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The evolutionary history of the human face
Open accessAbstract
The face is the most distinctive feature used to identify others. Modern humans have a short, retracted face beneath a large globular braincase that is distinctively different from that of our closest living relatives. The face is a skeletal complex formed by 14 individual bones that houses parts of the digestive, respiratory, visual and olfactory systems. A key to understanding the origin and evolution of the human face is analysis of the faces of extinct taxa in the hominin clade over the last 6 million years. Yet, as new fossils are recovered and the number of hominin species grows, the question of how and when the modern human face originated remains unclear. By examining key features of the facial skeleton, here we evaluate the evolutionary history of the modern human face in the context of its development, morphology and function, and suggest that its appearance is the result of a combination of biomechanical, physiological and social influences.
Overview
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Written from the paper’s full text.
TL;DR
The human face is a 14-bone skeletal complex whose modern short, retracted form evolved over the last ~4 million years through a mix of biomechanical, physiological, and social influences. The paper reviews fossil, developmental, genetic, and morphometric evidence and argues that a modern human-like face appears by ~300 Ka at Jebel Irhoud, with H. antecessor (~850 Ka) as a key possible ancestral morphology.
Key findings
- The modern human face is relatively small and nonprojecting, lacks pronounced supraorbital and midfacial prognathism, and has a canine fossa; these features do not appear all at once, but crucial elements are present in the earliest H. sapiens lineage, including Jebel Irhoud at ~300 Ka.
- H. antecessor specimen ATD6-69 (~850 Ka) shows a “modern” maxillary conformation, and growth simulations and remodeling analyses support its modern-like facial development; Sima de los Huesos fossils (~430 Ka) share facial features with Neanderthals and are regarded as Neanderthal ancestors.
- Facial evolution in hominins involved modular changes: early Ardipithecus and Australopithecus differ in masticatory structures, with Au. afarensis (~3.7–3.0 Ma) showing expansive zygomatic bones and robust masticatory anatomy, while Ar. ramidus (~4.4 Ma) has lightly built zygomatic bones and arches.
- Modern human facial variation reflects both population history and environmental factors; facial morphology carries a neutral genetic signal but is more strongly affected by climate than other cranial regions, especially in high-latitude populations, and nasal morphology is related to temperature and humidity.
Methods
- Review/synthesis of the hominin facial skeleton over the last ~4 Ma, focusing on australopiths, early Homo, Middle Pleistocene hominins, and H. sapiens, with key specimens listed in Table 1.
- Evidence includes comparative craniofacial morphology, developmental growth-remodeling maps of bone deposition and resorption, ancient DNA and quantitative genetic studies, and a principal components analysis of 19 facial three-dimensional landmarks in Pleistocene fossils and modern humans.
- This is a review; the analyses are drawn from cited studies and previously analyzed morphometric data rather than newly collected primary data.
Limitations
- The paper states that a detailed account of human facial evolution is difficult because of the intricate nature of facial development and the many pre- and post-natal factors influencing the face.
- The paper states that future work must integrate potentially conflicting evidence from craniofacial biomechanics, occlusal microwear, and stable isotopes, and must disentangle shared ancestry, climate adaptation, and subsistence effects.
- Evident from the text: the fossil record is fragmentary and uneven; H. naledi is too incomplete in the face to add much, the Harbin cranium is announced but “not yet published scientifically,” and the identity of the last common ancestor remains unresolved.
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Claims the paper makes
52 Not checked yet · 26 Own interpretation · 13 No basis given · 6 Own result · 1 Supported
Every source the paper cites for this claim was checked and supports it.
