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Authors : Dean Falk, Noriko Seguchi
Title: Professor C. Loring Brace: Bringing Physical Anthropology ("Kicking and Screaming") Into the 21st Century!
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Source: Professor C. Loring Brace: Bringing Physical Anthropology ("Kicking and Screaming") Into the 21st Century!
Dean Falk, Noriko Seguchi

vol. 16, no. 1, 2006
Issue title: Retrospectives: Works and Lives of Michigan Anthropologists
PDF: Link to full PDF [295kb ]

Professor C. Loring Brace: Bringing Physical Anthropology (“Kicking and Screaming”) Into the 21st Century!

Dean Falk

Department of Anthropology, Florida State University

Noriko Seguchi

Department of Anthropology, University of Montana, Missoula

On Christmas day, 1945, a fifteen-year-old boy was given a book written by Roy Chapman Andrews that was entitled Meet Your Ancestors. Thus began the career of C. Loring Brace IV who would one day become an internationally respected biological anthropologist. Born on December 19, 1930 in Hanover, New Hampshire to Gerald Warner Brace, a professor of English at Boston University, and Huldah Laird Brace, a trained biologist, Brace was descended from a long line that had deep roots in 17th–century New England. Foreshadowing the intellectual and humanitarian interests of a great-grandson he would never meet, his great-grandfather (the first Charles Loring Brace) read and was fascinated by an early copy of Darwin’s Origin of Species, founded the Children’s Aid Society, and was prominent in fighting against slavery (Ferrie 1997).

Eventually Brace earned a B.A. degree in geology from Williams College (1952). In the summer of 1953, Brace met wife-to-be Mary Louise Crozier (Mimi) at the last nickel Coke machine in Cambridge, near the entrance to the basement of the Peabody Museum. The daughter of biology professor William J. Crozier, Mimi was spending the summer washing and sorting potsherds, while Brace was spending his measuring chimpanzee teeth. Eventually they married and produced three sons, Charles Loring Brace V, Roger Crozier Brace, and Hudson Hoagland Brace. Based on mutual interests in nature, literature, cooking, science, and family, theirs was a marriage for life, and it is illustrative of their strong bond that until Mimi’s death on August 15, 2005, Loring read to her each evening as she prepared their dinner.

Brace went on to earn M.A. and Ph.D. degrees in anthropology from Harvard University (1958, 1962) where he studied with Earnest Albert Hooton and William W. Howells. His first faculty positions were at the University of Wisconsin–Milwaukee and the University of California, Santa Barbara (UCSB). In the 1960s, he decided to leave Santa Barbara because then Governor Ronald Reagan fired the president of the University of California, Clark Kerr, with words to the effect that “the State of California has no business subsidizing intellectual curiosity.” Margaret Hamilton, who was an undergraduate at UCSB at the time, recalls that there was an enormous reaction throughout the UC system when Kerr was fired:

Loring was teaching one of those monster Introduction to Physical Anthropology courses used in many schools to meet the general education requirement in the sciences. It just so happened that his class of 800 met at the noon hour on the day when a huge rally convened in support of Kerr. After the announcement of the firing, Loring led his entire class out of the classroom and across campus to join the rally. It was a wonderful moment.

Brace’s first doctoral student, Stephen Molnar, also recalls the early days at UCSB:

From my first encounters with Loring I was impressed with his scholastic breadth. Within just two years of receiving his doctorate, he published theoretical papers on structural reduction, a critical appraisal of the dogmas of racial taxonomy, and a paper evaluating the evolutionary relationship of the classic Neandertals to modern Homo sapiens. While these papers were being written a much needed text, Man’s Evolution: An Introduction to Physical Anthropology, was also in progress. Not a bad output for a new assistant professor!

Brace found a perfect haven in 1967 when he moved to the University of Michigan in Ann Arbor and became colleagues with Frank Livingstone, of “there are no races, there are only clines” fame (Livingstone 1962). Nearly four decades later, he remains at Michigan as Professor of Anthropology and the Curator of Biological Anthropology in the Museum of Anthropology. In addition to his close tie to Frank Livingstone, Brace had a special relationship with another physical anthropologist, Stanley Garn. Karen Rosenberg notes: “Stanley’s wife, Priscilla, and Mimi were half sisters. When I was Loring’s student and, simultaneously, Stanley’s research assistant, they would walk home together very often, Loring usually pushing his bike and the two of them talking animatedly as they walked down Washtenaw together.”

Student Recollections Brace also found the perfect home in Ann Arbor. Recalls Alan Ryan:

Loring’s home is located on a prestigious block near the University of Michigan. The house is extremely large and ornate, looking like a slightly modern version of the one owned by the Adam’s family. Once inside, there seemed to be several staircases, some leading to everywhere or nowhere. The dining room was magnificent and well lit by a chandelier that was handmade by one of the greatest heroes of the American Revolution and accomplished silversmith of Boston, Paul Revere! Mimi was a gourmet cook, and any dinner at their home was a three-hour affair, involving many courses of food and a large assortment of cordials and other beverages. Kindness, a key aspect of Loring’s personality, was evident throughout the meal. Many of the toasts that were made highlighted the few accomplishments of my graduate school career. Of course, the meal was magnificent.

Margaret Hamilton (who earned her Ph.D. at Michigan after graduating from UCSB) remembers that when he went home for the day Brace did not “leave his work at the office:”

Loring’s interest and delight in all things of natural history formed a bridge between his work and his avocations. It is not possible to mention this without mentioning their beloved parrots. I’m not sure how many they owned, but up until I graduated the house was always filled with them. You may recall that this interest actually resulted in a wonderful paper about the parrot family filling a niche similar to that of primates...Another area of interest overlapping or propelled by research was the topic of diet and food modification techniques. Loring’s early interest in dental evolution fed right into his and Mimi’s shared passion for ethnic cooking. This had as much to do with all of the effort and techniques that went into food preparation as it did with the actual preparation, serving, and consumption of wonderful meals enjoyed at Chez Brace. Loring and Mimi’s luau parties included all kinds of traditional food including a pig cooked in a traditional oven with hot stones in the ground.”

Bob Hinton says: “I remember the menagerie at the house. I ate most of a dinner there with a parrot perched on my shoulder. Neither Loring nor Mimi thought anything of it! I will also never forget the VW convertible ‘bug’ that served as a giant planter adjacent to the house.”

