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Ernst Mayr, Pioneer in Tracing Geography's Role in the Origin of Species, Dies at 100

By Carol Kaesuk Yoon
The New York Times, February 5, 2005.

Dr. Ernst Mayr, the leading evolutionary biologist of the 20th century, died on Thursday in Bedford, Mass. He was 100.

Dr. Mayr's death, in a retirement community where he had lived since 1997, was announced by his family and Harvard, whose faculty he joined in 1953.

He was known as an architect of the evolutionary or modern synthesis, an intellectual watershed when modern evolutionary biology was born. The synthesis, which was described by Dr. Stephen Jay Gould of Harvard as "one of the half-dozen major scientific achievements in our century," reconciled Darwin's theories of evolution with new findings in laboratory genetics and in fieldwork on animal populations and diversity.

One of Dr. Mayr's most significant contributions was his persuasive argument for the role of geography in the origin of new species, an idea that has won virtually universal acceptance among evolutionary theorists. He also established a philosophy of biology and founded the field of the history of biology.

"He was the Darwin of the 20th century, the defender of the faith," said Dr. Vassiliki Betty Smocovitis, a historian of science at the University of Florida.

In a career spanning nine decades, Dr. Mayr, a professor emeritus of zoology at Harvard, exerted a broad and powerful influence over the field of evolutionary biology. His most recent book, "What Makes Biology Unique?: Considerations on the Autonomy of a Scientific Discipline" (Cambridge University Press), was published in August, one month after his 100th birthday.

Prolific, opinionated, provocative and dynamic, Dr. Mayr had been a major figure and intellectual leader since the 1940's. Setting much of the conceptual agenda for the field, he put the focus just where Charles Darwin first placed it, on the question of how new species originate.

Though Dr. Mayr will be best remembered as a synthesizer and promoter of evolutionary ideas, he was also an accomplished ornithologist. In fact, it was with the sighting of a pair of unusual birds that Dr. Mayr's long career in biology began in 1923 at age 19.

Dr. Mayr was born in Kempten, Germany, in 1904. While still a boy, he was instructed in natural history by his father, Otto, a judge. He quickly became a skilled birdwatcher and naturalist. Intending to become a medical doctor like others in his family, Dr. Mayr was about to leave for medical school when he spotted a pair of red-crested pochards, a species of duck that had not been seen in Europe for 77 years.

Though he took detailed notes, he could not get anyone to believe his sighting. Finally, he met Dr. Erwin Stresemann, then the leading German ornithologist, who was at the Berlin Zoological Museum and who recognized his talents and invited him to work at the museum during school holidays.

After two years of medical studies at the University of Greifswald (chosen because it was in the most interesting German region for birdwatching), Dr. Mayr, like Darwin before him, opted for natural history. He completed his Ph.D. at the University of Berlin in just 16 months.

Dr. Mayr went on to fulfill what he called "the greatest ambition of my youth," heading off to the tropics. In the South Pacific, principally New Guinea and the Solomon Islands, Dr. Mayr collected more than 3,000 birds from 1928 to 1930. (He had to live off the land, and every bird, after being skinned for study, went into the pot. As a result, he is said to have eaten more birds of paradise than any other modern biologist.)

The South Seas experience, he once said, "had an impact on my thinking that cannot be exaggerated." For it was his detailed observations of the differences among geographically isolated populations that contributed to his conviction that geography played a crucial role in the origin of species.

Though Darwin titled his book "The Origin of Species," little in the book, in fact, addresses the question of how new species arise. Dr. Mayr determined that when populations of a single species are separated from one another, they slowly accumulate differences until they can no longer interbreed. Dr. Mayr called this allopatric speciation and detailed his arguments in his seminal book "Systematics and the Origin of Species," published in 1942. Today allopatric speciation ("allo," from the Greek for "other," and "patric," from the Greek for "fatherland") is accepted as the most common way in which new species arise.

"Organic diversity had at last received a convincing explanation," Dr. Jerry A. Coyne, an evolutionary biologist at the University of Chicago, wrote of Dr. Mayr's arguments. Dr. Coyne called the book "one of the greatest achievements of evolutionary biology."

Similarly, the most commonly held view of what constitutes a species remains the one that Dr. Mayr promoted more than 50 years ago, known as the biological species concept. First explicitly defined by Dr. Theodosius Dobzhansky, it states that populations that can successfully interbreed are the same species and those that cannot are different species. While numerous other species concepts have been proposed and debated, this one continues to reign supreme.

Dr. Mayr's focus on species, both their nature and their origins, appears to have derived from his experiences in the South Pacific. When he went to New Guinea, Dr. Mayr once explained in an interview with Omni magazine, there was a popular school of thinking known as the nominalist school of philosophy that held that species did not, in reality, exist. They were merely arbitrary categories, little more than names.

"But I discovered that the very same aggregations or groupings of individuals that the trained zoologist called separate species were called species by the New Guinea natives," Dr. Mayr said. "I collected 137 species of birds. The natives had 136 names for these birds - they confused only two of them. The coincidence of what Western scientists called species and what the natives called species was so total that I realized the species was a very real thing in nature."

Dr. Mayr eventually became a living symbol of the beginnings of the modern field of evolution, one of the last survivors of the handful of biologists, including Dr. Dobzhansky and Dr. George Gaylord Simpson, known as the architects of the evolutionary or modern synthesis.

In the evolutionary synthesis, neo-Darwinism took its place as today's dominant theory of evolution. Taking place between the 1920's and 50's, the synthesis is recognized as a period of conceptual unification, a time of "mutual education," as Dr. Mayr once described it. Laboratory geneticists, studying mutations and population genetics, began merging their views of evolution with those of field scientists like Dr. Mayr who studied the diversity and origins of different species. New findings, in genetics as well as other fields, were reconciled with Darwin's theories of evolution. Competing theories, including Lamarckism (the idea that acquired characteristics can be inherited), were tossed aside, producing a much more unified view of evolution at work.

