Charles Loring Brace IV was born on 19 December 1930 in Hanover, New Hampshire, into an illustrious family that advocated liberal and humanitarian principles. His great-great-grandfather, John Pierce Brace, was headmaster of Litchfield Female Academy, one of the most important schools for girls in the United States, and he later became the principal at the Hartford Female Seminary in Connecticut. His great-grandfather, Charles Loring Brace, was a prominent philanthropist, abolitionist, and social reformer who founded the Children’s Aid Society. He wrote books on social issues and corresponded with Charles Darwin about his theory of evolution. Brace’s father, Gerald Warner Brace, was a professor of English at Boston University and an author who wrote eleven novels portraying the seafaring culture of the eastern United States as well as academic life. Brace’s mother, Hulda Potter Laird, had an MA degree in biology and she was responsible for sparking her son’s interest in human paleontology by introducing him to Roy Chapman Andrews’ book Meet Your Ancestors (1945). This inspired Brace to read Charles Darwin’s On the Origin of Species when he was a student in high school.

Brace attended Williams College, a liberal arts college in Massachusetts, from 1948 to 1952. He was interested in studying human evolution, but the college did not have an anthropology department, so Brace constructed his own curriculum studying geology, vertebrate paleontology, and biology. After graduating with a BA in geology, Brace began graduate studies in anthropology at Harvard University in 1952. Initially he studied under the influential physical anthropologist Earnest Albert Hooton, but following Hooton’s death in May 1954 Brace continued his studies under William White Howells. Howells was known for using new techniques such as multivariate statistical analysis to study human populations, which convinced him that all living human populations represent a single homogeneous species, while the Neanderthals represented a quite distinct species. He was also known for his ideas about human evolution, which included criticism of Aleš Hrdlička’s Neanderthal phase hypothesis as well as the multiregional continuity model supported by Franz Weidenreich, which argued that geographically distinct populations of Homo erectus had evolved throughout the Old World into the various races of modern Homo sapiens.

By this time Brace was already working on metric and morphological studies of ape and human teeth with Elihu Leon Schuman, professor in the Harvard School of Dental Medicine (Schuman and Brace 1954). This influenced Brace’s early work on hominid teeth. Brace’s studies were interrupted when he was drafted by the U.S. Army during the Korean War where he served from 1954 to 1956. While in Korea he used his statistical skills to design a one-size-fits-all gas mask that would be able to fit a variety of different people. After the war, Howells encouraged Brace to visit The Jackson Laboratory, a biomedical research institution located in Maine, in order to analyze data relating to the relationship between the physical characteristics, physiology, behavior, and genetics of dogs raised at the laboratory. After returning to Harvard, Brace was influenced by Howells and others who were integrating the Modern Evolutionary Synthesis into paleoanthropology. Howells’s use of new multivariate statistical techniques, including factor analysis, influenced Brace’s research methods. Indeed, Brace utilized Howells’ measurements and analytical methods in much of his subsequent work.

Brace received a Sheldon Traveling Fellowship, which allowed him to travel to Europe in 1959 and 1960. There he participated in excavations conducted by the Harvard University Paleolithic archaeologist Hallam Movius at Abri Pataud in France. Brace also visited French archaeologist François Bordes’s excavations at Combe Grenal, examined the Krapina Neanderthal fossils in the museum in Zagreb (Croatia), and briefly worked in the animal behavior laboratory of Dutch biologist and ornithologist Niko Tinbergen at Oxford University. Brace obtained a master’s degree in anthropology in 1958 and completed his PhD in 1962 with a dissertation titled Physique, Physiology and Behavior: An Attempt to Analyse a Part of their Roles in the Canine Biogram. In 1957, Brace married Mary (Mimi) Louise Crozier, who was the daughter of Harvard physiologist William John Crozier. They collaborated on many of Brace’s published works, and she illustrated nearly all of his scholarly works, notably the Atlas of Human Evolution (Brace et al., 1979). Brace spent one year teaching at the University of Wisconsin-Milwaukee, during the 1960–61 academic year and he also taught courses at the University of Colorado during the summer session in 1964. He was hired as a professor of anthropology at the University of California at Santa Barbara where he taught from 1961 to 1967. Brace then took a position as professor of anthropology at the University of Michigan in 1967, where he also served as Curator of Biological Anthropology at the university’s Museum of Anthropology. Brace held these positions until he retired in 2008.

During the course of his career, Brace worked on a variety of topics including the concept of race in anthropology, evolutionary theory and the process of human evolution, the place of the Neanderthals in human evolution, and other anthropological questions. During the time that Brace was a graduate student at Harvard, anthropology and paleoanthropology were undergoing significant changes. Ernst Mayr, Theodosius Dobzhansky, George Gaylord Simpson, and others were arguing that paleoanthropologists needed to integrate the Modern Evolutionary Synthesis into the study of human evolution. Meanwhile, Sherwood Washburn was promoting the New Physical Anthropology (Washburn 1951; 1953). Both ideas rejected the typological conception of race that was prominent in nineteenth and early twentieth-century racial anthropology. Brace was very influenced by these ideas and when he arrived at the University of Michigan, Brace joined his colleague Frank B. Livingstone, an anthropologist who studied genetic variation in human populations, in promoting the view that biological human races do not exist (see Livingstone 1962). Brace also embraced the arguments made by the British-American anthropologist Ashley Montagu that “there are no races, only clines” (Brace 1964b; Brace and Montagu 1964); Brace and Montagu 1965). The word ‘cline’ derives from the Greek word for slope. When applied to anthropological studies of human variation it refers to a gradient in the expression of a trait across geographical regions.