In the paper
“Irhoud 1, found in Morocco and recently dated to ~300 Ka 31 , is likely an early member of the Homo sapiens lineage”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
What the sources say
- Supportedcites [31] Hublin, J. J. et al. New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens. Nature 546, 289–292 (2017). source
The Conclusion identifies the Irhoud fossils as evidence of the early phase of Homo sapiens evolution in Africa. Figure 3 specifically includes Irhoud 1 in the anatomical comparisons, supporting the claim that it was an early member of the lineage. (LLM, conf=0.92)
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In the paper
“Over time, the face became more gracile”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
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In the paper
“This is also evidenced in Asian specimens such as Nanjing.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
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In the paper
“since some sexually dimorphic structures (e.g.canines, browridges) reduced in size over time.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
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In the paper
“A striking feature observed in the MP hominins is that the face evolved more rapidly than other cranial components”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
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In the paper
“Figure 5: Morphometric analysis of the face discriminates Pleistocene fossils.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“These quantitative analyses also indicate the morphological proximity of some these fossils to modern humans.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“We here show a principal components analysis of nineteen facial three-dimensional landmarks (illustrated as black points on the faces shown in the left panel) of a number of Middle-Late Pleistocene fossils and a sample of modern humans, revealing clusters of fossils based on facial shape.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“In modern humans, there is a predominance of widely-spread resorptive fields variously spread over the maxilla, infraorbital and anterior zygoma as well as the mental region and coronoid process of the mandible, contributing to facial retraction (orthognathy) (see Fig. 2) 95 .”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
What the sources say
- Not checked yetcites [95] White, T. D. et al. Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature 423, 742 (2003). source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: Growth remodeling can be determined by analysis of the distribution of depository and resorptive fields on bone surfaces due to the activities of osteoblasts and osteoclasts.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“suggest that its appearance is the result of a combination of biomechanical, physiological and social influences”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Abstract
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- Probably not a substantial claim. 27
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In the paper
“In more recent fossil Homo, facial projection decreased further, with a moderate but cumulative increase in brain size.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“as illustrated by the facial characteristics of H. antecessor, a key LCA candidate, presenting modern facial features while retaining some primitive characters elsewhere in the cranium.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
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In the paper
“unlike Neanderthals and other MP crania (Petralona, Bodo, Broken Hill 1, Sima de los Huesos), Irhoud 1 already shows almost completely modern human-like facial morphology.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
plainSingle case- Asserted: passes — The authors put it forward as true.
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In the paper
“Modern humans are distinguished by a flat and anteriorly oriented face, angled zygomatic, and gracile browridge (top left), contrasting with the large browridge and 'puffy' midface of the Neanderthals (bottom left).”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“In prognathic faces such as in chimpanzees or early Australopithecus, this prognathism is characterized by a pattern of bone deposition.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
plaincorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
- Rests on an assumption: Growth remodeling can be determined by analysis of the distribution of depository and resorptive fields on bone surfaces due to the activities of osteoblasts and osteoclasts.
In the paper
“the main contributors to the development of the facial skeleton 2”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Introduction
What the sources say
- Not checked yetcites [2] Cordero, D. R. et al. Cranial neural crest cells on the move: their roles in craniofacial development. Am. J. Med. Genet. A. 155A, 270–279 (2011). source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Facial shape appears to be influenced by a limited set of candidate genes 3”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Introduction
What the sources say
- Not checked yetcites [3] Liu, F. et al. A genome-wide association study identifies five loci influencing facial morphology in Europeans. PLoS Genet. 8, e1002932 (2012). source
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In the paper
“signaling more than 20 different categories of emotion via the contraction or relaxation of muscles 1”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Introduction
What the sources say
