Evidence for deliberate burial of the dead by Homo naledi

  1. Lee R Berger
  2. Tebogo Makhubela
  3. Keneiloe Molopyane
  4. Ashley Krüger
  5. Patrick Randolph-Quinney
  6. Marina Elliott
  7. Becca Peixotto
  8. Agustín Fuentes
  9. Paul Tafforeau
  10. Vincent Beyrand
  11. Kathleen Dollman
  12. Zubair Jinnah
  13. Angharad Brewer Gillham
  14. Kenneth Broad
  15. Juliet Brophy
  16. Gideon Chinamatira
  17. Paul HM Dirks
  18. Elen Feuerriegel
  19. Alia Gurtov
  20. Nompumelelo Hlophe
  21. Lindsay Hunter
  22. Rick Hunter
  23. Kudakwashe Jakata
  24. Corey Jaskolski
  25. Hannah Morris
  26. Ellie Pryor
  27. Maropeng Mpete
  28. Eric Roberts
  29. Jacqueline S Smilg
  30. Mathabela Tsikoane
  31. Steven Tucker
  32. Dirk van Rooyen
  33. Kerryn Warren
  34. Colin D Wren
  35. Marc Kissel
  36. Penny Spikins
  37. John Hawks

  • Curated by eLife

    eLife logo

    eLife Assessment

    The authors study the context of the skeletal remains of three individuals and associated sediment samples to conclude that the hominin species Homo naledi intentionally buried their dead. Demonstration of the earliest known instance of intentional funerary practice - with a relatively small-brained hominin engaging in a highly complex behavior that has otherwise been observed from Homo sapiens and Homo neanderthalensis - would represent a landmark finding. The authors have revised their manuscript extensively in light of the reviews of their initial submission, with improved illustration, context, discussion, and theoretical frameworks, leading to an improved case supporting their conclusion that Homo naledi intentionally buried their dead. One of the reviewers concludes that the findings convincingly demonstrate intentional burial practices, while another considers evidence for such an unambiguous conclusion to be incomplete given a lack of definitive knowledge around how the hominins got into the chamber. We look forward to seeing the continued development and assessment of this hypothesis. It is worth noting that the detailed reviews (both rounds) and comprehensive author response are commendable and consequential parts of the scientific record of this study. The editors note that the authors' response repeatedly invokes precedent from previous publications to help justify the conclusions in this paper. While doing so is helpful, the editors also note that scientific norms and knowledge are constantly evolving, and that any study has to rest on its own scientific merit.

Reviewed by

Read the full article See related articles

Listed in

Log in to save this article

Abstract

In this study we describe new results of excavations in the Dinaledi Subsystem of the Rising Star cave system, South Africa. In two areas within the Hill Antechamber and the Dinaledi Chamber this work uncovered concentrations of abundant Homo naledi fossils including articulated, matrix-supported skeletal regions consistent with rapid covering by sediment prior to the decomposition of soft tissue. We additionally re-examine the spatial positioning of skeletal material and associated sediments within the Puzzle Box area, from which abundant H. naledi remains representing a minimum of six individuals were recovered in 2013 and 2014. Multiple lines of evidence exclude the hypothesis that skeletal remains from these three areas come from bodies that decomposed on the floor of the chamber or within a shallow depression prior to burial by sediments. The spatial positioning of skeletal material, the topography of the subsystem, and observations on sediments within and surrounding features exclude the hypothesis that rapid burial by sediment was a result of gravity-driven slumping or spontaneous movement of sediments. We present a minimal hypothesis of hominin cultural burial and test the evidence from all three areas, finding that this hypothesis is most compatible with the pattern of evidence. These results suggest that mortuary behavior including cultural burial was part of the repertoire of Homo naledi .

Article activity feed

  1. Version published to 10.7554/elife.89106.2 on eLife
  2. Version published to 10.7554/elife.89106 on eLife
  3. eLife

    eLife Assessment

    The authors study the context of the skeletal remains of three individuals and associated sediment samples to conclude that the hominin species Homo naledi intentionally buried their dead. Demonstration of the earliest known instance of intentional funerary practice - with a relatively small-brained hominin engaging in a highly complex behavior that has otherwise been observed from Homo sapiens and Homo neanderthalensis - would represent a landmark finding. The authors have revised their manuscript extensively in light of the reviews of their initial submission, with improved illustration, context, discussion, and theoretical frameworks, leading to an improved case supporting their conclusion that Homo naledi intentionally buried their dead. One of the reviewers concludes that the findings convincingly demonstrate intentional burial practices, while another considers evidence for such an unambiguous conclusion to be incomplete given a lack of definitive knowledge around how the hominins got into the chamber. We look forward to seeing the continued development and assessment of this hypothesis. It is worth noting that the detailed reviews (both rounds) and comprehensive author response are commendable and consequential parts of the scientific record of this study. The editors note that the authors' response repeatedly invokes precedent from previous publications to help justify the conclusions in this paper. While doing so is helpful, the editors also note that scientific norms and knowledge are constantly evolving, and that any study has to rest on its own scientific merit.

  4. eLife

    Reviewer #1 (Public review):

    Thank you for allowing me to review the paper "Evidence for deliberate burial of the dead by Homo naledi". This remains a very important site for paleoanthropology. I appreciate the work that the crew, especially the junior members of the team, put into this massive project. I appreciate that the authors did revise the paper since that is not a requirement of eLife. Extensive reviews by peer-reviewers have been provided for this paper, as well as professionally published replies (Martinón-Torres et al., 2023; Foecke et al., 2023). The composition, and citations of this version are much improved, though important information, some requested by reviewers, are buried in the supplementary section. It seems important that the authors make these sections more easily accessible to the general reader. The length of the paper is also unnecessary and impedes the readability of the work. Concise clarity is an expectation of most journals. The Netflix documentary was made to appeal to a mass audience, I would hope that the goal of the accompanying publication would be to enable readers to fully comprehend the work behind the claims.

    This version of the paper considers at great length many possibilities for how the H. naledi skeletal material came to rest in the cave system with some additional figures and data provided. However, quite a lot is still unclear. In my original review I stated, "The authors have repeatedly described how incredibly challenging it is to get into and out of this cave system and all of its chambers." This was a point emphasized in the Netflix documentary. In this version of the paper the authors have included within the supplementary section a brief discussion of other entrances. The work by Robbins et al. 2021 (a peer-reviewed paper in the impact factor rated journal Chemical Geology) is extremely relevant here. In this revision it is noted in the supplementary section that if the Postbox chamber was used as an opening, it would have reduced the length of the access to the system by 80 m. This fact seems important. This section should be moved out of the supplementary material and expanded because the conclusions published by Robbins et al. (2021) indicate a completely different route by which H. naledi accessed the cave, but this is hardly mentioned in the revision and deserves attention. To quote the Robbins et al.'s (2021) discussion section 6.3:

    "We acknowledge that additional data is required in order to confidently assess the relative timing of the Dragon's Back collapse and entry of H. naledi. Nonetheless, the stratigraphic and geochronologic observations presented here, together with those previously published (Dirks et al., 2017) are consistent with the following scenario. Prior to the collapse of the Dragon's Back, sometime before 241 ka (new minimum age for H. naledi from RS68), the cave could be entered by H. naledi via a shaft in the roof of the Postbox Chamber. From there H. naledi could walk along a straight passage that follows a gently descending, SW trending fracture into the Dragon's Back Chamber and, with the Dragon's Back block still attached to the roof, would have only needed to climb over a ~5 m high sill to access the Dinaledi Subsystem behind it. This sill and narrow fracture system behind the Dragon's Back block would have been a major impediment to any flood waters and most other fauna into the Dinaledi Subsystem, but it would have been a more accessible route than that today."

    The paper's conclusion continues, "The new dates further constrain the minimum age of H. naledi to 241 ka. Thus, H. naledi entered the subsystem between 241 ka and 335 ka, during a glacial period, when clastic sediment along the access route into the Dinaledi Subsystem experienced erosion. H. naledi would have probably entered the cave in the same way as the clastic sediments did, through an opening in the roof of the Postbox Chamber and may have entered via the Dragon's Back Chamber by climbing a 5 m high sill and passing below the Dragon's Back Block that was then still attached to the roof, to enter the Dinaledi Subsystem. In this context it is important to emphasize that it was not the Dragon's Back Block that prevented high-energy transport of coarse siliciclastic sediment from the Dragon's Back Chamber into the Dinaledi Subsystem, but rather the in situ floor block in the back wall of the Dragon's Back Chamber, against which the Dragon's Back Block slumped after it fell." This conclusion is very different from the complex pathway suggested by Berger et al. Martinón-Torres et al., 2023 also requested elaboration on this point in their reply by stating, "Moreover, recent studies by the Rising Star Cave team also point to a possible different and easier accesses for H. naledi into the fossil-bearing cave chambers than the current restricted access chute used by the research team, making clear that the degree of accessibility remains an open question (Robbins et al., 2021). Based on extensive dating studies of speleothem, this research (Robbins et al., 2021) implies that prior to 241 ka and the collapse of the Dragon's Back block hominins and other species could have more easily entered the cave via the Post Box Chamber and beneath the Dragon's Back Block before it fell. This gives access to a series of rifts that allow easier entry to the Dinaledi and other chambers beyond the present-day chute."

    Because this paper introduces very different sets of possibilities, it seems impossible to derive an understanding of the processes that occurred 335-241 ka throughout the cave system without going into detail on these other openings, especially openings that are hypothesized to have been used by the hominins in question.

    The world cares deeply about the H. naledi hominins and their story. I hope that in the coming years these issues are addressed, and perhaps other independent teams are allowed to do a full analysis since science is about replication. In any case, the excavation team has contributed important fossils to paleoanthropology.

    Literature cited:

    Foecke, Kimberly K., Queffelec, Alain, & Pickering, Robyn. (2023). No Sedimentological Evidence for Deliberate Burial by Homo naledi - A Case Study Highlighting the Need for Best Practices in Geochemical Studies Within Archaeology and Paleoanthropology. PaleoAnthropology, 2024. https://doi.org/10.48738/202x.issx.xxx

    Martinón-Torres, M., Garate, D., Herries, A. I. R., & Petraglia, M. D. (2023). No scientific evidence that Homo naledi buried their dead and produced rock art. Journal of Human Evolution, 103464. https://doi.org/10.1016/j.jhevol.2023.103464

    Robbins, J. L., Dirks, P. H. G. M., Roberts, E. M., Kramers, J. D., Makhubela, T. V., HilbertWolf, H. L., Elliott, M., Wiersma, J. P., Placzek, C. J., Evans, M., & Berger, L. R. (2021). Providing context to the Homo naledi fossils: Constraints from flowstones on the age of sediment deposits in Rising Star Cave, South Africa. Chemical Geology, 567, 120108. https://doi.org/10.1016/j.chemgeo.2021.120108

  5. eLife

    Reviewer #2 (Public review):

    Before providing my review of the revised version of this study by Berger et al., which explores potential deliberate burials of Homo naledi within the Rising Star Cave System, I would like to briefly summarize the key points from my previous review of the earlier version (in 2023). Summarizing my previous review will provide context for assessing how effectively the revised study addresses the concerns I raised previously (in 2023).

    In my earlier comments, I highlighted significant methodological and analytical shortcomings that, in my view, undermined the authors' claim of intentional burials by Homo naledi. While the study presented detailed geological and fossil data, I found the evidence for intentional burials unconvincing due to insufficient application of archaeothanatological principles and other methodological gaps.

    My key concerns included:

    (1) The absence of a comprehensive archaeothanatological analysis, particularly with respect to taphonomic changes, bone articulations, and displacement patterns such as the collapse of sediments and bone remains into voids created by decomposition.

    (2) Missing or unclear illustrations of bone arrangements, which are critical for interpreting burial positions and processes.

    (3) A lack of detailed discussion on the sequence of decomposition, joint disarticulation, sediment infill, and secondary bone displacement.

    To convincingly support claims of deliberate burial, I argued that the study must reconstruct the timeline and processes surrounding death and deposition while clearly distinguishing natural taphonomic changes from intentional human actions. I emphasized the importance of integrating established archaeothanatological frameworks, such as those outlined by Duday et al. or Boulestin et al., to provide the necessary analytical rigor.

    I will now explain how the revised version of this study has successfully addressed all the concerns raised in my previous review and why I now think that the authors provide sufficient evidence for the presence of "repeated and patterned" deliberate burials (referred to as "cultural burials" by the authors) by Homo naledi within the Rising Star Cave System.

    In their revised manuscript, the authors have implemented substantial improvements in methodology, analytical depth, and overall presentation, which have effectively resolved the critical issues I previously highlighted. These revisions greatly strengthen their argument for intentional funerary practices. Importantly, the authors remain cautious in their interpretation of the evidence, explicitly refraining from inferring "symbolic" behavior or complex cognitive motivations behind these burials. Instead, they focus on presenting clear evidence for deliberate, patterned practices while leaving the broader implications for Homo naledi's cultural and cognitive capacities open for further investigation. This cautious approach adds to the credibility of their conclusions and avoids overextending the interpretation of the data.

    The authors' enhanced application of archaeothanatological principles now offers a more comprehensive and convincing interpretation of the burial features. Key gaps in the earlier version, such as the absence of detailed reconstructions of taphonomic processes, bone articulations, and displacement patterns, have been addressed with thorough analyses and clearer illustrations. The study also now includes a well-structured timeline of events surrounding death and deposition, demonstrating an improved ability to differentiate between natural processes and deliberate human actions. These additions lend greater clarity and rigor to the evidence, making the argument for intentional burials both robust and persuasive.

    Furthermore, the revised study presents detailed data on skeletal arrangements, decomposition sequences, and spatial patterns. This information is now relatively well illustrated and contextualized, enabling readers to better understand the complex processes involved in these burial practices. Importantly, the authors provide a stronger theoretical framework, integrating established archaeothanatological methodologies and taphonomic studies that situate their findings within broader archaeological and anthropological discussions of funerary behavior.

    That being said, there remain relatively minor issues that could be refined further. Addressing these would help ensure the study is as clear and accessible as possible to the reader. Such adjustments would enhance the overall readability and reinforce the study's impact within the scientific community.

