The Middle Pleistocene Muddle

Roksandic, Mirjana et al.  “Resolving the "muddle in the middle":  The case for Homo bodoensis sp. nov.”  Evolutionary Anthropology 31(1):20–29.  2022.  The naming proposal, published online in October 2021.  It asks that Homo heidelbergensis and Homo rhodesiensis be dropped and a new species founded on the Bodo cranium;  no new fossil or new analysis is reported.  Mirror:  PMC

1. 1a.

   Page 25:  “The endocranial capacity was estimated to 1250 cm3 (i.e., between 1200 and 1325 cm3).  The series of cut marks situated on the facial and posterior parietal regions were interpreted as intentional postmortem defleshing.”  “Dated to ca. 600 ka by laser-fusion 40Ar/39Ar technique (0.64 ± 0.03 Ma), biostratigraphy and tephrochronology.” ↩ ↩

2. 1b.

   Page 22:  “Unfortunately, several problems are evident in applying BSC to fossil specimens:  (1) morphological variation does not necessarily reflect reproductive isolation;  (2) reproductive isolation is not absolute even in well-defined living primate and other mammal species and crossbreeding has been observed even at the genus level”.  BSC is the biological species concept.  The paper goes on to quote Clifford Jolly's suggestion that any hominine species whose ancestries diverged less than 4 million years earlier may have been able to produce fertile hybrids. ↩

3. 1c.

   Page 24:  “Here we introduce a new Middle Pleistocene (i.e., Chibanian Age/Stage, 774–129 ka) hominin species that represents the direct ancestor of H. sapiens”.  Page 25:  “Kabwe 1 could represent a late survivor of the taxon.”  The second sentence is footnoted to the 2020 redating of the Kabwe skull. ↩ ↩

4. 1d.

   Page 21:  “a mandible, without an associated cranium, was used as the holotype for the taxon, even though this bone is normally considered to be extremely plastic and may or may not reflect associated morphological changes in the crania.”  The paper assigns Mauer, with Sima de los Huesos and the other western European fossils once called Homo heidelbergensis, to Homo neanderthalensis as early Neanderthals. ↩ ↩

5. 1e.

   Page 21:  “Even to nonspecialists (e.g., biologists working in other realms, Palaeolithic archaeologists, etc.) H. heidelbergensis represents either (and sometimes paradoxically both) the generalized Middle Pleistocene hominin, or a chronospecies of Neanderthals.”  “in a single paper, one can find numerous descriptions of the taxon with incompatible hypodigms.” ↩

6. 1f.

   Page 26, section Discussion and conclusions:  “While not a true species in the strict biological sense (since there is strong and growing evidence of migrations as well as gene flow between these diverged groups) this newly defined taxon cuts through the obfuscating and inconsistent use of improperly named and defined Middle Pleistocene hominins in Europe and Africa”. ↩

7. 1g.

   Page 23:  “The name is associated with Cecil Rhodes and English mining colonialism … While these considerations are not at the root of our rejection of the name, they are not minor and should not be ignored. … Decolonizing palaeoanthropology is an important task that needs to take precedence over rigid taxonomic rules.” ↩ ↩

8. 1h.

   Page 23:  “As such, H. heidelbergensis sensu lato can no longer be considered the root of all African and European hominin lineages.”  Their reason is that the last common ancestor is now placed in the late Early or very early Middle Pleistocene, before the fossils assigned to the species. ↩

9. 1i.

   Page 20:  “In 2019, we dedicated an entire American Association of Biological Anthropology (formerly American Association of Physical Anthropology) conference session to defining Homo heidelbergensis.  The results of the meeting were:  (1) no one was happy with the taxon;  (2) different people assigned different meanings to the species and included different fossils in the hypodigm;  (3) ignoring this problem will not miraculously lead to a solution;  and (4) that in order to better understand Middle Pleistocene hominin systematics, it was critical to clear up this “muddle in the middle.””  Page 21:  “The Middle Pleistocene is no longer dismissed as the proverbial “muddle in the middle,” but is increasingly recognized as a key time frame that witnessed the appearance, on a global scale, of two critical traits of later human morphology:  greater encephalization and smaller teeth, and likely the differentiation of geographic groups.” ↩ ↩ ↩

10. 1j.

    Page 22:  “When it comes to Middle Pleistocene hominin evolution, we identify two main options, with the understanding that other possibilities may exist:  (1) we could consider entire Pleistocene Homo fossils as a single lineage of H. sapiens with separate subspecies and/or chronospecies, or (2) we could consider the observed morphological variation as taxonomically meaningful within the “practical” species concept, without assuming that they were biological species and therefore not interfertile.”  Option 1 is footnoted to Wolpoff, Thorne, Jelínek and Zhang's "The case for sinking Homo erectus" (1994), to Wolpoff's Paleoanthropology (1999), and to Hawks (2013).  The paper goes on with option 2 and gives no argument against option 1. ↩

11. 1k.

    Page 22:  “We find it unlikely that the observed Middle Pleistocene variability can be subsumed under a single taxon such as (nebulously defined) H. heidelbergensis.”  The sentence that follows the two options.  Its target is the catch-all species, not the single lineage of option 1. ↩

12. 1l.

    Page 23:  “H. rhodesiensis Woodward 1921 never gained a wide usage in palaeoanthropology.  Indeed, a quick search on the Web of Science provides 274 direct mentions of H. heidelbergensis while only 17 hits for H. rhodesiensis.” ↩

Arsuaga, J. L. et al.  “Neandertal roots:  Cranial and chronological evidence from Sima de los Huesos.”  Science 344(6190):1358–1363.  Jun 20, 2014.  The "2014 skulls paper" by the excavating team, thirty authors.  Seventeen crania, seven of them new, from one layer;  mean capacity 1,232 cm³ for fifteen;  a re-dating to about 430 ka;  and the formal proposal to remove the sample from Homo heidelbergensis.  The dating section also reports a bear tooth at 261 +26/-25 ka, read as a minimum, and two ESR ages on quartz of 0.9 to 1.5 Ma, read as maxima.  Mirror:  Cnrs.hal.science

1. 2a.

   Page 1359:  “The mean of the SH cranial capacities (1232 cm3;  n = 15, table S3) is clearly above the Asian Homo erectus mean.  It is also well below the Neandertal and Pleistocene H. sapiens means and similar to the mean of the non-H. erectus Middle Pleistocene sample”.  SH is Sima de los Huesos. ↩ ↩

2. 2b.

   Page 1362:  “the SH population differs from some other European MPHs, such as Ceprano and Arago, that do not exhibit the suite of derived Neandertal features seen in SH.  Thus, more than one evolutionary lineage appears to have coexisted during the European Middle Pleistocene”.  MPHs are Middle Pleistocene hominins. ↩ ↩

3. 2c.

   Page 1359:  “To date, more than 6500 human fossils from at least 28 individuals are represented in a single stratigraphic level that we have redated using a suite of numerical and relative dating methods”.  The abstract, p. 1358, counts seventeen crania, seven of them new.  The chamber lies at the foot of a shaft. ↩ ↩

4. 2d.

   Page 1362:  “we have long maintained that the SH hominins are members of the Neandertal lineage”.  The sentence goes on to say that the team had nonetheless assigned the fossils to Homo heidelbergensis in a broad sense.  What changed in 2014 was the species name, not the lineage. ↩

5. 2e.

   Page 1359:  “All six of the TT-OSL and pIR-IR ages (428 ± 27 to 441 ± 25 ka for K-feldspar and 396 ± 35 to 429 ± 32 ka for quartz …) and one of the ESR quartz ages (443 ± 90 ka …) for this unit are statistically indistinguishable and support sedimentological evidence for a short-lived depositional episode.  Although two additional ESR quartz samples taken from LU-7 yielded significantly older ages of 0.9 to 1.5 Ma, these are interpreted as maximum ages owing to potential methodological complications.”  The unit is LU-7, the sediment lying on the bone-bearing clay.  The same section gives a bear tooth from the bone layer a combined ESR and uranium-series age of 261 +26/-25 ka, read as a minimum. ↩

6. 2f.

   Page 1362:  “We argue that the SH p-deme is sufficiently different from that of H. neanderthalensis so as to be considered a separate taxon.  Whether this difference should be recognized on the specific or subspecific level is currently an open question.”  A p-deme is a palaeo-deme, a local breeding population recognised in the fossil record. ↩ ↩ ↩ ↩

7. 2g.