Alan Ryan adds:

Mimi had an incredible menagerie of pets, everything from the smallest snake to the largest bird. As a result, meals were sometimes interrupted by the flight of a bird above the dining room table or a visit from a friendly monkey begging for morsels of food. Nevertheless, everyone had a great time. Loring’s students viewed these gatherings as one of the many rites of passage that brought much cheer and comfort to their stressed lives.

Ken Weiss also remembers the Braces’ predilection for “pets:”

The Braces’ house was like a not-so-miniature zoo, as I’m sure his students well remember. Every vacant spot or chair was occupied by an animal or a dozing young Brace. For a number of weeks, I attended the Ann Arbor Zoo Club for the Brace kids and others from the neighborhood and even led a dissection of a rhesus monkey. Among the fauna was a litter of un-house-trained puppies trapped by a board in the master bedroom, some kind of surprise reptile in the black-lit bathroom to scare the deposits out of half-sloshed attendees at the many Brace parties, a few buried carcasses “maturing” into skeletons by the Braces’ invertebrate and microbial dwellers in the backyard, and the loud, foul-mouthed mynah bird that swore when the phone rang and then mimicked Mimi’s “hello” and shouted “Lorrrrring!!” made every call to contact one’s advisor an adventure.

In sum, Brace is a colorful professor who has been described more than once as a “legend on campus.” Says Ken Weiss:

An “acknowledgment” to NSF for not supporting his work appeared in at least one paper, and his advocacy of the Probable Mutation Effect (his version of the likely effect of mutation on traits no longer maintained by selection) and of the Competitive Exclusion idea of culture as an ecological niche that guaranteed a single hominid species at any time were forcefully argued. It was never clear that they were mainstream ideas or widely accepted, but controversy energized rather than inhibited Loring, and only he would have the nerve to get up and assert such contentious positions in doggerel at meetings. Always a treat to go to one of his presentations!

Brace’s doctoral students

An EmBracing Trilogette
Canto the First
Overflowing hair and beard
His blackboard writing all obscured
Yet when he moved, of what he’d writ
One never could decipher it
Canto the Second
To think of his menagerie of yore
Doth make an image nonpareil
Unmatched by old and venerated lore:
For even mustered all in file
The Ark could not have managed any more!
Canto the Third
This Professor was not at all boring
Religion and “races” abhorring
One Brace’d for the worst
Of opinions well versed
No doubt—there is only one Loring!
–Ken Weiss

Brace supervised 20 doctoral dissertations on various topics including the history of science, dental evolution, living primates, fossil hominids, and prehistoric peoples. [1] One of these dissertations echoed Brace’s fascination with the intellectual/historical underpinnings of biological and evolutionary theory;and six others focused on various aspects of dentition, reflecting Brace’s interests in functional morphology, biological variation in primates including humans, and the role of culture in shaping (biological) evolutionary responses. Three dissertations concentrated on morphology and behavior in nonhuman primates, in keeping with Brace’s dedication to analyzing human evolution within an unfettered Darwinian framework (discussed below). Brace also supervised three dissertations that bear on the “Neanderthal problem,” a controversial subject upon which he has taken a strong stance; and he directed seven dissertations that concern variation in pre-industrial or prehistoric peoples and the biological and cultural innovations that helped mold their evolution. In addition to supervising the above dissertations, Brace served on the dissertation committees of another 18 students. [1]

Many of Brace’s former students are now prominent anthropologists, and they owe much not only to his proactive direction of their research but also to the example he set as a thoughtful and intellectually driven researcher. As Holly Smith puts it:

Loring was and is a scholar and a gentleman; he took students seriously, and treated them like fellow scholars. He was solidly enthusiastic about any idea his students generated and was ready to give any degree of help one asked for. He encouraged me to make my research quite a bit more adventuresome than I would have alone. I had a nice blocked and balanced idea about comparing a few human groups (it was tooth wear patterns in hunter-gatherers vs. agriculturalists); soon Loring had piled on suggestions that ended up with me compiling data on 10 major human groups—I ended up traveling all over Europe and America for most of a year.

Margaret Schoeninger remembers her first conversation with Brace, which took place two years before she applied to graduate school:

He related the hypothesis that variation in diets of the South African australopithecines accounted for the variation in their morphologies. He mentioned the possibility of using the trace element composition of fossil bones and teeth as a proxy measure of australopithecine diets. This excited me so much that I spent the next year taking chemistry so that I could learn how to do the work. Loring was a living library with his incredible recall of references and his extensive card catalogue system. My strongest memories of graduate school involve my meetings with Loring in his museum office where I frantically scribbled down the references he pulled out for me. These sessions were always followed by intense hours in the library. All in all, it was an incredibly stimulating intellectual experience.

Karen Rosenberg recalls:

The thing I remember most fondly about Loring was his generosity with his time. I shared his lab for at least five years and I don’t think I ever heard him tell a student that he didn’t have time to talk to them and they should come back. He was always willing to stop what he was doing (reading, writing on the old selectric or most likely writing on or filing 3x5 cards in his amazing file system) to talk about a paper, dissertation, grant proposal or some other topic (often ranging quite far from biological anthropology but always interesting). Pat Bridges, Holly Smith, and I all had desk space in the “bone range” and essentially lived there and I know that even at the time, I thought he was remarkably tolerant of our often noisy presence.

Instead of imposing his views upon students as some professors do, Brace motivated them with enthusiasm that was contagious. Ken Weiss notes:

For the most part, while we were influenced by Loring’s ideas and scholarship, his training was not so much of protégés who would do the same kind of work that he did, but of scholars who went on mainly to do their own thing. But many of us were influenced and have been successful, no doubt due to his intellectual example.

Turning out doctoral students is a bit like being a parent, and given the number of students that he has mentored over the years one wonders how he found time to work on his own projects! But work he has and the results are highly significant for understanding human evolution. What became clear to us in preparing this retrospective is the interwoven nature of Brace’s research “threads” whose subjects range from philosophy and the history of science to the impact of culture and technology on human biological evolution. In keeping with the four-field approach to anthropology (entailing physical anthropology, archaeology, linguistics, and cultural anthropology) that unfortunately has fallen out of favor in some circles, Brace’s work is integrative and bridges vast amounts of time. One gets a sense of this from his book, Evolution in an Anthropological View (Brace 2000a), and from a delightful in-depth interview with him that was published in the prestigious journal Current Anthropology (Ferrie 1997).