    A - Suggested changes:

    While the revised version of this study marks a significant improvement, successfully addresses my previous major concerns and provides a convincing argument for deliberate burials by Homo naledi, I believe that including both one summary table + one summary figure for each of the three main locations and the-Hill Antechamber, and Dinaledi Chamber (Feature 1 and Puzzle Box)-would further enhance the clarity and accessibility of the findings. Such tables and figures would serve as a valuable reference, allowing readers to more easily follow how the detailed patterns observed at each site fit the criteria for distinguishing intentional from natural processes.

    The summary tables should consolidate key information for each location, such as:

    (1) Bone articulations: A comprehensive list of articulated skeletal elements, categorized by their anatomical relationships (e.g., labile vs. stable articulations).

    (2) Displacement patterns: Documentation of any spatial shifts in bone positions, noting directions and extents of disarticulation.

    (3) Sequence of decomposition: Observations regarding the sequence of decomposition, joint disarticulation and associated changes in bone arrangements.

    (4) Sediment interaction: Notes on sediment infill and its timing relative to decomposition, including evidence of secondary voids or delayed sediment deposition.

    (5) Distinguishing criteria: Clear indications of how each observed pattern supports intentional burial (e.g., structured placement, lack of natural transport mechanisms) versus natural processes (e.g., random dispersal, sediment-driven bone displacement).
    Including such tables would not only summarize the complex taphonomic and archaeothanatological data but also allow readers to quickly assess how the evidence supports the authors' conclusions. This approach would bridge the gap between the detailed narrative descriptions and the criteria necessary to differentiate deliberate funerary practices from natural occurrences.

    To streamline the main text further, many of the detailed descriptions of individual bones, specific displacement measurements, and other intricate observations could be moved to the supplementary data. This reorganization would maintain the richness of the data for those who wish to explore it in depth, while the summary tables would present the key findings concisely in the main text. This balance between accessibility and detail would ensure that the study appeals to both specialists requiring comprehensive data and readers looking for an overarching understanding of the findings.

    In addition to these structural changes, it is crucial to ensure that evidence is consistently illustrated throughout the text.

    Importantly the skeletal part representation is provided for Dinaledi Feature 1 in Figure 14, but similar data is not presented for the other burial features, such as those in the Hill Antechamber or Puzzle Box. This inconsistency could make it more challenging for readers to compare the features and fully appreciate the patterns of burial behavior across the different locations. Ensuring that similar types of evidence and analyses are presented uniformly for all features would strengthen the study and make its conclusions more cohesive and compelling.

    Adding supplementary figures to represent the skeletal part distribution (as in Figure 14) within each excavated area (i.e., not only for Dinaledi Feature 1 but also for Hill Antechamber and Puzzle Box) would significantly enhance the study's clarity and accessibility. These figures could provide a visual summary of skeletal part representation, allowing readers to easily understand the nature of human remains within each burial context.

    Specifically, such figures could:

    (1) Illustrate Skeletal Part Representation: By visually mapping the presence and location of various skeletal elements, the figures would make it easier for readers to assess the completeness and arrangement of remains in each feature. This is particularly important for interpreting patterns of bone articulation and disarticulation.
    For example, it is quite challenging to determine the exact number and characteristics of the human skeletal remains identified within the Puzzle Box and those recovered through the "subsurface collection" in its surrounding area. The authors state that "at least six individuals" were identified in this area (during "subsurface collection") but provide no further clarification. They simply mention that "most elements" were described previously, without specifying which elements or where this prior description can be found.

    (2) Highlight Articulations and Displacements: Figures could indicate which bones are articulated and their relative positions, as well as the spatial distribution of disarticulated elements. This would provide a clear visual context to support interpretations of taphonomic processes.

    (3) Facilitate Comparisons Across Locations: By presenting skeletal part representation consistently for each location, the figures would enable readers to directly compare features, reinforcing the argument for "repeated and patterned" behavior.

    (4) Simplify Complex Data: Instead of relying solely on textual descriptions, the visual format would allow readers to quickly grasp the key findings, making the study more accessible to a broader audience

    By including such figures alongside the proposed summary tables in the main text, the study would achieve a balance between detailed narrative descriptions and concise, visual representation of the data. This approach would strengthen the overall presentation and support the authors' conclusions effectively.

    Again, by presenting the data in a structured and comparative format, the new tables + figures could also highlight the differences and similarities between the three locations. This would reinforce the argument for "repeated and patterned" behavior, as the tables would make it easier to observe consistent burial practices across different contexts within the Rising Star Cave System.

    Adding these summary tables + figures, ensuring consistent presentation of evidence, and reallocating detailed descriptions to supplementary materials would not require significant new analysis. However, these organizational adjustments would greatly enhance the study's clarity, readability, and overall impact.

    B - A few additional changes are needed:

    Figure 8: This figure is critical but lacks clarity. Specifically:

    Panels 8a-c suffer from low contrast, making details difficult to discern.
    Panel 8d (sediment profile) is too small and lacks annotations that would aid interpretation.
    Figure S7: While this figure has significantly better contrast than Figures 8a-c, I am unable to identify the "articulated foot ... at right of frame," as mentioned in the caption. Please clarify this by adding annotations directly to the figure.

    Page 4, 2nd paragraph: In the sentence "Researchers thus have diverse opinions about how to test whether ...," the word "opinions" should be replaced with a more precise term, such as "approaches."

    C - In conclusion, I am impressed by the significant effort and meticulous work that has gone into this revised version of the study. The quality of the new evidence presented is commendable, and the findings now convincingly demonstrate not only clear evidence of intentional burial practices by Homo naledi but also compelling indications of post-depositional reworking. These advancements reflect a major improvement in the study's analytical rigor and the robustness of its conclusions, making it a valuable contribution to the understanding of early hominin funerary behavior.

  6. eLife

    Author response:

    The following is the authors’ response to the original reviews

    We extend our sincere thanks to the editor, referees for eLife, and other commentators who have written evaluations of this manuscript, either in whole or in part. Sources of these comments were highly varied, including within the bioRxiv preprint server, social media (including many comments received on X/Twitter and some YouTube presentations and interviews), comments made by colleagues to journalists, and also some reviews of the work published in other academic journals. Some of these are formal and referenced with citations. Others were informal but nonetheless expressed perspectives that helped enable us to revise the manuscript with the inclusion of broader perspectives than the formal review process. It is beyond the scope of this summary to list every one of these, which have often been brought to the attention of different coauthors, but we begin by acknowledging the very wide array of peer and public commentary that have contributed to this work. The reaction speaks to a broad interest in open discussion and review of preprints.

    As we compiled this summary of changes to the manuscript, we recognized that many colleagues made comments about the process of preprint dissemination and evaluation rather than the data or analyses in the manuscript. Addressing such comments is outside the scope of this revised manuscript, but we do feel that a broader discussion of these comments would be valuable in another venue. Many commentators have expressed confusion about the eLife system of evaluation of preprints, which differs from the editorial acceptance or rejection practiced in most academic journals. As authors in many different nations, in varied fields, and in varied career stages, we ourselves are still working to understand how the academic publication landscape is changing, and how best to prepare work for new models of evaluation and dissemination.

    The manuscript and coauthor list reflect an interdisciplinary collaboration. Analyses presented in the manuscript come from a wide range of scientific disciplines. These range from skeletal inventory, morphology, and description, spatial taphonomy, analysis of bone fracture patterns and bone surface modifications, sedimentology, geochemistry, and traditional survey and mapping. The manuscript additionally draws upon a large number of previous studies of the Rising Star cave system and the Dinaledi Subsystem, which have shaped our current work. No analysis within any one area of research stands alone within this body of work: all are interpreted in conjunction with the outcomes of other analyses and data from other areas of research. Any single analysis in isolation might be consistent with many different hypotheses for the formation of sediments and disposition of the skeletal remains. But testing a hypothesis requires considering all data in combination and not leaving out data that do not fit the hypothesis. We highlight this general principle at the outset because a number of the comments from referees and outside specialists have presented alternative hypotheses that may arguably be consistent with one kind of analysis that we have presented, while seeming to overlook other analyses, data, or previous work that exclude these alternatives. In our revision, we have expanded all sections describing results to consider not only the results of each analysis, but how the combination of data from different kinds of analysis relate to hypotheses for the deposition and subsequent history of the Homo naledi remains. We address some specific examples and how we have responded to these in our summary of changes below.

    General organization

    The referee and editor comments are mostly general and not line-by-line questions, and we have compiled them and treated them as a group in this summary of changes, except where specifically noted.

    The editorial comments on the previous version included the suggestion that the manuscript should be reorganized to test “natural” (i.e. noncultural) hypotheses for the situations that we examine. The editorial comment suggested this as a “null hypothesis” testing approach. Some outside comments also viewed noncultural deposition as a null hypothesis to be rejected. We do not concur that noncultural processes should be construed as a null hypothesis, as we discuss further below. However, because of the clear editorial opinion we elected to revise the manuscript to make more explicit how the data and analyses test noncultural depositional hypotheses first, followed by testing of cultural hypotheses. This reorganization means that the revised manuscript now examines each hypothesis separately in turn.

    Taking this approach resulted in a substantial reorganization of the “Results” section of the manuscript. The “Results” section now begins with summaries of analyses and data conducted on material from each excavation area. After the presentation of data and analyses from each area, we then present a separate section for each of several hypotheses for the disposition and sedimentary context of the remains. These hypotheses include deposition of bodies upon a talus (as hypothesized in some previous work), slow sedimentary burial on a cave floor or within a natural depression, rapid burial by gravity-driven slumping, and burial of naturally mummified remains. We then include sections to test the hypothesis of primary cultural burial and secondary cultural burial. This approach adds substantial length to the Results. While some elements may be repeated across sections, we do consider the new version to be easier to take piece by piece for a reader trying to understand how each hypothesis relates to the evidence.

    The Results section includes analyses on several different excavation areas within the Dinaledi Subsystem. Each of these presents somewhat different patterns of data. We conceived of this manuscript combining these distinct areas because each of them provides information about the formation history of the Homo naledi-associated sediments and the deposition of the Homo naledi remains. Together they speak more strongly than separately. In the previous version of the manuscript, two areas of excavation were considered in detail (Dinaledi Feature 1 and the Hill Antechamber Feature), with a third area (the Puzzle Box area) included only in the Discussion and with reference to prior work. We now describe the new work undertaken after the 2013-2014 excavations in more detail. This includes an overview of areas in the Hill Antechamber and Dinaledi Chamber that have not yielded substantial H. naledi remains and that thereby help contextualize the spatial concentration of H. naledi skeletal material. The most substantial change in the data presented is a much expanded reanalysis of the Puzzle Box area. This reanalysis provides greater clarity on how previously published descriptions relate to the new evidence. The reanalysis also provides the data to integrate the detailed information on bone identification fragmentation, and spatial taphonomy from this area with the new excavation results from the other areas.

    In addition to Results, the reorganization also affected the manuscript’s Introduction section. Where the previous version led directly from a brief review of Pleistocene burial into the description of the results, this revised manuscript now includes a review of previous studies of the Rising Star cave system. This review directly addresses referee comments that express some hesitation to accept previous results concerning the structure and formation of sediments, the accessibility of the Dinaledi Subsystem, the geochronological setting of the H. naledi remains, and the relation of the Dinaledi Subsystem to nearby cave areas. Some parts of this overview are further expanded in the Supplementary Information to enable readers to dive more deeply into the previous literature on the site formation and geological configuration of the Rising Star cave system without needing to digest the entirety of the cited sources.

    The Discussion section of the revised manuscript is differentiated from Results and focuses on several areas where the evidence presented in this study may benefit from greater context. One new section addresses hypothesis testing and parsimony for Pleistocene burial evidence, which we address at greater length in this summary below. The majority of the Discussion concerns the criteria for recognizing evidence for burial as applied in other studies. In this research we employ a minimal definition but other researchers have applied varied criteria. We consider whether these other criteria have relevance in light of our observations and whether they are essential to the recognition of burial evidence more broadly.

    Vocabulary

    We introduce the term “cultural burial” in this revised manuscript to refer to the burial of dead bodies as a mortuary practice. “Burial” as an unmodified term may refer to the passive covering of remains by sedimentary processes. Use of the term “intentional burial” would raise the question of interpreting intent, which we do not presume based on the evidence presented in this research. The relevant question in this case is whether the process of burial reflects repeated behavior by a group. As we received input from various colleagues it became clear that burial itself is a highly loaded term. In particular there is a common assumption within the literature and among professionals that burial must by definition be symbolic. We do not take any position on that question in this manuscript, and it is our hope that the term “cultural burial” may focus the conversation around the extent that the behavioral evidence is repeated and patterned.

    Sedimentology and geochemistry of Dinaledi Feature 1

    Reviewer 4 provided detailed comments on the sedimentological and geochemical context that we report in the manuscript. One outside review (Foecke et al. 2024) included some of the points raised by reviewer 4, and additionally addressed the reporting of geochemical and sedimentological data in previous work that we cite.

    To address these comments we have revised the sedimentary context and micromorphology of sediments associated with Dinaledi Feature 1. In the new text we demonstrate the lack of microstratigraphy (supported by grain size analysis) in the unlithified mud clast breccia (UMCB), while such a microstratigraphy is observed in the laminated orange-red mudstones

    (LORM) that contribute clasts to the UMCB. Thus, we emphasize the presence and importance

    of a laterally continuous layer of LORM nature occurring at a level that appears to be the maximum depth of fossil occurrence. This layer is severely broken under extensive accumulation of fossils such as Feature 1 and only evidenced by abundant LORM clasts within and around the fossils.

    We have completely reworked the geochemical context associated with Feature 1 following the comments of reviewer 4. We described the variations and trends observed in the major oxides separate from trace and rare-earth elements. We used Harker variations plots to assess relationships between these element groups with CaO and Zn, followed by principal component analysis of all elements analyzed. The new geochemical analysis clearly shows that Feature 1 is associated with localized trace element signatures that exist in the sediments only in association with the fossil bones, which suggests lack of postdepositional mobilization of the fossils and sediments. We additionally have included a fuller description of XRF methods.

    To clarify the relation of all results to the features described in this study, we removed the geochemical and sedimentological samples from other sites within the Dinaledi Subsystem. These localities within the fissure network represent only surface collection of sediment, as no excavation results are available from those sites to allow for comparison in the context of assessing evidence of burial. These were initially included for comparison, but have now been removed to avoid confusion.