Over his remarkably productive career, Dr. Mayr wrote or edited 20 books and wrote more than 600 journal articles. After his official retirement in 1975, he published more than 200 of the articles, more than many scientists do in their entire careers. He received awards including the National Medal of Science, the Balzan Prize and the International Prize. He once noted that Nobel Prizes were not given in evolutionary biology, saying, "Darwin wouldn't have won it either."

Dr. Mayr was an ardent promoter of the academic discipline of evolutionary biology, and perhaps its most energetic organizer, playing a critical role in founding the Society for the Study of Evolution in 1946, and serving as the first editor of its journal, Evolution, still the leading journal in the field.

Meanwhile, his birds were never forgotten. As a collector, ornithologist and curator, first at the University of Berlin, then the American Museum of Natural History in the 1930's and 1940's, and finally at the Museum of Comparative Zoology at Harvard, Dr. Mayr made his mark.

By the time he turned 90, in 1994, he had named more than 24 valid bird species, more than any other living ornithologist had at the time. He had named more than 400 subspecies and several new genuses of birds as well.

Dr. Mayr also took a serious interest in organisms other than birds, publishing work on species delineations in plants, hybrids formed by snail species, courtship behavior in fruit flies and the evolution of human blood groups.

Dr. Mayr may have taken the greatest pride in his theory of what he called peripatric speciation and genetic revolutions, an idea he called "perhaps the most original theory I have ever proposed." It was also his least successful.

According to this controversial theory, new species can be produced when very small populations are cut off from the rest of the species. Unlike the more general theory of allopatric speciation, holding that isolated populations slowly accumulate differences until they can no longer interbreed, peripatric speciation posits that extremely small populations, isolated in unusual habitats, undergo what Dr. Mayr termed a "genetic revolution." Undergoing drastic changes in their genome, populations evolve quickly to become new species.

Some scientists have said that this theory is unlikely, unsupported and untestable. Others have defended it as a proposal, saying that while the idea itself may not stand the test of time, it remains significant as one of the first explicit theoretical models of speciation and its genetic consequences.

Dr. Gould also credited Dr. Mayr with sowing the seeds for the "flowering of modern macroevolutionary theory."

While microevolutionary theory seeks to explain how species adapt to particular environments or how evolution among populations can give rise to new species, macroevolution theory encompasses a much bigger picture, examining how some species survive better than others and how likely or unlikely they are to give rise to other species. It was Dr. Mayr's concept of the species and its role in the evolutionary process, Dr. Gould said, that laid the foundations for many of the theories being tested by macroevolutionists today.

In addition to his several lifetimes' worth of work in evolution, Dr. Mayr also fathered an entirely new field of study, creating almost singlehandedly the field of history and philosophy of biology as a distinct discipline, apart from the history of physics and chemistry, Dr. Smocovitis of the University of Florida said.

As with modern evolutionary biology, Dr. Mayr nurtured the new discipline as organizer, mover and shaker. His own contributions include the defining book "The Growth of Biological Thought" (1982), as well as books on the philosophy of biology, Darwin and the evolutionary synthesis.

Dr. Mayr is survived by two daughters, Christa Menzel of Simsbury, Conn., and Susanne Harrison of Bedford; five grandchildren; and 10 great-grandchildren. His wife, Margarete Simon, died in 1990.

Dr. Mayr, a strong believer in the Hegelian dialectic as a way of advancing understanding, was known for his definitive proclamations, which often inspired as many heated rebuttals as nods of vigorous agreement. With so long to consider the great pageant of the history of life, he seemed to have taken on every subject of interest in evolutionary biology, and his views are an unavoidable point at which to begin nearly any argument of substance.

At the time of his 90th birthday, in 1994, when Dr. Mayr was as active and engaged in the field as ever, Dr. Douglas J. Futuyma, an evolutionary biologist at Stony Brook University, wrote, "No one will agree with all his positions, analyses, and opinions." But he added, "Anyone who has failed to read Mayr can hardly claim to be educated in evolutionary biology."

Ernst Mayr (1904–2005)

By Jared Diamond
Nature 433 (February 17, 2005): 700-701.

One Sunday in 1953, my father, a physician and haematologist at Harvard University, invited a newly recruited colleague to lunch at our house. Dad had just launched a study of possible associations between human blood groups and diseases with him. At that time, scientists widely assumed that blood groups were ‘selectively neutral’ — that is, that they had no effect on human survival. Our guest was an evolutionary biologist who suspected that they must have some effect, perhaps one far removed from blood’s familiar functions. Dad and his co-author went on to discover an association between ABO blood groups and stomach cancer, one of the first studies to show that blood groups are indeed influenced by natural selection. The co-author was Ernst Mayr, widely regarded as the greatest evolutionary biologist of the twentieth century, who died on 3 February 2005.

When I met Mayr that Sunday, I was a 16-year-old schoolboy. He later inspired me to launch a second career, parallel to my work as a membrane physiologist, on the evolutionary biology of New Guinea birds, his own early speciality. For 30 years he and I collaborated on analysing a mammoth database that he had accumulated on the distributions of island birds. The result was a co-authored 556-page book published soon after his 97th birthday. That Sunday lunch and its consequences illustrate many keys to Mayr’s greatness: his capacity for close friendships and collaborations with younger scientists as well as with peers; his broad perspective that let him recognize new significance in the work of many specialists; and his capacity for sustained hard work and complex analysis.

The achievements for which Mayr is best known fall into six areas. First, as an ornithologist he was the leading expert on birds of New Guinea and the tropical southwest Pacific; he described more species and subspecies of living birdsthan anyone else of his or subsequent generations. Second, as a systematist he was a principal architect of what is termed the ‘evolutionary synthesis’, which finally succeeded in showing how the adaptive changes that natural selection produces in single populations result in the evolution of biodiversity. That synthesis fused the hitherto separate research programmes of geneticists and field naturalists, and explained how evolution has given rise to organisms ranging from microscopic bacteria to redwood trees.