Throughout his career, Brace was a persistent critic of the concept of biological race, arguing that human features varied in a clinal pattern. This meant that supposed racial characteristics varied gradually across geographic regions, that anatomical features did not vary together, and that human traits were not consistent within so-called ‘races.’ Brace demonstrated that gradients of adaptive features such as skin color and nasal form are independent of each other. Thus, these features should not be used to classify human populations. He also showed that other kinds of features that do cluster in geographical regions originated and became prevalent by chance, due to random genetic drift, and do not possess adaptive significance. Brace also vigorously opposed the idea that race and IQ were associated traits. Brace believed that intelligence was the only human trait that is clearly adaptive but lacks the kind of clinal distribution that is seen in physical traits such as skin color. Thus, intelligence is similar in all human groups because the selective forces that shaped it were essentially identical during the approximately two million years that the genus Homo has existed. Brace also urged that terms introduced by anthropologists to designate human races should be abandoned and instead terms derived from geography should be used to refer to regional populations.

Brace was interested in the Neanderthals from the very beginning of his scientific career. Paleoanthropologists had been debating the role of the Neanderthals in human evolution and their phylogenetic relationship to modern humans since they were first discovered in 1856. Many considered the Neanderthals to be an extinct side branch of human evolution and rejected the notion that they were the direct ancestor of modern humans. Brace’s ideas about human evolution were influenced by the German paleoanthropologist Franz Weidenreich, who had studied the Peking Man (Homo erectus) fossils excavated from the site of Zhoukoudian in China. Weidenreich had also proposed a polycentric theory of human evolution, arguing that several racially distinct geographical populations of hominids throughout the Old World had evolved from Homo erectus into Homo sapiens, yet at any one time these populations remained part of a single species due to the small amounts of genetic exchange between these geographically separated populations. Brace was also familiar with Aleš Hrdlička’s theory of the Neanderthal phase of human evolution, which strongly influenced his interpretation of the Neanderthals. Beginning in the early 1960s, Brace began publishing papers rethinking the scientific interpretation of the Neanderthals.

Brace (1962a) argued that there is nothing in the skeletal morphology of Neanderthals to exclude them from being the direct ancestors of modern Homo sapiens. This paper did not shift the thinking of paleoanthropologists in the way that Brace thought it would so two years later he wrote a more forceful paper titled “The Fate of ‘Classic’ Neanderthals.” In this influential paper Brace argued that the failure of paleoanthropologists to accept the Neanderthals, as well as other early hominid species, as part of the human lineage was the result of the persistence of antievolutionary views in the discipline, which was influenced by a Neoplatonic essentialist mindset that encouraged researchers to accept ancestors only if they differed very little or not at all from their descendants (Brace 1964a). While visiting François Bordes’s excavations at Combe Grenal, a site inhabited during the Mousterian and Acheulean periods where Neanderthal fossils were found, Brace had noticed a rock-lined fire pit and concluded that Neanderthals had used earth ovens to thaw and cook meat. He realized that cooking food would reduce the selective pressure maintaining large tooth size, leading over time to the reduction in size of the teeth and jaws. In a paper titled “Cultural Factors in the Evolution of the Human Dentition,” Brace proposed that the distinctive wear patterns visible on many Neanderthal incisors indicated that they used their teeth as tools, which in turn affected the morphology of the Neanderthal face (Brace 1962b). This way of interpreting fossils was influenced by Sherwood Washburn’s New Physical Anthropology.

In these early papers, Brace rejected the argument that Neanderthals were too specialized to be the ancestors of modern humans. He instead believed that Neanderthals were the direct ancestors of modern humans. Brace criticized the widely held views of French paleoanthropologists Marcellin Boule and Henri-Victor Vallois that Neanderthals were an evolutionary cousin that became extinct and was replaced by Cro-Magnons. He compared Neanderthal skeletons to those of Homo erectus and Homo sapiens. Influenced by the new taxonomic ideas arising from the Modern Evolutionary Synthesis, Brace considered Neanderthals to be a subspecies of Homo sapiens (Homo sapiens neanderthalensis). Franz Weidenreich’s views about human evolution (Weidenreich 1949) prompted Brace to apply the concept of clines to the hominid fossil record and thus to view human morphology as a consequence of adaptation to local conditions, genetic drift (non-adaptive regional characteristics such as ear shape or nuances of cheekbone morphology), and interbreeding across populations. Like Weidenreich, Brace conceived of the relationship between hominid fossils and variation observed in modern human populations as one of Regional Continuity (Brace 1964a). Thus, current human populations are for the most part descendants of the hominid populations that inhabited the same geographical region in the past. As a result, Brace opposed the various versions of the Out of Africa hypothesis and argued that those who supported the replacement of Neanderthals by modern humans were un-Darwinian. He suggested that in the course of human history, when an immigrating population encountered an indigenous population with similar technology, admixture (interbreeding) would result, not replacement. He also argued that hominid evolution did not occur by rapid speciation at a periphery, but rather across an entire, globally dispersed species.