    Micromorphology of sediments

    Some referees (1, 3, and 4) and other commentators (including Martinón-Torres et al. 2024) have suggested that the previous manuscript was deficient due to an insufficient inclusion of micromorphological analysis of sediments. Because these commentators have emphasized this kind of evidence as particularly important, we review here what we have included and how our revision has addressed this comment. Previous work in the Dinaledi Chamber (Dirks et al., 2015; 2017) included thin section illustrations and analysis of sediment facies, including sediments in direct association with H. naledi remains within the Puzzle Box area. The previous work by Wiersma and coworkers (2020) used micromorphological analysis as one of several approaches to test the formation history of Unit 3 sediments in the Dinaledi Subsystem, leading to the interpretation of autobrecciation of earlier Unit 1 sediment. In the previous version of this manuscript we provided citations to this earlier work. The previous manuscript also provided new thin section illustrations of Unit 3 sediment near Dinaledi Feature 1 to place the disrupted layer of orange sediment (now designated the laminated orange silty mudstone unit) into context.

    In the new revised manuscript we have added to this information in three ways. First, as noted above in response to reviewer 4, we have revised and added to our discussion of

    micromorphology within and adjacent to the Dinaledi Feature 1. Second, we have included more discussion in the Supplementary Information of previous descriptions of sediment facies and associated thin section analysis, with illustrations from prior work (CC-BY licensed) brought into this paper as supplementary figures, so that readers can examine these without following the citations. Third, we have included Figure 10 in the manuscript which includes six panels with microtomographic sections from the Hill Antechamber Feature. This figure illustrates the consistency of sub-unit 3b sediment in direct contact with H. naledi skeletal material, including anatomically associated skeletal elements, with previous analyses that demonstrate the angular outlines and chaotic orientations of LORM clasts. It also shows density contrasts of sediment in immediate contact with some skeletal elements, the loose texture of this sediment with air-filled voids, and apparent invertebrate burrowing activity. To our knowledge this is the first application of microtomography to sediment structure in association with a Pleistocene burial feature.

    To forestall possible comments that the revised manuscript does not sufficiently employ micromorphological observations, or that any one particular approach to micromorphology is the standard, we present here some context from related studies of evidence from other research groups working at varied sites in Africa, Europe, and Asia. Hodgkins et al. (2021) noted: “Only a handful of micromorphological studies have been conducted on human burials and even fewer have been conducted on suspected burials from Paleolithic or hunter-gatherer contexts.” In that study, one supplementary figure with four photomicrographs of thin sections of sediments was presented. Interpretation of the evidence for a burial pit by Hodgkins et al. (2021) noted the more open microstructure of sediment but otherwise did not rely upon the thin section data in characterizing the sediments associated with grave fill. Martinón-Torres et al. (2021) included one Extended Data figure illustrating thin sections of sediments and bone, with two panels showing sediments (the remainder showing bone histology). The micromorphological analysis presented in the supplementary information of that paper was restricted to description of two microfacies associated with the proposed “pit” in that study. That study did carry out microCT scanning of the partially-prepared skeletal remains but did not report any sediment analysis from the microtomographic results. Maloney et al. (2022) reported no micromorphological or thin section analysis. Pomeroy et al. (2020a) included one illustration of a thin section; this study may be regarded as a preliminary account rather than a full description of the work undertaken. Goldberg et al. (2017) analyzed the geoarchaeology of the Roc de Marsal deposits in which possible burial-associated sediments had been fully excavated in the 1960s, providing new morphological assessments of sediment facies; the supplementary information to this work included five scans (not microscans) of sediment thin sections and no microphotographs. Fewlass et al. (2023) presented no thin section or micromorphological illustrations or methods. In summary of this research, we note that in one case micromorphological study provided observations that contributed to the evidence for a pit, in other cases micromorphological data did not test this hypothesis, and many researchers do not apply micromorphological techniques in their particular contexts.

    Sediment micromorphology is a growing area of research and may have much to provide to the understanding of ancient burial evidence as its standards continue to develop (Pomeroy et al. 2020b). In particular microtomographic analysis of sediments, as we have initiated in this study, may open new horizons that are not possible with more destructive thin-section preparation. In this manuscript, the thin section data reveals valuable evidence about the disruption of sediment structure by features within the Dinaledi Chamber, and microtomographic analysis further documents that the Hill Antechamber Feature reflects similar processes, in addition to possible post-burial diagenesis and invertebrate activity. Following up in detail on these processes will require further analysis outside the scope of this manuscript.

    Access into the Dinaledi Subsystem

    Reviewer 1 emphasizes the difficulty of access into the Dinaledi Subsystem as a reason why the burial hypothesis is not parsimonious. Similar comments have been made by several outside commentators who question whether past accessibility into the Dinaledi Subsystem may at one time have been substantially different from the situation documented in previous work. Several pieces of evidence are relevant to these questions and we have included some discussion of them in the Introduction, and additionally include a section in the Supplementary Information (“Entrances to the cave system”) to provide additional context for these questions. Homo naledi remains are found not only within the Dinaledi Subsystem but also in other parts of the cave system including the Lesedi Chamber, which is similarly difficult for non-expert cavers to access. The body plan, mass, and specific morphology of H. naledi suggest that this species would be vastly more suited to moving and climbing within narrow underground passages than living people. On this basis it is not unparsimonious to suggest that the evidence resulted from H. naledi activity within these spaces. We note that the accessibility of the subsystem is not strictly relevant to the hypothesis of cultural burial, although the location of the remains does inform the overall context which may reflect a selection of a location perceived as special in some way.

    Stuffing bodies down the entry to the subsystem

    Reviewer 3 suggests that one explanation for the emplacement of articulated remains at the top of the sloping floor of the Hill Antechamber is that bodies were “stuffed” into the chute that comprises the entry point of the subsystem and passively buried by additional accumulation of remains. This was one hypothesis presented in earlier work (Dirks et al. 2015) and considered there as a minimal explanation because it did not entail the entry of H. naledi individuals into the subsystem. The further exploration (Elliott et al. 2021) and ongoing survey work, as well as this manuscript, all have resulted in data that rejects this hypothesis. The revised manuscript includes a section in the results “Deposition upon a talus with passive burial” that examines this hypothesis in light of the data.

    Recognition of pits

    Referee 3 and 4 and several additional commentators have emphasized that the recognition of pit features is necessary to the hypothesis of burial, and questioned whether the data presented in the manuscript were sufficient to demonstrate that pits were present. We have revised the manuscript in several ways to clarify how all the different kinds of evidence from the subsystem test the hypothesis that pits were present. This includes the presentation of a minimal definition of burial to include a pit dug by hominins, criteria for recognizing that a pit was present, and an evaluation of the evidence in each case to make clear how the evidence relates to the presence of a pit and subsequent infill. As referee 3 notes, it can be challenging to recognize a pit when sediment is relatively homogeneous. This point was emphasized in the review by Pomeroy and coworkers (2020b), who reflected on the difficulty seeing evidence for shallow pits constructed by hominins, and we have cited this in the main text. As a result, the evidence for pits has been a recurrent topic of debate for most Pleistocene burial sites. However in addition to the sedimentological and contextual evidence in the cases we describe, the current version also reflects upon other possible mechanisms for the accumulation of bones or bodies. The data show that the sedimentary fill associated with the H. naledi remains in the cases we examine could not have passively accumulated slowly and is not indicative of mass movement by slumping or other high-energy flow. To further put these results into context, we added a section to the Discussion that briefly reviews prior work on distinguishing pits in Pleistocene burial contexts, including the substantial number of sites with accepted burial evidence for which no evidence of a pit is present.

    Extent of articulation and anatomical association

    We have added significantly greater detail to the descriptions of articulated remains and orientation of remains in order to describe more specifically the configuration of the skeletal material. We also provide 14 figures in main text (13 of them new) to illustrate the configuration of skeletal remains in our data. For the Puzzle Box area, this now includes substantial evidence on the individuation of skeletal fragments, which enables us to illustrate the spatial configuration of remains associated with the DH7 partial skeleton, as well as the spatial position of fragments refitted as part of the DH1, DH2, DH3, and DH4 crania. For Dinaledi Feature 1 and the Hill Antechamber Feature we now provide figures that key skeletal parts as identified, including material that is unexcavated where possible, and a skeletal part representation figure for elements excavated from Dinaledi Feature 1.

    Archaeothanatology

    Reviewer 2 suggests that a greater focus on the archaeothanatology literature would be helpful to the analysis, with specific reference to the sequence of joint disarticulation, the collapse of sediment and remains into voids created by decomposition, and associated fragmentation of the remains. In the revised manuscript we have provided additional analysis of the Hill Antechamber Feature with this approach in mind. This includes greater detail and illustration of our current hypothesis for individuation of elements. We now discuss a hypothesis of body disposition, describe the persistent joints and articulation of elements, and examine likely decomposition scenarios associated with these remains. Additionally, we expand our description and illustration of the orientation of remains and degree of anatomical association and articulation within Dinaledi Feature 1. For this feature and for the Hill Antechamber Feature we have revised the text to describe how fracturing and crushing patterns are consistent with downward pressure from overlying sediment and material. In these features, postdepositional fracturing occurred subsequent to the decomposition of soft tissue and partial loss of organic integrity of the bone. We also indicate that the loss by postdepositional processes of most long bone epiphyses, vertebral bodies, and other portions of the skeleton less rich in cortical bone, poses a challenge for testing the anatomical associations of the remaining elements. This is a primary reason why we have taken a conservative approach to identification of elements and possible associations.

    A further aspect of the site revealed by our analysis is the selective reworking of sediments within the Puzzle Box area subsequent to the primary deposition of some bodies. The skeletal evidence from this area includes body parts with elements in anatomical association or articulation, juxtaposed closely with bone fragments at varied pitch and orientation. This complexity of events evidenced within this area is a challenge for approaches that have been developed primarily based on comparative data from single-burial situations. In these discussions we deepen our use of references as suggested by the referee.

    Burial positions

    Reviewer 2 further suggests that illustrations of hypothesized burial positions would be valuable. We recognize that a hypothesized burial position may be an appealing illustration, and that some recent studies have created such illustrations in the context of their scientific articles. However such illustrations generally include a great deal of speculation and artist imagination, and tend to have an emotive character. We have added more discussion to the manuscript of possible primary disposition in the case of the Hill Antechamber Feature as discussed above. We have not created new illustrations of hypothesized burial positions for this revision.

    Carnivore involvement

    Referee 1 suggests that the manuscript should provide further consideration of whether carnivore activity may have introduced bones or bodies into the cave system. The reorganized Introduction now includes a review of previous work, and an expanded discussion within the Supplementary Information (“Hypotheses tested in previous work”). This includes a review of literature on the topic of carnivore accumulation and the evidence from the Dinaledi and Lesedi Chamber that rejects this hypothesis.

    Water transport and mud

    The eLife referees broadly accepted previous work showing that water inundation or mass flow of water-saturated sediment did not occur within the history of Unit 2 and 3 sediments, including those associated with H. naledi remains. However several outside commentators did refer specifically to water flow or mud flow as a mechanism for slumping of deposits and possible sedimentary covering of the remains. To address these comments we have added a section to the

    Supplementary Information (“Description of the sedimentary deposits of the Dinaledi Subsystem”) that reviews previous work on the sedimentary units and formation processes documented in this area. We also include a subsection specifically discussing the term “mud” as used in the description of the sedimentology within the system, as this term has clearly been confusing for nonspecialists who have read and commented on the work. We appreciate the referees’ attention to the previous work and its terminology.

    Redescription of areas of the cave system

    Reviewer 1 suggests that a detailed reanalysis of all portions of the cave system in and around the Dinaledi Subsystem is warranted to reject the hypothesis that bodies entered the space passively and were scattered from the floor by natural (i.e. noncultural) processes. The referee suggests that National Geographic could help us with these efforts. To address this comment we have made several changes to the manuscript. As noted above, we have added material in Supplementary Information to review the geochronology of the Dinaledi Subsystem and nearby Dragon’s Back Chamber, together with a discussion of the connections between these spaces.

    Most directly in response to this comment we provide additional documentation of the possibility of movement of bodies or body parts by gravity within the subsystem itself. This includes detailed floor maps based on photogrammetry and LIDAR measurement, where these are physically possible, presented in Figures 2 and 3. In some parts of the subsystem the necessary equipment cannot be used due to the extremely confined spaces, and for these areas our maps are based on traditional survey methods. In addition to plan maps we have included a figure showing the elevation of the subsystem floor in a cross-section that includes key excavation areas, showing their relative elevation. All figures that illustrate excavation areas are now keyed to their location with reference to a subsystem plan. These data have been provided in previous publications but the visualization in the revised manuscript should make the relationship of areas clear for readers. The Introduction now includes text that discusses the configuration of the Hill Antechamber, Dinaledi Chamber, and nearby areas, and also discusses the instances in which gravity-driven movement may be possible, at the same time reviewing that gravity-driven movement from the entry point of the subsystem to most of the localities with hominin skeletal remains is not possible.

    Within the Results, we have added a section on the relationship of features to their surroundings in order to assist readers in understanding the context of these bone-bearing areas and the evidence this context brings to the hypothesis in question. We have also included within this new section a discussion of the discrete nature of these features, a question that has been raised by outside commentators.

    Passive sedimentation upon a cave floor or within a natural depression

    Reviewer 3 suggests that the situation in the Dinaledi Subsystem may be similar to a European cave where a cave bear skeleton might remain articulated on a cave floor (or we can add, within a hollow for hibernation), later to be covered in sediment. The reviewer suggests that articulation is therefore no evidence of burial, and suggests that further documentation of disarticulation processes is essential to demonstrating the processes that buried the remains. We concur that articulation by itself is not sufficient evidence of cultural burial. To address this comment we have included a section in the Results that tests the hypothesis that bodies were exposed upon the cave floor or within a natural depression. To a considerable degree, additional data about disarticulation processes subsequent to deposition are provided in our reanalysis of the Puzzle Box area, including evidence for selective reworking of material after burial.