   Page 1359:  “U-series dating of a cave raft speleothem deposited directly on a hominin cranium (cranium 4) from LU-6 yielded a mean age of 434 +36/-24 ka (n = 5 …).  We interpret this age to be pene-contemporaneous with the deposition of the cranium.” ↩

8. 2h.

   Page 1362:  “the difficulty with identifying derived Neandertal features in the Mauer mandible, the type specimen of H. heidelbergensis, contrasts strongly with the presence of numerous Neandertal apomorphies in the SH mandibles.  On this basis, we suggest that the SH sample be removed from the H. heidelbergensis hypodigm.” ↩ ↩

9. 2i.

   Page 1361:  “provides a minimum age of ~430 ka for the SH human fossils, which is some 100 ka younger than previously reported”. ↩ ↩

Delson, Eric, and Chris Stringer.  “The naming of Homo bodoensis by Roksandic and colleagues does not resolve issues surrounding Middle Pleistocene human evolution.”  Evolutionary Anthropology 31(5):233–236.  2022.  The first published rejection of Homo bodoensis, by two senior workers who agree with much of the biology and reject the name on the rules of zoological nomenclature.  Stringer is the author of the standard 2012 paper on Homo heidelbergensis.

1. 3a.

   Page 234:  “the facial shape of Bodo is derived by comparison to that of the inferred ancestor of H. sapiens, which was probably more like that of H. antecessor.”  Given as the argument of Stringer, Lacruz and colleagues, among others. ↩ ↩

2. 3b.

   Page 234:  “Wu et al. (whose lead author is a coauthor of Roksandic et al.) have suggested that the Hualongdong fossil from China is also a Middle Pleistocene ancestor for H. sapiens, which may well add further to the muddle.” ↩

3. 3c.

   Page 234:  “About the same time, many researchers took up the idea of “archaic H. sapiens” as a “taxon” for almost all post‐erectus specimens without clear subdivision, so that (sub)specific nomina such as rhodesiensis were rarely mentioned in the 1980s and 1990s … During the 2000s, the “archaic H. sapiens” concept was rejected in favor of recognizing several species or subspecies of Homo younger than 1 Ma.”  At the first omission the sentence continues "whereas Stringer and Rightmire included H. rhodesiensis in their conception of H. heidelbergensis".  The second rejection in print is Sarmiento and Pickford's, in the same issue. ↩ ↩

4. 3d.

   Page 233, section Abstract:  “We reject H. bodoensis as a junior synonym, with no value from its inception.” ↩

5. 3e.

   Page 233, section Abstract:  “It is not permitted to change a name solely because of variable (or erroneous) later use once it has been originally defined correctly, nor can a name be modified because it is offensive to one or more authors or to be politically expedient.” ↩

6. 3f.

   Page 233:  “Roksandic et al. surveyed Middle Pleistocene Homo (especially in Africa) and came to three questionable conclusions”.  “they suggested that the latter population/sample/species was the likely direct ancestor of Homo sapiens.”  The second quotation is the third of the three conclusions listed. ↩ ↩ ↩

7. 3g.

   Page 235:  “One can no more remove H. rhodesiensis from the history of the Kabwe/Broken Hill fossil than one can remove the name Rhodesia from the history of the countries of Zambia and Zimbabwe”. ↩

8. 3h.

   Page 234:  “If for some reason an author wished to recognize a separate taxon for the earlier Middle Pleistocene Africans, Homo saldanensis Drennan, 1955, would be available.” ↩

9. 3i.

   Page 234:  “Some authors (e.g., Delson & Baab, Rightmire, Hublin [the latter two offered several alternatives as well]) considered H. heidelbergensis as an early stage of H. neanderthalensis while recognizing H. rhodesiensis as a possible predecessor of H. sapiens, essentially the same picture as put forward by Roksandic et al. with a different name.” ↩

10. 3j.

    Page 234:  “We agree that H. heidelbergensis has at times been utilized for too broad a constellation of specimens/populations, but that is not a valid reason to suppress it”. ↩

Butzer, Karl W., and Glynn Ll. Isaac.  After the Australopithecines:  Stratigraphy, Ecology, and Culture Change in the Middle Pleistocene.  Mouton, The Hague.  1975.  Proceedings of a Wenner-Gren symposium held at Burg Wartenstein, Austria, in July 1973, edited by Butzer and Isaac, who wrote the Preface.  Isaac's closing chapter is "Sorting out the Muddle in the Middle:  An Anthropologist's Post-Conference Appraisal", pp. 875–887.  The earliest print use of the phrase found for this article.

1. 4a.

   Page 876, chapter Sorting out the Muddle in the Middle:  “Shackleton laid before the symposium clear evidence from oxygen isotope analysis of deep-sea sediments, that since the Brunhes-Matuyama polarity change there have been AT LEAST eight interglacials … terrestrial stratigraphers can finally lay to rest the time-honored four-fold climato-stratigraphic scheme of Gunz, Mindel, Riss, and Würm, plus correlatives.”  “Particular hazards exist with the dating procedure aptly described by Kukla as the “famous count-from-top” method, since many sections contain hidden gaps.”  The capitals are Isaac's. ↩

2. 4b.

   Page viii, section Preface:  “It is within this “floating” stratigraphic and paleoenvironmental framework that rare, informative archeological occurrences and even rarer hominid fossils rest, with little precision possible as to temporal or spatial interrelationships.” ↩

3. 4c.

   Page 881, chapter Sorting out the Muddle in the Middle:  “Detailed discussion of the morphology of hominid fossils and their evolutionary significance lay beyond the scope and ambitions of the symposium”.  Pages 881–882:  “Abiding uncertainty about the relative and absolute age of most Middle Pleistocene hominid fossils has been a great hindrance in the formulation and testing of adequate evolutionary hypotheses.”  Isaac is speaking.  No species name is debated anywhere in his chapter. ↩ ↩

4. 4d.

   Page viii, section Preface:  “Yet, between these two extremities, stratigraphic correlation and chronological resolution of the mid-Pleistocene time range is at present unsatisfactory.  Sedimentary sequences are characteristically brief, incomplete, and isolated, with rare opportunities for isotopic dating of any kind.” ↩ ↩

5. 4e.

   Page 875, chapter Sorting out the Muddle in the Middle:  “Instead of having to fit our fragments of evidence into a frame formed by a few hundred thousand years, we are free to move them about within a span of time approaching a million years.” ↩

6. 4f.

   Page ix, section Preface:  “If the “muddle in the middle” is to be resolved there must be awareness, problem-formulation, and interdisciplinary collaboration.”  “there has been little attempt to formulate the research questions to be asked or to specify the particular kinds of data needed.”  The phrase is already in quotation marks here, and is the title of Isaac's closing chapter.  The second quotation is from the paragraph before it. ↩ ↩ ↩

Stringer, Chris.  “The status of Homo heidelbergensis (Schoetensack 1908).”  Evolutionary Anthropology 21(3):101–107.  2012.  The standard statement of the case for keeping Homo heidelbergensis, by the Natural History Museum's palaeoanthropologist, and the paper that asked for the Sima de los Huesos fossils to be taken out of it.  Table 2 lists his hypodigm:  Mauer, Petralona, Arago, Vértesszöllös, Broken Hill, Bodo, Elandsfontein, Ndutu and Kapthurin, with queried entries including Ceprano, Dali, Jinniushan and Yunxian.

1. 5a.

   Page 104:  “Although Mounier and coworkers felt able to include the SH material in their heidelbergensis hypodigm, I believe that this inclusion has led to the most serious confusion surrounding the status of the species.” ↩ ↩

2. 5b.

   Page 103, table 2:  “In my view, the inclusion of fossils such as Steinheim and Montmaurin is still doubtful, while Narmada, Dali and Jinniushan may alternatively represent early “Denisovans.””  From the table's footnote. ↩ ↩ ↩ ↩ ↩

3. 5c.

   Page 105:  “reclassifying the SH material as an early form of H. neanderthalensis on the basis of its derived Neanderthal features and dating it to no earlier than 400 ka would remove most of the data supporting a European chronospecies of H. heidelbergensis-H. neanderthalensis.  This would open the possibility of a less inclusive diagnosis for the species that is ancestral to modern humans and Neanderthals, which, in my view, is still most reasonably named Homo heidelbergensis.” ↩

4. 5d.