Brace argued that cultural factors, especially the increased use of tools by Neanderthals, produced morphological changes that led the ‘classic’ Neanderthals to evolve into modern humans. He linked the exclusion of Neanderthals as a direct human ancestor to the tradition of catastrophism in the sciences and he believed this was incompatible with Darwinian evolution. He argued that no individual Neanderthal specimen displayed the full set of supposedly ‘Neanderthal’ features identified by previous generations of paleoanthropologists. Instead, he suggested that Neanderthals display a complement of traits, some shared by modern humans, while many so-called ‘Neanderthal’ traits are visible in Upper Paleolithic human fossils. Brace also suggested that the difference between early Neanderthals, such as the fossils excavated from Krapina, in Croatia, and later Neanderthals was greater than the differences between some Neanderthals and early Homo sapiens. He also addressed the evidence for Homo sapiens specimens that were thought to be older than some Neanderthal specimens. Although the Qafzeh and Skhūl specimens excavated in Israel were usually represented as ‘anatomically modern’ Homo sapiens, Brace examined these fossils and found that the Qafzeh skulls displayed a ‘modern’ cranial morphology and large tooth size, while the Skhūl specimens possess an intermediate cranial shape and tooth size. In other words, the Skhūl fossils were intermediate between ‘classic’ Neanderthal and ‘modern human’ morphology. When Qafzeh and the ‘classic’ Neanderthals are compared, the average size of their teeth are indistinguishable, although the craniofacial morphology of the Qafzeh hominids was different from that of the ‘classic’ Neanderthals of Western Europe.

Beyond these contributions to the study and interpretation of Neanderthals, Brace also established a research program that focused on the role that cultural innovation played in changing human anatomy. He was a strong supporter of the idea that culture played a vital role in hominid morphological evolution. Brace believed culture was a unique human adaptation and was the most distinctive human attribute and that it was the driver of physical evolution. His research demonstrated that humans evolved mostly by the structural reduction of certain parts of the body (the major exception being the size of the brain). These include reductions in both tooth size and skeletal robusticity. Brace also argued that hominid evolution did not occur by rapid speciation at a periphery, but among the entire globally dispersed species. He published papers on the reduction of tooth size and the selective pressures that changed the hominid cranium during the Pleistocene and he suggested that the introduction of pottery and changes in food preparation led to a significant reduction in tooth size during the post-Pleistocene period (Brace 1967; Brace and Mahler 1971). As previously mentioned, he applied this research to Neanderthal evolution, arguing that Neanderthals used their front teeth as tools, which is what produced their distinctive skull shape and the large size of their incisors.

Related to this, Brace proposed what he called the Probable Mutation Effect. This is the likely effect of genetic mutations on physical traits that are no longer maintained by selection, or in other words, the effect of the disuse of anatomical structures. Brace observed that under most circumstances the reduction in size or the elimination of certain structures is detrimental to an organisms’ survival. However, if the circumstances under which an anatomical structure evolved should disappear, then the structure in question becomes free to vary without having any influence upon the organism’s chances of survival. Subsequent variation is at the mercy of random mutation and because of the Probable Mutation Effect, reduction of the structure is inevitable (Brace 1963; 1964c). Brace was influenced by American geneticist Sewall Wright’s work on genetic drift, the idea that some biological change does not result from natural selection and are not necessarily adaptive. Applying these ideas to hominid evolution resulted in Brace’s hypothesis that modern humans evolved as a consequence of structural reductions in both tooth size and skeletal robustness. As humans adopted cooking, invented more sophisticated tools, developed projectiles that allowed them to kill prey from a distance, and as they invented pounding and grinding tools around 15,000 years ago and the use of pottery around 12,000–10,000 years ago, selection for a robust body was decreased.

Brace spent decades collecting and analyzing data on dentition and facial morphology. This led him to amass one of the largest databases of craniofacial and dental measurements of hominids over the course of his career. He discovered that during the last 200,000 years, teeth and faces had gradually decreased in size, but to different degrees in various parts of the world. The archaeological record reveals that the distribution of small tooth size occurs in areas where cooking had emerged, such as parts of Europe and the Middle East, some areas in eastern Asia, and pre-agricultural Japan. Brace continued to explore dental and skeletal size reduction and their relationship to behavioral change in humans throughout his career (Brace 1973; 1979). He was also interested in the role that various stresses affecting the body during growth and development played in determining the form of the adult body. One result of this research was Brace’s insight that the overbite was not a normal configuration of the jaws but instead emerged only after forks and chopsticks began to be used (Brace and Nagai, 1982). Before eating utensils were developed, a piece of meat held in the hand and gripped by the incisors was cut by a knife. Such repeated edge-to-edge gripping would affect the growth of the jaw, producing the edge-to-edge bite. When edge-to-edge gripping came to an end as a cultural practice with the introduction of eating utensils that allowed food to be cut into small pieces, an overbite emerged. Brace believed that the emergence of the ‘F’ phoneme in some languages arose as a result of the overbite (Brace 1986).