    Postdepositional movement and floor drains

    Reviewer 3 notes that previous work has suggested that subsurface floor drains may have caused some postdepositional movement of skeletal remains. The hypothesis of postdepositional slumping or downslope movement has also been discussed by some external commentators (including Martinón-Torres et al. 2024). We have addressed this question in several places within the revised manuscript. As we now review, previous discussion of floor drains attempted to explain the subvertical orientation of many skeletal elements excavated from the Puzzle Box area. The arrangement of these bones reflects reworking as described in our previous work, and without considering the possibility of reworking by hominins, one mechanism that conceivably might cause reworking was downward movement of sediments into subsurface drains. Further exploration and mapping, combined with additional excavation into the sediments beneath the Puzzle Box area provided more information relevant to this hypothesis. In particular this evidence shows that subsurface drains cannot explain the arrangement of skeletal material observed within the Puzzle Box area. As now discussed in the text, the reworking is selective and initiated from above rather than below. This is best explained by hominin activity subsequent to burial.

    In a new section of the Results we discuss slumping as a hypothesis for the deposition of the remains. This includes discussion of downslope movement within the Hill Antechamber and the idea that floor drains may have been a mechanism for sediment reworking in and around the Puzzle Box area and Dinaledi Feature 1. As described in this section the evidence does not support these hypotheses.

    Hypothesis testing and parsimony

    Referees 1 and 3 and the editorial guidance all suggested that a more appropriate presentation would adopt a null hypothesis and test it. The specific suggestion that the null hypothesis should be a natural sedimentary process of deposition was provided not only by these reviewers but also by some outside commentators. To address this comment, we have edited the manuscript in two ways. The first is the addition of a section to the Discussion that specifically discusses hypothesis testing and parsimony as related to Pleistocene evidence of cultural burial. This includes a brief synopsis of recent disciplinary conversations and citation of work by other groups of authors, none of whom adopted this “null hypothesis” approach in their published work.

    As we now describe in the manuscript, previous work on the Dinaledi evidence never assumed any role for H. naledi in the burial of remains. Reading the reviewer reports caused us to realize that this previous work had followed exactly the “null hypothesis” approach that some suggested we follow. By following this null hypothesis approach, we neglected a valuable avenue of investigation. In retrospect, we see how this approach impeded us from understanding the pattern of evidence within the Puzzle Box area. Thus in the revised manuscript we have mentioned this history within the Discussion and also presented more of the background to our previous work in the Introduction. Hopefully by including this discussion of these issues, the manuscript will broaden conversation about the relation of parsimony to these issues.

    Language and presentation style

    Reviewer 4 criticizes our presentation, suggesting that the text “gives the impression that a hypothesis was formulated before data were collected.” Other outside commentators have mentioned this notion also, including Martinón-Torres et al. (2024) who suggest that the study began from a preferred hypothesis and gathered data to support it. The accurate communication of results and hypotheses in a scientific article is a broader issue than this one study. Preferences about presentation style vary across fields of study as well as across languages. We do not regret using plain language where possible. In any study that combines data and methods from different

    scientific disciplines, the use of plain language is particularly important to avoid misunderstandings where terms may mean different things in different fields.

    The essential question raised by these comments is whether it is appropriate to present the results of a study in terms of the hypothesis that is best supported. As noted above, we read carefully many recent studies of Pleistocene burial evidence. We note that in each of these studies that concluded that burial is the best hypothesis, the authors framed their results in the same way as our previous manuscript: an introduction that briefly reviews background evidence for treatment of the dead, a presentation of results focused on how each analysis supports the hypothesis of burial for the case, and then in some (but not all) cases discussion of why some alternative hypotheses could be rejected. We do not infer from this that these other studies started from a presupposition and collected data only to confirm it. Rather, this is a simple matter of presentation style.

    The alternative to this approach is to present an exhaustive list of possible hypotheses and to describe how the data relate to each of them, at the end selecting the best. This is the approach that we have followed in the revised manuscript, as described above under the direction of the reviewer and editorial guidance. This approach has the advantage of bringing together evidence in different combinations to show how each data point rejects some hypotheses while supporting others. It has the disadvantage of length and repetition.

    Possible artifact

    We have chosen to keep the description of the possible artifact associated with the Hill Antechamber Feature in the Supplementary Information. We do this while acknowledging that this is against the opinion of reviewer 4, who felt the description should be removed unless the object in question is fully excavated and physically analyzed. The previous version of the manuscript did not rely upon the stone as positive evidence of grave goods or symbolic content, and it noted that the data do not test whether the possible artifact was placed or was intentionally modified. However this did not satisfy reviewer 4, and some outside commentators likewise asserted that the object must be a “geofact” and that it should be removed.

    We have three arguments against this line of thinking. First, we do not omit data from our reporting. Whether Homo naledi shaped the rock or not, used it as a tool or not, whether the rock was placed with the body or not, it is unquestionably there. Omitting this one object from the report would be simply dishonest. Second, the data on this rock are at 16 micron resolution. While physical inspection of its surface may eventually reveal trace evidence and will enable better characterization of the raw material, no mode of surface scanning will produce better evidence about the object’s shape. Third, the position of this possible artifact within the feature provides significant information about the deposition of the skeletal material and associated sediments. The pitch, orientation, and position of the stone is not consistent with slow deposition but are consistent with the hypothesis that the surrounding sediment was rapidly emplaced at the same time as the articulated elements less than 2 cm away.

    In the current version, we have redoubled our efforts to provide information about the position and shape of this stone while not presupposing the intentionality of its shape or placement. We add here that the attitude expressed by referee 4 and other commentators, if followed at other sites, would certainly lead to the loss or underreporting of evidence, which we are trying to avoid.

    Consistency versus variability of behavior

    As described in the revised manuscript, different features within the Dinaledi Subsystem exhibit some shared characteristics. At the same time, they vary in positioning, representation of individuals and extent of commingling. Other localities within the subsystem and broader cave system present different evidence. Some commentators have questioned whether the patterning is consistent with a single common explanation, or whether multiple explanations are necessary. To address this line of questioning, we have added several elements to the manuscript. We created a new section on secondary cultural burial, discussing whether any of the situations may reflect this practice. In the Discussion, we briefly review the ways in which the different features support the involvement of H. naledi without interpreting anything about the intentionality or meaning of the behavior. We further added a section to the Discussion to consider whether variation among the features reflects variation in mortuary practices by H. naledi. One aspect of this section briefly cites variation in the location and treatment of skeletal remains at other sites with evidence of burial.

    Grave goods

    Some commentators have argued that grave goods are a necessary criterion for recognizing evidence of ancient burial. We added a section to the Discussion to review evidence of grave goods at other Pleistocene sites where burial is accepted.

    References

    Dirks, P. H., Berger, L. R., Roberts, E. M., Kramers, J. D., Hawks, J., Randolph-Quinney, P. S., Elliott, M., Musiba, C. M., Churchill, S. E., de Ruiter, D. J., Schmid, P., Backwell, L. R., Belyanin, G. A., Boshoff, P., Hunter, K. L., Feuerriegel, E. M., Gurtov, A., Harrison, J. du G., Hunter, R., … Tucker, S. (2015). Geological and taphonomic context for the new hominin species Homo naledi from the Dinaledi Chamber, South Africa. eLife, 4, e09561. https://doi.org/10.7554/eLife.09561

    Dirks, P. H., Roberts, E. M., Hilbert-Wolf, H., Kramers, J. D., Hawks, J., Dosseto, A., Duval, M., Elliott, M., Evans, M., Grün, R., Hellstrom, J., Herries, A. I., Joannes-Boyau, R., Makhubela, T. V., Placzek, C. J., Robbins, J., Spandler, C., Wiersma, J., Woodhead, J., & Berger, L. R. (2017). The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa. eLife, 6, e24231. https://doi.org/10.7554/eLife.24231

    Elliott, M., Makhubela, T., Brophy, J., Churchill, S., Peixotto, B., FEUERRIEGEL, E., Morris, H., Van Rooyen, D., Ramalepa, M., Tsikoane, M., Kruger, A., Spandler, C., Kramers, J., Roberts, E., Dirks, P., Hawks, J., & Berger, L. R. (2021). Expanded Explorations of the Dinaledi Subsystem,Rising Star Cave System, South Africa. PaleoAnthropology, 2021(1), 15–22. https://doi.org/10.48738/2021.iss1.68

    Fewlass, H., Zavala, E. I., Fagault, Y., Tuna, T., Bard, E., Hublin, J.-J., Hajdinjak, M., & Wilczyński, J. (2023). Chronological and genetic analysis of an Upper Palaeolithic female infant burial from Borsuka Cave, Poland. iScience, 26(12). https://doi.org/10.1016/j.isci.2023.108283

    Foecke, Kimberly K., Queffelec, Alain, & Pickering, Robyn. (n.d.). No Sedimentological Evidence for Deliberate Burial by Homo naledi – A Case Study Highlighting the Need for Best Practices in Geochemical Studies Within Archaeology and Paleoanthropology. PaleoAnthropology, 2024. https://doi.org/10.48738/202x.issx.xxx

    Goldberg, P., Aldeias, V., Dibble, H., McPherron, S., Sandgathe, D., & Turq, A. (2017). Testing the Roc de Marsal Neandertal “Burial” with Geoarchaeology. Archaeological and Anthropological Sciences, 9(6), 1005–1015. https://doi.org/10.1007/s12520-013-0163-2

    Maloney, T. R., Dilkes-Hall, I. E., Vlok, M., Oktaviana, A. A., Setiawan, P., Priyatno, A. A. D., Ririmasse, M., Geria, I. M., Effendy, M. A. R., Istiawan, B., Atmoko, F. T., Adhityatama, S., Moffat, I., Joannes-Boyau, R., Brumm, A., & Aubert, M. (2022). Surgical amputation of a limb 31,000 years ago in Borneo. Nature, 609(7927), 547–551. https://doi.org/10.1038/s41586-022-05160-8

    Martinón-Torres, M., d’Errico, F., Santos, E., Álvaro Gallo, A., Amano, N., Archer, W., Armitage, S. J., Arsuaga, J. L., Bermúdez de Castro, J. M., Blinkhorn, J., Crowther, A., Douka, K., Dubernet, S., Faulkner, P., Fernández-Colón, P., Kourampas, N., González García, J., Larreina, D., Le Bourdonnec, F.-X., … Petraglia, M. D. (2021). Earliest known human burial in Africa. Nature, 593(7857), Article 7857. https://doi.org/10.1038/s41586021-03457-8

    Martinón-Torres, M., Garate, D., Herries, A. I. R., & Petraglia, M. D. (2023). No scientific evidence that Homo naledi buried their dead and produced rock art. Journal of Human Evolution, 103464. https://doi.org/10.1016/j.jhevol.2023.103464

    Pomeroy, E., Bennett, P., Hunt, C. O., Reynolds, T., Farr, L., Frouin, M., Holman, J., Lane, R., French, C., & Barker, G. (2020a). New Neanderthal remains associated with the ‘flower burial’ at Shanidar Cave. Antiquity, 94(373), 11–26. https://doi.org/10.15184/aqy.2019.207

    Pomeroy, E., Hunt, C. O., Reynolds, T., Abdulmutalb, D., Asouti, E., Bennett, P., Bosch, M., Burke, A., Farr, L., Foley, R., French, C., Frumkin, A., Goldberg, P., Hill, E., Kabukcu, C., Lahr, M. M., Lane, R., Marean, C., Maureille, B., … Barker, G. (2020b). Issues of theory and method in the analysis of Paleolithic mortuary behavior: A view from Shanidar Cave. Evolutionary Anthropology: Issues, News, and Reviews, 29(5), 263–279. https://doi.org/10.1002/evan.21854

    Robbins, J. L., Dirks, P. H. G. M., Roberts, E. M., Kramers, J. D., Makhubela, T. V., HilbertWolf, H. L., Elliott, M., Wiersma, J. P., Placzek, C. J., Evans, M., & Berger, L. R. (2021). Providing context to the Homo naledi fossils: Constraints from flowstones on the age of sediment deposits in Rising Star Cave, South Africa. Chemical Geology, 567, 120108. https://doi.org/10.1016/j.chemgeo.2021.120108

    Wiersma, J. P., Roberts, E. M., & Dirks, P. H. G. M. (2020). Formation of mud clast breccias and the process of sedimentary autobrecciation in the hominin-bearing (Homo naledi) Rising Star Cave system, South Africa. Sedimentology, 67(2), 897–919. https://doi.org/10.1111/sed.12666

  7. Version published to 10.1101/2023.06.01.543127v2 on bioRxiv
  8. Version published to 10.7554/elife.89106.1 on eLife
  9. eLife

    eLife assessment

    The authors study the context of the skeletal remains of three individuals and associated sediment samples to conclude that the hominin species Homo naledi intentionally buried their dead. Demonstration of the earliest known instance of intentional funerary practice – with a relatively small-brained hominin engaging in a highly complex behavior that has otherwise been observed from Homo sapiens and Homo neanderthalensis – would be a landmark finding. However, the evidence for these claims is considered inadequate in the current version of the study. The four reviewers were in strong consensus that the methods, data, and analyses do not support the primary conclusions. Without full excavations, the study is missing crucial geoarchaeology (especially micromorphology) and taphonomic components, among other limitations, that make premature the conclusion that H. naledi intentionally buried their dead. The null hypothesis must be that these skeletons accumulated naturally and the research must then reject the null hypothesis and robustly exclude equifinality in order to justifiably draw the remarkable conclusions made in the present version of the paper.

  10. eLife

    Reviewer #1 (Public Review):

    The discovery of Homo naledi fossils and the rising star cave system is unquestionably important for paleoanthropology. The fossils themselves hold a wealth of information about the diversity and complexity of morphological and evolutionary change on the hominin family tree. It is a truly amazing find and important site and it is important that information about this site continues to be produced so that more can be known. It is equally important that the papers produced from the site be fully reviewed for scientific rigor. I hope to assist with this to the best of my ability.

    In its current form the paper, "Evidence for the deliberate burial of the dead by Homo naledi," does not meet the standards of our field. The paper is hard to follow. It lacks key citations, contextual background information to inform the reader about the geological and depositional structure of the caves, and concise understandable descriptions of the methods and the significance of the results.