- Not checked yetcites [1] Du, S., Tao, Y. & Martinez, A. M. Compound facial expressions of emotion. Proc. Natl Acad. Sci. USA 111, E1454–E1462 (2014). source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“The modern human face is distinct from that of earlier hominin species in several important ways (Fig. 3h): it is relatively small and nonprojecting, shows a depression -the 'canine fossa' -below the orbit and lacks the pronounced supraorbital structures and the alveolar and midfacial prognathism exhibited by MP hominins.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
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In the paper
“a shift towards a strong C4 dietary signature occurred in Homo with the appearance of early African H. erectus (≤ 1.7 Ma), rather than in the earlier species H. habilis and H. rudolfensis, whose modal carbon stable isotope signal is similar to that of Au. afarensis and Au.africanus, suggesting a mixed C3/C4 diet 19”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Patterns of diversity in the early Homo face
What the sources say
- Not checked yetcites [19] Cerling, T. E. et al. Stable isotope-based diet reconstructions of Turkana Basin hominins. Proc. Natl Acad. Sci. USA 110, 10501–10506 (2013). source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“the earliest known stone tools, at 3.3 Ma 26 , coincide in time with Au. afarensis (and the australopith-grade taxon K. platyops of eastern Africa), predating the earliest fossils of Homo by several hundred thousand years”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Patterns of diversity in the early Homo face
What the sources say
- Not checked yetcites [26] Harmand, S. et al. 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature 521, 310–315 (2015). source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“The younger species Au. sediba (~2.0 Ma) from South Africa 23 shares derived facial features with Au. africanus 10 , but its facial remodeling differs from all other Australopithecus facial skeletons (see Box 2, Fig. 2) 15”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Australopithecus (incl. Paranthropus) craniofacial diversity: Patterns and causes
What the sources say
- Not checked yetcites [10] Kimbel, W. H. & Rak, Y. Australopithecus sediba and the emergence of Homo: questionable evidence from the cranium of the juvenile holotype MH 1. J. Hum. Evol. 107, 94–106 (2017). source
- Not checked yetcites [15] Lacruz, R. S. et al. Distinct growth of the nasomaxillary complex in Au. sediba. Sci. Rep. 5, 15175 (2015). source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“using recently published estimates of the autosomal human mutation rate, it has been suggested that the divergence date of the H. neanderthalensis and H. sapiens lineages could indeed be placed earlier -between 550 and 765 Ka 48”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > The LCA, ancient genomes and the earliest Homo sapiens face
What the sources say
- Not checked yetcites [48] Meyer, M. et al. Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins. Nature 531, 504–507 (2016). source
hedgedconditional- Asserted: fails — The authors may not put it forward as true.
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In the paper
“Stringer 32 has argued that shape resemblances between the Petralona and Broken Hill 1 crania indicated the existence of a widespread MP population which, if the Mauer mandible (Germany) is included, would be called H. heidelbergensis.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo heidelbergensis as the LCA?
What the sources say
- Not checked yetcites [32] Stringer, C. B. Some further notes on the morphology and dating of the Petralona hominid. J. Hum. Evol. 12, 731–742 (1983). source
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This morphology is similar to that shown by Florisbad (South Africa) and Herto (Ethiopia) (Fig. 3g) at a somewhat later date, Ndutu and Thomas Quarry at an earlier date, and in the currently undated Eliye Springs (Kenya) and Broken Hill 2 fossils.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > FIGURE 4 HERE
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In the paper
“One implication of a H. heidelbergensis-like LCA would be that the zygomaxillary morphology found in these large MP crania would have undergone gracilisation to become the form found in recent H. sapiens, and a second trajectory, featuring increased midfacial projection and maxillary inflation, that led to the facial configuration found in the Neanderthals.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo heidelbergensis as the LCA?
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The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“the face of early H. erectus is more modern human-like than that of either H. habilis or H. rudolfensis, with a more vertical profile featuring a retracted subnasal plate and a prominent nasal bridge”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Patterns of diversity in the early Homo face
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In the paper
“Thus, sexual dimorphism could also be a factor if Thomas Quarry, Ndutu and Broken Hill 2 represent female individuals, and Petralona, Broken Hill 1 and Bodo much larger males.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo heidelbergensis as the LCA?
hedgedconditional- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“It is possible that allometric factors influence the zygomaxillary morphology when comparing smaller and larger individuals 37,38 given the association between body size and facial size in primates 39”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo heidelbergensis as the LCA?