Brace was interested in the mechanisms and process of evolutionary change and he was influenced by the ideas of the Modern Evolutionary Synthesis and Sherwood Washburn’s New Physical Anthropology. During the 1950s evolutionary biologist Ernst Mayr, geneticist Theodosius Dobzhansky, paleontologist George Gaylord Simpson and others had begun to apply the new ideas from the Modern Synthesis to the problem of human evolution. Dobzhansky argued that humans should be understood as a single polytypic species, with races representing populations displaying geographical variations in morphology. Likewise, Washburn sought to replace the old typological and descriptive physical anthropology of the early twentieth century, which relied upon anthropometry to identify and characterize static human races. He imagined instead a new kind of physical anthropology that would rely upon the study of genetics, morphology, and function in order to produce a dynamic view of humans. Washburn saw human races as populations and not as natural morphological types, a perspective that was consistent with the Modern Synthesis’s view of biological species. Dobzhansky and Mayr also believed that in the course of human evolution only one species of hominid had existed at any one time. Hominids consisted of a single evolving lineage that possessed considerable geographical variation. This idea came to be known as the single species hypothesis. They recognized that geographical (racial) variation existed among humans and very likely among earlier hominids, but they considered these variations to be taxonomically trivial. Mayr and Simpson sought to reform hominid taxonomy according to the principles of the Modern Synthesis. They argued that the large number of hominid genera and species recognized at that time should be dramatically reduced. Mayr went as far as to suggest that all known hominids should be classified within just three species: Homo transvaalensis (comprising the australopithecines), Homo erectus, and Homo sapiens. Brace agreed with the need for taxonomic reform consistent with the principles of the Modern Synthesis and in Man’s Evolution: An Introduction to Physical Anthropology (1965) Brace supported Mayr’s argument for including the australopithecines in the genus Homo. However, by 1977 he came to accept Australopithecus as a genus but wanted to include all the known Australopithecus fossils within the single species Australopithecus africanus.

Brace adopted many of these ideas in his own thinking about human evolution. In his influential book Stages of Human Evolution, which went through five editions between 1967 and 1995, Brace supported the single species hypothesis. He proposed that hominids had evolved through four morphological stages: australopithecine, pithecanthropine, Neanderthal, and modern human. He believed the transitions between stages had occurred essentially simultaneously (from an evolutionary time scale) across different geographic areas, as a result of cultural diffusion and gene flow. Brace’s conception of human evolution relied upon something called the competitive exclusion principle. This is the idea that culture was the best way to exploit any ecological niche, and since culture allowed humans to exploit all ecological niches there would be no cause for speciation events in the course of human evolution. Once hominids adopted culture, selective pressure led them to evolve in similar ways and rates everywhere, since any important cultural and technological developments would be shared by all other groups, thus maintaining the unity of the species at each stage of evolution. The ‘cultural ecological niche’ consisted of all the learned behaviors of a group, which were passed from one generation to the next. This would result in a powerful survival strategy that diminished the effects of natural selection. Thus, the effects of the competitive exclusion principle were broad enough and dominant enough to preclude the emergence of new species during the course of hominid evolution. Brace dismissed the notion of geographical ‘racial’ hominid populations that were present in the evolutionary schemes of Franz Weidenreich and later adopted by American anthropologist Carleton Coon in the 1960s and even retained by Dobzhansky. He also vigorously opposed Carleton Coon’s suggestion that different human races had evolved at different rates.

As paleoanthropologists discovered new hominid fossils, Brace integrated them into his theory, but he also had to revise his ideas in the face of the new evidence. Since he believed that modern humans had evolved from Neanderthals, Brace challenged the dates for Homo sapiens fossils from Europe, the Middle East, and Africa that appeared to be older than the dates for Neanderthal fossils. He embraced the KNM-ER 1470 (Homo rudolfensis) fossil discovered by Richard Leakey’s team at Lake Turkana in 1972 as representing one of the steps by which Australopithecus evolved into Homo. Brace emphasized the effect that sexual dimorphism had on variation in early hominids, such as the australopithecines, and argued that earlier stages of human evolution were characterized by greater differences in tooth and body size between males and females than are observed in humans today. This led him to argue that all the australopithecine fossils known at that time belonged to a single species (Brace 1969, 1973). This view was ridiculed by F. Clark Howell and other paleoanthropologists.