   Page 105:  “Such gene flow is bound to raise serious issues about the validity of specific distinctions between heidelbergensis and its daughter species, particularly as such hybridization events may even have occurred in Africa.  Nevertheless, on morphological grounds, I believe that a pragmatic case can still be made for such species distinctions, especially bearing in mind the extensive evidence for interspecific hybridization even in living primates.” ↩ ↩

Bermúdez de Castro, J. M. et al.  “A Hominid from the Lower Pleistocene of Atapuerca, Spain:  Possible Ancestor to Neandertals and Modern Humans.”  Science 276(5317):1392–1395.  May 30, 1997.  The naming of Homo antecessor by the Atapuerca team, on fossils of at least six individuals from Gran Dolina level TD6.  The case rests chiefly on ATD6-69, the partial face of a child of 10 to 11.5 years.

1. 6a.

   Page 1392, section Abstract:  “The fully modern midfacial morphology of the fossils antedates other evidence of this feature by about 650,000 years.”  “Homo antecessor may represent the last common ancestor for Neandertals and modern humans.” ↩ ↩ ↩

2. 6b.

   Page 1394:  “the exclusive common ancestor of Neandertals and modern humans is not represented in the currently available European middle Pleistocene record.  We suggest that all the European middle Pleistocene fossils are ancestors (only) of the late Pleistocene Neandertals.  Moreover, the holotype, the Mauer mandible, shows clear derived neandertal traits”. ↩

Stringer, Chris.  “The origin and evolution of Homo sapiens.”  Philosophical Transactions of the Royal Society B 371(1698):20150237.  Jul 5, 2016.  Stringer's reconsideration of his 2012 position.  He asks what follows if the fossils he assigns to Homo heidelbergensis are too young and too derived in the face to be the last common ancestor of Neanderthals and modern humans, and draws an alternative tree with a hypothetical "Ancestor X" at the root.  He does not say the older view is refuted.  Mirror:  PMC

1. 7a.

   Page 9:  “An alternative would be to consider a H. antecessor-like morphology as more likely for the LCA of H. sapiens and H. heidelbergensis”.  “If heidelbergensis (sensu Stringer) is not the LCA for sapiens and neanderthalensis because of a derived facial morphology and because the known fossils post-date the actual divergence date, then what did the LCA look like (Ancestor X in figure 2c).  It may have displayed a morphology of the lower face more like antecessor than heidelbergensis, but what about the rest of the cranium and the dentition?”  LCA is last common ancestor. ↩

2. 7b.

   Page 8:  “As long as the biological species concept (which does not operate well for many closely related extant species of birds and mammals) is not imposed, species can be recognized in the fossil record as evolutionary lineages which maintain their identity through significant periods of time (and in the face of small amounts of introgression).”  “However, that pragmatic use of the term ’species’ must be tempered with a recognition that these ‘species’ were not genetically impermeable.” ↩

3. 7c.

   Page 9:  “using the latest estimates of the autosomal human mutation rate, the divergence date of the neanderthalensis and sapiens lineages can indeed be placed earlier, between 550 and 765 ka, which would be consistent with only the oldest suggested examples of heidelbergensis as potentially representing the LCA”. ↩

4. 7d.

   Page 7, figure 2:  “H. helmei and H. steinheimensis as intermediate species between each terminal species and LCA, here suggested to be H. heidelbergensis.”  “A tree which uses the new date and Neanderthal-like morphology of the Sima sample, plus an inferred deeper divergence date based on new genomic mutation rate estimates.  Here, a hypothetical and older ‘Ancestor X’ replaces heidelbergensis as the LCA.”  From the caption.  The first quotation describes tree (a), the second tree (c).  The abstract says:  "I reconsider the nature of the middle Pleistocene ancestor of these lineages". ↩ ↩ ↩

5. 7e.

   Page 9:  “Bermúdez de Castro & Martinón-Torres concluded that antecessor was a side-branch of a lower Pleistocene radiation of lineages in Eurasia that eventually gave rise to Neanderthals in Europe and to H. sapiens in Africa.”  The work cited is dated 2014.  Bermúdez de Castro is the first author of the 1997 paper that named the species. ↩ ↩

6. 7f.

   Page 2:  “Since 1983, I have built the case that shape resemblances between the Broken Hill and Petralona crania indicate the existence of a widespread middle Pleistocene population which can be called Homo heidelbergensis if the Mauer mandible is also included”. ↩

Welker, Frido et al.  “The dental proteome of Homo antecessor.”  Nature 580(7802):235–238.  Apr 9, 2020.  Mass spectrometry of enamel from one Homo antecessor molar fragment, ATD6-92, by a Copenhagen-led team with the Atapuerca excavators.  Seven proteins, 22 percent of sites, one substitution unique to the specimen;  the last authors are Bermúdez de Castro, Willerslev and Cappellini.  Quotations are located by section.  Mirror:  PMC

1. 8a.

   Section Main text:  “The enamel protein sequences do not resolve the relationships between HNDs due to the low number of informative single amino acid polymorphisms (SAPs).”  “The concatenated gene tree may suffer from incomplete lineage sorting, and we have too little sequence data to discard this possibility at the moment.  If we were, however, to use the concatenation of available gene trees as a best guess for the population tree, and assuming such a population tree is a good descriptor of the relationships among ancient hominins, then our results support the placement of Homo antecessor as a closely related sister taxon of the last common ancestor of HNDs.”  HNDs are modern humans, Neanderthals and Denisovans collectively. ↩

2. 8b.

   Section Methods:  “We used sequence information for seven proteins (ALB, AMBN, AMELX, AMELY, COL17α1, ENAM and MMP20) for the Homo antecessor individual and six proteins for the Homo erectus individual … with a total of 22.08% and 22.14% non-missing sites, respectively (Tab. S9).  We were able to recover a unique SAP for Homo antecessor, however, for Homo erectus no unique SAP was detected”.  A SAP is a single amino-acid polymorphism.  The Homo erectus tooth is from Dmanisi, 1.77 million years on the mainstream scale. ↩

3. 8c.

   Page 235, section Abstract:  “We demonstrate that Homo antecessor is a close sister lineage to subsequent Middle and Late Pleistocene hominins such as modern humans, Neanderthals, and Denisovans.  This placement implies that the modernlike face of Homo antecessor may have a considerably deep ancestry in the genus Homo, and that the Neanderthal cranial morphology represents a derived form.”  Section Main text:  “In these trees, the Homo antecessor sequence represents a sister taxon closely related to, but not part of, the group composed of Late Pleistocene hominins for which molecular data is available”. ↩ ↩ ↩

4. 8d.

   Section Main text:  “Homo antecessor was tentatively proposed as the last common ancestor of Neanderthals and modern humans.  The modern-like face of some immature individuals, and particularly that of the more complete specimen ATD6-69, as well as the zygomaxillary fragment ATD6-58 of one adult individual, were key in this proposition.”  “It is still contentious whether Homo antecessor could represent the last common ancestor of Homo sapiens, Neanderthals, and Denisovans, or whether it represents a sister lineage to the last common ancestor of these species (here collectively called HNDs).”  The 1997 proposal is Bermúdez de Castro et al., Science 276:1392–1395. ↩ ↩

Hutson, Jarod M. et al.  “Revised age for Schöningen hunting spears indicates intensification of Neanderthal cooperative behavior around 200,000 years ago.”  Science Advances 11(19):eadv0752.  May 9, 2025.  Re-dates the Schöningen Spear Horizon by amino-acid racemization in snail opercula from the find layer, supported by ostracods and horse teeth that the authors call still experimental, together with a re-reading of the regional interglacial sequence.  The result moves the spears from about 300 to about 200 ka and from Homo heidelbergensis to Neanderthals.

1. 9a.

   Section Introduction:  “lays claim to the world’s oldest complete wooden hunting weapons, including nine spears, one lance, and six double-pointed sticks.  This arsenal is joined by a small assemblage of about 1500 flint artifacts scattered around the butchered remains of more than 50 horses …, altogether representing the spoils of repeated ambush hunting episodes along the shoreline of a former lake.  Nowhere else in the Middle Pleistocene record are such hunting encounters so well preserved in undisturbed contexts.”  The first paragraph of the paper.  It mentions no human bones from the site. ↩ ↩

2. 9b.