What the sources say
- Not checked yetcites [37] Maddux, S. D. & Franciscus, R. G. Allometric scaling of infraorbital surface topography in Homo. J. Hum. Evol. 56, 161–174 (2009). source
- Not checked yetcites [38] Freidline, S. E., Gunz, P., Harvati, K. & Hublin, J. J. Evaluating developmental shape changes in Homo antecessor subadult facial morphology. J. Hum. Evol. 65, 404–423 (2013). source
- Not checked yetcites [39] Dobson, S. D. Allometry of facial mobility in anthropoid primates: implications for the evolution of facial expression. Am. J. Phys. Anthropol. 138, 70–81 (2009). source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“The modern appearance of ATD6-69 is also substantiated by growth simulations of this specimen based on both the Neanderthal and modern human facial growth trajectories 38 , and by the analysis of the facial growth and remodeling of ATD6-69, which identified resorptive fields on that maxilla similar to those found in sub-adult H. sapiens 16”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo antecessor as the LCA?
What the sources say
- Not checked yetcites [38] Freidline, S. E., Gunz, P., Harvati, K. & Hublin, J. J. Evaluating developmental shape changes in Homo antecessor subadult facial morphology. J. Hum. Evol. 65, 404–423 (2013). source
- Not checked yetcites [16] Lacruz, R. S. et al. Facial morphogenesis of the earliest europeans. PLoS One 8, e65199 (2013). source
plaincorrelationalSingle case- Asserted: passes — The authors put it forward as true.
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A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These MP hominins are a possible ancestral pool for the emergence of Homo sapiens.Hence, MP taxa could include the last common ancestor (LCA) of H. sapiens and other now extinct taxa.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > The faces of the Middle Pleistocene hominins and the LCA
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A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This model implies that a H. antecessor-like facial morphology was retained in the descendant H. sapiens lineage, but was modified in that of the Neanderthals.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo antecessor as the LCA?
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“it was argued that this species represented the best candidate for the LCA of the H. sapiens and H. neanderthalensis lineages (see also 35 )”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Homo heidelbergensis as the LCA?
What the sources say
- Not checked yetcites [32] Stringer, C. B. Some further notes on the morphology and dating of the Petralona hominid. J. Hum. Evol. 12, 731–742 (1983). source
- Not checked yetcites [35] Rightmire, G. P. Human evolution in the Middle Pleistocene: the role of Homo heidelbergensis. Evol. Anthropol. 6, 218–227 (1998). source
plain- Asserted: passes — The authors put it forward as true.
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- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Adaptive explanations for an enlarged midface and large brows have focused either on enlargement of the nasal cavity and paranasal sinuses emphasizing respiratory/energetic demands 56 and climatic adaptation 57,58 , or on mechanical adaptations to diet, paramasticatory activity 59,60 , or increase in body size [61][62][63][64]”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [56] García-Martínez, D. et al. Ribcage measurements indicate greater lung capacity in Neanderthals and Lower Pleistocene hominins compared to modern humans. Commun. Biol. 1, 117 (2018). source
- Not checked yetcites [57] Churchill, S.E. in Neanderthals Revisited: New Approaches and Perspectives (eds. Havarti, K. & Harrison, T.) 113–133 (Springer, 2006). source
- Not checked yetcites [58] Wroe, S. et al. Computer simulations show that Neanderthal facial morphology represents adaptation to cold and high energy demands, but not heavy biting. Proc. R. Soc. Lond. B 285, 20180085 (2018). source
- Not checked yetcites [59] Rak, Y. The Neanderthal: a new look at an old face. J. Hum. Evol. 15, 151–164 (1986). source
- Not checked yetcites [60] Demes, B. Another look at an old face: biomechanics of the Neandertal facial skeleton reconsidered. J. Hum. Evol. 16, 297–303 (1987). source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“An alternative would be to consider a H. antecessorlike facial morphology as more likely for the LCA of H. sapiens and H. heidelbergensis, with the H. heidelbergensis group exemplified by Petralona, Bodo and Broken Hill 1 having more in common facially with the Sima de los Huesos fossils and subsequent Neanderthals.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > The LCA, ancient genomes and the earliest Homo sapiens face
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The evidence instead suggests that the large and non-H.sapiens-like faces of Bodo and Broken Hill 1 (Figs.3b & c), represent taxonomic diversity in the African MP record, which could exclude the large fossils assigned to H. heidelbergensis/H.rhodesiensis as representing an ancestral morph for H. sapiens.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > FIGURE 4 HERE