Brace initially believed that the australopithecines must have possessed tools and a rudimentary culture in order to survive, but he later had to change his thinking about this. After Donald Johanson’s discovery of Australopithecus afarensis in Ethiopia in 1974, Brace defended the single species hypothesis by arguing that since the australopithecines were the first terrestrial hominids and because the australopithecines lacked culture (tool-use) and probably lived by scavenging, a number of local species could have evolved. Richard Leakey’s discovery of KNM-ER 406 (Paranthropus boisei) in 1969 and KNM-ER 3733 (Homo ergaster) in 1975 at Lake Turkana, two clearly different hominid species that lived at the same time, posed a serious problem for the single species hypothesis. These discoveries forced Brace in later editions of Stages of Human Evolution to accept one speciation event early in hominid evolution and to argue that the competitive exclusion principle had been less effective during the australopithecine phase of hominid evolution because culture was less prevalent at that time and had a less adaptive effect. However, he continued to argue that during the course of the Pleistocene, from the emergence of Homo erectus to the appearance of Homo sapiens, hominid evolution consisted of a single evolving lineage with only one species living at any one time, with no extinctions or side branches. Brace believed that once hominids began to hunt and use tools, selective pressures led them to evolve in similar ways and at similar rates everywhere, since any important cultural and technological development would be passed on to other groups, thus maintaining the unity of the species at each stage of hominid evolution. In fact, he suggested that it was the emergence of hunting that led to the transformation of Australopithecus into Homo.

Brace was critical of the theory of punctuated equilibria that Stephen J. Gould and Niles Eldredge proposed in 1972. Punctuated equilibria challenged some central tenants of the Modern Synthesis. While the Modern Synthesis envisioned evolution as gradually evolving lineages with fuzzy essentially arbitrary boundaries separating an ancestral species from its descendant, Gould and Eldredge argued that evolution was better represented by periods of rapid (punctuated) evolution separated by long periods of stasis. They argued that species are real, not just conventions. Gould rejected the ‘ladder of progress’ conception of human evolution and he promoted instead a view of hominid phylogeny that was more akin to a bush than a tree (Gould and Eldredge 1972; Gould 1976). They explicitly criticized Brace’s linear hominid phylogeny based upon the single species hypothesis. The discovery of many new hominid fossils during the 1950s and 1960s led Ian Tattersall and Niles Eldredge to assert that the Modern Synthesis’ view of evolution did not match the evidence from the hominid fossil record. They argued instead that the punctuated equilibria view of evolution was a better fit (Eldredge and Tattersall 1975; Delson, Eldredge and Tattersall 1977). By this time German biologist Willi Hennig’s phylogenetic systematics (cladistics) was also beginning to influence paleoanthropologists, especially after the appearance of an English translation of his book (Hennig 1950; 1966). Tattersall and Eldredge introduced cladistics and punctuated equilibria into hominid systematics and soon other paleoanthropologists began to apply these new approaches to the problem of hominid evolution and hominid taxonomy.

Brace was wary of this enthusiasm for punctuated equilibria among paleoanthropologists. He thought the theory’s claims about long periods of evolutionary stasis and its rejection of transitional fossils linking ancestor and descendant species was a manifestation of essentialist, non-evolutionary thinking. He also disagreed with the ‘bushy’ hominid phylogeny promoted by punctuated equilibria because it led to the identification of too many hominid species. Brace also criticized the growing influence of cladistics by paleoanthropologists because he thought its approach to hominid phylogeny reinforced the perspective of punctuated equilibria and he criticized cladistics’ inclination toward typological thinking. Brace’s attitude about hominid taxonomy is reflected in his perspective on Homo habilis. During excavations at Olduvai Gorge in 1960, Louis and Mary Leakey unearthed fossils (Olduvai Hominid 7) that Leakey, Phillip Tobias, and John Napier determined belonged to a new species of hominid that they called Homo habilis (Leakey, Tobias, Napier 1964). There was immediate criticism from some paleoanthropologists, notably Wilfrid Le Gros Clark and John Robinson, about the creation of this new species. Some critics thought there was insufficient morphological space between the gracile australopithecines and Homo erectus to accommodate this new species. Brace also criticized the validity of Homo habilis as a taxon on the basis of an examination of its dentition (Brace, Mahler, and Rosen 1973).

Brace coauthored a textbook with the British-American anthropologist Ashley Montagu titled Man’s Evolution: An Introduction to Physical Anthropology in 1965. A second edition, with the new title of Human Evolution: An Introduction to Biological Anthropology was published in 1977. They covered the basic principles of evolutionary theory, primate taxonomy, the hominid fossil record, and the anthropological study of living human populations. The first edition already introduced the notion of australopithecine, pithecanthropine, Neanderthal, and modern human stages of human evolution. Montagu, like Brace, rejected the biological concept of race and the type of racial anthropology that was common in late nineteenth and early twentieth century scientific and political thought. Montagu had served on the UNESCO committee, later known as the Committee of Experts on Race Problems, and he coauthored the Statement on Race in 1950. Unsurprisingly then, Brace and Montagu also addressed the history of the concept of race in anthropology and offered critiques of it in their textbook. Brace also coauthored, with Harry Nelson and Noel Korn, the Atlas of Human Evolution (1979). This presented a general overview of hominid evolution and contained descriptions of hominid fossils from around the world as well as illustrations drawn by Mary Brace.