In that process lies Mayr’s third major achievement, his assembly of overwhelming evidence that most species are not collections of individuals arbitrarily delineated by taxonomists but real entities: “a group of actually or potentially interbreeding natural populations reproductively isolated from other such populations”, to quote his widely cited formulation. He also demonstrated that new species of birds and mammals arise through allopatric speciation (geographical isolation of initially conspecific populations) — thereby in effect solving the problem of the origin of species that had eluded Charles Darwin despite the title of Darwin’s great book.

Fourth, as a biogeographer Mayr’s studies of a fauna’s composition, origins, history and boundaries have served subsequent biogeographers as models for testing their own results. Fifth, as an evolutionary biologist, Mayr traced in detail the combined operation of population genetics and evolutionary processes in diverse phenomena throughout the animal kingdom, as illustrated by the study of blood groups that resulted in my meeting him.

Finally, as a historian and philosopher of science, in recent decades Mayr clarified the regularly misunderstood central concepts of biology: teleology; the foundations of biological classification; proximate and ultimate causation; the special problems posed by historical sciences to which experimental methods cannot be applied; and the distinctness (autonomy) of biology as a science.

The facts of Mayr’s career can be briefly summarized. He was born in Germany on 5 July 1904, at a time when evolution’s co-discoverer, Alfred Russel Wallace, was still alive. On weekends, his parents took Ernst and his brothers on walks to observe animals, plants and fossils, among which birds in particular kindled his interest. After a rigorous high-school education in Dresden, Ernst obeyed Mayr family expectations by preparing for a medical career and completed his preclinical studies in 1925. However, his observation and careful description of a pair of a rare duck, last recorded in Germany 77 years earlier, led to his introduction to the Berlin ornithologist Erwin Stresemann. Recognizing Mayr’s talent, and also his thirst to visit the tropics, Stresemann offered Mayr two irresistible enticements: a position in the Berlin Museum, and prospects of a birdcollecting trip to the tropics, if Mayr could complete an entire PhD programme within 16 months.

Mayr accepted the challenge, worked 16 to 18 hours a day to receive his PhD in 1926, and took up the promised museum position. In 1928 Stresemann, now armed with money from Lord Rothschild and from the American Museum of Natural History (AMNH) in New York, delivered on his second promise by sending Mayr to the southwest Pacific for more than two years. The instructions given to Mayr were to explore five New Guinea mountain ranges, to solve the long-standing mystery of New Guinea’s apparently rarest birds of paradise (he did, and they proved to be hybrids), and to collect birds on islands in the Solomon group that had been considered too dangerous to visit by previous collectors. Mayr succeeded beyond everyone’s expectations.Having re-explored six of those mountain ranges and islands between 1974 and 2004, under the less-threatening conditions of the late twentieth century, I can testify that they are physically gruelling even today. Mayr managed to amass comprehensive bird collections there from 1928 to 1930, despite the perils of diseases, capsized canoes, forced descents of waterfalls and periodic threats of natives to kill him.

Soon after his return from New Guinea, in 1931 Mayr was appointed by the AMNH to curate the museum’s overflowing collections of Pacific island birds. For the next decade, all of his publications were technical taxonomic studiesof birds, giving few signs of his broader interests, until the publication in 1942 of his first book, Systematics and the Origin of Species, which completed the evolutionary synthesis. In 1953, a desire for contact with students and for wider intellectual horizons led Mayr to move to Harvard, as Agassiz Professor of Zoology, where he also served as director of the Museum of Comparative Zoology from 1961 to 1970. Following his official retirement from Harvard in 1975, he continued to publish with undiminished productivity. Fourteen of his 25 books were published after the age of 65, including one of his most important ones, The Growth of Biological Thought, which appeared when he was 78.He celebrated his 100th birthday with the publication of What Makes Biology Unique? in 2004. Whenever I talked to him during his nineties, I would ask him: “How many books are you working on now?” The answer was never less than two nor more than four.

What accounted for Mayr’s remarkable originality and productivity? I came to realize that there wasn’t a single explanation but the combination of a dozen of them — cognitive, organizational, emotional and social. Among the cognitive ones, he had an outstanding memory. When, in 1965, 24 years after the peak of Mayr’s work on New Guinea birds, John Terborgh and I asked him to identify the stuffed bird specimens that we had just collected in New Guinea, we saw that, for each of the 1,400 species and subspecies of birds that he had discussed in his 1941 Checklist of New Guinea Birds, Mayr still remembered who had described it — and when and in what journal, its differences from its relatives, and its alternative names. To that memory for facts wereallied outstanding visual recall (for example, he was alert to slight subspecific differences between bird specimens seen at different times in different museums) and auditory recall (the ethologist Klaus Immelmann related how, while he and Mayr were sitting on a garden bench in Germany in the 1970s, Mayr correctly identified a brief call note of an unseen bird as a grey wagtail,which he had not encountered since leaving Germany 40 years previously).

Mayr was also a quick learner: in the month before he reached New Guinea in 1928, he learned to speak Malay and Neo-Melanesian, to shoot a gun, and to skin and stuff birds. Like Darwin, he was a constantly curious field observer; also like Darwin, his wide interests let him reinterpret the work of specialists, as he did with my father’s data on blood groups. In my own collaboration with him, I was struck by his comfort with complexity: unlike many other scientists, he did not force facts into a one-factor explanation, but acknowledged the possibility of different multi-factor outcomes (such as different evolutionary trajectories for different bird populations).

Mayr himself spoke of his Sitzfleisch or capacity to stick to a job, just as the composer J. S. Bach attributed his prodigious musical output to mere Fleiss (industriousness). During Mayr’s years as a museum director at Harvard, a job that absorbed his daytime hours, he maintained his scientific output by writing each morning from 4:30 until 7:30 a.m., then spending the evening reading. In the 16 months that it took him to complete his PhD by age 21, he took all of the required courses in zoology, learned and passed an exam in botany, completed a minor in philosophy, and researched and wrote his thesis. When he arrived at the AMNH on 20 January 1931, he was given a one-year appointment with the understanding that reappointment would depend on productivity. He published his first paper two months later (a reclassification of kingfisher subspecies based on measuring hundreds of specimens), and finished 11 more papers by the year’s end. (That convinced the AMNH to renew his appointment.)