   Section Discussion:  “With claimed ages of ~400 or ~300 ka for the Spear Horizon, Schöningen stood as an outlier among sites attributed to the Lower Paleolithic and to Homo heidelbergensis, the probable last common ancestor of early modern humans that evolved in Africa and the Neanderthals of western Eurasia.  Our dating evidence for the Spear Horizon corrects this mismatch and aligns the Schöningen spears within the timeframe of European Neanderthals and the Middle Paleolithic.” ↩ ↩ ↩

Grün, Rainer et al.  “Dating the skull from Broken Hill, Zambia, and its position in human evolution.”  Nature 580(7803):372–375.  Apr 16, 2020.  The first direct dating of the Kabwe skull:  uranium-series by laser ablation on a piece of the skull base, with ESR on enamel for an upper limit, by the geochronologist Rainer Grün with Chris Stringer.  Quotations located by line number are from the authors' submitted manuscript and may differ in wording from the printed text;  the abstract and the figure of 299 ± 25 ka are from the published paper.

1. 10a.

   Page 372, section Abstract:  “the original site has since been completely quarried away … the cranium is often estimated to be around 500 thousand years old … Here we carried out analyses directly on the skull and found a best age estimate of 299 ± 25 thousand years (mean ± 2σ).  The result suggests that later Middle Pleistocene Africa contained multiple contemporaneous hominin lineages (that is, Homo sapiens, H. heidelbergensis/H. rhodesiensis and Homo naledi)”.  The skull is museum number E686.  The authors' manuscript gives the ESR upper limit as 322 ka and says faunal comparisons had suggested an age "perhaps as old as 500 ka". ↩ ↩

2. 10b.

   Page 372, section Abstract:  “The age estimate also raises further questions about the mode of evolution of H. sapiens in Africa and whether H. heidelbergensis/H. rhodesiensis was a direct ancestor of our species.” ↩

3. 10c.

   Lines 181–182:  “Such diversity might even have included gene flow between populations that would normally be considered distinct species.” ↩

4. 10d.

   Lines 169–171:  “Although the new age determination for the Broken Hill skull lies at a date when an intermediate morphology might be expected, the skull shows no significant anatomically modern derived traits.”  Lines 174–176:  “age estimates of >200 ka for the fossils from Florisbad (South Africa) and Guomde (Kenya), and of ~300 ka for the Jebel Irhoud (Morocco) material, all of which display more "modern" traits than E686.”  E686 is the Kabwe skull. ↩ ↩

Carbonell, Eudald, and Marina Mosquera.  “The emergence of a symbolic behaviour:  the sepulchral pit of Sima de los Huesos, Sierra de Atapuerca, Burgos, Spain.”  Comptes Rendus Palevol 5(1–2):155–160.  2006.  The excavators' reading of a handaxe found with the Sima bones, and of the heap of bodies itself, as symbolic.  Read for this article in its French abstract only.

1. 11a.

   Section Abstract:  “Il a fourni les restes de 28 hominidés datant d'environ 400 000 ans.  Ceci représente la plus riche collection d'Homo heidelbergensis du Pléistocène moyen dans le monde.  Sima de los Huesos n'a jamais été un lieu d'occupation humaine, puisqu'il n'y a pas de trace d'habitat”.  “C'est une hache sur un fin éclat de quartzite, qui est associée à l'assemblage des restes humains.  La nature particulière du dépôt du point de vue taphonomique, paléontologique et technologique indique une signification symbolique à la fois de l'outil et de l'accumulation des restes humains.”  In English:  it has yielded the remains of 28 hominids about 400,000 years old, the richest collection of Homo heidelbergensis of the Middle Pleistocene in the world;  it was never a place of human occupation, since there is no trace of habitation.  The tool is a handaxe on a thin quartzite flake, found with the human remains;  the particular nature of the deposit, taphonomically, palaeontologically and technologically, indicates a symbolic meaning for both the tool and the accumulation of human remains. ↩ ↩ ↩ ↩

Meyer, Matthias et al.  “Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins.”  Nature 531(7595):504–507.  2016.  Nuclear DNA from two Sima specimens, which places the population on the Neanderthal side, confirming the 2014 skull-based reassignment.

1. 12a.

   Page 504, section Abstract:  “Here we recover nuclear DNA sequences from two specimens, which show that the Sima de los Huesos hominins were related to Neanderthals rather than to Denisovans, indicating that the population divergence between Neanderthals and Denisovans predates 430,000 years ago.  A mitochondrial DNA recovered from one of the specimens shares the previously described relationship to Denisovan mitochondrial DNAs, suggesting, among other possibilities, that the mitochondrial DNA gene pool of Neanderthals turned over later in their history.” ↩ ↩

Fu, Qiaomei et al.  “The proteome of the late Middle Pleistocene Harbin individual.”  Science 389(6761):704–707.  Aug 14, 2025.  Ninety-five proteins recovered from the Harbin skull.  The abstract counts three Denisovan-derived variants;  the body of the paper calls them one highly confident and two potential.

1. 13a.

   Page 704, section Abstract:  “we report 95 endogenous proteins retrieved from a nearly complete cranium from Harbin, China, dating to at least 146,000 years ago and previously assigned to a new species, Homo longi.  This individual has three Denisovan-derived amino acid variants and clusters with Denisova 3, suggesting that the Harbin individual belongs to a Denisovan population.” ↩ ↩ ↩

2. 13b.

   Page 706:  “the identification of one highly confident and two potential Denisovan-derived amino acid variants supports the conclusion that the Harbin individual was from a Denisovan population.”  Page 707, section Discussion and conclusions:  “Through proteomic analysis, we were able to definitively identify this individual as Denisovan.” ↩

Fu, Qiaomei et al.  “Denisovan mitochondrial DNA from dental calculus of the >146,000-year-old Harbin cranium.”  Cell 188(15):3919–3926.e9.  Jul 24, 2025.  Mitochondrial DNA recovered from tartar on the Harbin skull's teeth, after a tooth and the petrous bone yielded none.  Pääbo's laboratory.  The paper states its own limit:  a small amount of mitochondrial DNA and no nuclear DNA.  Mirror:  Hdl.handle.net

1. 14a.

   Page 3919, section Summary:  “Although no DNA could be retrieved from a tooth or the petrous bone, mitochondrial DNA (mtDNA) could be isolated from dental calculus.  The mtDNA falls within Denisovan mtDNA variation and is related to an mtDNA branch carried by early Denisovan individuals in southern Siberia, previously observed in Denisova Cave.” ↩ ↩

2. 14b.

   Page 3922, section Discussion:  “fossils from Dali, Jinniushan, and Hualongdong exhibit morphological traits similar to those in the Harbin cranium, making it plausible that they also represent Denisovan populations.”  Section Limitations of the study:  “This study is based on the limited amount of hominin mtDNA that could be retrieved from the dental calculus of the Harbin cranium.” ↩ ↩ ↩ ↩

Bischoff, J. L. et al.  “Geology and preliminary dating of the hominid-bearing sedimentary fill of the Sima de los Huesos Chamber, Cueva Mayor of the Sierra de Atapuerca, Burgos, Spain.”  Journal of Human Evolution 33(2–3):129–154.  1997.  The first dating of the Sima fossils, by the excavating team's geochronologist.  Uranium-series on sixteen human and 25 bear bones, and ESR on six bear bones.

1. 15a.

   Section Abstract:  “Electron spin resonance (ESR) analyses of six selected bear bones indicates dates of 189 +/- 28 ka, for which each is cordant with their corresponding U-series date (181 +/- 41 ka).  Combined ESR and U-series dates for these samples yielded 200 +/- 4 ka.  Such agreement is highly suggestive that uranium uptake in these bones was close to the early-uptake (EU) model, and the dates are essentially correct.”  “The dating results, therefore, seem to provide a firm minimum age of about 200 ka for the human entry: and suggestive evidence of entry before 320 ka.”  "cordant" is as printed in the abstract.  The second quotation is the abstract's last sentence;  the authors claimed a minimum, and the accepted age is now about 430 ka. ↩ ↩

Barham, L. et al.  “Evidence for the earliest structural use of wood at least 476,000 years ago.”  Nature 622(7981):107–111.  Oct 5, 2023.  Excavation report from Kalambo Falls, Zambia.  Sixteen sand samples were dated by luminescence, the older ones by pIR-IR on potassium feldspar;  the ages fall in three clusters in stratigraphic order, 476, 390 and 324 ka.  No human bones were found.  Mirror:  PMC

1. 16a.

   Page 107, section Abstract:  “Waterlogged deposits at the archaeological site of Kalambo Falls, Zambia, dated by luminescence to at least 476 ± 23 kyr ago (ka), preserved two interlocking logs joined transversely by an intentionally cut notch.  This construction has no known parallels in the African or Eurasian Palaeolithic.” ↩ ↩

2. 16b.