Brace continued to think about evolutionary processes and to respond to new developments in biology, including discoveries in genetics and molecular biology, throughout the 1970s and 1980s. During the early part of his career, Brace emphasized the role of natural selection in evolution and as a result he doubted that biological traits could be non-adaptive. However, advances in genetics had demonstrated that many biological traits were simply neutral with respect to the functioning of that biological organism. The ‘hyperselectionist’ approach of the biologists who had promoted the Modern Evolutionary Synthesis were being criticized by biologists such as Stephen Jay Gould and Richard Lewontin. Brace responded to this critique in the 1980s, perhaps because it fit with his conception of the Probable Mutation Effect, by becoming less hyperselectionist in his approach to hominid evolution. For example, in his research investigating the connection between the Ainu (an indigenous population in Japan) and the modern Japanese, Brace came to believe that closely related groups were better differentiated using traits that probably were nonadaptive, or were not subject to strong selection.

Brace continued to study prehistoric changes in human dentition during the 1980s (Brace 1980; Brace and Ryan 1980; Brace 1984; Brace, Rosenberg, and Hunt 1987; Brace, Smith, and Hunt 1991), but he also began to investigate the differences in facial morphology and human dentition in Asian populations (Brace 1980; Brace and Nagai 1982; Brace, Shao, and Zhang 1984; Brace and Tracer 1992). He traveled to China and Japan to examine human cranial and dental specimens. He found similarities between living Japanese Ainu and the prehistoric Jōmon people who inhabited Japan from approximately 13,000 BCE to 300 BCE. Brace suggested that the Ainu were the remnants of a widespread Jōmon-Pacific people who not only populated Japan and Taiwan but also much of the Pacific rim. He found that non-Ainu Japanese people resembled populations in Korea and South China and Brace argued that these ancient people had migrated into Japan bringing with them wet-rice agriculture, which allowed them to displace the earlier hunter-gatherer Jōmon people. In the 2000s, Brace and his students extended this research on craniofacial morphology to link the Pleistocene peopling of the New World to their work on the Jōmon and Ainu. Brace and his colleagues argued that the approximately 9000 year old Kennewick Man fossil found in Washington state in 1996, as well as living indigenous American populations, could trace their ancestry to a prehistoric Jōmon-Pacific dispersal (Brace, Brace, and Leonard 1989; Brace et al. 2001; Brace, Seguchi, and Brace 2008; Brace et al. 2014).

During the years that Brace was involved in studying human evolution, the creationist movement was gaining influence in the United States. Brace was a vocal critic of creationism and its rejection of evolutionary theory and the evidence for human evolution. He debated leading creationists, including Henry Morris in Auckland, New Zealand in 1973 and Duane Gish at the University of Michigan in 1982. Brace gave a lecture titled “Confronting a Creationist” at Central Michigan University in 1983. He also wrote several articles attacking creationist ideas. These include “Humans in Time and Space” (Brace 1983), “Creationists and the Pithecanthropines” (Brace 1986b), and “Human Emergence: Natural Process or Divine Creation” in the book Scientists Confront Intelligent Design and Creationism (Brace 2007). Brace argued that so-called scientific creationism “can only be maintained … by either ignoring or denying virtually all of the data and their implications accumulated by biologists, paleontologists, archaeologists, and anthropologists as a result of a century and more of increasingly carefully checked and substantiated work” (Brace 1983).

Throughout his scientific career, Brace’s scientific thinking was influenced by his interest in the history and philosophy of science. As paleoanthropologists were quarreling over hominid taxonomy in the aftermath of the effects of cladistics and punctuated equilibria in paleoanthropology, Brace wrote “Tales of the Phylogenetic Woods: The Evolution and Significance of Phylogenetic Trees” tracing the historical and philosophical issues related to hominid phylogeny (Brace 1981; see also Brace 1988). He also wrote on the history of the concept of race and racial anthropology (Brace 1982), culminating in the book “Race” Is a Four-Letter Word: The Genesis of the Concept (Brace 2005). Brace’s lifelong interest in evolutionary theory resulted in the publication of Evolution in an Anthropological View (Brace 2000), which supported Darwin’s theory of evolution, its methodology and philosophical perspectives, and its application to explaining human evolution. The book criticized those anthropologists and paleoanthropologists who failed to understand the mechanisms driving evolution, particularly the implications of the Modern Synthesis, as well as criticizing their lack of interest in investigating the processes by which modern humans evolved from earlier hominids.

Brace was active in several scientific institutions. He served as a Fellow at the Center for Advanced Study in the Behavioral Sciences at Stanford University in 1977–1978 and again in 1980–1981. He was a member of the American Association of Physical Anthropologists; of the American Anthropological Association; and the Society for Social Biology. He was also a member of the Dental Anthropology Association and served as its president from 1988–1990. He was elected a member of the American Association for the Advancement of Science. Brace was awarded the Charles R. Darwin Lifetime Achievement Award by the American Association of Physical Anthropologists in 2006. Kevin Hunt and Lucia Allen Yaroch organized “Evolution, History and Biological Anthropology: A Symposium in Honor of C. Loring Brace” at the 64th annual meeting of the American Association of Physical Anthropologists, which was held in Oakland, California, on 30 March 1995.