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The initial changes were focused in the central and upper portions of the face, perhaps in response to changes in the function of the canines and supraorbital region in social signaling, whereas subsequent changes mainly involved the masticatory apparatus.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Ardipithecus vs. Australopithecus: Principal morphological shifts -canines vs. mastication
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“These derived features have long been interpreted as adaptations to mechanically tough or hard plant foods that dominated in African environments during a period of aridification in the late Pliocene (≤ 2.8-2.7 Ma) 20,21”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Australopithecus (incl. Paranthropus) craniofacial diversity: Patterns and causes
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“while fully fledged H. sapiens likely had an African beginning, the evolutionary origins of traits characteristic of the modern face, represented by the LCA, may be found elsewhere”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > The faces of the Middle Pleistocene hominins and the LCA
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“both the external nasal morphology and nasal cavity are related to temperature and humidity 80,82,85”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Population history and climatic adaptation shape the face
What the sources say
- Not checked yetcites [80] von Cramon-Taubadel, N. Evolutionary insights into global patterns of human cranial diversity: population history, climatic and dietary effects. J. Anthropol. Sci. 92, 43–77 (2014).
- Not checked yetcites [82] Hubbe, M., Hanihara, T. & Harvati, K. Climate signatures in the morphological differentiation of worldwide modern human populations. Anat. Rec. (Hoboken) 292, 1720–1733 (2009). source
- Not checked yetcites [85] Noback, M. L., Harvati, K. & Spoor, F. Climate-related variation of the human nasal cavity. Am. J. Phys. Anthropol. 145, 599–614 (2011). source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“midfacial reduction may have arisen simply as a result of loss of pre-existing selective pressures to maintain a large midface, with cultural adaptations to climate, feeding and lifestyle being possible factors in reducing these pressures and so allowing drift and other neutral processes to impact on midfacial form”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“modern humans have a high degree of cranial variation exceeding that found in other primate species, with the face being an important contributor to this variation 75,76”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Population history and climatic adaptation shape the face
What the sources say
- Not checked yetcites [75] Harvati, K. The Neanderthal taxonomic position: models of intra- and inter-specific craniofacial variation. J. Hum. Evol. 44, 107–132 (2003). source
- Not checked yetcites [76] Harvati, K., Frost, S. R. & McNulty, K. P. Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences. Proc. Natl Acad. Sci. USA 101, 1147–1152 (2004). source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“it has also been argued, but is less likely, that they play a role in resisting loading of the jaws (masticatory loading hypothesis) 68”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [68] Russell, M. D. et al. The supraorbital torus: "A Most Remarkable Peculiarity". Curr. Anthropol. 26, 337–360 (1985). source
hedgedconditional- Asserted: fails — The authors may not put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it.
In the paper
“While brow ridges have been considered to arise as a structural consequence of fitting a large face under a retracted frontal (spatial hypothesis) 67”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [67] Moss, M. L. & Young, R. W. A functional approach to craniology. Am. J. Phys. Anthropol. 18, 281–292 (1960). source
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The earliest changes during the australopith to early Homo transition may have been driven, in part, by abiotic environmental factors but also by the social context”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“it has been argued 71 that H. sapiens facial skeletons show increased bite force relative to MP hominins 72,73 , which could be an advantage in fracturing hard objects”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [71] Wroe, S., Ferrara, T. L., McHenry, C. R., Curnoe, D. & Chamoli, U. The craniomandibular mechanics of being human. Proc. R. Soc. Lond. B 277, 3579–3586 (2010). source
- Not checked yetcites [72] Anton, S. C. Neandertals and the anterior dental loading hypothesis: a biomechanical evaluation of bite force production. Kroeber Anthropological Society Papers 71–72, 67–76 (1990).