Despite not visiting an English-speaking country until his twenties, Mayr mastered English as a second language to the point where his English prose style was widely admired for its clarity. In addition to publishing 25 books and more than 700 papers and directing Harvard’s museum for nine years, he designed the AMNH’s bird exhibit hall, integrated the Rothschild collection of 280,000 bird specimens into the AMNH collection, and edited the last eight volumes of the Checklist of the Birds of the World (a critical reassessment of all bird taxa down to the subspecies level). Each of those ‘additional’ achievements was a mammoth undertaking in itself.

Mayr was self-confident without being overconfident. And he could change strongly held views when presented with new evidence, as when he abandoned his initially lamarckian belief in the inheritance of acquired characteristics. His confidence in his abilities included recognition of their limitations: for instance, he resisted friends’ suggestions that he expand his 1963 book Animal Species and Evolution to include plants and microorganisms, because of his insufficient familiarity with them. Those limitations also involved mathematics beyond algebra, which he did not use. He maintained a low opinion of the value of the cladistic methods now dominant among taxonomists. That contributed to the distance, in his later years, between his views and those of some other evolutionary biologists now active. They felt that his work belonged to the past; he felt with exasperation that they ignored much of the knowledge already gained in the past.

A widespread misconception is that great scientists tend to be loners. Actually, outstanding success in most areas of science requires outstanding social skills, as illustrated by Mayr’s relationships with a wide variety of people. He achieved such good understanding with New Guinea and Solomon tribespeople in the 1920s that they not only led him in and out of areas where other Europeans feared being killed,but they also taught him their local names for birds and brought him hundreds of specimens of bird species missed by other European collectors. He once explained to me that a secret of living happily past age 90, after most friends of the same generation have died, is the continued willingness to forge friendships with younger people. All of these qualities contributed to Ernst Mayr’s scientific greatness and his productivity. They also lie at the root of the love felt for him by several generations of colleagues and friends.

Jared Diamond

Jared Diamond is in the Department of
Geography, University of California, Los Angeles,
California 90095-1524, USA.

Obituary: Ernst Mayr 1904-2005

By Michael Ruse
Biology and Philosophy, 20 (2005): 623–631.

Ernst Mayr has died at the very old age of one hundred. It is hard to imagine that he is gone. I remember back in 1970 at a conference in Indiana, where Mayr rather dominated the discussion. I thought at the time: "Well, we won't have to put up with this much longer." It was not the last time or the only way in which I was very much mistaken about Ernst Mayr.

Ernst Mayr was born in Germany, in Bavaria. His father was a lawyer, but the family tradition was medicine and it was that profession towards which young Ernst was directed. He was a keen bird watcher and, through this, was diverted to a lifetime career as a professional biologist. German universities when Mayr was a young man were incredibly hierarchical. His professor (Erwin Stresemann) told him that, because there was another bright young scholar (Bernhard Rensch) ahead of him, there was little chance of the coveted professorship, and that hence he should seek other pastures. Mayr went west, and for the next two decades worked at the American Museum of Natural History in New York.

For many reasons, this was an incredibly fortunate move. First, he was now working with one of the greatest collections of bird skins in the world. He was able to study variation in detail and range in a way that was impossible for virtually anyone else. He met and became close friends with a group of men who were determined to put evolutionary studies on a new and proper foundation. Most important of all was the Russian-born geneticist Theodosius Dobzhansky, closely followed by the brilliant paleontologist George Gaylord Simpson. He met Greta—a fellow German—who was to become his wife. And, more than anything, he missed the rise of the Nazis, and the corruption that that brought to his homeland.

As a student in Germany, Mayr had been rather disdainful of evolution. I do not mean that he was not an evolutionist—he never had any doubts on that score. It was just that he did not really think it a proper subject for a professional biologist. This may seem strange, and even stranger is the fact that the man who was to become one of the dominating figures of neo-Darwinism was at this time a committed Lamarckian—he believed that the main causal force was the inherited of acquired characteristics. In fact, the odd thing would have been if Mayr had thought evolution worthy of study by an ambitious young biologist, or had he been other than a Lamarckian. For all that Darwin, back in the mid-nineteenth century, had discovered the mechanism of natural selection—the survival of the fittest—evolution had been captured by those (Herbert Spencer, Ernst Haeckel, Thomas Henry and grandson Julian Huxley) who were determined to make evolution into a kind of secular alternative to the dominant Christianity. It was at best an inferior science and at worst a vehicle on which any enthusiast could hang wild metaphysical theories and hypotheses. Selection was thought a minor factor in significant change, and Lamarckism was one of the favored alternatives.

Theodosius Dobzhansky changed all of that. He was captivated by the adaptive landscape metaphor of Sewall Wright, and used it as the basis of Genetics and the Origin of Species, his paradigm-making survey of the forces of organic change, first given as lectures at Columbia University in 1936 and then published as a book the year following. As everyone knows, Dobzhansky was an enthusiast, gathering up co-workers and students. Mayr was brought into the circle, and at Dobzhansky's urgings gave his own lectures, and published the result as Systematics and the Origin of Species in 1942.

By this time, everyone was starting to see selection as the most important force for change, and Mayr's book reflected that. His main contribution was to demonstrate beyond doubt the variation that exists in nature, and how everything points to gradual change—a key plank in the Darwinian program. Particularly important was the evidence of groups that range all the way from good species (interbreeding populations, isolated from others), to species-in the-making, with subgroups starting to break apart, and on to fully defined separate species. Although Mayr's actual examples have been challenged by later biologists, really significant were so-called "rings of races" where on has a chain of groups (sub species) touching and interbreeding, except at the ends that come together (completing the circle) where the touching groups behave as distinct species.

The work of Dobzhansky and Mayr and Simpson (who published the paleontological contribution, Tempo and Mode in Evolution, in 1944) collectively founded what came to be known in America as the "Synthetic Theory of Evolution" and in Britain as "Neo-Darwinism." (The botanical contribution was delayed. Edgar Anderson was supposed to write it—he gave lectures at Columbia at the same time as Mayr—but proved unable to produce the book. Dobzhansky then recruited G. Ledyard Stebbins, who published Variation and Evolution in Plants in 1950.)