Charles Loring Brace IV died on 7 September 2019 at Ann Arbor, Michigan. The C. Loring Brace papers, which contain material from 1954 to 2009, are held at the University of Michigan Bentley Historical Library.

Selected Bibliography

L. Schuman and C. L. Brace, “Metric and Morphologic Variations in the Dentition of the Liberian Chimpanzee; Comparisons with Anthropoid and Human Dentitions.” Human Biology 26 (1954): 239–268.

“Refocusing on the Neanderthal Problem.” American Anthropologist 64 (1962a): 729–741.

“Cultural Factors in the Evolution of the Human Dentition.” In M.F.A. Montagu, (ed.), Culture and the Evolution of Man. Pp. 343–354. Oxford University Press, New York, 1962b.

“Structural Reduction in Evolution.” The American Naturalist 97 (1963): 39–49.

“The Fate of the ‘Classic’ Neanderthals: A Consideration of Hominid Catastrophism.” Current Anthropology 4 (1964a): 3–43.

“On the Race Concept.” Current Anthropology 4 (1964b): 313–320.

“The Probable Mutation Effect.” The American Naturalist 98 (1964c): 453–455.

L. Brace and M. F. A. Montagu, “A Non-racial Approach towards the Understanding of Human Diversity.” In M. F. A. Montagu (ed.), The Concept of Race. Pp. 103–152. Glencoe, IL: The Free Press, 1964.

L. Brace and M. F. A. Montagu, Man’s Evolution: An Introduction to Physical Anthropology. New York: Macmillan, 1965.

“Environment, Tooth Form, and Size in the Pleistocene.” Journal of Dental Research 46 (1967): 809–816.

The Stages of Human Evolution. Englewood Cliffs, NJ: Prentice Hall, 1967 (5th ed. 1995).

L. Brace and Paul Mahler, “Post-Pleistocene Changes in the Human Dentition.” American Journal of Physical Anthropology 34 (1971): 191–203.

L. Brace, Harry Nelson, and Noel Korn, Atlas of Human Evolution. New York: Holt, Rinehart & Winston, 1971.

“Sexual Dimorphism in Human Evolution.” Yearbook of Physical Anthropology 16 (1973): 31–49.

L. Brace, P. Mahler, and R. Rosen, “Tooth Measurements and the Rejection of the Taxon ‘Homo habilis’.” Yearbook of Physical Anthropology 16 (1973): 50–68.

L. Brace and M. F. A. Montagu, Human Evolution: An Introduction to Biological Anthropology. 2nd ed. New York: Macmillan, 1977.

“Krapina, ‘Classic’ Neanderthals, and the Evolution of the European Face.” Journal of Human Evolution 8 (1979): 527–550.

L. Brace, Harry Nelson, Noel Korn, and Mary Brace (illustrator), Atlas of Human Evolution. 2nd ed. New York: Holt, Rinehart & Winston, 1979.

“Australian Tooth Size Clines and the Death of a Stereotype.” Current Anthropology 21 (1980): 141–164.

L. Brace and Alan Ryan, “Sexual Dimorphism and Human Tooth Size Differences.” Journal of Human Evolution 9 (1980): 417–435.

“Tales of the Phylogenetic Woods: The Evolution and Significance of Phylogenetic Trees.” American Journal of Physical Anthropology 56 (1981): 411–429.

L. Brace and M. Nagai. “Japanese Tooth Size: Past and Present.” American Journal of Physical Anthropology 59 (1982): 399–411.

“The Roots of the Race Concept in American Physical Anthropology.” In Frank Spencer (ed.), A History of American Physical Anthropology, 1930–1980. Pp. 11-29. Academic Press, 1982.

“Humans in Time and Space.” In L. Godfrey (ed.), Scientists Confront Creationism. Pp. 245–282. New York: W. W. Norton, 1983.

“Rates of Hominid Dental Reduction in the Late and Post- Pleistocene.” American Journal of Physical Anthropology 63 (1984): 140–141.

L. Brace, Shao Xiang-qing, and Zhang Zhen-biao, “Prehistoric and Modern Tooth Size in China.” In F. Spencer and F. Smith (eds.), The Origins of Modern Humans: A World Survey of the Fossil Evidence. Pp. 485–516. New York: Alan Liss, 1984.

“Creationists and the Pithecanthropines.” Creation/Evolution 19 (1986): 16–23.

L. Brace, K. R. Rosenberg, Kevin D. Hunt, “Gradual Change in Human Tooth Size in the Late Pleistocene and Post-Pleistocene.” Evolution 41 (1987): 705–720.

“Punctuationism, Cladistics and the Legacy of Medieval Neoplatonism.” Human Evolution 3 (1988): 121–138.

L. Brace, M. L. Brace, W. R. Leonard, “Reflections on the Face of Japan: A Multivariate Craniofacial and Odontometric Perspective.” American Journal of Physical Anthropology 78(1989): 93–113.

L. Brace, S. L. Smith, and Kevin Hunt, “What Big Teeth You Had Grandma! Human Tooth Size, Past and Present.” In M. A. Kelley and C. S. Larsen (eds.) Advances in Dental Anthropology: Pp. 33–57. New York: Wiley-Liss, 1991.