    The main point of the paper is to describe three possible burial features. The working hypothesis is that the features are intentional burials, and the authors seek to support this hypothesis throughout rather than test it. The authors do this by noting mineralogical differences in sediment and possible bowl-shaped sedimentological distinctions where fossil bones occur. As stated above, this evidence needs to be elaborated on the in text, contextualized, and edited for clarity. In addition, throughout the paper, the authors only consider two depositional scenarios for burial and body decomposition: 1) a body was intentionally buried in a pit that was dug into the cave sediments, and then buried in sediment (without detailing in the main text what sediment was used to backfill the pit); and 2) the body was left in a natural pit and decayed in the open. A major problem with only considering these two scenarios for body decomposition is that previous reports about cave geology and sedimentology show that it is a dynamic system involving erosion, sediment slumping and drainage, and contraction of clay, which is a major component of the sediment, etc. The authors are very clear that flooding is not a viable option for the movement of skeletal elements in the cave. However, they do not mention other processes such as erosion or sediment slumping, that are known to occur and could be responsible for moving sediment and fossils in each chamber of the cave. They also do not consider carnivore involvement which has been suggested by Val (2016) and Egeland et al. (2018). Such processes could naturally transport bodies, shift them around, and sediment erosion could bury them. The articulation of some skeletal elements is a major argument for intentional burial, yet within the cave substructure, articulated bones are often commingled with disarticulated elements from the same or different individuals. This same situation exists in the features included in this paper. It does appear that some skeletal material was covered in sediment before decomposition and remains in articulation, but bodies decompose at different rates, and can decompose slowly, especially in environments that lack insects (see Simmons et al. 2010 Journal of Forensic Sciences https://doi-org.aurarialibrary.idm.oclc.org/10.1111/j.1556-4029.2009.01206.x). Wiersma et al., 2019 describe the cave system as very humid, but dry due to little standing water, mildly acidic, with an average temperature today of 18{degree sign}C and a minimum of 12{degree sign}C over the last million years. The starting null hypothesis should be that the bodies were naturally covered in sediment. Intentional burial requires extraordinary circumstances and requires multiple lines of solid evidence to support the hypothesis. In testing for natural burial processes, the rate of body decomposition should be reconstructed given the environmental parameters of the cave.

    In keeping with supporting their starting hypothesis that Homo naledi intentionally buried individuals in the cave, the authors conclude that "A parsimonious explanation for this configuration of skeletal remains is that these remains may be a palimpsest of burials that have sequentially disrupted each other. In this hypothesis, early burials were disturbed when pits were dug for subsequent burials. Other occurrences of remains outside of the Dinaledi Chamber and Hill Antechamber (Hawks et al., 2017; Brophy et al., 2021) are discussed as possible evidence of mortuary practices in SI 4.2. Instances where parts of individuals occur in remote narrow passages cannot be explained as a result of carnivore or water transport (Elliott et al., 2021; Brophy et al., 2021), making it necessary to consider that H. naledi may have placed these partial remains in these locations, possibly representing a form of funerary caching." After reviewing the evidence presented in the current manuscript, it is not clear why this is a parsimonious explanation. The authors have repeatedly described how incredibly challenging it is to get into and out of this cave system and all of its chambers. How could any species, even small bodied species, drag/pull/shove dead bodies through small crevasses, shove or drop them down a narrow shoot, continue to move through the hill antechamber to the Dineledi chamber and bury bodies? It is not impossible but given the previously published descriptions of the dynamic process of sedimentation movement in the cave it is certainly not a parsimonious explanation. To support this will take many more lines of evidence than presented here such as micromorphological analysis of the overall cave system and each feature (discussed in the supplementary information but briefly), full detailed reconstruction of sediment, water, fossil, and debris movement throughout the cave system coupled with reconstructions of body decomposition rates. Scientifically precise computer-generated reconstructions of all of this are possible working with specialists affiliated with National Geographic. An analysis also needs to start by testing a null hypothesis, not deciding on the conclusion and setting out to "prove" it.

  11. eLife

    Reviewer #2 (Public Review):

    In this study (Berger et al.), geological and fossil data from the Rising Star Cave System in South Africa are presented to provide evidence for intentional burials of Homo naledi individuals. The authors focus on describing and interpreting what they refer to as "delimited burial features." These features include two located on the floor of the Dinaledi Chamber (referred to as 'Dinaledi Features' 1 and 2) and one from the floor of the Hill Antechamber.

    'Dinaledi Feature 1' consists of a collection of 108 skeletal elements recovered from sub-unit 3b deposits. These remains are believed to primarily represent the remains of a single adult individual, along with at least one additional juvenile individual. Although additional anatomical elements associated with 'Dinaledi Feature 1' are mentioned, they are not described as they remain unexcavated. The study states that the spatial arrangement of the skeletal remains is indicative of the primary burial of a fleshed body. On the other hand, 'Dinaledi Feature 2' is not extensively discussed, and its complete extent was not thoroughly investigated.

    Regarding the Hill Antechamber feature, it was divided into three separate plaster jackets for removal from the excavation. Through micro-CT and medical CT scans of these plaster jackets, a total of 90 skeletal elements and 51 dental elements were identified. From these data, three individuals were identified, along with a fourth individual described as significantly younger. Individuals 1 and 2 are classified as juveniles.

    I feel that there is a significant amount of missing information in the study presented here, which fails to convince me that the human remains described represent primary burials, i.e. singular events where the bodies are placed in their final resting places. Insufficient evidence is provided to differentiate between natural processes and intentional funerary practices. In my opinion, the study should include a section that distinguishes between taphonomic changes and deliberate human modifications of the remains and their context, as well as reconstruct the sequence and timeline of events surrounding death and deposition. A deliberate burial involves a complex series of changes, including decomposition of soft tissues, disruption of articulations between bones, and the sequence of skeletonization. While the geological information is detailed, the archaeothanatological reasoning (see below) is largely absent and, when presented, it lacks clarity and unambiguousness.

    My main concern is that the study does not apply or cite the basic principles of archaeothanatology, which combines taphonomy, anatomy, and knowledge of human decomposition to interpret the arrangement of human bones within the Dinaledi Chamber and the Hill Antechamber. Archaeothanatology has been developed since the 1970s (see Duday et al., 1990; Boulestin and Duday, 2005; Duday and Guillon, 2006) and has been widely used by archaeologists and osteologists to reconstruct various aspects such as the original treatment of the body, associated mortuary practices, the sequence of body decomposition, and the factors influencing changes in the skeleton within the burial.

    Specifically, the study lacks a description of the relative sequence of joint disarticulation during decomposition and the spatial displacement of bones. A detailed assessment of the anatomical relationships of bones, both articulated and disarticulated, as well as the direction and extent of bone displacement, is missing. For instance, while it is mentioned that "many elements are in articulation or sequential anatomical position," a comprehensive list of these articulated elements and their classification (as labile or not) is not provided.

    Furthermore, the patterns described are not illustrated in sufficient detail. If Homo naledi was deliberately buried, it would be crucial to present illustrations depicting the individuals in their burial positions, as well as the representation and proportions of the larger and smaller anatomical elements for each individual. While Figure 2B provides an overall view of 'Dinaledi Feature 1,' it is challenging to determine the relationships of bones, whether articulated or disarticulated, in Figures 2C or 2D. Such information is essential to determine whether the bones are in a primary or secondary position, differentiate between collective and multiple burials, ascertain the body's stage of decomposition at the time of burial, identify postmortem and post-depositional manipulation of the body and grave (e.g., intentional removal of bodies/body parts), and establish whether burial occurred immediately after death or was delayed.

    Moreover, the study does not address bone displacements within secondary voids created after the decomposition of soft tissues, nor does it provide assessments of the position of bones within or outside of the original body volume. Factors such as variations in soft tissue volume between individuals of different sizes/corpulence, and the progressive filling (i.e., sediment continually fills newly formed voids) or delayed filling (causing the 'flattening' of the ribcage and 'hyper-flexed' burials, for instance) of secondary open spaces with sediment over time should also be discussed.

    In conclusion, while I acknowledge the importance of investigating potential deliberate burials in Homo naledi, I do not think that in its present form, the evidence presented in this study is as robust as it should be.

  12. eLife

    Reviewer #3 (Public Review):

    This paper provides new information on the Dinaledi Chamber at the Rising Star Cave System. In short, a previously excavated area was expanded and resulted in the discovery of a cluster of bones appearing to be of one individual, a second similar cluster, and a third cluster with articulated elements (though with several individuals). Two of these clusters are argued to be intentionally buried individuals (the third one has not been investigated) and thus Homo naledi not only placed conspecifics in deep and hard to reach parts of caves but also buried them (apparently in shallow graves). This would be the oldest evidence of intentional burial. The main issue with the paper is that the purported burials were not fully excavated. Two are still in the ground, and one was removed in blocks but left unexcavated. As burials are mostly about sediments, it means the authors are lacking important lines of evidence. Instead, they bring other lines of argument as outlined below. While their preferred scenario is possible, there are important issues with the evidence as presented and they are severely hampered by the lack of detailed archaeological and geoarchaeological information both from the specific skeletal contexts and more generally from the chamber (because in fact the amount of excavation conducted here is still quite limited in scope). I also found that while the presentations of the various specialists in the team was quite good, the integration of these contributions into the main text was not. In particular, the geology of the cave system and the chamber need (especially what is known of the depositional and post-depositional processes) need to be better integrated into the presentation of the archaeology and the interpretation of the finds.

    Often times the presence of articulated or mostly articulated skeletons is used to argue for intentional burial. This argument, however, is based on the premise that if not buried, these skeletons would have otherwise become disarticulated. Normally disarticulation would happen as a result of subsequent use of the site by hominins (e.g. purported burials in Neandertal cave sites) or by carnivores scavenging the body. Indeed this latter point is why bodies are buried so deeply in many Western societies (i.e. beyond the reach and smell of carnivores). Bodies can also be disarticulated by natural processes of deposit and erosion.

    However, here in the case of the Dinaledi Chamber, we apparently don't have any of these other processes. The chamber was not used by carnivores and it was not a living area where H. naledi would have frequently returned and cleared out the space. As for depositional processes, it is more complex, but it is clear from Wiersma et al. that there is a steady, constant movement of these sediments towards drains. They also think that this process can account for the mix of articulated and non-articulated elements in the cave. Importantly, that same paper makes the argument that the formation of these sediments is not the result of water movement and that the cave has been dry since the formation of this deposit. So bodies lying on the surface and slowly covered by the formation of the deposit and slowly moving towards the drains could perhaps account for the pattern observed, meaning burial is not needed to account for articulations (note that more information on fabrics would be good in this context - orientation analysis of surface finds or of excavated finds is either completely lacking or minimal - see figure 13b and c report orientations on 79 bones of unknown context that appear to show perhaps elevated plunge angles and some slightly patterning in bearing but there is no associated statistics or text explaining the significance).

    So, unless the team can provide some process that would have otherwise disarticulated these skeletons after the bodies arrived here and decomposed, their articulated state is not evidence of burial (no more than finding an articulated or mostly articulated bear skeleton deep in a European cave would suggest that it was buried).

    As for the elemental analysis, what I understood from the paper is that the sediment associated with bones is different from the sediment not associated with bones. It is therefore unsurprising that the sediment associated with the reported skeletons clusters with sediments with bones. The linking argument for why this makes this sediment pit fill is unclear to me. Perhaps it is there, but as written I didn't follow it.

    What the elemental analysis could suggest, I think, is that there has not been substantial reworking of the sediments (as opposed to the creep suggested by Wiersma et al.) since the bones leached these minerals into the sediment. What I don't know, and what is not reported, is how long after deposition we can expect the soil chemistry to change. If this elemental analysis were extended in a systematic way across the chamber (both vertically and horizontally) after more extensive excavations, I could see it perhaps being useful for better understanding the site formation processes and depositional context. As it is now, I did not see the argument in support of a burial pit.

    The other line of evidence here is that some bones are sediment supported. The argument here is that when a body decomposes, bones that were previously held in place by soft tissues will be free to move and will shift their position. How the bones shift will differ depending on whether the body is surrounded by matrix (as they argue here in an excavated burial pit) or whether it is in the open (say, for instance, in a coffin) (and there are other possibilities as well - for instance wrapped in a shroud). Experiments have also shown the order in which the tendons, for instance, decompose and therefore which bones are likely to be free to move first or last.

    I will note that this literature is poorly cited. I think the only two papers cited for how bodies decompose are Roksandic 2002 and Mickleburgh and Wescott 2019. The former is a review paper that summarizes a great many contexts that are clearly not appropriate here, and it generally makes the point that it is difficult to sort out, and it notes that progressively filled is an additional alternative to not buried/buried. The other looks at experimental data of bodies decomposing without being buried. In the paper here, this citation is used to argue that the body must have been buried. I don't see the linking argument at all. And the cited paper is mostly about how complicated it is to figure this all out and how many variables are still unaccounted for (including the initial positioning of the body and the consumption of the body by insects - something that is attested to at Naledi - plus snails - see not just Val but also Wiersma et al. and I think the initial Dirk et al. paper).

    So the team here instead simply speaks of how the body decomposes in burials as if it is known. For the Feature 1 skeleton, the authors note that the ribs are "apparently" sediment supported and that a portion of the partial cranium is vertical or subvertical and sediment supported. For both of these, the figures show it very poorly. We really have to take their word for it. Second, I would have liked to have seen some reference and comparison to the literature for how the ribs should be in sediment burial cases. For the cranium, seems like a broken cranium resting on a surface will have vertical aspects regardless of sediment support. To the contrary, the orientation of the cranium will change depending on whether there is sediment holding it in place or not. But that argument is not made here. It is very hard from the figures to have a detailed idea of how these skeletons are oriented in the sediments, to know which elements are in articulation, which are missing, etc.

    In the case of the Hill Antechamber Feature, an additional argument is made about the orientation of the finds in relation to the natural stratigraphy in this location. The team argues that the skeleton is lying more horizontally than the sediments and that in fact the foot is lying against the slope. First, there is no documentation of the slope of the layers here (e.g. a stratigraphic profile with the layers marked or a fabric analysis). There is a photo in the SI that says it shows sloping, but it needs some work. Second, this skeleton was removed in three blocks and then scanned. So the position of the skeleton is being worked out separate from its context. This is doable, but I would have liked to have seen some mention of how the blocks were georeferenced in the field and then subsequently in the lab and of how the items inside the block (i.e. the data coming from the CT scanner) were then georeferenced. I can think of ways I would try to do this, but without some discussion of this critical issue, the argument presented in Figure 10c is difficult to evaluate. Further, even if we accept this work, it is hard for me to see how the alignment of the foot is 15 degrees opposite the slope (the figure in the SI is better). It is also hard to understand the argument that the sediment separating the lower limb from the torso means burial. The team gives the explanation that if the body was in an open pit it would have been flat with no separation. Maybe. I mean I guess if the pit was flat. But there is no evidence here of a pit (at all). And what if the body was stuffed down the chute and was resting on a slope and covered with additional sediments from the chute (or additional bodies) as it decomposed? It seems that this should be the starting point here rather than imagining a pit.