A “true” Darwinian scholar

Punctured Cladomania
Jargon phrase that’s so seductive;
Denies what might be called inductive
For the view,
The known can never be productive
Of the new.
But science viewed in such a way
Is nothing more than idle play;
For though the sun arose today,
Yet to our sorrow,
Deduction cannot let us say
’Twill rise tomorrow.
–I. Wright Drivell
(see Brace 1988:133 for full verse)

Brace first read Charles Darwin’s On the Origin of Species by Means of Natural Selection, or, the Preservation of Favoured Races in the Struggle for Life (1859, hereafter Origin) when he was in high school. It made a lasting impression and influenced the development of his research on a number of levels. As an historian of science, Brace traced Darwin’s intellectual underpinnings to the Scottish Enlightenment (Brace 1997), and defended his writing style and use of metaphor (Brace 2000a). He showed that certain criticisms of Darwin’s philosophical point of view were grounded in debates among philosophers about the relative merits of inductive versus deductive reasoning, and concluded that “Darwin was indeed using an inductive–deductive approach quite in the manner that characterizes the best of science being done at the present time” (Brace 2000a:51).

Brace adopted the Darwinian approach of making observations and collecting data from around the globe in order to formulate and test hypotheses about human evolution. Since 1959, he has collected measurements on cranial and dental material reposited in Europe, Africa, Southeast Asia, Australia, New Zealand, China, Japan, Thailand, and America, and has used these data to formulate and test hypotheses about human evolution. As a chronicler of contemporary science, Brace is understandably irked by current misperception of Darwin’s work by biological anthropologists and even paleoanthropologists (Brace 2000a). Rather than grasping the importance of a methodology that entailed extensive observation, Darwin’s critics continue to belittle him as something of a dullard who apparently got lucky when he came up with the concept of “descent with modification” (read: evolution). Those same critics tend to misunderstand the melding of Darwinian theory with modern molecular genetics (called the “New Synthesis” or “Synthetic [Neo-Darwinian] Theory of Evolution”) and, worse, they fail to fully grasp and apply the main mechanism that Darwin offered to account for evolution, natural selection: “Certainly a concern for the processes by which the form of living human beings emerged from non-modern-appearing ancestors is almost entirely missing from paleoanthropology today” (Brace 2000a:7). These are serious criticisms, and Brace has substantiated them with extensive documentation. An overarching goal of his own research has been to bring the outlook established by Charles Darwin and recognized by the other biological sciences to professional anthropology.

Above all else, Brace appreciated Darwin’s focus on natural selection as the “main but not exclusive means of modification” (1859:6). He was particularly impressed with Darwin’s repeated observations of traits that had formerly been adaptive but had become reduced as a consequence of relaxed selection (Brace 2000a). For example, Darwin attributed the loss of pigment and functional eyes in cave fish to disuse. Remarkably, Darwin articulated the theory of natural selection without understanding genes or anything about molecular genetics. Over a century after Origin was published, Brace formulated a hypothesis about the genetic mechanism whereby disuse led not only to those reductions described by Darwin, but also to the well known skeletal and dental reductions that are associated with the evolution of anatomically modern people (Brace 1963). That mechanism is called the Probable Mutation Effect, or PME.

Structural reduction and the Probable Mutation Effect

“In another extension of Loring’s research interests into his home interests, I remember that part of their domestic zoo included a blind cave fish named PME, short for probable mutation effect.”
–Margaret Hamilton

In 1963, Brace observed that, under most circumstances, the reduction or elimination of organic structures is detrimental to organisms’ survival. If, however, the circumstances under which a structure evolved should suddenly (or even gradually) cease to exist, then the structure in question is free to vary without having any influence upon the possessor’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, 2000a). The genetic underpinnings of PME have recently been described by Brace (2003):

When natural selection is removed from its control over the manifestation of a given trait, then the DNA segments that are responsible for producing the trait should accumulate nucleotide changes comparable to those that occur in the non-coding parts of the genome. To be sure, the trait in question is still produced at first, but with mutations accumulating in its DNA, modifications in that trait should become apparent. Nucleotide changes are most likely to alter the nature of the amino acids in the protein being coded, and random changes in the amino acid sequence in a protein molecule are much more likely to interfere with the function of that protein than they are to improve or increase that function. The accumulation of random mutations then should produce a reduction and disintegration of the phenotypic features under their control.

An important caveat is that the mechanisms of natural selection and selection relaxation (PME) apply only to traits such as skin-color, tooth size, and tooth morphology that have functional or adaptive significance for humans. One of Brace’s more genetically sophisticated former students, Ken Weiss, says:

I think the PME stands in principle, but it is not clear how to evaluate the effects or nature of relaxed selection, and some biologists still resist thinking of any aspect of DNA being free of selection...Anthropologists still have a rather simplistic notion of selection, derived from the Modern Synthesis without much serious consideration of the complexity of genetic control that has been discovered in the recent decade or two.

Brace, of course, concurs, as illustrated by his observation that not all anthropological data, either biological or cultural, can be tied to the operation of selective forces. Although, like Darwin, he recognizes that non-adaptive traits (such as certain aspects of craniofacial form) exist and may even have evolved relatively quickly as a result of neutral mutations, he has focused a good deal of research on the study of clinal distributions of adaptive traits.

There are no races, only clines

As noted above, one of the big attractions that U of M held for Brace was the presence of another physical anthropologist, Frank Livingstone, who championed clinal analyses of human variation. The word “cline” derives from the Greek word for slope and, in studies of variation, refers to a gradient in the expression of a trait across geographical regions. To quote Brace (2000a:22–23):

Quite simply, a trait that is under selective force control will grade without break from one region to another, responding only to the change in intensity of the selective force that controls its expression. The most obvious such instance is skin color. The skin pigment melanin is at a maximum near the equator where the ultraviolet component of sunlight is at its most intense. Melanin prevents the penetration of cancer-inducing ultraviolet rays, and it grades off to the north and south, with that reduction being in proportion to the distance away from the tropics and to the length of time the population in question has remained at the latitude where it is found today... Human traits that are distributed as separate and independent clines clearly show the effects of the distribution of the relevant selective forces. Skin color has one pattern of distribution, hemoglobin variants display a different one, the ABO blood group system yet another one, and tooth size has a gradient that cuts across all the others. And the pattern—gestalt—made by their intersections has no meaning in and of itself.