- Not checked yetcites [73] O’Connor, C. F., Franciscus, R. G. & Holton, N. E. Bite force production capability and efficiency in Neandertals and modern humans. Am. J. Phys. Anthropol. 127, 129–151 (2005). source
hedgedconditional- Asserted: fails — The authors may not put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“While the modern human cranium, overall, appears to be mainly shaped by neutral evolutionary processes [80][81][82] , the modern human face reflects both phylogenetic and environmental factors [81][82][83][84]”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Population history and climatic adaptation shape the face
What the sources say
- Not checked yetcites [80] von Cramon-Taubadel, N. Evolutionary insights into global patterns of human cranial diversity: population history, climatic and dietary effects. J. Anthropol. Sci. 92, 43–77 (2014).
- Not checked yetcites [81] Harvati, K. & Weaver, T. D. Human cranial anatomy and the differential preservation of population history and climate signatures. Anat. Rec. A Discov. Mol. Cell. Evol. Biol. 288, 1225–1233 (2006). source
- Not checked yetcites [82] Hubbe, M., Hanihara, T. & Harvati, K. Climate signatures in the morphological differentiation of worldwide modern human populations. Anat. Rec. (Hoboken) 292, 1720–1733 (2009). source
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“while facial morphology does carry a neutral genetic or population history signal, it is more strongly affected by climatic conditions than other parts of the cranium, especially in high latitude populations 81,82,83”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Population history and climatic adaptation shape the face
What the sources say
- Not checked yetcites [81] Harvati, K. & Weaver, T. D. Human cranial anatomy and the differential preservation of population history and climate signatures. Anat. Rec. A Discov. Mol. Cell. Evol. Biol. 288, 1225–1233 (2006). source
- Not checked yetcites [82] Hubbe, M., Hanihara, T. & Harvati, K. Climate signatures in the morphological differentiation of worldwide modern human populations. Anat. Rec. (Hoboken) 292, 1720–1733 (2009). source
- Not checked yetcites [83] Reyes-Centeno, H., Ghirotto, S. & Harvati, K. Genomic validation of the differential preservation of population history in the human cranium. Am. J. Phys. Anthropol. 162, 170–179 (2017). source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“In addition, the size of the jaws is restricted by the housing of teeth.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“It is also important to recognize the role of gene flow associated with migrations across the globe, which will likely affect the pace of evolutionary change.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
In the paper
“because H. sapiens shows decreased ability to resist masticatory loading despite increased bite force, the increased bite force may simply be a by-product of facial reduction, driven by other factors 74”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [74] Godinho, R. M. et al. The biting performance of Homo sapiens and Homo heidelbergensis. J. Hum. Evol. 118, 56–71 (2018). source
hedgedconditional- Asserted: fails — The authors may not put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Integration, or covariation, among morphological units can be quantitated, enabling the development of mathematical models that predict how changes of individual units may occur in response to changes in other units 92 .”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 1. Modularity and Integration of the Craniofacial Complex:
What the sources say
- Not checked yetcites [92] Conroy, G. C., Jolly, C. J., Cramer, D. & Kalb, J. E. Newly discovered fossil hominid skull from the Afar depression, Ethiopia. Nature 276, 67 (1978). source
plainconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“the large midface, chinless jaws and enlarged brow ridges of MP hominins make up a suite of features adapted to masticatory or paramasticatory uses 59,60,65,66”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
What the sources say
- Not checked yetcites [59] Rak, Y. The Neanderthal: a new look at an old face. J. Hum. Evol. 15, 151–164 (1986). source
- Not checked yetcites [60] Demes, B. Another look at an old face: biomechanics of the Neandertal facial skeleton reconsidered. J. Hum. Evol. 16, 297–303 (1987). source
- Not checked yetcites [65] Gröning, F., Liu, J., Fagan, M. J. & O’Higgins, P. Why do humans have chins? Testing the mechanical significance of modern human symphyseal morphology with finite element analysis. Am. J. Phys. Anthropol. 144, 593–606 (2011). source
- Not checked yetcites [66] Pampush, J. D. & Daegling, D. J. The enduring puzzle of the human chin. Evol. Anthropol. 25, 20–35 (2016). source
plaindirect- Asserted: fails — The authors may not put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The authors may not put it forward as true. The paper’s conclusions may not depend on it.