   Page 107, section Abstract:  “At Kalambo we also recovered four wood tools from 390 ka to 324 ka, including a wedge, digging stick, cut log and notched branch.  The finds show an unexpected early diversity of forms and the capacity to shape tree trunks into large combined structures.”  The joined logs are in the lowest cluster at 476 ± 23 ka;  the tools lie in the two younger clusters above. ↩ ↩

White, Tim D.  “Cut marks on the Bodo cranium:  a case of prehistoric defleshing.”  American Journal of Physical Anthropology 69(4):503–509.  Apr 1986.  The description of the stone-tool cut marks on the Bodo skull, by the palaeoanthropologist who later led the Middle Awash project.

1. 17a.

   Page 503, section Abstract:  “Cut marks were discovered on the Middle Pleistocene Bodo cranium from Ethiopia.  The cut marks most closely resemble experimental damage caused by the application of stone tools to fresh bone.  This discovery constitutes the earliest solid evidence for intentional defleshing of a human ancestor and offers new research avenues for the investigation of early hominid mortuary practices.” ↩ ↩

Sala, Nohemi et al.  “Lethal interpersonal violence in the Middle Pleistocene.”  PLOS ONE 10(5):e0126589.  May 27, 2015.  Forensic analysis of Cranium 17 from Sima de los Huesos, which carries two matching depression fractures struck from different directions around the time of death.  The paper calls the sample part of the Neandertal clade.  Mirror:  PMC

1. 18a.

   Page 9, section Conclusions:  “The only possible manner by which a deceased individual could have arrived at the SH site is if its cadaver were dropped down the shaft by other hominins.  Thus, the interpretation of the SH site as a place where hominins deposited deceased members of their social groups seems to be the most likely scenario to explain the presence of human bodies at the site.  This interpretation implies this was a social practice among this group of Middle Pleistocene hominins and may represent the earliest funerary behavior in the human fossil record.” ↩ ↩

2. 18b.

   Page 1, section Abstract:  “Given that either of the two traumatic events was likely lethal, the presence of multiple blows implies an intention to kill.” ↩

3. 18c.

   Page 2, section Introduction:  “hominin fossils belonging to the Neandertal clade and corresponding to a minimum of 28 individuals.  During the time the hominin bones accumulated at the SH site, the only possible access route to the site were through a deep (13 m) vertical chimney.  Given the skeletal part representation in the collection, it is likely that entire bodies were deposited at the site.  There are no cutmarks on any of the 6700+ hominin bones recovered to date, and carnivore manipulation (tooth marks) of the bones is rare.”  "were" is as printed. ↩ ↩ ↩

Wood, Todd Charles.  “Baraminological analysis places Homo habilis, Homo rudolfensis, and Australopithecus sediba in the human holobaramin.”  Answers Research Journal 3:71–90.  2010.  Statistical baraminology, baraminic distance correlation and multidimensional scaling, on eight published hominin character matrices, by the creationist biologist who developed the method.  One matrix, Zeitoun's, scores individual skulls, which is how Kabwe, Petralona and Dali were tested. ↩

1. 19a.

   Page 79, section Results:  “BDC results for Zeitoun’s dataset show two obvious clusters (fig. 10).  One consists of 38 specimens referred to Homo and the other consists of chimp, gorilla, P. boisei (KNM-ER 406) and Au. africanus (Sterkfontein 5).  …  Otherwise most Homo taxa share significant, positive BDC with other members of Homo.”  Page 80, figure 10:  BDC is baraminic distance correlation.  The 38 specimens labelled in the figure include Broken Hill 1 (the Kabwe skull), Petralona and Dali.  Bodo 1, two Arago fossils and Saldanha were dropped before the run because more than 60 percent of their characters were unknown. ↩

Wagner, Günther A. et al.  “Radiometric dating of the type-site for Homo heidelbergensis at Mauer, Germany.”  PNAS 107(46):19726–19730.  Nov 16, 2010.  Dates the find horizon of the Mauer jaw by infrared radiofluorescence on sand grains and combined ESR and uranium-series on herbivore teeth.  Before this the jaw was placed by its fauna at 478–533 or 563–621 ka.  Mirror:  PMC

1. 20a.

   Page 19726, section Abstract:  “we show that two independent techniques, the combined electron spin resonance/U-series method used with mammal teeth and infrared radiofluorescence applied to sand grains, date the type-site of Homo heidelbergensis at Mauer to 609 ± 40 ka.” ↩ ↩

2. 20b.

   Page 19727:  “the results obtained should be treated as minimum ages and cannot be considered in the geochronological interpretation.  The ages obtained for the four other teeth are, respectively, 624 +79/−73 and 627 +73/−71 ka for the lower sands and 458 +39/−37 and 502 +43/−41 ka for the upper sands”.  Eight teeth were analysed, five from the lower sands and three from the upper. ↩

3. 20c.

   Page 19727:  “The hypodigm and evolutionary significance of Homo heidelbergensis are currently uncertain, but this species is generally considered to be the ancestor of Neanderthals in Europe”. ↩

Woodward, Arthur Smith.  “A New Cave Man from Rhodesia, South Africa.”  Nature 108(2716):371–372.  Nov 17, 1921.  The original description of the Kabwe skull and the naming of Homo rhodesiensis, by the Keeper of Geology at the British Museum (Natural History).

1. 21a.

   Page 371:  “a nearly complete human skull, a fragment of the upper jaw of another, a sacrum, a tibia, and the two ends of a femur”.  Woodward says they were dug out of the earth in a remote part of the cave. ↩

2. 21b.

   Page 371:  “Its brain-case is typically human, with a wall no thicker than that of the average European, and its capacity, though still not determined, is obviously well above the lower human limit.  Its large and heavy face is even more simian in appearance than that of Neanderthal man”. ↩

Rightmire, G. Philip.  “Homo in the Middle Pleistocene:  hypodigms, variation, and species recognition.”  Evolutionary Anthropology 17(1):8–21.  2008.  The case for one Afro-European species argued from measurements:  ten crania tabulated and their coefficients of variation compared with those of Homo erectus and living humans.  Table 1 lists the endocranial capacities used in this article.

1. 22a.

   Page 18:  “comparisons of CV and V for several measurements suggest that variation within the mid-Pleistocene assemblage is not excessive relative to that observed in other taxa.  Especially for some facial dimensions, values of CV and V are higher than in samples of modern humans.  However, magnitudes of variation within the mid-Pleistocene group are evidently not greater than in Homo erectus (sensu lato).  On the strength of these findings, it can be proposed that the crania from Africa and Europe constitute a single, species-level hypodigm.  If all of the earlier Middle Pleistocene fossils, including the Mauer mandible, belong to a single lineage, then this species should be called Homo heidelbergensis.” ↩

2. 22b.

   Page 18:  “Broken Hill is very much like Petralona in many of its features.  All of the crania share a morphology that sets them apart from Homo erectus and differentiates them from modern humans.” ↩

3. 22c.

   Page 9, table 1:  Endocranial capacities in cm³:  Bodo 1250;  Ndutu 1100;  Broken Hill 1280;  Arago 1166;  Sima de los Huesos 4, 1390;  Sima de los Huesos 5, 1125;  Steinheim 1100;  Petralona 1230.  Omo 2 and Elandsfontein are listed without a capacity.  Page 12:  “At close to 1,100 cm3, SH 5 is rather small, but SH 4, with a capacity of 1,390 cm3, is one of the largest of all Middle Pleistocene specimens.” ↩ ↩ ↩ ↩ ↩

Clark, J. D. et al.  “African Homo erectus:  old radiometric ages and young Oldowan assemblages in the Middle Awash Valley, Ethiopia.”  Science 264(5167):1907–1910.  Jun 24, 1994.  The source of the Bodo date that later papers repeat.  The argon-argon ages are on volcanic material in the sand unit, not on the skull, which was a surface find.

1. 23a.

   Section Abstract:  “Ar/Ar ages, biostratigraphy, and tephrachronology from this area indicate that the Pleistocene Bodo hominid cranium and newer specimens are approximately 0.6 million years old.”  "tephrachronology" is the abstract's spelling.  Roksandic and colleagues give the figure as 0.64 ± 0.03 Ma by laser-fusion argon-argon. ↩

Falguères, Christophe et al.  “New U-series dates on the Petralona cranium, a key fossil in European human evolution.”  Journal of Human Evolution 206:103732.  Sep 2025.  Uranium-series on the calcite that grew directly on the Petralona skull, giving a minimum age of 286 ± 9 ka.  Quotations are located by the line numbers of the authors' manuscript.