Certain other traits, such as lower arm and leg proportions, and length and elevation of the nose, are also distributed in a graded fashion that reflects the extent of the selective forces that influence their appearance. In fact, Brace has identified only one trait that is clearly of major adaptive significance but which does not show a distribution graded with the intensity of its selective force, namely intelligence. He reasons that intelligence lacks a clinal distribution and is similar in all human groups because the selective forces that shaped it were essentially identical during the two million years that Homo is known to have existed. Having focused much of his career on clarifying human variation by identifying and analyzing human clines (Brace 1964a, 1996, 2005), Brace demonstrated that gradients of adaptive features such as skin color and nasal form are independent of each other and that, therefore, these features should not be used (as they have been) to classify 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 are without adaptive significance, and that:

It is by an appraisal of features such as these that we can recognize the portion of the world to which the ancestors of a given person are likely to have belonged...It would be a mistake to say that this is just another name for “race,” since nothing held in common by the members of any cluster allows us to say anything about the adaptations of its components. The only thing that ties members of a given cluster together is the common inheritance of traits of no adaptive value whatsoever. [Brace 2000a:316–317]

Brace concludes that major regional cluster names are simply convenient labels for describing people who differ in “utterly unimportant ways,” and recommends that geographical names such as Africa, Amerind, Pacific-Ainu (Jomon-Pacific?), and Southeast Asia be used when labeling such clusters. He is keenly aware of racist agendas to which typological/biological concepts of race have been (and are being) applied. After four decades of research on the complexities of human variation, Brace published a major synthesis of his contributions on January 18, 2005, in his book colorfully titled “Race” Is a Four-Letter Word. Although his work on PME and human variation would, in and of itself, constitute a long and distinguished career, Brace’s unquenchable thirst for comprehending the mechanisms of evolution led him to do more.

Dental reduction as a correlate of culture

Nature: What scientific paper changed your career path?

Roberto Macchiarelli: Loring Brace’s papers on dental size reduction in hominid evolution. Papers such as “Post-Pleistocene changes in the human dentition” (Am. J. Phys. Anthropol. 34, 191-203; 1971) were a tentative application of the “probable mutation effect” model published in 1964 in the American Naturalist. Coarsely, large teeth means hard food, small teeth means soft food. Just three days before our wedding, my fiancée said that I seemed more interested in these papers than our marriage. She left, and we never saw each other again. [2]

Stephen Molnar notes that early in his career Brace took the remarkable step of proposing experimental research to test his hypothesis that tooth wear was a natural process that could provide evidence of adaptation to a particular diet, food preparation, and tool use. Says Molnar:

I ended up building a device (or “contraption” as my Australian colleagues called it) with cables and cams that could move a mandibular model through motions approximating human chewing. Loring insisted on calling the device a “Cam Actuated Nibbling Appliance” or CANIBAL for short. Some promising results were obtained and the machine went through several design changes (Brace and Molnar 1967; Molnar 1968). I am just thankful that the CANIBAL was never the subject of one of Loring’s limericks.

Brace’s decades of collecting and analyzing data on dentition and facial morphology permitted him to document dental size clines in Australia, address the peopling of Oceania, assess ancestral continuity in Japan, and examine dental reduction in China. He discovered that during the last 200,000 years (or more), dentitions and faces (the supporting structure for teeth) had gradually reduced in size, but to different degrees in various parts of the world (Brace 1978, 1980, 1984, 1995a, 1995b; Brace, Brace, and Leonard 1989; Brace and Hunt 1990; Brace and Nagai 1982; Brace, Rosenberg and Hunt 1987; Brace, Shao, and Zhang 1984; Brace, Smith, and Hunt 1991; Brace and Tracer 1992; Brace and Vitzthum 1984). Brace hypothesized that PME was responsible and began to explore the nature of the selective pressures that had been removed or relaxed. The answer was both startling and satisfying: dietary change and the development of cooking technology had relaxed selection for large dentitions and their supporting facial architecture. In other words, cultural practices ultimately altered the human skull!

As Brace and his coauthors articulated, hunters and gatherers that lived during the onset of glacial conditions in the temperate zone could not have survived without fire that allowed them to thaw and heat food that had frozen. Obligatory cooking first started some 250,000 years ago in the northwestern hemisphere (Brace 1995b), when “earth ovens” may have been used to make frozen food edible. The archaeological record reveals that the distribution of small tooth size occurs in areas where obligatory cooking had started, such as the north temperate zone between Europe and the Middle East, some areas in eastern Asia, and pre-agricultural Japan (Brace, Brace, and Leonard 1989; Brace and Nagai 1982; Brace, Rosenberg, and Hunt, 1987; Brace and Tracer 1992). This culinary practice had the effect of reducing the amount of chewing needed to process food and, consequently, it relaxed selection for maintaining tooth size. Dental reduction then proceeded at a rate of 1% every 2,000 years (Brace, Hunt, and Smith 1991).

Other significant dietary changes were made possible by the development of food processing techniques, such as pounding and grinding with tools that had been invented around 15,000 years ago, and the use of pottery, which had occurred 12,000–10,000 years ago. After the adoption of pottery, the rate of dental reduction increased to a rate of 2% every 2,000 years (Brace, Brace, and Leonard 1989; Brace, Smith, and Hunt 1991). For instance, Norwegians, whose ancestors might have been the population that first started obligatory cooking, and the Ainu in Japan, show some of the smallest tooth size in proportion to body size of any population among modern Homo sapiens (Brace and Tracer 1992). The Ainu are the direct descendants of the Jomon population who invented Jomon pottery possibly around 16,000 years ago.