One must take some care however to understand the synthetic theory and Mayr's contribution to it. Most obviously the theory is an updating of Darwin's theory in the Origin. There, Darwin offered what his mentor, the great historian and philosopher of science William Whewell, called a "consilience of inductions." First Darwin presented and justified his causal heart, evolution through natural selection. Then he used this to explain biological phenomena across a wide range of subjects—instinct, paleontology, geographical distribution, morphology, embryology, and more. He argued that the success in explanation in turn provides confirmation of the unifying mechanism.

Following Sewall Wright's work in population genetics, showing how Mendelism blends with selection, apparently Dobzhansky and the others worked in the tradition of Darwin, using population genetics (especially as incorporated in the adaptive landscape) as the updated version of Darwin's presentation of natural selection. Then, Dobzhansky having put empirical flesh on Sewall Wright's theoretical skeleton, the other synthesizers worked on the range—Mayr on biogeographical distribution, Simpson on paleontology, and so forth.

There is a lot of truth in this picture, although one should not over-emphasize the extent to which Wright and the early Dobzhansky were enthused by natural selection. Sewall Wright was (following both his own father and his Harvard teacher, L. J. Henderson) an ardent Spencerian, and for him upward progress and the balance between homogeneity and heterogeneity was what really counted. Selection was secondary, drift was highlighted, and the same was true for Dobzhansky in the first edition of Genetics and the Origin of Species. It was around 1940 that everything became a lot more selectionist, when Dobzhansky discovered synchronic chromosome changes in separate populations of fruitflies. Selection can explain these and drift cannot.

Mayr fits into this picture, but rather uncomfortably. The adaptive landscape model is there, but it is not dominant. Mayr used to deny (to me, very often) that he really cared about the model at all—he said that he just dropped it in to keep others happy. This is not quite true, but it is true that Mayr was not doing what Simpson was doing, namely trying to complete the neo-Darwinian picture as sketched above. Mayr was still fighting the battles of the 1920s in Germany. There, the big enemies of naturalists like himself were the geneticists. They were arguing that really significant change is caused by macromutations, jumps from one form to another. Mayr and his fellows knew that this could not be true—nature showed them that there was gradual change.

This was the battle that Mayr was still fighting in Systematics and the Origin of Species. In fact, Germany had come to New York. One of the leading scientific refugees from the Nazi oppression was Richard Goldschmidt, who was arguing strongly for macromutations and jumps—what are known in the trade as "saltations." Mayr took him on and wrestled him to the ground. In fact, there was a satisfying personal element here. Goldschmidt was a distinguished Herr Professor. Mayr was an upstart, a museum worker. (Hard to imagine him in this role, but it is true.) Personally they got on well—Mayr loved to tell the tale of taking Goldschmidt home one Saturday lunch for soup prepared by Greta—but fighting this battle was for Mayr very much (to change metaphors) very much a matter of winning his spurs.

The battle also helps to explain what always struck me as a rather curious feature of Mayr's attitude to his science. For all that genetics was fundamental to the synthetic theory, and for all that in the early 1950s Mayr himself extended Sewall Wright's thinking—Sewall Wright said that Mayr pinched Sewall Wright's thinking—about the significance for evolution of isolated groups that undergo genetic revolutions and rapidly become new species (the so-called "founder principle"), Mayr was forever rude about and belittling of genetics. Not molecular genetics, so much, as Mendelian genetics. He spoke of it contemptuously as "beanbag genetics" (causing J. B. S. Haldane to speak in favour of beanbag genetics), he excoriated its reductionistic nature, and somewhat pettily when in the early 1970s he organized two major conferences on the founding of the synthetic theory, he did not invite Sewall Wright.

It would be easy to say simply that it was all simply personal—for the record, I think the founder principle does come out of Sewall Wright's work, but that Mayr deserves full credit for its development and application (and, for all that today evolutionists are questioning its significance, that the principle has proved itself a very important guide to productive research). It would be truer to say that the roots of Mayr's hostility lies back in the battles when he was a student and genetics was the enemy of naturalists, rather than the foundation of modern evolutionary theory.

It would be unfair to say that after Systematics and the Origin of Species, the rest of Mayr's career was simply footnotes and appendices. Apart from the founder principle, he was a very important systematist, working on the theory of classification. It is true that around him grew up the new taxonomic approach of phylogenetic systematics or cladism, and Mayr could never buy into this. In fact, in respects he became the Goldschmidt for the new generation of Young Turks. But without genetics, Mayr could not have been what he is, and I suspect that without Mayr modern taxonomy could not be what it is. However, one does sense that after Systematics and the Origin of Species, to use yet another metaphor Mayr was skiing down the other side. In 1963 he published a major synthesis, Animal Species and Evolution, but it was more a synthesis of what had been done than a work to inspire for the future.

Conceptually that is. Because Mayr was now about to make an even more significant contribution to twentieth-century, evolutionary biology. The 1940s saw major moves by the evolutionists to bring workers together and to make a formal discipline from and for evolutionary studies—a journal, a society, grants, students, jobs, and the rest. Mayr was in the thick of things. He was the first editor of the new journal Evolution, and showing that he was German in more things than one, whether accepting papers or rejecting them, he never wrote a sentence when a paragraph would do. He ran the society. Once, somewhat sardonically he said to me: "Unlike my good friends Dobzhansky and Simpson, I never thought that the only position I could take in an organization was that of president." He moved to Harvard—the immigrant boy made good, as Dobzhansky once said of himself and Mayr—and worked unceasingly to promote organismic biology at that institution, at a time when the molecular biologists were all conquering, and not very nice about it either.

To quote Ed Wilson: "Jim Watson? The most unpleasant man I ever knew." (Actually, one of Mayr's daughters dated Watson for a while. Although more on this topic in a moment.) The fact is that without Ernst Mayr we would not have the professional discipline of evolutionary biology that we have today. I do not mean that nothing would exist—others were organizing, including Dobzhansky with his many students and (over in England) E B "Henry" Ford was also showing himlsef a master of organization and funding finding. But without Mayr, things would simply not be as well developed as they are now.