1. 24a.

   Section Abstract, lines 17–20:  “Previous results obtained on various types of samples yielded a large age range between about 170 and 700 ka, precluding any consensus on the age of the human fossil.  On the other hand, the original stratigraphic position of the cranium also remains enigmatic”. ↩

2. 24b.

   Section Conclusions, lines 320–323:  “According to this hypothesis and from a morphological point of view, the Petralona hominin forms part of a distinct and more primitive group than H. sapiens and Neanderthals, and the new age estimate further supports the co-existence of this population alongside the evolving Neanderthal lineage in the later Middle Pleistocene of Europe.” ↩

3. 24c.

   Section The site, lines 49–50:  “without any stratigraphic context, apparently just cemented to the chamber wall”.  Of the skull's discovery in 1960.  The paper shows that the crust on the skull is not the same age as the wall. ↩

4. 24d.

   Section The site, lines 51–52:  “attributing it variously to Homo erectus, H. neanderthalensis or ‘archaic H. sapiens’”.  Lines 55–56:  “this has often led to the inclusion of both Petralona and Kabwe in an extended hypodigm of H. heidelbergensis”.  Lines 57–60:  “There is currently much debate about the validity of the extended hypodigm H. heidelbergensis sensu lato and the evolutionary position of the associated fossils, particularly whether any of them might have been ancestral to H. sapiens or to H. neanderthalensis, ancestral to both, or ancestral to neither”. ↩ ↩

Arnold, Lee J. et al.  “Luminescence dating and palaeomagnetic age constraint on hominins from Sima de los Huesos, Atapuerca, Spain.”  Journal of Human Evolution 67:85–107.  2014.  Two extended-range luminescence methods, pIR-IR on potassium feldspar and TT-OSL on single quartz grains, applied to the sediment lying on the bone-bearing clay, by the Atapuerca team's own geochronologists.  The paper also withdraws the team's earlier "more than 530 ka".

1. 25a.

   Page 85, section Abstract:  “Post-infrared-infrared stimulated luminescence (pIR-IR) dating of K-feldspars and thermally transferred optically stimulated luminescence (TT-OSL) dating of individual quartz grains provide weighted mean ages of 433 ± 15 ka (thousands of years) and 416 ± 19 ka, respectively, for allochthonous sedimentary horizons overlying the hominin-bearing clay breccia.  The six replicate luminescence ages obtained for this deposit are reproducible and provide a combined minimum age estimate of 427 ± 12 ka for the underlying hominin fossils.” ↩

Fernández-Jalvo, Yolanda et al.  “Human cannibalism in the Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain).”  Journal of Human Evolution 37(3–4):591–622.  1999.  The taphonomic study of the human bones from the 1994 to 1996 test pit in level TD6, by members of the Atapuerca team.

1. 26a.

   Section Abstract:  “Human remains belonging to at least six individuals were found in an exploratory excavation … The area of the exploratory excavation is about 7 m(2). … There is no specific distribution, treatment, or arrangement of the human remains, which were found randomly mixed with abundant faunal remains and stone tools.  Most of the faunal and human fossil bones from the Aurora Stratum have human induced damage.  Stone tool cutmarks are frequent”.  “Both nonhuman and human remains show similar intensive exploitation.”  Read in abstract only.  "7 m(2)" is 7 square metres as the abstract prints it. ↩ ↩ ↩

2. 26b.

   Section Abstract:  “The characteristics of this fossil assemblage suggest that it is solely the result of consumptive activities as there is no evidence of ritual or other intention.” ↩

Saladié, Palmira et al.  “Intergroup cannibalism in the European Early Pleistocene:  the range expansion and imbalance of power hypotheses.”  Journal of Human Evolution 63(5):682–695.  2012.  A re-analysis of the butchered human and deer remains from Gran Dolina TD6 by the Atapuerca team, comparing the age profile of the victims with cannibalism in chimpanzees.  Mirror:  Docta.ucm.es

1. 27a.

   Section Abstract:  “The cannibalism documented in level TD6 mainly involves the consumption of infants and other immature individuals.  The human induced modifications on Homo antecessor and deer remains suggest that butchering processes were similar for both taxa, and the remains were discarded on the living floor in the same way.” ↩

2. 27b.

   Section Abstract:  “it is similar to the age profiles seen in cannibalism associated with intergroup aggression in chimpanzees.” ↩

3. 27c.

   Section Results, The assemblage:  “a minimum number of individuals (MNI) of 11 has been established for H. antecessor”.  “Six were children, four of whom were younger than five years of age and two of whom were between five and nine years of age at the time of death, three were between 10 and 15 years of age, and two were young adults”.  Section Discussion:  “At TD6, most of the cannibalized individuals were children at the time of death.” ↩ ↩

Duval, Mathieu et al.  “The first direct ESR dating of a hominin tooth from Atapuerca Gran Dolina TD-6 (Spain) supports the antiquity of Homo antecessor.”  Quaternary Geochronology 47:120–137.  2018.  The only direct date on a Homo antecessor fossil:  ESR on the enamel of molar fragment ATD6-92, by the Atapuerca team with the geochronologist Rainer Grün.  Section 2.3 tabulates every earlier date on the layer.

1. 28a.

   Page 120, section Abstract:  “The analysis of a tooth (ATD6-92) from TD6 unit of Atapuerca Gran Dolina (Spain) following a “semi non-destructive” procedure provides a final age estimate ranging from 624 to 949 ka, which covers all possible uranium uptake scenarios.  Last, the additional magnetostratigraphic data collected within TD6 enables to further constrain the initial ESR chronology and to propose an age of between 772 and 949 ka for Homo antecessor, in agreement with previous dating works.  Although our new results do not refine the existing chronology of TD6 unit, they nevertheless support the antiquity of H. antecessor”. ↩

Ji, Qiang et al.  “Late Middle Pleistocene Harbin cranium represents a new Homo species.”  The Innovation 2(3):100132.  2021.  The two-page paper that names Homo longi on the Harbin cranium and proposes Homo daliensis for Dali and Hualongdong.  The journal uses article numbers, so quotations are located by section.  Mirror:  PMC

1. 29a.

   Section Comparative morphology:  “The endocranial capacity is estimated as 1,420 ml, falling in the range of H. sapiens and Neanderthals, and larger than other Homo species such as H. erectus, H. naledi, H. floresiensis, and even some H. heidelbergensis/H. rhodesiensis.” ↩ ↩ ↩

2. 29b.

   Section Remarks:  “The Dali cranium was initially proposed as a subspecies of H. sapiens (H. s. daliensis) by Xinzhi Wu, but Wu abandoned the subspecies name and called the cranium “archaic H. sapiens” in his later publications.  It was also suggested to be a subspecies of H. heidelbergensis (H. h. daliensis), or should be raised to the species level (H. daliensis).” ↩ ↩

3. 29c.

   Section Comparative morphology:  “The Jinniushan cranium has a similar cranial capacity (1,390 ml) to the Harbin, but is more gracile.”  Other figures for Jinniushan, about 1,260 to 1,300 cm³, are also in print;  the skeleton is usually described from Rosenberg, Lü and Ruff 2006. ↩

4. 29d.

   Section Locality and age:  “Middle Pleistocene, upper part of the Upper Huangshan Formation (138–309 ka), near the Dongjiang Bridge in Harbin City, Heilongjiang Province, China.  A reliable minimum U-series age is 148±2 ka.” ↩

Feng, Xiaobo et al.  “The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans.”  Science 389(6767):1320–1324.  Sep 25, 2025.  A digital reconstruction of the crushed Yunxian 2 cranium, with parts taken from Yunxian 1, and a tip-dated tree of 57 fossils that places it in the Homo longi clade.  The last two authors named that clade in 2021.  Earlier papers in its reference list call the Yunxian skulls Homo erectus.  A preprint of March 2025 differs from the printed text in wording, not in any number quoted here.  Mirror:  bioRxiv

1. 30a.

   Page 1320, section Abstract:  “Morphometric and phylogenetic analyses suggest that it is an early member of the Asian H. longi clade, which includes the Denisovans and is the main part of the sister group to the H. sapiens clade.  Both the H. sapiens and H. longi clades have deep roots extending beyond the Middle Pleistocene and probably experienced rapid early diversification.”  The abstract opens:  "Diverse forms of Homo coexisted during the Middle Pleistocene.  Whether these fossil humans represent different species or clades is debated." ↩ ↩

2. 30b.

