodrigo et al., 2014a; Domínguez-Rodrigo et al., 2014c; Organista et al., 2019; Organista et al., 2017; Organista et al., 2016). At the very same sites, the stone artifactual assemblages, as well as the site dimensions, are substantially larger than those documented in the Bed I Oldowan sites (Diez-Martín et al., 2024b; Diez-Martín et al., 2009b; Diez-Martín et al., 2014; Diez-Martín et al., 2017). Whether this reflects larger hominin groups or more prolonged and time-averaged intervention in more productive environments remains to be determined (it will likely vary according to site).

A recent discovery of a couple of hippopotamus partial carcasses at the 3.0–2.6 Ma site of Nyayanga (Kenya), spatially concurrent with stone artifacts, has been argued to be causally linked by the presence of cut marks on some bones (Plummer et al., 2023). The only evidence published thereof is a series of bone surface modifications on a hippo rib and a tibial crest, which we suggest they might also be the result of byproduct of abiotic abrasive processes; the marks contrast noticeably with the well-defined cut marks found on smaller mammal bones in the same localities (Plummer et al., 2023: Figure 3C and D) associated with the hippo remains (Plummer et al., 2023). The two trenches from which both hippo remains derive are located in the stratigraphic unit NY1, consisting of a channel containing cobble conglomerates and coarse sands in addition to silts. Gravel and coarse sand can generate the same types of marks seen in the figures of the Nyayanga hippo bones (Domínguez-Rodrigo et al., 2009a; Domínguez-Rodrigo et al., 2010b). Hippopotamidae are the most abundant taxa in the Nyayanga excavations, indicating a productive taphocoenotic environment, in which hippopotamus remains co-occur with a large diversity of mammal taxa, most of which could have been naturally deposited and mixed. No green fractures have been reported on the hippopotamus bones as found in some other mammal fauna. Although Nyayanga could potentially be one of the earliest examples of megafaunal butchery, the taphonomic evidence remains ambiguous. Butchery experiments show that complete butchery of animals is reflected in a large percentage of remains bearing traces of butchery marks. In T3, these are documented in <0.9% and in T5 in 1%. This is a magnitude of 10–30 times lower than expected using experimental butchery frameworks (even if using only well-preserved bone specimens; Domínguez-Rodrigo, 1997; Domínguez-Rodrigo et al., 2014b; Pizarro-Monzo et al., 2021).

The 1.8 Ma Elephas recki of FLK North (level 6, Olduvai Bed I) has the potential of being the oldest proboscidean butchery site (Leakey, 1971). It was argued that cut marks existed on its complete bones (Bunn, 1982; Delagnes et al., 2006), but a subsequent taphonomic study of those marks suggested that they might have been made by trampling (Domínguez-Rodrigo et al., 2007). Additionally, it is unclear if the spatial association of stone tools and elephant bones is functional or artificial. Leakey, 1971 did not report on the third dimension of her excavations, and this is crucial. In our excavations of FLK N6, we documented that the level was a deposit spanning more than half a meter depth and fossils and stone tools were vertically scattered throughout (Domínguez-Rodrigo et al., 2010a). It would have been essential to document that the FLK N6 tools associated with the elephant were either on the same depositional surface as the elephant bones and/or on the same vertical position.

The ambiguity about the FLK N6 elephant renders EAK, potentially, the oldest proboscidean butchery evidence at Olduvai, and also probably one of the oldest in the early Pleistocene elsewhere in Africa. FLK North Deinotherium occupied a position above the EAK stratigraphic level, and evidence of hominin intervention there is missing (Leakey, 1971). A cut-marked bone fragment of a large animal (probably a hippopotamus) was documented at El-Kherba (Algeria) dated to 1.78 Ma (Sahnouni et al., 2013). These findings are suggestive of the beginning of megafaunal exploitation by humans in similar times both in East and North Africa.