But Mayr had many more years of active life. Even last year he was scrounging one of my books from our shared publisher, Harvard University Press, so that he could put the boot into me one more time before he was done. By about 1960, Mayr was starting to move interests from pure science to its history and its philosophy. I know only second-hand about the history, although I do know that he was much involved in the development of the History of Science department at Harvard, he worked hard to promote the careers of young scholars (Frank Sulloway, notably), he himself organized the above mentioned conferences on the history of the synthetic theory (and then co-edited with Will Provine the collection of the proceedings) and Mayr was one of those who worked hard to get the Journal of the History of Biology off the ground.

Mayr himself was much interested in the history of biology and eventually was to write a massive work on the history of evolutionary biology, the only volume of what had been intended to be a complete history of the life sciences. When he had finished the evolutionary side, he realized that he had said what he wanted to say. For one should never think of Mayr as interested in history for its own sake. He was always writing for a purpose—a purpose that I would describe as philosophical.

As a student in Germany, Mayr had studied the classics of philosophy—and he came from the cultured middle-classes, where discussion of philosophical issues would have been normal rather than a sign of pretension—and so, at one level, philosophy was part of his very being. But, by about 1960, it became much more. It became a tool in his fight for his vision of the life sciences, where evolution took an honored and central place. (When Dobzhansky said that nothing in biology makes sense except in the light of evolution, he was not just making an epistemological claim. He was making a political statement. A war cry to rally the troops.)

I have said that I never sensed in Mayr's attitude towards molecular biology, the deep hostility he felt to the genetics of the 1920s. Early fights and hatred run more deeply and last forever. I have disliked people in my life, but never with the visceral hatred that I still feel towards my headmaster. (Mark you, he was late middle-aged, and I suspect his feelings about me prove the exception to what I have just said about early experiences causing the really deep antagonism.) However, whatever the ordering of the threats that Mayr felt, it is undoubtedly true that with good reason he could see that by 1959—the year of the centenary of the Origin and of the celebration of the triumph of neo-Darwinism—the molecular biologists were grabbing all of the goodies.

Not just the Nobel prizes, but the students and grants and research space and much more. The biology of whole organisms was being pushed aside. (Read Jim Watson's Double Helix if you want more proof.) Mayr was ever a fighter. He was not about to sit by and let this happen. He turned to philosophy to prove that organismic biology will always have a place in the sciences, that it cannot simply be subsumed under—absorbed into—the molecular sciences, of physics and chemistry. In other words, he started a fight against what he saw as unwarranted reductionism. Not because he was a Marxist (like Steve Gould and Dick Lewontin in the decades later), but because he wanted to preserve the autonomy of those parts of the life sciences that had evolutionary thought as central. If he could make his case, then biology as he knew it could continue to survive and flourish.

And so from Mayr, as from others working alongside him (notably Simpson), we got a stream of articles and writings and conference interjections promoting the autonomy of whole-organism biology. We learnt that (unlike the physical sciences) it has a differentiating historical dimensions, that it deals with unique phenomena, and that above all it is teleological—forward-looking. Although, in a clever move, realizing that teleology has a bad name in biology—Mayr himself battered would-be contributors to Evolution for teleological tendencies—he took up the name of "teleonomy." This is sanitized teleology—forward-looking, dealing with functions, without the unneeded vital forces and so forth.

Above all, Mayr was a holist, meaning that he thought breaking things down to small components is not only not necessarily the right way to go in biological science, it is often positively exactly the wrong way to go. For this reason, when Michael Ghiselin and David Hull in the 1970s began arguing that species are not classes—groups of member organisms—but individuals—integrated wholes—Mayr embraced their thesis with enthusiasm. And it was for this reason that Mayr wrote a lot of his history.

It is true that old battles were not forgotten. Those dreadful geneticists had ignored variation, the basis of Mayr's gradualism, and so a lot of the history was devoted to showing that that rotter Plato had illicitly introduced essentialism—the idea that groups have no variation—into biological thought. Only slowly and gradually, thanks primarily to Darwin and to a certain immigrant to the United States of America, had population thinking finally triumphed. But this history telling was only part of the story, for the important underlying message was that whole-organism thinking has a grand tradition and, as one recognizes this, one recognizes that such thinking has its own autonomous problems and ways of solution. Molecular biology is important, but only a molecular biologist would think it all important. And that in itself tells you something about their limitations.

Was Mayr successful in his efforts? Well, yes I think he was, although how much was due to Mayr himself and how much to others, and what the nature of that success are perhaps questions that yield answers Mayr would not entirely have liked. On the one hand, Mayr's labours in the history and philosophy of science—always as much organizational as conceptual—paid off greatly in the development of both the history and philosophy of biology as thriving enterprises as we have them today.

Mayr encouraged many of us, and helped in so many ways. He may have irritated me—I have never been much of a holist—but he encouraged me and said very kind things when my first book appeared. When I founded Biology and Philosophy, I knew that Ernst Mayr (and David Hull) would always write a report on a paper. I generally knew what he would say before I sent it to him, but it always came back with lots of useful comments. And I know that others have had the same experiences.

At the same time however, in history particularly, my suspicion is that generally the (justified) feeling is that Mayr's extra-historical motives distort his thinking to the point where what he wrote is of minimal value. Certainly the feeling among historians is that not only is the essentialism thesis just plain false, but that we have wasted a lot of valuable time getting over it. Perhaps this is just a reaction of the present, but I suspect there is some truth to it.

On the other hand, today organismic biology does have a full and honored place in the biological spectrum. Far from the molecular biologists swallowing up evolutionary biology, the molecular biologists have been turned on by the problems of evolution. Witness the thriving success of the field of evolutionary development (evo-devo). But this in a way is the very point. Evolutionary biology today is so exciting because of molecular biology, not in the face of it. Bring on the molecules, we all say. A rather different cry from that of the Ernst Mayr of the 1960s.