In the paper
“such a shift in eyebrow mobility could have enabled a wider range of subtler social signals, enhancing social communication.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > A cultural/social component for the evolution of the face?
What the sources say
- Not checked yetcites [87] Godinho, R. M., Spikins, P. & O’Higgins, P. Supraorbital morphology and social dynamics in human evolution. Nat. Ecol. Evol. 2, 956–961 (2018). source
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
- Rests on an assumption: Brow ridge reduction accompanied midfacial reduction.
None of the sources cited for this claim have been checked yet.
In the paper
“One aspect of this organization in the craniofacial skeleton is the presence of bone growth centers 22,55 , which are situated to optimize the organism's physiological requirements through the actions of integrating factors over time.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 1. Modularity and Integration of the Craniofacial Complex:
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“The mammalian skull is developmentally complex and a highly integrated structure 91 .”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 1. Modularity and Integration of the Craniofacial Complex:
What the sources say
- Not checked yetcites [91] Wood, B.A. & Boyle, E.K. Hominin taxic diversity: fact or fantasy? Am. J. Phys. Anthropol. 159, S37–S78 (2016). source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“the modern human face is more sculpted and remodeled, has more complex topography than any other hominin face 16”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > A cultural/social component for the evolution of the face?
What the sources say
- Not checked yetcites [16] Lacruz, R. S. et al. Facial morphogenesis of the earliest europeans. PLoS One 8, e65199 (2013). source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Compared to modern humans, the region that most differs is the nasal cavity itself, suggesting reduction of this is the primary underlying cause of midfacial reduction in modern humans.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Adaptation in MP and modern human faces
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Growth remodeling (bone formation and bone resorption) is an integral process of craniofacial growth that relates to maintaining the shape and proportions of the face during development 94 .”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
What the sources say
- Not checked yetcites [94] Stringer, C. & Galway-Witham, J. Palaeoanthropology: on the origin of our species. Nature 546, 212 (2017). source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“potentially gaining an ability to generate more diverse facial expressions, likely enhancing non-verbal communication.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These new faces continued to evolve during challenges by their environment, impacted increasingly by culture and social factors.”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Conclusion
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds. It may state two separate claims as one.
The paper relies on this without support.
The human and chimpanzee/bonobo lineages shared a most recent common ancestor.
“That the human and chimpanzee/bonobo lineages shared a most recent common ancestor is beyond dispute”
The paper relies on this without support.
The last common ancestor of H. sapiens and other extinct taxa should predate the appearance of a modern face.
“the LCA should predate the appearance of a modern face”
The paper relies on this without support.
Brow ridge reduction accompanied midfacial reduction.
“It is of interest in this regard that brow ridge reduction accompanied midfacial reduction.”
The paper relies on this without support.
Biomolecular and mechanical signals trigger coordinated bone forming and bone resorbing activity during growth.
“Such factors include biomolecular and mechanical signals that trigger coordinated bone forming and bone resorbing activity during growth (see Box 2).”
The paper relies on this without support.
Growth remodeling can be determined by analysis of the distribution of depository and resorptive fields on bone surfaces due to the activities of osteoblasts and osteoclasts.
“Growth remodeling can be determined by analysis of the distribution of depository and resorptive fields on bone surfaces due to the activities of osteoblasts and osteoclasts, respectively.”