    One of the key pieces of evidence for demonstrating deliberate burial is the recognition of a pit. Pits can be identified because of the rupture they create in the stratigraphy when older sediments are brought to the surface, mixed, and then refilled into the pit with a different color, texture, compaction, etc. In some homogenous sediments a pit can be hard to detect and in some instances post-depositional processes (e.g. burrowing) can blur the distinction between the pit and the surrounding sediments. But the starting point of any discussion of deliberate burial has to be the demonstration of a pit. And I don't see it here. It might just be that the figures need to be improved. But I am skeptical because the team has taken the view that these finds can't be excavated. While I appreciate the scanning work done on the Antechamber find, it is not the same as excavating. Same comment for Features 1 and 2.

    In short, my view is that they have an extremely interesting dataset. That H. naledi buried their dead here can't be excluded based on the data, but neither is it supported here. My view is that this paper is premature and that more excavation and the use of geoarchaeological techniques (especially micromorphology) are required to sort this out (or go a long way towards sorting it out).

  13. eLife

    Reviewer #4 (Public Review):

    Berger et al. 2023a argues that Homo naledi intentionally buried their dead within the Rising Star cave system by digging pits and covering the bodies with infilled sediment. The authors identified two burials: Dinaledi Feature 1 from the Dinaledi Chamber, and the Hill Antechamber Feature from the Hill Antechamber. The evolutionary and behavioral implications for such behavior are highly significant and would be the first instance of a relatively small-brained hominin engaging is complex behavior that is often found in association with Homo sapiens and Homo neanderthalensis. Thus, the scientific rigor to validate these findings should be of the highest quality, and thus, provide clear documentation of intentional burial. In an attempt to meet these standards, the authors stated a series of tests that would support their hypothesis of intentional burials in the Rising Star Cave system:

    "The key observations are (1) the difference in sediment composition within the feature compared to surrounding sediment; (2) the disruption of stratigraphy; (3) the anatomical coherence of the skeletal remains; (4) the matrix-supported position of some skeletal elements; and (5) the compatibility of non-articulated material with decomposition and subsequent collapse." (page 5)

    To find support for the first (1) test, the authors collected sediment samples from various locations within the Rising Star Cave system, including sediment from within and outside Dinaledi Feature 1. However:

    • The authors did not select sediment samples from within the Hill Antechamber Feature, so this test was only used to assess Dinaledi Feature 1.

    • The sediment samples were analyzed using x-ray diffraction (XRD) and x-ray fluorescence (XRF) to test the mineralogy and chemistry of the samples from within and outside the feature. The XRF results were presented as weighted percentages (not intensities) with no control source reported. The weighted percentages were analyzed using a principal components analysis (PCA) while the particle-size distribution was analyzed using GRADISTAT statistics package and the Folk and Ward Method to summarize "mean grain size, sorting, skewness and kurtosis in addition to the percentages of clay, silt and sand in each sample." (page 28).

    • The PCA results were reported solely as a biplot without showing the PC scores projected into the loading space, which is unusual and does not present the data accurately. Instead, the authors present the scores of a single component (PC2, figure 3) because the authors interpreted this component as "distinctly delineates fossil-bearing sediments from sterile sediments based on the positive loadings of P and S" (Page 6). However, the supplementary table that reports XRF bulk chemistry results as a weighted percentage of minerals within each sample (SI Table 1) shows mostly an absence of data for both Na and S. Since Na is at the lower end of detection limits for the method, and S seems to just be absent from the list, the intentions of the authors for showing the inclusion of these elements in their PCA results is unclear. Given that this is the author's primary method for demonstrating a burial, this issue is particularly concerning and requires additional attention.

    • Regardless of the missing data, this reviewer attempted to replicate the XRF PCA results using the data provided in SI Table 1 and was unsuccessful. The samples that were collected from within the feature (SB) cluster with samples collected from sterile sediments and other locations around the cave system. Thus, these results are not replicable as currently reported.

    • Visual comparisons of sediment grain size, shape, and composition were qualitatively summarized. Grain size was plotted as a line graph and is buried as supplemental Figure S13 showing sample by color and area, but these results do not distinguish samples from WITHIN the burial compared to OUTSIDE the burial as the authors state in the methods as a primary goal.

    To test the second (2) aim, the "stratigraphy" was primarily described in text.

    • For Dinaledi Feature 1, the authors state that the layer around Feature 1 "is continuous in the profile immediately to the east of the feature; it is disrupted in the sediment profile at the southern extent of the feature (fig. 3b)." Upon examination of figure 3b, the image shows an incredibly small depiction of the south (?) profile view with an extremely large black box overlaying a large portion of the photograph containing a small 5 cm scale. Visually, there is no difference in the profile that would suggest a disruption in the form of a pit. The LORM (orange-red mud layer) does seem to become fragmentary, but no micromorphological analysis was conducted on this section to provide an evaluation of stratigraphic composition. Also, by only excavating a portion of the feature, the authors were unable to adequately demonstrate the full extent of this feature.

    • The authors attempt to describe "a bowl-shaped concave layer of clasts and sediment-free voids make up the bottom of the feature" (page 13) and refer to figures and supplementary information that do not depict any stratigraphic profile. Moreover, the authors state that "the leg, foot, and adjacent [skeletal?] material cut across stratigraphy" indicating that the skeleton is orientated on a flat plane against the surrounding stratigraphy that is "30{degree sign} slope of floor and underlying strata" (page 51, fig. 10c captions). There is no mention of infilled sediment from a pit and how this relates to the skeleton or the slope of the floor. It is therefore extremely unclear what the authors are meaning to describe without any visual or micromorphological supplementation to demonstrate a "bowl-shaped concave layer".

    The third (3) test was to evaluate the anatomical coherence of the skeletal remains using macro- and micro-CT (computed tomography) of the Hill Antechamber Feature that was removed during excavation. To visually assess the anatomy of the Dinaledi Feature 1 burial, the authors describe the spatial relationship of skeletal elements as they were being excavated but halted partway through the excavation.

    • The authors do not provide any documentation (piece-plotting, 3D rendering of stages of excavation, etc.) of the elements that were removed from the Dinaledi Feature. Figure 4 and SI Fig. S22 show the spatial relationship between identifiable skeletal elements that remain in the Feature. However, in Fig. 4, it is unclear why the authors chose to plot 2023-2014 excavated material along with material reported here, and it's even more difficult to understand the anatomical positioning of the elements given their color and point size choices. Although, the authors do provide a 3D rendering of the unexcavated remains showing some skeletal cohesion, apart from the mandible and teeth being re-located near the pelvis (Fig. 9). That said, it is very difficult to visually confirm the elements from this model or understand the original placement of the skeleton.

    • 3D renderings of the Hill Antechamber feature skeletal material is clearly shown in SI Fig. S26. Contrary to what the authors state in text, there is a rather wide dispersal and rearrangement of elements for a "burial" that is theoretically protected from scavengers and other agents that would aid in dispersing bone from the surface. The authors do not offer any alternatives to explain disturbance, such as human activity, which clearly took place.

    • Moreover, there does not appear to be any intentional arrangement of limbs that may suggest symbolic orientation of the dead (another line of evidence often used to support intentional burial but omitted by the authors). Thus, skeletal cohesion is not enough evidence to support the hypothesis of an intentional burial.

    The fourth (4) test was attempted by evaluating whether some elements were vertically aligned from 3D reconstructed models of Hill Antechamber Feature and a photogrammetric model of the Dinaledi Feature 1. The authors state that "the spatial arrangement of the skeletal remains is consistent with primary burial of the fleshed body" (page 8 in reference to Dinaledi Feature 1) without providing any evidence, qualitative or quantitative, that this is the case for either burial.

    Since this reviewer was unable to understand the fifth (5) test as it was written by the authors, I am unable to comment on the evidence to support this test and will default to the other reviewers for evaluation of this claim.

    In addition to a lack of evidence to support the claims of intentional burial, this paper was also written extremely poorly. For example, the authors often overused 'persuasive communication devices' (see eLife article, https://elifesciences.org/articles/88654) to mislead readers:

    "During this excavation, we recognized that the developing evidence was suggestive of a burial, due to the spatial configuration of the feature and the evidence that the excavated material seemed to come from a single body." (page 5)

    As an opening statement to introduce Dinaledi Feature 1, the authors state the interpretation and working hypothesis as fact before the authors present any evidence. This is known as "HARKing" and "gives the impression that a hypothesis was formulated before data were collected" (Corneille et al. 2023). This type of writing is pervasive throughout the manuscript and requires extensive editing. I recommend that the authors review the article provided by eLife (https://elifesciences.org/articles/88654) and carefully review the manuscript. Moreover, as this text demonstrates, the authors’ word choice is indicative of storytelling for a popular news article instead of a scientific paper. I highly suggest that the authors review the manuscript carefully and present the data prior to giving conclusions in a clear and concise manner.

    Moreover, the writing structure is inconsistent. Information that should be included in results is included in the methods, text in the results should be in discussions, and so forth. This inconsistency is pervasive throughout the entire manuscript, making it incredibly difficult to adequately understand what the authors had done and how the results were interpreted.

    Finally, the "artifact" that was described and visualized using CT models is just that - a digitally colored model. The object in question has not been analyzed. Until this object is removed from the dirt and physically analyzed, this information needs to be removed from the manuscript as there is nothing to report before the object is physically examined.

    Overall, there is not enough evidence to support the claim that Homo naledi intentionally buried their dead inside the Rising Star Cave system. Unfortunately, the manuscript in its current condition is deemed incomplete and inadequate, and should not be viewed as finalized scholarship.

  14. eLife

    Author Response:

    We would like to thank the eLife reviewers for the considerable time and effort they have invested to review these manuscripts. We have also benefited from a previous round of review of the manuscript describing the proposed burial features, which underwent two rounds of revisions in a high-impact journal over a period of approximately 8 months during 2022 and early 2023. Both sets of reviews have reflected mixed responses to the evidence we have presented, with one reviewer recommending acceptance with minor editorial revisions, two recommending acceptance with minor revisions and the fourth recommending rejection based upon similar arguments to those reflected by some of the reviewers in this current round of reviews in eLife. Ultimately the managing editor of this first journal took the decision that the review process could not be completed in a timely manner and rejected the manuscript although the submission here reflected our consideration of these reviewers suggestions.

    We have chosen in this initial response to the eLife reviews to include some references to the previous anonymous reviews in order to illustrate differences of opinion and differences in revision suggestions within the review process. Our goal is to offer maximal insight into our decision-making process and to acknowledge the considerable time and effort put into the assessment of these manuscripts by reviewers (for eLife and in the case of the earlier review process). We hope that this approach will assist the readers, and reviewers, of our manuscripts in understanding why we are proceeding with certain decisions during the revision process.

    This is a new process for us and the reviewers, and one way in which it significantly differs from more traditional review is that both the reviews and our reply will be public well in advance of our revisions to the manuscript. Indeed, considering the scope of the reviews, some of those revisions may take considerable time, although many can be accomplished fairly easily. Thus, we are not in a position to say that we have solved every issue raised by the reviewers. Instead, we will examine what appear to be the key critical issues raised regarding the data and the analyses and how we propose to address these as we revise the papers. We will also address several philosophical and ethical issues raised by the reviews and our proposal for dealing with these. More specific editorial and citational recommendations will be dealt with on a case-by-case basis, and we do not address these point-by-point in this reply. Please note, this response to the reviewers is not the revision of the manuscript and is only the initial opinion of the corresponding authors with some guidance from the larger group of authors of all three papers. Our final submitted revision will reflect the input of all authors included on those submissions.

    We took the decision to submit three separate papers consciously. The two different categories of evidence, burials and engravings, involve different kinds of analysis and different (although overlapping) teams of researchers, and we recognized that each deserved their own presentation and assessment. Meanwhile, together they inform the context of H. naledi in a way that requires some synthetic discussion, in which both kinds of evidence are relevant, leading to a third paper. But the mutual relevance of these different kinds of evidence and their review by a common set of reviewers naturally raises cross-cutting issues, and the reviewers have cross-referenced the three articles. This has sometimes led to suggestions about one manuscript based on the contents of another. Considering the situation, we accepted the recommendation that it would be clearer to consider all three articles in a single reply. Thus, while each of the three papers will proceed separately during the revision process, it will be necessary to highlight across all three papers occasionally in our responses.

    Scientific Issues:

    In reading the reviews, we feel there are 9 critical points/assertions raised by one or more of the reviewers that present a problem for, or challenge to, our hypothesis that the observed evidence (bone accumulations and engravings) described in the Dinaledi subsystem are of intentional naledigenic origin. These are:

    1. The evidence presented does not demonstrate a clear interruption of the floor sediments, thus failing to demonstrate excavated holes.

    2. The sediments infilling the holes where the skeletal remains are found have not been demonstrated to originate from the disruption of the floor sediments and thus could be part of a natural geological process (e.g. water movement, slumping) or carnivore accumulations.

    3. Previous geological interpretations by our research group have given alternative geological explanations for formation of the bony accumulations that contradict the present evidence presented here and result in alternative origins hypotheses.

    4. Burial cannot be effectively assessed without complete excavation of the features and site.

    5. The skeletal remains as presented do not conform clearly to typical body arrangement/positions associated with human (Homo sapiens) burials.

    6. There is no evidence of grave goods or lithic scatters that are typically associated with human burials.

    7. Humans may have been involved with the creation of either the Homo naledi bone accumulations, the engravings, or both.