   Page 1320:  “Given its geological age of 0.94 to 1.10 million years (Ma) and the presence of mosaic primitive and derived features, we argue that Yunxian 2 probably lies close to the last common ancestor of the clades containing H. sapiens and H. longi”.  The age is taken from earlier dating work, which the paper cites and does not repeat. ↩

3. 30c.

   Page 1321:  “Yunxian 2 has a moderate endocranial capacity of about 1143 cm3, similar to Dali, Hualongdong, Arago 21, Petralona, and Ceprano.”  The paper says the site has yielded three crania, the third still unprepared. ↩

4. 30d.

   Page 1323:  “Previous estimates of the divergence time between Neanderthals and H. sapiens are about 0.5 to 0.7 Ma.  However, recent evolutionary genetic studies based on a large number of modern and ancient genomes of H. sapiens, Neanderthals, and Denisovans have suggested a much deeper ancestry and population divergence in H. sapiens. … The divergence between the longi clade and the sapiens clade is at about 1.32 Ma.  The monophyletic Neanderthal clade, widely thought to be sister to H. sapiens, diverged from the longi and sapiens clades at about 1.38 Ma in our analysis.” ↩ ↩

5. 30e.

   Page 1322:  “most Asian Chibanian hominins, previously often referred to as “archaic Homo sapiens,” including Dali, Jinniushan, Xujiayao, and Hualongdong, are grouped together with the Xiahe and Penghu mandibles to form the monophyletic longi clade.  This clade, plus H. antecessor, is the sister clade to the monophyletic sapiens clade.  Recently, Xujiayao, Xuchang, Xiahe, Penghu, and Denisovans were proposed as a new species of Homo.  Our phylogenetic analysis shows that these Homo samples (except for Xuchang) instead belong to the monophyletic longi clade.”  The "new species" is Bae and Wu's Homo juluensis;  their 2024 Comment is the reference given. ↩ ↩ ↩ ↩ ↩

6. 30f.

   Pages 1322–1323:  “Analyses of mitochondrial DNA place Denisovans with the Sima de los Huesos fossils and outside the divergence between H. sapiens and Neanderthals, whereas nuclear genome sequences suggest that Denisovans are a sister group to Neanderthals.  With only three clades (sapiens, Neanderthals, and Denisovans), both possible phylogenetic relationships, as reflected in the form of a tree topology (Fig. 4), are equally logical and depend on the choice of the rooting point of the tree”. ↩

Ni, Xijun et al.  “Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage.”  The Innovation 2(3):100130.  2021.  The companion paper to the naming of Homo longi:  a parsimony and Bayesian analysis of 55 fossil skulls scored individually, with the matrix in its supplement.  Quotations are located by section.  Mirror:  PMC

1. 31a.

   Section Results and discussion:  “it probably represents a male individual”.  “the ectocranial suture composite scores would suggest an old adult around 50 years old.  However, the tooth wear seems to suggest a younger age.” ↩

2. 31b.

   Section Phylogenetic position of the Harbin cranium:  “When backbone constraints are used to reflect the results from palaeoproteomic and ancient DNA research by forcing the Xiahe mandible as the sister group of Neanderthals and H. antecessor outside of the H. sapiens-Neanderthal clade, Chinese late Middle Pleistocene humans, including the Harbin cranium, form a monophyletic clade as the sister group of Neanderthals (Figure S20).” ↩

3. 31c.

   Section Phylogenetic position of the Harbin cranium:  “The Neanderthal-H. sapiens divergence time in our analysis was 1,007 ka (1,114–919 ka).  This estimation falls in the range based on mtDNAs for the split between the basal Neanderthal (Sima de los Huesos) and the H. sapiens lineage, but is much older than the estimation based on nuclear DNAs for the splits between the Neanderthal and H. sapiens lineages.  However, it is possible that this younger estimated divergence date is an artifact of statistical averaging between “super-archaic” and “recent gene flow” events.” ↩ ↩

4. 31d.

   Section Phylogenetic position of the Harbin cranium:  “The Harbin cranium and Xiahe mandible form a sister group, and they, plus the Dali, Hualongdong, Jinniushan specimens, the European H. antecessor partial cranium, the African Eliye Springs cranium, and Rabat palate, form a monophyletic group.  This clade forms the sister group of the similarly monophyletic H. sapiens clade.  The specimens traditionally grouped in H. heidelbergensis/H. rhodesiensis do not constitute a monophyletic group and the Asian and African H. erectus specimens similarly form a paraphyletic group.” ↩ ↩

Shao, Qingfeng et al.  “Geochemical provenancing and direct dating of the Harbin archaic human cranium.”  The Innovation 2(3):100131.  2021.  Matches the sediment inside the Harbin skull to a lake bed by X-ray fluorescence, rare-earth elements and strontium isotopes, and runs uranium-series on the bone, which gave apparent ages scattered from 62 ± 3 to 148 ± 2 ka;  the oldest is taken as a minimum.  Mirror:  PMC

1. 32a.

   Section Introduction:  “The Harbin cranium was reportedly discovered in 1933 during construction work when a bridge (Dongjiang Bridge) was built over the Songhua River in Harbin City in Northeastern China (Figure 1).  Because of a long, difficult, and confused history since the discovery, the information about the exact geographic origin and stratigraphical context of the cranium has been lost, impeding its accurate dating.”  The naming paper by Ji and colleagues adds that the skull was donated to Hebei GEO University in 2018. ↩

Tu, Hua et al.  “The oldest in situ Homo erectus crania in eastern Asia:  The Yunxian site dates to ~1.77 Ma.”  Science Advances 12(8):eady2270.  Feb 20, 2026.  A cosmogenic aluminium-beryllium burial dating study of the gravels in the layer that yielded the Yunxian skulls in central China.  Nine of ten quartz clasts fall on one line and give about 1.77 million years, against the 0.8 to 1.1 million years of earlier palaeomagnetic and tooth dates.  If it holds, the large Yunxian 2 skull is the oldest large brain known.  Mirror:  PMC

1. 33a.

   Section Results:  “For the sample set C3, 9 of the 10 26Al and 10Be concentrations yield an isochron age of 1.77 ± 0.08 Ma”.  Section Discussion:  “The set C3 was taken from the layer where the hominin crania were unearthed”.  Section Introduction:  “Given this range of dates, Yunxian has been widely cited as 0.8 to 1.0 Ma”.  The total uncertainty is ±0.11 million years;  the dated section lies about 75 m from where the first two skulls were found. ↩ ↩

Wu, Xiu-Jie et al.  “Archaic human remains from Hualongdong, China, and Middle Pleistocene human continuity and variation.”  PNAS 116(20):9820–9824.  2019.  Description of the adolescent skull and jaw HLD 6 from a collapsed cave in Anhui, dated 275 to 331 ka by uranium-series on the breccia.  Mirror:  PMC

1. 34a.

   Page 9820, section Abstract:  “Newly found ∼300,000-y-old human remains from Hualongdong (HLD), China, including a largely complete skull (HLD 6), share East Asian Middle Pleistocene (MPl) human traits of a low vault with a frontal keel (but no parietal sagittal keel or angular torus), a low and wide nasal aperture, a pronounced supraorbital torus (especially medially), a nonlevel nasal floor, and small or absent third molars.” ↩

2. 34b.

   Page 9821, section The HLD 6 Cranium and Mandible:  “The estimated endocranial capacity (∼1,150 cm3;  SI Appendix, section 5) is unexceptional for its age and context [global MPl, 1,179 ± 146 cm3, n = 49;  east Asia MPl (eAMPl), 1,109 ± 131 cm3, n = 18”.  MPl is Middle Pleistocene.  The skull is an adolescent's, 13 to 15 years by its teeth. ↩

Bae, Christopher J., and Xiujie Wu.  “Making sense of eastern Asian Late Quaternary hominin variability.”  Nature Communications 15:9479.  Nov 2, 2024.  A three-page Comment by two of the four authors of Homo bodoensis.  It sorts the later Asian fossils into four species, including the newly named Homo juluensis, to which it assigns the Denisova fossils on teeth and jaws.  Mirror:  PMC

1. 35a.