The paper relies on this without support.
Such developmental signatures are best characterized in sub-adult individuals.
“Such developmental signatures are best characterized in sub-adult individuals.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The cited source says what the claim attributes to it.
In the citing paper
“given the evidence that the Neanderthal face also displays derived characteristics (42)”
From Dosage analysis of the 7q11.23 Williams region identifies BAZ1B as a major human gene patterning the modern human face and underlying self-domestication (2019) · cites this paper as [42]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > The faces of the Middle Pleistocene hominins and the LCA
“Broken Hill 1 (Zambia) (Fig. 3c)”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
“b) Broken Hill 1 (Zambia) dated to ~250-300 Ka.”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
“Nanjing H. erectus?~500 Ka”
§ THE EVOLUTIONARY HISTORY OF THE HUMAN FACE > Box 2. Facial Growth and Remodeling of the Hominin Face:
“d) Nanjing, China, dated to ~400 Ka.”
Citations
Cites
New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens.
Hublin, J.-J., Ben-Ncer, A., Bailey, S. E., Freidline, S. E., Neubauer, S., Skinner, M. M., Bergmann, I., Le Cabec, A., Benazzi, S., Harvati, K., et al. (2017) · Nature · doi:10.1038/nature22336 · Cited by 1,347
Self-citationPlio-Pleistocene African Climate
deMenocal, P. B. (1995) · Science · doi:10.1126/science.270.5233.53 · Cited by 1,241
3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya
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Mid-face Morphometrics and Sexual Dimorphism
Adefisan, I. E., Iboro, E. E., & Adebori, O. D. (2024) · Journal of bioanthropology · doi:10.54062/jb.4.3.1
Olsen, S. T., & White, S. (2025) · Journal of Human Evolution · doi:10.1016/j.jhevol.2024.103645
Metadata
- Authors
- Lacruz, R. S., Stringer, C. B., Kimbel, W. H., Wood, B., Harvati, K., O’Higgins, P., Bromage, T. G., & Arsuaga, J.-L.
- Affiliations
- New York Consortium in Evolutionary Primatology (US)
- Natural History Museum (GB)
- Arizona State University (US)
- George Washington University (US)
- Senckenberg Centre for Human Evolution and Palaeoenvironment (DE)
- University of York (GB)
- New York University (US)
- Corresponding author
- Lacruz, Rodrigo S.
- ORCID
- Venue
- Nature Ecology & Evolution
- Published
- 2019-04-15
- Volume
- 3
- Issue
- 5
- Pages
- 726-736
- ISSN
- 2397-334X (ISSN-L)
- OpenAlex
- W2936204499
- Cited by
- 108
- References
- 101
- Citations by year
- 2026: 6 · 2025: 16 · 2024: 17 · 2023: 17 · 2022: 17 · 2021: 19 · 2020: 11 · 2019: 5
- FWCI
- 16.67
- Type
- journal article
- Language
- en
- Open access
- Licence
- Locations
- Nature Ecology & Evolution (publishedVersion)
- PubMed (publishedVersion)
- White Rose Research Online (University of Leeds, The University of Sheffield, University of York) (submittedVersion)
- Docta Complutense (Universidad Complutense de Madrid) (submittedVersion)
- Zenodo (CERN European Organization for Nuclear Research) (submittedVersion, cc-by-nc-nd)
- Funders
- Fundación Ramón Areces
- Deutsche Forschungsgemeinschaft (DFG‐FOR 2237, FOR2237)
- Topics
- Social Sciences › Anthropology › Pleistocene-Era Hominins and Archaeology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Morphological variations and asymmetry
- Social Sciences › Psychology › Experimental and Cognitive Psychology › Evolutionary Psychology and Human Behavior
- Keywords
- facial skeleton, fossil hominins, craniofacial morphology, facial growth, human origins
- MeSH terms
- Adaptation, Biological, Animals, Biological Evolution, Face, Humans, Hominidae, Genome, Human
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