Our landscape study of the megafaunal remains at Olduvai shows that elephants and hippopotamuses overlap on the same clustering areas. In all of them, the geology is indicative of extensive alluvial zones. At the base of Bed II, the gorge junction is characterized by extensive wetlands (Ashley, 2003; Ashley et al., 2009; Deocampo et al., 2002; Hay, 1976; Liutkus and Ashley, 2003). In the overlying LAS, wetlands are crossed by important fluvial systems (Domínguez-Rodrigo et al., 2023; Uribelarrea et al., 2017). In the middle and upper Bed II fluvial and wetland systems intertwined as the lake was reduced and moved away from its former shoreline in the Bed I-Bed II transition (Arráiz et al., 2017; Garrett, 2017; Uribelarrea del Val and Domínguez-Rodrigo, 2017). In all cases, these wet environments must have been preferred places for water-dependent megafauna, like elephants and hippos, and their overlapping ecological niches are reflected in the spatial co-occurrence of their carcasses. Both types of megafauna show traces of hominin use through either cutmarked or percussed bones, green-broken bones, or both (Appendix 1).

Green-broken elephant bones are documented at Olduvai at a landscape scale during the LAS deposition (1.7 Ma; Diez-Martín et al., 2015; Domínguez-Rodrigo et al., 2023; Uribelarrea et al., 2017). This record is now added to the newly discovered EAK site, showing tight spatial association (both vertically and horizontally) between bones and stone tools. This trend of spatial association of stone artifacts and proboscidean bones continues through the deposition of Bed II. Although no cut marks have been documented on the proboscidean bones that we identified (part of which had cortical surfaces impacted by biostratinomic and diagenetic modification), the presence of green-broken bones at the sites, the butchery of carcasses as indicated by spatial statistical analyses at sites like EAK and TK (Domínguez-Rodrigo et al., 2014a; Organista et al., 2016; Panera et al., 2019), and the presence of hammerstone-broken bones on the surrounding paleo-landscapes of these and other sites support the occasional exploitation of megafaunal remains by humans. In most green-broken limb bones, we document the presence of a medullary cavity, despite the continuous presence of trabecular bone tissue on its walls. We do not know if the bone breakage was intended to obtain marrow or to create raw material for making artifacts or tools with functional purposes or both. We have been able to identify only one potential bone tool in our sample (Figure 6, Appendix 1—figures 4–23); the scarcity of bone implements supports the former option. The exploitation of megafauna is not anecdotal, since at sites like BK or TK (upper Bed II), the taphonomic analysis of other megafaunal remains (namely, Sivatherium and Pelorovis) clearly shows intensive exploitation of complete carcasses in both of these taxa, especially at BK (Domínguez-Rodrigo et al., 2014a; Organista et al., 2016; Panera et al., 2019). The food surplus generated by the exploitation of some of these animals would have been a significant advantage for the survival and adaptation of the hominin groups.

Figure 6

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These results of widespread conspicuous evidence of hammerstone-breaking behaviors during Bed II partially confirm the recent report on bone artifacts discovered in Bed II by another research team (de la Torre et al., 2025). Excavations at the T69 Complex area unearthed a wealth of megafaunal (proboscidean and hippopotamus) remains displaying hammerstone green breakage and hominin modifications. These remains seem to be spatially connected with a highly productive paleolandscape, since at its northern end, it connects with BBS, a recently discovered site with megafaunal (Hippopotamus) remains. Green breakage at BBS also reflects hominin intervention in the palustrine environment congregating so many megafaunal carcasses. This unit would be equivalent to our second unit in Bed II.

We are not confident that the artifacts reported by dela Torre et al. at the T69 Complex are indeed tools. They are similar to the green-broken specimens that we have reported here. They are characterized by repeated number of flake scars along the edges; however, this is not a diagnostic feature, since multiple scars along edges also indirectly result from the repeated battering until the final breakage of the bone during butchery (Haynes et al., 2021; Haynes et al., 2020; work in progress). In large bovids, it can also be derived from the gnawing of durophagous carnivores (Villa and Bartram, 1996; Appendix 1—figure 32). Carnivores capable of breaking elephant bones and producing similar break shapes go back to at least the early Middle Miocene (Appendix 1—figure 33), although no carnivore today can effectively break the dense long bone mid-shafts of adult elephant bones. Another argument used by the authors is that the purported tools are longer and bear more flake scars compared to other faunal fragments (approximate average 13 vs 4). Again, this is documented among durophagous carnivores (Villa and Bartram, 1996; see examples in Appendix 1). A higher scar number in the T69 artifact collection could result from a correlation of different variables (i.e. longer dimension of the compared specimens, or different carcass sizes) and be, therefore, unrelated to anthropic intentionality. A third argument provided is that the items share consistent technological features, such as the production of morphologically similar, elongated, pointed, and notched bone tools, suggesting a patterned behavior. Again, this is equifinal with unintentional breaking of megafaunal remains during butchery (Haynes et al., 2021; Haynes et al., 2020; Appendix 1—figures 6–19; experimental work in progress). There are several arguments that could be used to support this alternative:

1. The scars in the T69 artifacts exhibit an asymmetric distribution, characterized by stepped or reflected, incomplete fractures predominantly on the medullary surface, with minimal invasiveness. This contrasts with the expected conchoidal and complete scarring distributed equally across both cortical (i.e. periosteal) and medullary surfaces, as would be anticipated in cases of intentional flaking. Given that bone is more easily flaked than metamorphic rock, and that bifacial knapping of stone typically produces symmetrical, conchoidal scars on both surfaces, this would also be expected in the T69 bone specimens.

2. This bilateral symmetry principle is observed in Middle Pleistocene bone handaxes, where intentional modification is inferred based on the equal treatment of both sides and evidence of artificial shaping (Zutovski and Barkai, 2016). It is also documented on the similarly aged 1.4 Ma bone handaxe from Konso (Sano et al., 2020), and a purported handaxe from FC (Bed II, Olduvai; Backwell and d’Errico, 2004), but it is absent from the reported items in T69, as well as those of the present study. Why, otherwise, would scars on the medullary surface be mostly limited to the wall of the breakage plane?

3. The repeated presence of incomplete, stepped scars on the medullary surfaces of the artifacts found at T69 aligns more closely with damage resulting from deliberate battering associated with breakage rather than subsequent edge retouching. The latter process would be expected to generate more controlled and complete conchoidal scars indicative of intentional shaping (Backwell and d’Errico, 2004; Haynes et al., 2021; Haynes et al., 2020). This reflected scarring is also documented in Bos/Bison bones broken by hyenas, caused by redundancy in gnawing and the resulting overlapping micro-cracking leading to final collapse (Villa and Bartram, 1996; Appendix 1—figure 31).

4. Scar frequency is not a taphonomically valid criterion to attribute agency, since elephant bones can be broken in multiple ways, thus impacting experimentally derived scar patterns. Sequential battering of initial crack lines with handheld hammerstones can generate overlapping notches and scars (work in progress). This is not necessarily reproduced when other experimental studies use alternative methods (such as throwing or bashing against immobile anvils) (Backwell and d’Errico, 2004; Haynes et al., 2021).

5. In several of the artifacts from T69, most of the overlapping scarring occurs in association with notches (and, in some cases, associated percussion marks) derived from dynamic loading. This reinforces that they may have resulted from repeated battering during bone breaking.

6. The T69 Complex bone artifacts and those reported here are large-sized. If their potential function was for heavy-duty activities, those should have left any visible traces (at the macro and microscales) on the scarred edges. These are absent. A faunal assemblage that has good preservation of cut and percussion marks should certainly have good preservation conditions for microwear use and macro-damage scars.

7. If applying the same criteria defining bone tools at T69 Complex to Bos/Bison modified bones, hyenas would also create similar artifacts (Appendix 1—figure 32).

A final argument used by the authors to justify the intentional artifactual nature of their bone implements is that the bone tools were found in situ within a single stratigraphic horizon securely dated to 1.5 million years ago, indicating systematic production rather than episodic use. This is taphonomically unjustified. In a productive paleosurface where megafauna is abundant and hominin intervention is intense, the presence of this type of highly scarred green-broken bones can be fairly common, since they result from butchery and carnivore post-depositional ravaging on non-proboscidean megafaunal bones (Domínguez-Rodrigo et al., 2014a; Domínguez-Rodrigo et al., 2014c; Domínguez-Rodrigo et al., 2009b; Villa and Bartram, 1996). Here, we have shown that most green-broken elephant bones cluster in three main areas of the Olduvai gorge.