    8. Without a date of the engravings, the null hypothesis should be the engravings were created by Homo sapiens.

    9. The null hypothesis for explanation of the skeletal remains in this situation should be “natural accumulation”.

    Our analysis of the Dinaledi Feature 1 leads us to accept that the laminated orange-red mudstone (LORM) sedimentary layer is interrupted, indicating a non-natural intervention, and that the hole created by the interruption was then filled by both a fleshed body (and perhaps parts of other bodies) which were then covered by sediment that originated from the hole that was dug. We recognize that the four eLife reviewers are not convinced that our presentation is sufficient to establish this. Interestingly, this was not the universal opinion of earlier reviewers of the initial manuscript several of whom felt we had adequately supported this hypothesis. The lack of clarity in this current version of the burial manuscript is our responsibility. In the upcoming revision of this paper to be submitted, we will take the reviewers’ critiques to heart and add additional figures that illustrate better the disruption of the LORM and clarify the sedimentological data showing the material covering the skeletal remains in the hole are the disrupted sediments excavated from the same hole. We are proposing to isolate this most critical evidence for burial into a separate section in the revised submission based on the reviewers’ comments. The fact that the LORM layer is disrupted, a fleshed body was placed in the hole created by this disruption, and the body (and perhaps parts of other bodies) was/were then covered by the same sediments from the hole is the central feature of our hypothesis that the bone accumulations observed reflect a burial and not a natural process.

    The possibility of fluvial transport or involvement in the subsystem is a topic that we have addressed extensively in past work, and it is clear from these reviews that we must enhance our current manuscript to discuss this issue at greater length. Our previous work (Dirks et al. 2015; Dirks et al. 2017) emphasized that fluvial transport of whole bodies into the subsystem was precluded by several lines of sedimentological evidence. We excavated a rich accumulation of skeletal remains, including articulated limbs and other elements in subvertical orientations inconsistent with slow sedimentary infill, which were difficult to explain without positing either a large and dense pile of bodies and/or sediment movement. We encountered fractured chunks of laminated orange-red mudstone (LORM) in random orientations within our excavation area, within and among skeletal remains, which directly refuted that the remains were inundated with water at the time of burial, and this limited the possibility of fluvial transport. Water flow sufficient to displace bodies or complete skeletal evidence would also transport large and course sediment, which is absent from the subsystem, and would sort the commingled skeletal material that we found by size, which we do not observe. But our excavation only covered less than a square meter at very limited depth, and this was the limit to our knowledge of subsurface sediment. We thus were left with uncertainty that led us to suggest the possibility of sediment slumping or movement into subsurface drains, although these were not observed near our excavation. Our current work expands our knowledge of the subsurface and presents an alternative explanation for the disposition of skeletal remains from our earlier excavation. But we acknowledge that this new explanation is vulnerable to our own previous published proposals, and we must do a better job of explaining how the new information addresses our previous suggestions. By not clearly creating a section where we explained how these previous hypotheses were now nullified by new evidence, we clearly confused the reviewers with our own previous work. We will revise the manuscript by enhancing the review of the significant geological evidence demonstrating that there is no significant fluvial action in the system and making it clear how the burial hypothesis provides a clearer explanation for the situation of skeletal remains from our previous excavation work.

    One of the central issues raised by reviewers has been a perceived need to excavate these features completely, totally exhuming all skeletal remains from them. Reviewers have written that it is necessary to identify every skeletal element that is present and account for any missing elements. On this point, we have both ethical and scientific differences from these reviewers. We express our ethical concerns first. Many of the best-preserved possible burials ever discovered by archaeologists were subjected to total excavation and exhumation. Cases like La Chapelle-aux-Saints, La Ferrassie, and Skhūl were fully excavated at a time when data recording and excavation methods did not include the range of spatial and geomorphological approaches that later became routine. The judgment of early investigators that these situations were intentional burials was challenged by later workers, and the kind of information that might enable better tests had been irrevocably lost (Gargett 1999; Dibble et al. 2015; Rendu et al. 2014).

    Later, improved excavation standards have not sufficed to remove uncertainty or debate about possible burials. For example, it was long presumed that well-preserved remains of young children were by themselves diagnostic of intentional burial, such as those from Dederiyeh, Border Cave, or Roc de Marsal. Such cases were also fully excavated, with adequate documentation of the positioning of skeletal remains and their surrounding stratigraphic situation, but such cases were later challenged on several bases and the complete exhumation of material has confused or precluded testing of new hypotheses (e.g. Gargett 1999). The case of Roc de Marsal is one in which data from the initial excavation combined with data from the initial excavation combined with re-excavation and geoarchaeological analysis led to a naturalistic interpretation of the skeletal material (Sandgathe et al. 2011; Goldberg et al. 2017). But even in this case, the researchers erred in their interpretation of the skeleton’s situation due to a lack of identification of parts of the infant’s skeleton (Gómez-Olivencia and García-Martinez 2019). That is to say, it is not only the burial hypothesis but other hypotheses that suffer from complete excavation. Researchers concerned with preserving all possible information have sometimes taken extraordinary measures to remove and study possible burials at high-resolution in the laboratory. Such was the case of the Shanidar IV burial removed from the site and transported in plaster jacket by Solecki, which led to the disruption and loss of internal stratigraphic information (Pomeroy et al. 2020). Arguably, the current state of the art is full excavation with partial preparation, such as that undertaken at Panga ya Saidi (Martinón-Torres et al. 2021). But again, any future attempt to reinterpret or test the hypothesis of burial must rely on the adequacy of documentation as the original context has been removed.

    In our decision to leave material in place as much as possible, we are expanding upon standard practice to leave witness sections and unexcavated areas for future research. The situation is novel, representing possible burials by a nonhuman species, and that makes it doubly important in our opinion to be conservative in not fully exhuming the skeletal material from its context. We anticipate that many other researchers, including future investigators, will suggest additional methods to further test the hypothesis of burial, something that would be impossible if we had excavated the features in their entirety prior to publishing a description of our work. We believe strongly that our ethical responsibility is to publish the work and the most likely interpretation while leaving as much evidence in place as possible to enable further testing and replication. We welcome the suggestions of additional methods/analyses to test the H. naledi burial hypothesis.

    This being said, we also observe that total exhumation would not resolve the concerns raised by the reviewers. The recommendation of total exhumation is in pursuit of a full account of all skeletal material present and its preservation and spatial situation, in order to demonstrate that they conform to body positions comparable to human burials. As has been highlighted in forensic casework, the excavation of an inhumation feature does not necessarily provide an accurate spatial or anatomical manifest of the stratigraphical relationships between the body, encapsulating matrix, and any cut present due to preservational, taphonomic and operational factors (Dirkmaat and Cabo, 2016; Hunter, 2014). In particular, in cases where skeletal elements are highly fragmented, friable, or degraded (such as through bioerosion) then complete excavation—even under controlled laboratory conditions—may destroy bone and severely limit skeletal identification (Henderson, 1997; Hochrein, 2002; Owsley and Compton, 1997), particularly in elements where the ratio of trabecular to cortical bone is high (Darwent and Lyman, 2002; Lyman, 1994). As such, non-invasive methods of 3D and 4D modelling (preservation in situ) are often considered preferable to complete necropsy or excavation (preservation by record) where appropriate (Bolliger and Thali, 2009; Dell’Unto and Landeschi, 2022; Randolph-Quinney et al., 2018; Silver, 2016).

    The test of burial is not primarily positional, but taphonomic and geological. The position and number of bones can elaborate on process-driven questions of decay and destruction in the burial environment, or post-mortem modification, but are not singularly indicative of whether the remains were intentionally buried – the post-mortem narrative of all the processes affecting the cadaveric island is required (Knüsel and Robb, 2016). In previous cases, researchers have disputed or accepted the hypothesis of intentional hominin burial based upon assumptions about how modern humans or Neandertals would have positioned bodies, with the idea that some positions reflect ritual intent while others do not. But applying such assumptions is unjustifiable, particularly for a species like H. naledi, whose culture may have differed fundamentally from our own. Our work acknowledges that the present evidence does not enable a full reconstruction of the burial positions, but it does show that fleshed remains were encased in sediment prior to decomposition of soft tissue, and that subsequent spatial changes can be most parsimoniously explained by natural decomposition within sedimentary matrix contained within a burial feature (after Green, 2022; Mickleburgh and Wescott, 2018; Mickleburgh et al., 2022). If the argument is that extraordinary claims require extraordinary evidence, we feel that the evidence documents excavation and interment (and will do so more clearly in the revision) and the fact of the remains do not match a “typical” human burial in body positioning is not in itself evidence that these are not H. naledi burials.

    We feel that the reviewers (in keeping with many palaeoanthropologists) have a clear idea of what they “think” a burial should look like in an idealised sense, but this platonic ideal of burial form is not matched by the extensive literature in archaeothanatology, funerary archaeology and forensic science which indicates enormous variability in the activity, morphology and post-mortem system experienced by the human body in cases of interment and body disposal (e.g. Aspöck, 2008; Boulestin and Duday, 2005 and 2006; Connelly et al., 2005; Channing and Randolph-Quinney, 2006; Cherryson, 2008; Donnelly et al., 1995; Finley, 2000; Hunter, 2014; Parker Pearson, 1999; Randolph-Quinney, 2013). Decades of experience in the identification, recovery and interpretation of clandestine, deviant, and non-formal burials indicates the platonic ideal is rare, and in many contexts, the exception (Cherryson, 2008; Parker Pearson, 1999). This variability is particularly relevant to morphological traits in burial context, such as the informal nature of the grave cut in plan and section, shallow burial depth, and initial disposition of body (placement) during the early post-mortem period. These might run counter to the expectations of reviewers or others referencing the fossil hominin record, but are well accepted within the communities of researchers investigating Holocene archaeological sites and forensic contexts.

    It is encouraging to see reviewers beginning to incorporate the extensive (often experimentally derived) literature from archaeothanatology and forensic taphonomy in their deliberations, and we will be taking these comments on board going forward. In particular, we acknowledge reviewers’ comments and the need to construct a more detailed post-mortem narrative, accounting for joint disarticulation (labile versus persistent joints etc), displacement, and final disposition of elements within the burial space. As such we will incorporate the hierarchy of decomposition (rank order disarticulation), associations between regions of anatomical association, areas of disassociation, and the voids produced during decomposition (after Mickleburgh and Wescott, 2018; Mickleburgh et al., 2022) into our narrative. In doing so we acknowledge the tensions between the inductive archaeolothanatological narrative-driven approach (e.g. Duday, 2005 & 2009) versus robust decomposition data derived from human forensic taphonomic experimentation recently articulated by Schotsmans and colleagues (2022) - noting that we will highlight comparative data based on forensic experimental casework and actualistic modelling over inductive intuitive approaches which come with significant evidential shortcomings (Bristow et al. 2011).

    Finally, from a taphonomic perspective it is worth pointing out to reviewers that we have already addressed the issue of lack of taphonomic evidence for carnivore involvement in the formation of the Dinaledi assemblage (Dirks, et al., 2016). Absence of any carnivore-induced bone surface modifications, patterns of skeletal part representation, and a total absence of any carnivore remains found within the Dinaledi chamber (following Kuhn and colleagues, 2010) lead us to reject carnivores as possible vectors of body accumulation within the Dinaledi Chamber and Hill Antechamber.

    Reviewers suggest that without a date derived from geochronological methods, the engravings cannot be associated with H. naledi, and that it is possible (or probable) that the engravings were done in the recent past by H. sapiens. This suggestion neglects the context of the site. We have previously documented the structure and extremely limited accessibility of the Dinaledi subsystem. This subsystem was not recorded on maps of the documented Rising Star Cave system prior to our work and its discovery by our teams. Furthermore, there is no evidence of prehistoric human activity in the areas of the cave related to possible subterranean entrances There is no evidence that humans in the past typically ventured into such extreme spaces like those of Rising Star. It is clear from the presence of the remains of many individuals that H. naledi ventured into these spaces again and again. It is likely that H. naledi moved through these spaces more easily than humans do based on their physique. We show that the engravings overlay each other suggesting multiple engraving events. These engravings took time and effort and the only evidence for use of the Dinaledi subsystem by any hominin is by H. naledi. The context leads to the null hypothesis that H. naledi made the marks. In our revision, we will elaborate on this argument to clarify the evidence for our stance on this hypothesis. Several reviewers took issue with the title of the engraving paper as we did not insert a qualifier in front of the suggested date range for the engravings. We deliberately left out qualifying language so that the title took the form of a testable hypothesis rather than a weak assertation. Should future work find the engravings were not produced within this time range, then we will restate this hypothesis.

    Finally, with regards to the engravings we have chosen to report them because they exist. Not reporting the presence of engraved marks on the walls of a cave above hypothesized burials would be tantamount to leaving relevant evidence out of the description of an archeological context. We recognize and state in our manuscript that these markings require substantial further study, including attempts at geochronological dating. But the current evidence is clearly relevant to the archaeological context of the subsystem. We take a similar stance with reporting the presence of the tool shaped artefact near the hand of the H. naledi skeleton in the Hill Antechamber. It is evident that this object requires further study, as we stated in our manuscript, but again omitting it from our study would be leaving out relevant evidence.

    Some have suggested that the null hypothesis should be that all of these observed circumstances are of natural origin. Our team took this approach in our early investigation of the Dinaledi subsystem (Dirks et al. 2015). We adopted the null hypothesis that the geological processes involved in the accumulation of H. naledi skeletal remains were “natural” (e.g., non-naledigenic involvement), and we were able to reject many alternative explanations for the assemblage, including carnivore accumulation, “death trap” accumulation, and fluvial transport of bodies or bones (Dirks et al. 2015). This led us to the hypothesis that H. naledi were involved in bringing the bodies into the spaces where they were found. But we did not hypothesize their involvement in the formation of the deposit itself beyond bringing the bodies to the location.

    This approach seems conservative. It followed the traditional view that small-brained hominins do not engage in cultural practices. But we recognize in hindsight that this null hypothesis approach did harm to our analyses. It impeded us from recognizing within our initial excavations of the puzzle box area and other excavations between 2014 – 2017 that we might be encountering remains that were intrusive in the sedimentary floor of the chamber. If we had approached the accumulation of a large number of hominins from the perspective of the null hypothesis being that the situation was likely cultural, we perhaps would have collected evidence in a slightly different manner. We certainly note that if the Dinaledi system had been full of the remains of modern humans, there would have been little doubt that the null hypothesis would have been that this was a cultural space and not a “natural space”. We therefore respectfully disagree with the reviewers who continue to support the idea that we should approach hominin excavations with the null hypothesis that they will be natural (specifically non-cultural) in origins. If excavations continue with this mindset we believe that potential cultural evidence is almost certain to be lost.