In Australia, where the use of pottery never penetrated and the earth oven was a late entrant, Australians show the least amount of dental reduction. Although in Australia dental reduction has only been going on for about 15,000 years (Brace 1980), there is a remarkable North–South gradient:

Cooking traditions evidently spread slowly from a northern point of entry and eventually were adopted everywhere so that by the time of first European contact in the 18th century, fire was used in food preparation throughout the entire continent as well as in Tasmania. Even the largest-toothed of living Australians in the Murray Valley of the south have somewhat smaller teeth than their ancestors of 15,000 years ago, who had fully Middle Pleistocene tooth-size levels but who had only recently adopted the techniques of earth oven cookery...That reduction is less apparent towards the South, where incoming cultural elements had much less time depth. Australian tooth size reduction proceeded from that time at the same rate of reduction that produced the change from Neanderthal to early “modern” levels at the western edge of the Old World. [Brace and Seguchi 2003:67; 2004]

At this writing, Brace has obtained worldwide craniofacial data from 8,530 individuals and world tooth size data from more than 19,680 individuals, and he will continue to analyze these data for years to come. His thoughtful synthesis and disentanglement of selective pressures, genetic mechanisms, clinal observations, and cultural constraints is an enormous contribution to the field. As Ken Weiss notes:

Brace’s evidence of dental reduction as a correlate of culture seems reasonable, and his integration of culture and biological evolution was, and is, a valuable aspect of anthropology that is part of our professional ancestry...Of the various aspects of the Michigan school of the time, I believe this integration—real anthropology—was the greatest strength and a legacy that should be nurtured or rejuvenated.

Brace’s influence on Japanese anthropology

Brace’s research has strongly influenced the field of biological anthropology in Japan. One area in which he has made a major impact is in supporting a model of Jomon–Ainu continuity (Brace and Nagai 1982), and the Ainu–Samurai (in this context, the word “Samurai” represents the inhabitants of the Kanto Plain area around the 14th century) connection via odontometric and craniofacial perspectives (Brace, Brace, and Leonard 1989; Brace and Tracer 1990). The latter has been controversial in Japan, and a public talk entitled “Brace’s Ainu–Samurai Hypothesis from 1989 Until Today” was presented by one of us (Seguchi) at the 2005 Annual Meeting of the Anthropological Society of Nippon, in Yokohama, Japan.

Brace’s influential research on the invalidity of the biological “race” concept is now beginning to spread in biological anthropology in Japan. Japanese anthropologists still do research on human variation among Japanese and Asians from a “racial” approach. For example, “Mongoloids” are thought to be of a younger stock than the “Caucasoids” or “Negroids,” and the “archaic Mongoloids” are supposed to possess archaic characteristics in common with their European and African contemporaries in addition to having distinct “Mongoloid” features (Hanihara 1986, 1991, 1996). Japanese anthropologists assume that more recent Japanese ancestors were “typical or Neo-Mongoloids” who derived from “archaic Mongoloids” who had adapted to an extremely cold climate about 20,000–10,000 B.P. Thus Japanese anthropologists believe that “typical Mongoloids” developed the following physical characteristics in a questionably short span of time: medium height, relatively short limbs, medium pigmentation, less body hair, flat face, expanded cheekbones, thick subcutaneous facial fat, heavy eyelids, and narrow eyes (Hanihara 1986, 1996, 1997; Matsumura 1996).

Although Japanese anthropologists still accept Coon’s (1965) description of a “typical Mongoloid type,” this profile is now considered to be old-fashioned and is largely discredited in the U.S. Because many Japanese anthropologists have accepted the hypothesis that “typical” physical characteristics were caused by natural selection, for example via a frostbite selection model (Coon, Garn, and Birdsell 1950), further examination as to why or how these traits developed and spread has been stifled. This scheme in which all Asians and their ancestors are “Mongoloids” has been extremely influential within contemporary Japan, although it is impossible to determine some physical features such as the shape of eyelids, hair texture, and skin color from skulls.

For example, Southern “Mongoloids” exhibit large eyes, wide noses, square faces, and thick eyebrows and beard in contrast to the Northern “Mongoloids” (Hanihara 1991, 1996, 1997). This stereotype, however, is not universally true and several new studies now claim that modern Japanese represent a mixed population from Northern and Southern groups. Thus the use of racial terminology is not considered to be a bad thing within anthropology in Japan, and it is thought that racial terms can be used for convenience and scientific priority, although Japanese anthropologists often state that they do not believe in the biological concept of “race.”

Brace and Seguchi presented a paper entitled “Race Is Not a Valid Biological Concept” (2003) at the 2002 Inter-Congress Symposia of the International Union of Anthropological and Ethnological Sciences in Tokyo. Brace described the realm of human variation by clinal distribution of several adaptive traits (tooth size, hemoglobin S, and skin color) and non-adaptive traits (craniofacial morphology and mtDNA) which form clusters as “family resemblance writ large,” and presented the case that the concept of “race” did not emerge from the study of human biological variation and could only hinder our attempts to understand the various aspects of human biology (Brace and Seguchi 2003). The presentation strongly impacted Japanese anthropology. For example, in this same symposium a Japanese geneticist, Dr. Naruya Saitou, recalled that Brace had criticized him for the use of racial terminology when he described the Asian people as “Mongoloid” at the Pacific-rim archaeological international conference in 1989. He said that Brace’s criticism had made him think more about the use of the concept of “race” and that, since then, he has not used racial terminology and now refers to people by their geographic locale.

In addition to showing that the Jomon of prehistoric Japan cluster with the Ainu and deserve recognition as their ancestors, Brace’s cranial and dental data demonstrate that the Polynesians also cluster with the Jomon and Ainu, and that Bronze Age Mongolians fit into this cluster along with Northern Plain Native Americans. His data further suggest that there was a northern continuity across the Old World that goes back to pre-modern roots in the Pleistocene, which explains why the Ainu look somewhat like Europeans, as do northern Plains Native Americans and some Polynesians (Brace et al. 2001). Happily, Brace continues to engage in research on the peopling of the New World.