L. Brace and D. P. Tracer. “Craniofacial Continuity and Change: A Comparison of Late Pleistocene and Recent Europe and Asia.” In T. Akazawa, K. Aoki, and T. Kimura (eds.), The Evolution and Dispersal of Modern Humans in Asia. Pp. 439–471. Tokyo: Hokusen-sha, 1992.

“Bio-Cultural Interaction and the Mechanism of Mosaic Evolution in the Emergence of ‘Modern’ Morphology.” American Anthropologist 97 (1995): 711–721.

“Cro-Magnon and Qafzeh—Vive la Difference.” Dental Anthropology Newsletter 10 (1996): 2–9.

Evolution in an Anthropological View. Walnut Creek, CA: Altamira Press, 2000.

L. Brace, A. Russell Nelson, Noriko Seguchi, Hiroaki Oe, Leslie Sering, Pan Qifeng, Li Yongyi, Dashtseveg Tumen, “Old World Sources of the First New World Human Inhabitants.” Proceedings of the National Academy of Sciences 98 (2001): 10017–10022.

“Race” Is a Four-Letter Word: The Genesis of the Concept. Oxford: Oxford University Press, 2005.

“Human Emergence: Natural Process or Divine Creation.” In Andrew J. Petto and Laurie R. Godfrey (eds.), Scientists Confront Intelligent Design and Creationism. New York: W.W. Norton & Company, 2007.

L. Brace, N. Seguchi, and M. L. Brace, “Exploring the Kennewick Connection.” In C. Smith, L. Zimmerman, J. Watkins and D. Lippert (eds.) Kennewick Man: Perspectives on the Ancient One. Pp. 153–68. Walnut Creek, Calif.: The Left Coast Press, 2008.

“‘Physical’ Anthropology at the Turn of the Last Century.” In Michael Little and Kenneth Kennedy (eds.), Histories of American Physical Anthropology in the Twentieth Century. Pp. 25–53. Lanham, Mass.: Rowman & Littlefield, 2010.

L. Brace, Noriko Seguchi, A. Russell Nelson, Pan Qifeng, Hideyuki Umeda, Margaret Wilson, and Mary L. Brace, “The Ainu and Jomon Connection.” In Douglas Owsley and Richard Jantz (eds.), Kennewick Man: The Scientific Investigation of an Ancient American Skeleton. Pp. 463–471. College Station: Texas A&M Press, 2014.

Other Sources

Delson, N. Eldredge and I. Tattersall, “Reconstruction of Hominid Phylogeny: A Testable Framework Based on a Cladistic Analysis.” Journal of Human Evolution 6 (1977): 263–278.

Eldredge and I. Tattersall, “Evolutionary Models, Phylogenetic Reconstruction, and Another Look at Hominid Phylogeny.” In F. S. Szalay (ed.), Approaches to Primate Paleobiology. Pp. 218–243. Basel: S. Karger, 1975.

J. Gould and N. Eldredge, “Punctuated Equilibria: An Alternative to Phyletic Gradualism.” In T. J. M. Schopf (ed.), Models in Paleobiology. Pp 82–115. San Francisco: Freeman, Cooper & Co, 1972.

J. Gould, “Ladders, Bushes and Human Evolution.” Natural History 85 (1976): 24–31.

Willi Hennig, Grundzüge einer Theorie der Phylogenetischen Systematik. Berlin: Deutscher Zentralverlag, 1950.

Willi Hennig, Phylogenetic Systematics. Chicago: University of Chicago Press, 1966.

Louis Leakey, Phillip Tobias, and John Napier, “A New Species of the Genus Homo from Olduvai Gorge.” Nature 202 (1964): 7–9.

B. Livingstone, “On the Non-existence of Human Races.” Current Anthropology 3 (1962): 279–281.

Sherwood Washburn, “The New Physical Anthropology.” Transactions of the New York Academy of Sciences 13 (1951):298–304.

Sherwood Washburn, “The Strategy of Physical Anthropology.” In Alfred Kroeber (ed.), Anthropology Today. Pp. 714–727. Chicago: University of Chicago Press, 1953.

Secondary Sources

Helke Ferrie, “An Interview with C. Loring Brace.” Current Anthropology 38 (1997): 851–869.

Dean Falk and Noriko Seguchi. “Professor C. Loring Brace: Bringing Physical Anthropology (‘Kicking and Screaming’) into the 21st Century!” Michigan Discussions in Anthropology 16 (2006): 175–211.

Shelley Smith, “Single (Sub)species Then and Now: An Examination of the Nonracial Perspective of C. Loring Brace.” American Journal of Physical Anthropology 165 (2018): 104–125.

Kevin Dean Hunt, “Obituary of Charles Loring Brace, IV (1930–2019).” Journal of Human Evolution 142 (2020):

Kevin Dean Hunt, “C. Loring Brace IV (1930–2019).” Journal of Anthropological Research (2020): 157–163.

Shelley L. Smith and Kevin D. Hunt, “Charles Loring Brace, IV (1930–2019).” American Journal of Biological Anthropology 171 (2020): 567–568.