   Page 2:  “The Hualongdong fossils date to the late Middle Pleistocene (~300,000 years BP) and display a mosaic of characteristics that cannot be easily fitted into any one lineage.  In this case, Hualongdong does not fit neatly into H. juluensis or H. longi and certainly not into H. floresiensis or H. luzonensis”. ↩ ↩

2. 35b.

   Page 1:  “Importantly, we have assigned the enigmatic Denisova, along with the Xiahe and Penghu fossils, to H. juluensis based on comparative study of the present dentognathic remains.” ↩

3. 35c.

   Page 1:  “Since the 1920s with the formal introduction of Sinanthropus pekinensis … and the formal designation of Homo longi in 2021, all (>100 sites) Late Quaternary hominin fossils in China were considered to either represent H. erectus, a transitional archaic version of H. sapiens, or modern H. sapiens.” ↩

Sarmiento, Esteban E., and Martin Pickford.  “Muddying the muddle in the middle even more.”  Evolutionary Anthropology 31(5):237–239.  2022.  The second published rejection of Homo bodoensis, harsher than Delson and Stringer's, in the same issue. ↩

1. 36a.

   Section Abstract:  “Roksandic et al. are unable to account for some European fossils (i.e., Petralona and Arago) and provide no evidence as to how their new species meets their objectives.  Fatally, they overlook the priority rule and fail to realize that H. bodoensis is both a junior synonym of Homo rhodesiensis and Homo saldanensis.  Roksandic et al. conflate taxonomy with phylogeny, present hypotheses as facts, and harbor many systematic and evolutionary misconceptions.” ↩

Lubenow, Marvin L.  Bones of Contention:  A Creationist Assessment of Human Fossils.  Baker Books, Grand Rapids.  revised and updated edition.  2004.  The standard young-earth treatment of the human fossil record.  Chapter 16, "Rhodesian Man:  The Fastest-Aging Man in the World", pp. 159–164, covers Kabwe;  the Sima de los Huesos discussion is at pp. 199–203.

1. 37a.

   Page 199:  “Recently, a more respectful category was devised for these European fossils:  Homo heidelbergensis, named after Heidelberg Man.  However, this did nothing to solve the problem of their relationship to Homo erectus or to the Neandertals.”  “Michael Day, looking for that transition from Homo erectus to modern humans, referred to these fossils as “the muddle in the middle.”” ↩

Rightmire, G. Philip.  “Brain size and encephalization in early to Mid-Pleistocene Homo.”  American Journal of Physical Anthropology 124(2):109–123.  Jun 2004.  The measured case for Homo heidelbergensis as a real species, by its main defender:  thirty Homo erectus skulls averaging 973 cm³ against ten Middle Pleistocene skulls averaging 1,206 cm³, with the increase holding after correction for body size.

1. 38a.

   Page 109, section Abstract:  “Ten of these mid-Quaternary hominins have brains averaging 1,206 cm(3) in volume, and many fall beyond the limits of size predicted for Homo erectus of equivalent age.”  Read in abstract only.  The abstract names the sites as "such as Arago Cave in France, the Sima de los Huesos in Spain, Bodo in Ethiopia, Broken Hill in Zambia, and perhaps Dali in China" and does not list the ten. ↩

2. 38b.

   Page 109, section Abstract:  “These changes in absolute and relative brain size can be taken as further corroborative evidence for a speciation event, in which Homo erectus produced a daughter lineage.  It is probable that Homo heidelbergensis originated in Africa or western Eurasia and then ranged widely across the Old World.” ↩

Roksandic, Mirjana et al.  “Homo bodoensis and why it matters.”  Evolutionary Anthropology 31(5):240–244.  2022.  The authors' reply to Delson and Stringer and to Sarmiento and Pickford, in the same issue.  They do not withdraw the name.

1. 39a.

   Section Abstract:  “(1) we are encouraged that the primary proposal in our paper, to discontinue the use of H. heidelbergensis (as a junior synonym to Homo neanderthalensis) due to its' nomenclatural problems, is acknowledged.  (2) we provide additional clarification about the rules governing taxonomic nomenclature as outlined by the International Code of Zoological Nomenclature and join the growing calls for a revision to these rules.  (3) we discuss further why H. rhodesiensis should be abandoned, particularly in light of the current sensitivity to using culturally inappropriate names.  We conclude that H. bodoensis is a better solution than the proposed alternatives.”  "its'" is the abstract's. ↩

Line, Peter.  “Fossil evidence for alleged apemen—Part 1:  the genus Homo.”  Journal of Creation 19(1):22–32.  Apr 2005.  A survey of the genus Homo for Creation Ministries International by its principal writer on human fossils.  Quotations are located by the section headings of the web posting.

1. 40a.

   Section Homo antecessor:  “Surely room could have been found in the heidelbergensis ‘bushel basket’ to include the Grand Dolina remains.  One suspects, with an evolutionary age (about 0.78 Ma) of the remains that was older than any member of heidelbergensis from Europe, the temptation to give these ‘oldest known Europeans’ a new name was just too much”.  "Grand Dolina" is the source's spelling. ↩

Meyer, Matthias et al.  “A mitochondrial genome sequence of a hominin from Sima de los Huesos.”  Nature 505(7483):403–406.  2014.  The first Sima de los Huesos DNA, a near-complete mitochondrial genome from one femur, from Svante Pääbo's laboratory.

1. 41a.

   Page 403, section Abstract:  “The skeletal remains share a number of morphological features with fossils classified as Homo heidelbergensis and also display distinct Neanderthal-derived traits.  Here we determine an almost complete mitochondrial genome sequence of a hominin from Sima de los Huesos and show that it is closely related to the lineage leading to mitochondrial genomes of Denisovans, an eastern Eurasian sister group to Neanderthals.” ↩

Bonmatí, Alejandro et al.  “Middle Pleistocene lower back and pelvis from an aged human individual from the Sima de los Huesos site, Spain.”  PNAS 107(43):18386–18391.  2010.  Description of Pelvis 1 and associated vertebrae, a man over 45 with a bent lower spine and a slipped vertebra.  The paper still calls the Sima people Homo heidelbergensis.  Mirror:  PMC

1. 42a.

   Page 18386, section Abstract:  “As a result, the individual's daily physical activities would have been restricted to some extent.”  “The presence of degenerative pathological lesions and the advanced age-at-death of this individual make it the most ancient postcranial evidence of an aged individual in the human fossil record.”  The abstract gives his age as over 45 and lists a bent lower spine, a slipped vertebra and Baastrup disease. ↩ ↩

Thomas, Brian.  “Eight Spears Found in German Coal Mine.”  Institute for Creation Research.  Oct 8, 2012.  A news item on the Schöningen spears by the Institute for Creation Research's science writer.

1. 43a.

   Section Article text:  “Such an assemblage of various animals and human remains clearly indicate that a severe Ice Age storm washed them into a low-lying area and quickly covered them with mud.” ↩

Gracia, Ana et al.  “Craniosynostosis in the Middle Pleistocene human Cranium 14 from the Sima de los Huesos, Atapuerca, Spain.”  PNAS 106(16):6573–6578.  2009.  Description of Cranium 14, a child whose left lambdoid suture fused before birth, deforming the skull, and who lived past five.  Mirror:  PMC

1. 44a.

   Page 6573, section Introduction:  “a Middle Pleistocene case of a serious congenital skull deformation that may have required extra conspecific care for the individual to survive for a number of years before he/she died at the end of childhood.”  The paper puts the age at death above 5 to 8 years and below 12.5. ↩

Line, Peter.  “Developments in paleoanthropology no. 2.”  Journal of Creation 36(1):89–98.  Apr 2022.  A review of recent papers, with a section on the Homo bodoensis proposal written before the published replies appeared.  Its Roksandic quotations match the paper.

1. 45a.

   Section Homo bodoensis and the muddle in the middle:  “Whether they are called Homo heidelbergensis, Neanderthals, ‘archaic’ Homo sapiens, Homo bodoensis, or some other name, the fossil specimens assigned to these categories do not represent evolutionary ‘apemen’ stages between Homo erectus and modern Homo sapiens.  Rather, they were fully human, that is, descendants of Adam and Eve, as also were the fossil specimens assigned to Homo erectus.  As such, they belonged to various subgroups of the same species, Homo sapiens, whose individuals were capable of interbreeding with one another.  The few differences were caused by factors such as human variation, genetic drift, as well as environmental influences.”  “For as long as the fossil specimens are interpreted using an incorrect model (evolution), ‘the muddle in the middle’ will remain in a confusing state.” ↩