The “problem” with Neanderthals

“When Mimi and Loring traveled in Yugoslavia to study the Neanderthal remains from Krapina, Mimi claimed that the potent national plum brandy was so flammable and cheap that it made an ideal substitute for very expensive anti-freeze for their automobile.”
–Margaret Hamilton
“Yes, we did use Slivovica as anti-freeze on occasion; but most often we could not bear to lose it to the cooling system in our little French car, and wound up drinking it instead.”
–Loring Brace

Brace brought his expertise in the history of science, evolutionary processes including natural selection and PME, and cultural evolution to bear in his famous analysis of “classic” Neanderthals (Brace 1964b); he continues to view the general theoretical model that he presented in that paper as valid (Brace 2000a). The “classic” Neanderthals lived in Western Europe from about 125,000–35,000 years ago. Dramatic features of the skull gave Neanderthals a rugged, some say “brutish,” look. These included thickened bony arches over the eye sockets (brow ridges), the lack of an indentation (known as the “canine fossa”) below the cheek bone, and a protuberance at the back of the skull (the “occipital bun”). Neanderthal faces were also characterized by large, worn anterior teeth in jaws that jutted forward (“midfacial prognathism”), and they had very large brains as reflected in their cranial capacities. From the start, these odd-looking hominids were interpreted as an evolutionary dead end because of biases of European workers, especially the French Professor, Marcellin Boule, who unwittingly focused his 1911–1913 interpretations of all Neanderthals on an inept analysis of one badly reconstructed specimen known as La Chapelle-aux-Saints (Brace 1964b).

Brace notes that Boule and numerous subsequent workers were not concerned with the processes by which the characteristics of modern man developed because their primary purpose was to demonstrate that Neanderthals could not have left any descendants. They also hypothesized that Neanderthals were completely replaced by other, more presentable, hominids who swept into Western Europe from some remote, mysterious place. Brace rejected this view, which for historical reasons he dubbed “hominid catastrophism,” and chose instead to compare Neanderthal morphology to that of both earlier Homo erectus and later Homo sapiens, and to interpret the results in light of the archaeological evidence regarding cultural development. This led him to conclude that Neanderthals represent a stage of human evolution that preceded the transformation of the modern face, which transpired after Neanderthals invented tools that took over tasks formerly handled by their teeth. These tools relaxed selection for rugged dental/facial architecture, and the resulting PME resulted in smaller anatomical features.

Workers who balk at the idea of Neanderthals as our ancestors claim that anatomically modern people pre-date them in the fossil record, and Brace has used metric data to address this criticism. For example, although the Qafzeh and Skhûl samples from Israel are usually represented as “anatomically modern” Homo sapiens, Brace analyzed their dental metrics and found that the Qafzeh skulls show “modern” cranial morphology and large tooth size, while Skhûl shows intermediate cranial form and tooth size. In other words, the Skhûl forms are intermediate between “classic” Neanderthal and “moderns.” When Qafzeh and the “classic” Neanderthals are compared, the average gross dental dimensions are not distinguishable, although Qafzeh craniofacial morphology is different from that of the “classic” Neanderthals of Western Europe. The Qafzeh sample actually has a summary tooth size figure (ÓTS) of 1503 mm2, and it has larger teeth than “classic” Neanderthals with a ÓTS of 1415 mm.2 Brace notes that the tooth size reduction between Qafzeh and modern West Africans is 17%, exactly the same as the percentage reduction between “classic” Neanderthal tooth size and the Upper Paleolithic in Europe. Both dental and craniofacial form indicate that Qafzeh is a model for the ancestors of modern sub-Saharan Africans (Brace 1995a, 1995b, 1996a, 2000a). Once again, Brace explains the mechanism of mosaic evolution in the emergence of “modern” forms by biocultural interaction; cooking preparation and the use of projectiles lead to post-cranial reduction.

Brace details how Boule’s unflattering portrait and assertion that Neanderthals became “extinct without issue” made a lingering impression on the field that inspired not only the negative caricature of cave men one sees in popular culture, but also the paradigms that dominate paleoanthropology today. Thus Boule’s notion of sudden replacement by a species that arose elsewhere is recapitulated in the concept of “punctuated equilibria” (PE), offered by its proponents as an alternative to Darwin’s hypothesis of gradual change through natural selection (Eldredge and Gould 1972). Although PE is sometimes applicable (e.g., it worked for Gould’s snails), it fails to address the details of how and why new species arise elsewhere. Nevertheless, the concept has gained a toehold and is now overused to such an extent that “as was true over a generation ago, most of the students of human ‘evolution’ today spend far more time trying to deny the possibility of evolution than in trying to figure out what actually happened and why” (Brace 2000b:327). This realization was not a sudden (punctuated?) epiphany for Brace, but rather one that continued to develop as more examples came to light over the years (Brace 1981:423):

The current in modern paleontology that accepts the discontinuous nature of the stratigraphic record as indicative that the history of organic life has been discontinuous (Gould 1965), the emphasis on sudden, dramatic change (Gould 1974, 1978a, 1978b), and the stress on speciation events in small isolated groups remote from the area of consideration followed by sweeping take-overs, all are remarkably similar to the outlook that characterized nineteenth-century French paleontology. A century ago this contributed to the French rejection of the idea of evolution by means of natural selection. Its reemergence today in the writings of a generation of paleontologists who have not looked carefully into the traditions of the field in which they have been trained could well lead to its being designated “neocatastrophism” (Brace 1978:983). In view of what we can suspect is the lurking French connection, it would be appropriate here to recall what the French would say in this regard: Plus ça change, plus c’est la meme chose.

The idea that modern species that evolved elsewhere swept into areas to replace others is not only associated with the “Neanderthal problem.” Neocatastrophism is also alive and well among the advocates of the “OutofAfrica” or “Eve” hypothesis regarding the origins of anatomically modern Homo sapiens, as opposed to the view of “regional continuity,” of which Brace is an adherent. The latter model is also known as “multiregional continuity,” and because Brace outlined it in 1964 he has been credited with being “the intellectual ‘father’ of the multiregional continuity position” (Clark 1992:189). Nevertheless, Brace continues to prefer the term “regional continuity,” having noted that “the addition of two syllables to make it ‘multiregional’ and the removal of the mechanism previously suggested adds nothing to our ability to understand the dynamics by which those changes take place” (Brace 1992:1819).