What sort of man was Ernst Mayr? I trust that Ed Wilson of Harvard and Guy Bush of Michigan State, both men who encountered Mayr at Harvard when they were very junior, will not think I am betraying confidences when I say that he was not always much fun for the young and less established. He could be formidable to the point of—beyond the point of—bullying. As is often the case, one suspects that this concealed certain insecurities—the foreigner in the new world, in a field that is under siege.

But it does not make for easiness, whatever. At times, Mayr was desperately—almost pathetically—eager to make his own place in history. He truly loved Dobzhansky and admired him greatly, but worried that he (Mayr) would always have second place. It is easy to joke about this, as I did above, but it has an edge nevertheless. He was deeply hurt that, at the Origin centenary celebration in Chicago in 1959, Dobzhansky, Simpson, and Julian Huxley were given honorary degrees and not him.

However, insecure or not, no one could deny that Ernst Mayr had a deep love of the living world. To walk with him around his cottage in New Hampshire was to see a man who simply loved animals and plants—loved them for their own sake, loved them because he was trying to understand them. Mayr was a man who could think of no higher calling. He told me that reading Ernst Haeckel as a child had turned him away from Christianity. He joked, when I admired a photo of him in his twenties in New Guinea, where he was wearing a beard, that people took him for a missionary and he shaved at once! He was much influenced by Julian Huxley's Religion without Revelation. He told me that he felt with Huxley and with others (including, very much, Wilson) that it is possible to be a deeply religious person in the complete absence of theology. He meant it and proved it.

It is certainly true that, although Mayr may have been rough on people like Wilson and Bush, he respected them and other biologists in a way that he never felt about others, including historians and philosophers. I always felt a bit like a court eunuch when I was around him. Useful and interesting, but ultimately not a real man. Or perhaps court jester. Mayr was much easer on us than on the biologists, but that was because we existed only to help biology. This did not preclude real friendship. Over the years he and I built up a close relationship. I did earn some respect. At a discussion group in the 1980s at Harvard, I told him to his face that I thought the species-as-individuals thesis is nonsense. He came at me across the room, shaking his finger, and yelling. I stood my gourd, and got invited to lunch the next day!

I never sensed that Mayr was a man deeply engaged in politics. He told me once that as a young man, like everyone in his class, he was mildly anti-Semitic, and then mentioned the Holocaust and remarked how far we had moved since then. After Mayr retired, one of his secretaries was a gay man, who died of AIDS. I saw genuine concern and affection. He liked kids—my children thought he was a terrific old man—and he (unlike a lot of academics) always treated my wife Lizzie with affection. He remembered her name without prompting. Some of my fondest memories are of lunchtime soup with the Mayrs and of his wife pushing him around and making him go for an after-lunch walk.

My favorite Mayr story is of an event about twenty years ago. A French film company decided to do a one-hour biography of Mayr. When finished, flyers were sent far and wide, advertising this film. Memorably, it described Mayr as a man of intelligence, a man of humility, a man of humor. I wrote to Ernst saying that I certainly thought of him as a man of intelligence, and that although he was a German I was sure that at least once in his life he had cracked funny joke, but that—think goodness—he was not a man of humility. He was a man of pride and determination and no nonsense about falsely putting himself down.

Mayr was absolutely livid. What did I mean by saying that he had no sense of humor? He was the funniest man that he knew. The next time we met, he would order me to listen to his jokes and to laugh. (Actually, his favorite joke with me was to call me at about ten in the morning and apologize for getting me out of bed. Mayr was famously an early riser—four in the morning, every day. He told me that at the beginning of the War, when he was an enemy alien—he had not then taken out citizenship—and fearful for his job at the Museum, he had decided to be a milkman if all else failed. He liked getting up early in the morning, and he liked the outdoors life.)

My final memory is four years old. I had just retired from thirty-five years of teaching in Canada, and moved to Florida State University here in Tallahassee. With my professorship comes money for (and expectations of) putting on conferences. So for the first, in 2001, I organized a conference on the philosophy of biology. Mayr was still in the habit of spending the winter at a small college in central Florida, and Betty Smocovitis at the University of Florida—an old friend of both Mayr and me—arranged to bring Ernst along. All sorts of prior warnings were sent that Ernst was old and frail and would need lots of time out. Although the conference was from Friday to Sunday, he would need to leave on Saturday afternoon. Unfortunately he would not be able to go on the field trip to Wakulla Springs, a nature reserves full of birds and alligators (and the site of the original Tarzan movies as well as of The Creature from the Black Lagoon and Airport 77.)

Along came Betty and Ernst. Inevitably as soon as the paper was over, Ernst was on his feet. This continued for the rest of Friday and all day Saturday. Who, on Saturday at Wakulla, was at the front of the boat? You guessed it! Ernst Mayr having a wonderful time, telling us that this was his first visit to Wakulla since 1931. And who saw the Saturday barbecue out to the bitter end, until finally I said: "Ernst go to bed." None other than our frail old friend and evolutionist.

Mayr once told me that the greatest influence on his life had been his mother, who had impressed upon him the obligation of people with his talents and background and education to serve others. I wish I could tell his mother how proud she could be of him. I wish I could tell his father, a deeply educated and cultured man, how proud he could be of his son. The truly great person is not he or she who simply does things easily, without strain. The truly great person triumphs above his or her limitations, and makes them seem trivial and unimportant. Ernst Mayr was a truly great man. What a life. What vitality. What service. What a privilege to have known him.

Ernst Mayr, 1904–2005: Remembrances & Tribute

By Frank J. Sulloway

Ernst Mayr was, without doubt, the most important intellectual figure in my life. He was my closest mentor and a towering model for anyone to try to live up to. He was always remarkably generous with his time to younger scholars and scientists. He was well known at the Museum of Comparative Zoology for his open-door policy, which effectively invited people to drop in unannounced, so that they could chat with Ernst about scientific matters.