    There has been a gradient across paleoanthropological excavations, archaeological work, and forensic investigation, with increasing precision of context. The reality is that the recording precision and frame of approach is typically different in most paleontological excavations than in those related to contemporary human remains. If anything comes from the present discussion of whether the Dinaledi system is a burial site for H. naledi or not, we hope that by taking seriously the possibility of deep cultural dynamics of hominins, we will encourage other teams to meet the highest standards of excavation in order to preserve potential cultural evidence. Given H. naledi’s cranial capacity we suggest that even very early hominin skeletal assemblages should be re-examined, if there is sufficient evidence or records available. These would include examples such as the A.L. 333 Au. afarensis site (the so called First Family site in Hadar Ethiopia), the Dikika infant skeleton, WT 15000 (Turkana Boy) and even A.L. 288 (Lucy) as such unusual taphonomic situations where skeletons are preserved cannot be simply explained away as “natural” in origin, based solely on the cranial capacity and assumed lack of cognitive and cultural complexity of the hominins as emphasized by us in Fuentes et al. (2023). We are not the first to observe that some very early hominin situations may represent early mortuary activity (Pettitt 2013), but we would advocate a step further. We suggest it may be damaging to take “natural accumulation” as the standard null hypothesis for hominin paleoanthropology, and that it is more conservative in practice to engage remains with the null hypothesis of possible cultural formation.

    We are deeply grateful for the time and effort all of the 8 reviewers (across three reviews) have taken with this work. We also acknowledge the anonymous reviewers from previous submissions who’s opinions and comments will have made the final iterations of these manuscripts better for their efforts. As this process is rather public and includes commentary outside of the eLife forum, we ask that the efforts of all 37 authors and 8 reviewers involved be respected and that the discourse remain professional in all venues as we study this fascinating and quite complex occurrence. We appreciate also the efforts of members of the public who have engaged with this relatively new process where preprints are posted prior to the reviews allowing comments and interactions from colleagues and the public who are normally not part of the internal peer review process. We believe these interactions will make for better final papers. We feel we have met the standards of demonstrating burials in H. naledi and that the engraving are most likely associated with H. naledi. However, given the reviews we see many areas where our clarity and context, and analyses, were less strong than they can be. With the clarifications and additions taken on board through these review processes the final papers will be stronger and clearer. We, recognize that this is an ongoing process of scientific investigation and further work will allow continued, and possibly better, evaluation of these hypothesis and others.

    Lee R Berger, Agustín Fuentes, John Hawks, Tebogo Makhubela

    Works cited:

    • Aspöck, E. (2008). What Actually is a ‘Deviant Burial’?: Comparing German-Language and Anglophone Research on ‘Deviant Burials.’ In E. M. Murphy (Ed.). Deviant Burial in the Archaeological Record. Oxford: Oxbow Books. pp 17–34.

    • Bolliger, S.A. & Thali, M.J. (2009). Thanatology. In S.A. Bolliger and M.J. Thali (eds) Virtopsy Approach: 3D Optical and Radiological Scanning and Reconstruction in Forensic Medicine. Boca Raton: CRC Press. pp 187-218.

    • Boulestin, B. & Duday, H. (2005). Ethnologie et archéologie de la mort: de l’illusion des références à l’emploi d’un vocabulaire. In: C. Mordant and G. Depierre (eds) Les Pratiques Funéraires à l’Âge du Bronze en France. Actes de la table ronde de Sens-en-Bourgogne. Paris: Éditions du Comité des Travaux Historiques et Scientifiques. pp. 17–30.

    • Boulestin, B. & Duday, H. (2006). Ethnology and archaeology of death: from the illusion of references to the use of a terminology. Archaeologia Polona 44: 149–169.

    • Bristow, J., Simms, Z. & Randolph-Quinney, P.S. Taphonomy. In S. Black and E. Ferguson (eds.) Forensic Anthropology 2000-2010. Boca Raton, FL: CRC Press. pp 279-318.

    • Channing, J. & Randolph-Quinney, P.S. (2006). Death, decay and reconstruction: the archaeology of Ballykilmore Cemetery, County Westmeath. In J. O’Sullivan and M. Stanley (eds.) Settlement, Industry and Ritual: Archaeology. National Roads Authority Monograph Series No. 3. Dublin: NRA/Four Courts Press. pp 113-126.

    • Cherryson, A. K. (2008). Normal, Deviant and Atypical: Burial Variation in Late Saxon Wessex, c. AD 700–1100. In E. M. Murphy (Ed.). Deviant Burial in the Archaeological Record. Oxford: Oxbow Books. pp 115–130.

    • Connolly, M., F. Coyne & L. G. Lynch (2005). Underworld : Death and Burial in Cloghermore Cave, Co. Kerry. Bray, Co. Wicklow: Wordwell.

    • Darwent, C. M. & R. L. Lyman (2002). Detecting the postburial fragmentation of carpals, tarsals and phalanges. In M. H. Sorg and W. D. Haglund (eds). Advances in Forensic Taphonomy: Method, Theory and Archeological Perspectives. Boca Raton, FL, CRC Press. pp 355-378.

    • d’Errico, F., & Backwell, L. (2016). Earliest evidence of personal ornaments associated with burial: The Conus shells from Border Cave. Journal of Human Evolution, 93, 91–108.

    • De Villiers. H. (1973). Human skeletal remains from Border Cave, Ingwavuma District, KwaZulu, South Africa. Annals of the Transvaal Museum, 28(13), 229–246.

    • Dell’Unto, N. and Landeschi, G. (2022). Archaeological 3D GIS. London: Routledge.

    • Dibble, H. L., Aldeias, V., Goldberg, P., McPherron, S. P., Sandgathe, D., & Steele, T. E. (2015). A critical look at evidence from La Chapelle-aux-Saints supporting an intentional Neandertal burial. Journal of Archaeological Science, 53, 649–657.

    • Dirkmaat, D. C., & Cabo, L. L. (2016). Forensic archaeology and forensic taphonomy: basic considerations on how to properly process and interpret the outdoor forensic scene_. Academic Forensic Pathology_ 6, 439–454.

    • Dirks, P. H., Berger, L. R., Roberts, E. M., Kramers, J. D., Hawks, J., Randolph-Quinney, P. S., Elliott, M., Musiba, C. M., Churchill, S. E., de Ruiter, D. J., Schmid, P., Backwell, L. R., Belyanin, G. A., Boshoff, P., Hunter, K. L., Feuerriegel, E. M., Gurtov, A., Harrison, J. du G., Hunter, R., … Tucker, S. (2015). Geological and taphonomic context for the new hominin species Homo naledi from the Dinaledi Chamber, South Africa. ELife, 4, e09561.

    • Dirks, P.H.G.M., Berger, L.R., Hawks, J., Randolph-Quinney, P.S., Backwell, L.R., and Roberts, E.M. (2016). Comment on “Deliberate body disposal by hominins in the Dinaledi Chamber, Cradle of Humankind, South Africa?” [J. Hum. Evol. 96 (2016) 145-148]. Journal of Human Evolution 96: 149-153.

    • Dirks, P. H., Roberts, E. M., Hilbert-Wolf, H., Kramers, J. D., Hawks, J., Dosseto, A., Duval, M., Elliott, M., Evans, M., Grün, R., Hellstrom, J., Herries, A. I., Joannes-Boyau, R., Makhubela, T. V., Placzek, C. J., Robbins, J., Spandler, C., Wiersma, J., Woodhead, J., & Berger, L. R. (2017). The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa. ELife, 6, e24231.

    • Donnelly, S., C. Donnelly & E. Murphy (1999). The forgotten dead: The cíllíní and disused burial grounds of Ballintoy, County Antrim. Ulster Journal of Archaeology 58, 109-113.

    • Duday, H. (2005). L’archéothanatologie ou l’archéologie de la mort. In: O. Dutour, J.-J. Hublin and B. Vandermeersch (eds) Objets et Méthodes en Paléoanthropologie. Paris: Comité des Travaux Historiques et Scientifiques. pp. 153–215.

    • Duday, H. (2009). Archaeology of the Dead: Lectures in Archaeothanatology. Oxford: Oxbow Books.

    • Finley, N. (2000). Outside of life: Traditions of infant burial in Ireland from cillin to cist. World Archaeology 31, 407-422.

    • Gargett, R. H. (1999). Middle Palaeolithic burial is not a dead issue: The view from Qafzeh, Saint-Césaire, Kebara, Amud, and Dederiyeh. Journal of Human Evolution, 37(1), 27–90.

    • Goldberg, P., Aldeias, V., Dibble, H., McPherron, S., Sandgathe, D., & Turq, A. (2017). Testing the Roc de Marsal Neandertal “Burial” with Geoarchaeology. Archaeological and Anthropological Sciences, 9(6), 1005–1015.

    • Gómez-Olivencia, A., & García-Martínez, D. (2019). New postcranial remains from the Roc de Marsal Neandertal child. PALEO. Revue d’archéologie Préhistorique, 30–1, 30–1.

    • Green, E.C. (2022). An archaeothanatological approach to the identification of late Anglo-Saxon burials in wooden containers. In C.J. Knüsel and E.M.J. Schotsmans (eds.) The Routledge Handbook of Archaeothanatology. London: Routledge. pp 436-455.

    • Henderson, J. (1987). Factors determining the state of preservation of human remains. In A. Boddington, A. Garland and R. Janaway (eds). Death, Decay and Reconstruction: Approaches to Archaeology and Forensic Science. Manchester: Manchester University Press. pp 43-54.

    • Hunter, J. R. (2014). Human remains recovery: archaeological and forensic perspectives. In C. Smith (ed). Encyclopedia of Global Archaeology. New York: Springer New York. pp 3549-3556.

    • Hochrein, M. (2002). An Autopsy of the Grave: Recognizing, Collecting and Preserving Forensic Geotaphonomic Evidence. In M. H. Sorg and W. D. Haglund (eds). Advances in Forensic Taphonomy: Method, Theory and Archeological Perspectives. Boca Raton, FL, CRC Press: 45-70.

    • Knüsel, C.K. & Robb, J. (2016). Funerary taphonomy: An overview of goals and methods. Journal of Archaeological Science: Reports 10, 655-673.

    • Kuhn, B.F., Berger, L.R. & Skinner, J.D. (2010). Examining criteria for identifying and differentiating fossil faunal assemblages accumulated by hyenas and hominins using extant hyenid accumulations. International Journal of Osteoarchaeology 20, 15-35.

    • Lyman, R. (1994). Vertebrate Taphonomy. Cambridge, Cambridge University Press.

    • Martinón-Torres, M., d’Errico, F., Santos, E., Álvaro Gallo, A., Amano, N., Archer, W., Armitage, S. J., Arsuaga, J. L., Bermúdez de Castro, J. M., Blinkhorn, J., Crowther, A., Douka, K., Dubernet, S., Faulkner, P., Fernández-Colón, P., Kourampas, N., González García, J., Larreina, D., Le Bourdonnec, F.-X., … Petraglia, M. D. (2021). Earliest known human burial in Africa. Nature, 593(7857), 7857.

    • Mickleburgh, H.L & Wescott, D.J. (2018). Controlled experimental observations on joint disarticulation and bone displacement of a human body in an open pit: implications for funerary archaeology. Journal of Archaeological Science: Reports 20: 158-167.

    • Mickleburgh, H.L., Wescott, D.J., Gluschitz, S. & Klinkenberg, V.M. (2022). Exploring the use of actualistic forensic taphonomy in the study of (forensic) archaeological human burials: An actualistic experimental research programme at the Forensic Anthropology Center at Texas State University (FACTS), San Marcos, Texas. In C.J. Knüsel and E.M.J. Schotsmans (eds.) The Routledge Handbook of Archaeothanatology. London: Routledge. pp 542-562.

    • Owsley, D. & B. Compton (1997). Preservation in late 19th Century iron coffin burials. In W. Haglund and M. Sorg (eds). Forensic Taphonomy: The Postmortem Fate of Human Remains. Boca Raton, FL, CRC Press: 511-526.

    • Parker Pearson, M. (1999). The Archaeology of Death and Burial. College Station: Texas A&M University Press.

    • Pettitt, P. (2013). The Palaeolithic Origins of Human Burial. Routledge.

    • Pomeroy, E., Bennett, P., Hunt, C. O., Reynolds, T., Farr, L., Frouin, M., Holman, J., Lane, R., French, C., & Barker, G. (2020). New Neanderthal remains associated with the ‘flower burial’ at Shanidar Cave. Antiquity, 94(373), 11–26.

    • Randolph-Quinney, P.S. (2013). From the cradle to the grave: the bioarchaeology of Clonfad 3 and Ballykilmore 6. In N. Brady, P. Stevens and J. Channing (eds.). Settlement and Community in the Fir Tulach Kingdom. Dublin: National Roads Authority Press. pp A2.1-48.

    • Randolph-Quinney, P.S., Haines, S. and Kruger, A. (2018). The use of three-dimensional scanning and surface capture methods in recording forensic taphonomic traces: issues of technology, visualisation, and validation. In: W.J. M. Groen and P. M. Barone (eds). Multidisciplinary Approaches to Forensic Archaeology. Berlin: Springer International Publishing, pp. 115-130.

    • Rendu, W., Beauval, C., Crevecoeur, I., Bayle, P., Balzeau, A., Bismuth, T., Bourguignon, L., Delfour, G., Faivre, J.-P., Lacrampe-Cuyaubère, F., Tavormina, C., Todisco, D., Turq, A., & Maureille, B. (2014). Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints. Proceedings of the National Academy of Sciences, 111(1), 81–86.

    • Sandgathe, D. M., Dibble, H. L., Goldberg, P., & McPherron, S. P. (2011). The Roc de Marsal Neandertal child: A reassessment of its status as a deliberate burial. Journal of Human Evolution, 61(3), 243–253.

    • Silver, M. (2016). Conservation Techniques in Cultural Heritage. In E. Stylianidis and F. Remondino (eds) 3D Recording, Documentation and Management of Cultural Heritage. Dunbeath: Whittles Publishing. pp 15-106.

    • Schotsmans, E.M.J., Georges-Zimmermann, P., Ueland, M. and Dent, B.B. (2022). From flesh to bone: Building bridges between taphonomy, archaeothanatology and forensic science for a better understanding of mortuary practices. In C.J. Knüsel and E.M.J. Schotsmans (eds.) The Routledge Handbook of Archaeothanatology. London: Routledge. pp 501-541.

  15. Version published to 10.1101/2023.06.01.543127v1 on bioRxiv