According to the neocatastrophic school, in order for there to be species to do the “sweeping in,” there must be more rather than fewer species. Thus, as Brace points out (1988), PE enthusiasts envision more rather than fewer branches (known as clades) on evolutionary trees. The preference for such “bushiness” has been facilitated by the assent of cladistic analysis as a preferred method for studying phylogeny, a predilection for typological thinking, and, some would say, the ego-driven desire of paleoanthropologists to name new species. Brace has traced these ideas back to Boule in what he terms “the great leap backwards” (Brace 2000a:150), and has shown how they ultimately resulted in recognition of too many “new” species of hominids—termed “nominophilia” by Brace—by the “splitters” as opposed to the “lumpers” in the profession. Brace’s good friend from the Department of Homopathic Anthropopoetics of the University of Southern North Dakota at Hoople, I. Doolittle Wright, succinctly sums up the matter (Brace 2000a:151152):

Consider the volatile splitter,
Excited and all of a twitter
At the fun in the game
Of adding a name
For each little variant critter.
If you make up a different name
For forms that just might be the same,
You can use the confusion
To support the illusion
That the effort will add to your fame.
Combining a fragment of face,
With a jaw from a far-off place,
They say they are able
To warrant a label,
Creating a whole new race.
But it’s not always praxis that leads
To the multiplication of breeds,
Nor the glory of God,
But conceit of the clod
With his ego-aggrandizing needs.
Now a lumper’s a different kind;
He thinks the world is designed
Of similar bits
That won’t addle the wits
In the limited scope of his mind.
“Specify!” somebody flutters;
“Simplify!” one of them mutters;
The lumpers and splitters
Will never be quitters,
So the field is a patchwork of splutters.
As names blossom forth anew,
There’s much to be said for the view
That puts its store
In “less is more”
And errs on the side of the few.

The stages of human evolution

The decades that Brace has spent attempting “to introduce an element of Darwinian processual thinking into a field that was traditionally dominated by a pre-Darwinian, typological mind-set” (Brace 2000b:332) have resulted in an overview of hominid evolution that is original, thoughtful, grounded in extensive data, and synthetic. Brace is truly a four-field anthropologist, and he writes with admirable clarity and good humor. Development of his ideas about what happened in the past and why may be traced through the five editions (so far) of his textbook, The Stages of Human Evolution (Brace 1995b), and many other publications.

As the title of his textbook implies, Brace envisions major developments, or stages, in the lifeways of hominids during the course of their evolution. It is now generally accepted that hominids split from the arboreal ancestors of chimpanzees around five to seven million years ago, and Brace believes that australopithecines, the first terrestrial hominids, from that time until about two million years ago were divided into a series of separate local species. “At that point, however, the rules of the game changed” and hominids that had previously scavenged carcasses of large animals actually began to hunt them, i.e., they joined the “large carnivore guild” (Brace 2000b:338). This development opened up a whole new cultural ecological niche that involved persistence hunting, with associated hairlessness and sweaty skin, and, importantly, the invention and spread of butchering tools. Brace interprets both the distribution of stone tools over the Old World and the many lifeway similarities between wolves and hunting-and-gathering human populations as strong support for believing that at any one time from the late Pliocene (Homo erectus) up to the present (Homo sapiens) hunting hominids constituted a single species. Thus:

Entering the niche of the large carnivore guild as a diurnal hunter provided circumstances that led to the transformation of Australopithecus into Homo, and that unlikely event almost certainly occurred only once. Subsequently it was the creation of and entry into the realm of the cultural ecological niche that generated what we would now regard as the fully human. The manufacture of tools is not simply a manifestation of a genetically encoded set of instructions designed to result in their fabrication. Tools are not independently invented and produced by each generation. Instead, they represent the continuity of learned behavior transmitted from one generation to the next. In this sense, tools are cultural elements in the anthropological sense. [Brace 2000b:345]

Brace’s third stage is that of Neanderthals (Homo sapiens neanderthalensis) and, as discussed above, he suggests that their distinctive facial morphology was correlated with biological and cultural factors that influenced dentition. Over time, the adaptive significance of their large dentition was reduced, thus bringing PME into play and ultimately transforming the once-rugged face into that of more gracile Homo sapiens sapiens, the fourth of Brace’s stages. But cooking, use of fire, and inventions of more sophisticated tools such as hand-axes are not the only cultural innovations that caused the transformation from erectus into sapiens. Brace notes that around 200,000 years ago human brain size had reached its modern level, fire was controlled, and “local stylistic embellishments became evident on what was functionally the same kind of tool kit” (2000b:361). He also thinks that, although it did not occur in a “flick of a switch,” language had begun by then, and that “at this point we can recognize the beginning of Homo sapiens. I am also suggesting that this marks the beginning of culture as a full-scale ecological niche in and of itself” (Brace 2000b:362).

Brace’s legacy

Brace is frequently quoted as saying that his goal is to bring physical anthropologists “kicking and screaming” into the (now) 21st century (Ferrie 1997). So how is he doing? Our assessment is that physical anthropologists are definitely kicking and screaming, and that Brace is slowly but surely making progress. For example, on the Darwin front, a recent appraisal from the point of view of current philosophy fully concurs with Brace’s observations about Darwin’s current standing and his defense of Darwin as a solid scientist (Dennett 1995).

Anthropologists have written several partial critiques of the concept of PME, and Brace has responded to these in various publications that account for the reduction that occurred during human dental evolution, such as the wonderfully titled “What Big Teeth You Had, Grandma! Human Tooth Size, Past and Present” (Brace, Smith, and Hunt 1991). Although anthropologists may have been less than enthusiastic, PME has fared much better with geneticists (recall Ken Weiss’s belief that “PME stands in principle”). One of the most persuasive arguments in favor of PME is that:

[S]o far, there has been no plausible alternative explanation proposed to account for why so many aspects of the human physique have undergone reduction immediately after specific tools and procedures had been devised that reduced the amount of sheer brute force that had formerly been necessary for human survival. [Brace 2000a:154 155]

Building on earlier work with Frank Livingstone (Brace and Livingstone 1975) and Ashley Montagu (Brace and Montagu 1977), Brace’s views on “race” have, to date, perhaps had the greatest impact of all of his contributions to the field. Simply put, many anthropology textbooks have changed the way that human variation is described and have incorporated Brace’s suggestion that geographical names be used to identify different human groups in place of typological labels. With the 2005 publication of his book, “Race” Is a Four-Letter Word, his views are gaining even wider acceptance in the United States and are in the process of becoming the norm elsewhere, such as in Japan.