Leakey collected more than a hundred purported bone tools from the same locations of Bed II where we collected our hammerstone-broken bones (Leakey, 1971; Appendix 1—figure 33). All of them derive from Bed II (Backwell and d’Errico, 2004). A recent taphonomic review cautiously identified six potential bone tools in that collection (Pante et al., 2020). Some of these authors, using the much smaller T69 Complex artifact sample, now argue about systematic bone tool production. Bone artifacts (i.e. hominin-made items) that are used as tools should display evidence thereof through the macroscopic scarring, the polishing of the actively used surface, and the presence of microabrasive modifications on it (Backwell and d’Errico, 2008, Backwell and d’Errico, 2001; d’Errico and Backwell, 2009; LeMoine, 1994). None of that has been documented in the T69 assemblage. We do not deny that the items reported by de la Torre et al., 2025 have the potential of having been used as tools, but proof is currently missing. The same applies to our reported sample (Figure 6; Appendix 1—figures 4–25). Only one artifact shows modifications that are potentially derived from its use as a tool. If confirmed, it is interesting to note the convergence in the point shape targeted in such an implement and those intentionally shaped in the earliest Acheulian implements documented at Olduvai Gorge (Figure 6, Appendix 1—figures 6–8). If such a bone implement was a tool, it would be the oldest bone tool documented to date (>1.7 Ma).

All data is included in the paper.

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Author details

1. Manuel Dominguez-Rodrigo

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Department of Anthropology, Rice University, Houston, United States

   Contribution

   Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Writing – original draft, Project administration

   For correspondence

   mdr@rice.edu

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-7233-331X

2. Enrique Baquedano

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Archaeological and Paleontological Museum of Madrid, Madrid, Spain

   Contribution

   Resources, Funding acquisition, Validation, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

3. Abel Moclan

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. PALEVOPRIM laboratory, UMR CNRS 7262, and Université de Poitiers, Poitiers, France

   Contribution

   Formal analysis, Validation, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

4. David Uribelarrea

   Department of Geodynamics, Stratigraphy and Paleontology, Complutense University, Madrid, Spain

   Contribution

   Formal analysis, Validation, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

5. Jose Angel Corre-Cano

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Archaeological and Paleontological Museum of Madrid, Madrid, Spain

   Contribution

   Validation, Investigation, Methodology, Writing – review and editing

   Competing interests

   No competing interests declared

6. Fernando Diez-Martin

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Department of Prehistory and Archaeology, University of Valladolid, Valladolid, Spain

   Contribution

   Resources, Validation, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

7. Alejandro Velazquez-tello

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Archaeological and Paleontological Museum of Madrid, Madrid, Spain

   Contribution

   Investigation

   Competing interests

   No competing interests declared

8. Elia Organista

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Osteoarkeologiska forskningslaboratoriet och antikens kultur, Stockholms Universitet, Stockholm, Sweden

   Contribution

   Validation, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

9. Eduardo Mendez-Quintas

   1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
   2. Universidade de Vigo, MAPAS Lab, Vigo, Spain
   3. UNIARQ – Centre for Archaeology, School of Arts and Humanities, University of Lisbon, Lisbon, Portugal

   Contribution

   Validation, Investigation, Writing – review and editing

   Competing interests

   No competing interests declared

10. Marina Vegara-Riquelme

    1. Institute of Evolution in Africa (IDEA), Rice University, Archaeological and Paleontological Museum of Madrid and Foundation of the University of Alcalá, Madrid, Spain
    2. Area of Prehistory, Department of History and Philosophy, University of Alcalá de Henares, Alcalá de Henares, Spain

    Contribution

    Validation, Investigation, Writing – review and editing

    Competing interests

    No competing interests declared

11. Agness Gidna

    Ngorongoro Conservation Area Authority, Department of Cultural Heritage, Karatu, United Republic of Tanzania

    Contribution

    Validation, Investigation, Writing – review and editing

    Competing interests

    No competing interests declared

12. Audax Mabulla

    Department of Archaeology, University of Dar es Salaam, Dar es Salaam, United Republic of Tanzania

    Contribution

    Project administration

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0001-9591-5497

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