Alas, the reception for Brace’s view of Neanderthals as ancestors of contemporary Homo sapiens has met with less enthusiasm, although some workers have back-pedaled a bit from earlier catastrophic models and now hold that Neanderthals may have been gradually displaced to more marginal environments, i.e., that they “probably went out with a whimper, not a bang” (Stringer and Grün 1991). One reason Neanderthals continue to be regarded by many as evolutionary dead-ends is because of recent studies on mitochondrial DNA that was extracted from one bone of the original Neanderthal specimen (Krings et al. 1997, 1999). Brace, however, has done an admirable job of addressing these studies (2000a:97–98).

The concept of punctuated equilibria also continues to be trendy, for lack of a better word, no doubt because it helps justify naming new species, Brace’s “nominophilia.” Although Brace’s hypothesis of one widespread species of Homo erectus, his second stage of human evolution, flies in the face of nominophilia, it is worth noting that other workers are beginning to agree that “less is more” when it comes to Homo (Asfaw et al. 2003; Conroy 2002, 2003). Thus, there is a curious kind of disconnect in the reception of Brace’s stages of human evolution: Although the Neanderthal (third) stage is viewed by many with skepticism, the first australopithecine stage is generally accepted, the second Homo erectus (formerly Pithecanthropine) stage is gaining more credibility, and the fourth Homo sapiens sapiens stage is universally embraced (emBraced?).

This brings up another disconnect, which has to do with the application of Brace’s findings regarding clinal variation for numerous adaptive traits among humans. First of all, Brace’s enormous contribution in diligently amassing the world’s largest single database of craniofacial and dental measurements has been widely acknowledged. W. W. Howells put it succinctly: “Brace can be heartily thanked for producing a large amount of primary data” (1980:154); and to that Brace has added data collected during an additional quarter century! As noted, the field has accepted and incorporated Brace’s suggestions that resulted from his findings regarding clinal variation when it comes to naming living human groups. What is puzzling, however, is the extent to which this same field resists applying clinal thinking vertically within the fossil record (i.e., viewing traits as graduated over time rather than horizontally across geographical regions). But then nobody ever said it was going to be easy to drag those kicking and screaming anthropologists into the 21st century!

As graduate students, we knew of Brace’s groundings in philosophy and the history of science, his elucidation of the probable mutation effect (PME), his infatuation with teeth and skulls, his love of cooking technology (and cooking), and, perhaps most of all, his seminal work on hominid catastrophism and Neanderthals. But what was not clear to some of us at the time was an understanding of the logical relationship between these apparently diverse interests and their implications for achieving a comprehensive overview of the mechanisms and causes of human evolution. As we learned more about the scope of Brace’s contributions during preparation of this retrospective, it dawned on us that such an overview is exactly what he has achieved. This brings to mind those telling words attributed, rightly or wrongly, to Mark Twain: “When I was a boy of fourteen, my father was so ignorant I could hardly stand to have the old man around. But when I got to be twenty-one, I was astonished at how much the old man had learned in seven years.” Brace’s first doctoral student, Stephen Molnar, observes that “Loring was way ahead of his time when he published the hominid catastrophism paper—and he still is today.” Fittingly, on March 10, 2006, C. Loring Brace received the Charles R. Darwin Lifetime Achievement Award from the American Association of Physical Anthropologists. 


1. Brace directed the following dissertations: Patricia S. Bridges (1985): A Biomechanical Analysis of Two Prehistoric Amerindian Groups: Changes in Habitual Activities and the Division of Labor with the Transition from Hunting–and–Gathering to Agriculture; Dean Falk (1976): External Neuroanatomy of the Cercopithecoidea; Sonia E. Guillen (1992): The Chinchorro Culture: Mummies and Crania in the Reconstruction of Preceramic Coastal Adaptation in the South Central Andes; Margaret E. Hamilton (1975): Variation Among Five Groups of Amerindians in the Magnitude of Sexual Dimorphism of Skeletal Size; Robert J. Hinton (1979): Influence of Dental Function on Form of the Human Mandibular Fossa; Kevin D. Hunt (1989): Positional Behavior in Pan Troglodytes at the Mahale Mountains and the Gombe Stream National Parks, Tanzania; Carol J. Lauer (1976): Determinants of Troop Movement Patterns in Free-Ranging Rhesus Monkeys; Paul E. Mahler (1973): Metric Variation in the Pongid Dentition; Stephen Molnar (1968): Some Functional Interpretations of Tooth Wear in Prehistoric and Modern Man; A. Russell Nelson (1998): A Craniofacial Perspective on North American Indian Population Affinity and Relations; Conrad B. Quintyn (1999): The Populational Affinities of the Qafzeh and Skhul Hominids; Karen R. Rosenberg (1986): The Functional Significance of Neandertal Pubic Morphology; Alan S. Ryan (1980): SEM Studies of Anterior Tooth Microwear; Margaret J. Schoeninger (1980): Bone Strontium Levels and Diet in Prehistoric Human Populations; Noriko Seguchi (2000): Secular Change in the Japanese Occlusion: The Frequency of the Overbite and its Association With Food Preparation Techniques and Eating Habits; B. Holly Smith (1983): Dental Attrition in Hunter-Gatherers and Agriculturalists; Frank Spencer (1979): Ales Hrdlicka M.D., 1869–1943. A Chronicle of the Life and Work of an American Physical Anthropologist; Kenneth M. Weiss (1972): Model Life Tables for Pre-Industrial Populations (dissertation committee co-chaired by Brace and F. B. Livingstone); Richard G. Wilkinson (1970): Prehistoric Biological Relationships in the Great Lakes Region; and Lucia Allen Yaroch (1994): Characterization of Neanderthal Cranial Shape Using the Method of Thin-Plate Splines.

Brace also served on the doctoral committees of the following people: William Babler (1977), Stephen M. Bailey (1980), Kari Leigh Brandt (1996), John W. Eaton (1969), Robert B. Eckhardt (1970), David W. Frayer (1976), Lillian K. Gleibermann (1975), Leonard O. Greenfield (1977), Gregg F. Gunnell (1986), William L. Jungers (1976), William R. Leonard (1987), Marquisa La Velle Moerman (1981), Lorna Grindlay Moore (1973), Marcia Robertson (1984), Fred H. Smith (1976), Shelley L. Smith (1990), David P. Tracer (1991), and Timothy D. White (1977).

2. From a lifelines interview in Nature 425:349 (2003).

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