Ernst dutifully read and commented on every paper that I ever gave him to read, supplying excellent advice regarding corrections and revisions. He read my undergraduate thesis on “Darwin and the Beagle Voyage” (1969), although he was not required to do so, and he voluntarily wrote a report about it for the History of Science Department, which I was very flattered to be able to read later, because it was so positive and thoughtful. Others were often surprised by the fact that Ernst would read papers sent to him by mail, by people he did not even know, and he would supply important comments and suggestions. Once I tried to thank Ernst for reading a paper of mine, by presenting him with a bottle of cognac. But Ernst would not accept it, saying that it was a pleasure for him to read such manuscripts and that I should drink the bottle myself. How he had time to read all these manuscripts, and to write and proof read everything he published himself, remains a mystery to me.

I first got to know Ernst in 1967, when I was just 20 years old and organizing the Harvard-Darwin film expedition to retrace Darwin’s route in South America. Ernst agreed to be the chairman of my little film advisory group, which I had assembled to give this project a semblance of legitimacy. Thanks in part to Ernst’s name and prestige, I was able to raise $25,000 for this film expedition—a considerable sum in those days. While in South America for four months, doors opened at the very mention of Ernst’s name, and local scientists eagerly offered their services as guides into the jungles of Brazil, the pampas of Argentina, the channels of Tierra del Fuego, and the mountains of the Chilean Andes. Because of my own association with Ernst, people often thought I was a Ph.D., but I had yet to obtain even my bachelor’s degree!

After I wrote a paper for Ernst’s graduate seminar in evolutionary theory, in the fall of 1970, Ernst took me under his wing. He was very impressed by this paper, which showed that Darwin had mistaken the various forms of Darwin’s finches in the Galapagos Islands, for the forms they mimic through convergent evolution, and hence that Darwin had not been an evolutionist during his visit to these islands. I showed that it was the case of the Galapagos mockingbirds that finally converted Darwin to evolution, after his return to England and a meeting, in March 1837, with ornithologist John Gould. (Gould, it turned out, understood Darwin’s Galapagos birds much better than Darwin did.) Ernst always dutifully cited me for these historical discoveries.

After I took his seminar in 1970, Ernst used to invite me to informal seminars at his house. I also used to drop by on an occasional basis just to chat, since I lived nearby. Mostly Ernst brought me up to date about his latest ideas, or talked about the things that interested him, and I just listened. Many times, in subsequent years, Ernst brought up how much he had enjoyed these conversations and how much he missed them. But I never felt that I was contributing much, although I think I was rather good at knowing just enough about whatever was being discussed to be able to make some comment that allowed Ernst to expand to a new or related topic. In short, I was good at keeping him talking (and I did enjoy these encounters). I also taught two seminar courses with Ernst in the history of biology, in the early 1970s, and this was a great learning experience for me.

I owe much of the success of my career to Ernst and his unflagging support for me. In 1973 he nominated me for a Junior Fellowship at Harvard, and when the Senior Fellows did not see things his way, he nominated me again the next year. This time I got the fellowship, one of the most prestigious that a young scholar could possibly receive. Ernst was like that—he did not take no for an answer when he believed strongly in something or someone. Other letters of recommendation that he wrote for me were doubtless largely responsible for my receiving subsequent fellowships.

There are so many ways that Ernst’s intellectual style has influenced my own scholarship. His thinking was so logical, his scholarship so meticulous, and his intellectual sweep so impressive. In my own career, I always tried to live up to this stellar example and to make Ernst proud of the fact that he had nurtured my scholarship along and had supported me so generously with his time, recommendations, and advice.

Ernst’s influence on me continues as I write here in the Galapagos Islands. I recently read a manuscript by a scientist visiting these islands who works on Darwin’s finches. I thought the conclusions of the paper were basically wrong because they violated Ernst’s fundamental ideas about the role of geographical isolation in the emergence of new species. So I rewrote the conclusion to the paper, showing that the interesting case, involving Geospiza fuliginosa (the Small Ground Finch) that scientist had studied was actually consistent with Ernst’s model of allopatric speciation, although the scientist’s findings perhaps added a new wrinkle to that model. My corrections were entirely accepted, and now I am a coauthor on the paper. But it is really to Ernst that I owe such a basic understanding of the origin of species.

I have heard Ernst say, several times, how much his own career was enabled by luck, such as the wonderful episode of seeing a pair of birds with a red bill in Germany, that had not been seen in that region for nearly a century—and how this chance observation led to his meeting Stresemann and his subsequent career in science. Well, the greatest stroke of good fortune in my career was my meeting Ernst as a young undergraduate in 1967, and the considerable interest he took, thereafter, in my own career. On so many levels, then, I shall miss Ernst.

Frank J. Sulloway, Visiting Scholar
Department of Psychology
University of California, Berkeley, CA

Seminal Evolutionist Ernst Mayr Dies

by Patricia Sullivan
Washington Post, February 5, 2005. Page B05.

Ernst Mayr, an evolutionary biologist who connected Charles Darwin's theories on natural selection to the science of genetics and in doing so helped create the field of evolutionary biology, died Feb. 3 of liver cancer at a retirement community in Bedford, Mass., where he lived. He was 100.

A seminal figure who began his career searching for birds in New Guinea, Dr. Mayr developed "evolutionary synthesis," making the origin of species diversity the central question in biology for the past six decades.

"I do consider him the Darwin of the 20th century," said Vassiliki Betty Smocovitis, a historian of science at the University of Florida. "He carried over the naturalist tradition, starting out in ornithology, then moving into evolution, biology. And in the last 40 years of his career, he became a historian, philosopher and writer."

Although not as well known as such scientists as E.O. Wilson, Stephen Jay Gould and Jared Diamond, Dr. Mayr was in many ways their intellectual predecessor, and each publicly acknowledged his leadership. His reputation was made with his 1942 book, "Systematics and the Origin of the Species from the Viewpoint of a Zoologist," which proposed that Darwin's theory of natural selection could explain all of evolution, including why genes evolve at the molecular level.

Species originate, he said, when a population is separated from the main group by time or geography. The separated groups eventually evolve different traits. Those traits are called "isolating mechanisms," and they discourage the two populations from interbreeding.

Thus, species are organisms that have the ability to interbreed, he said. That definition has remained the most widely accepted one in the field.

Dr. Mayr was the first scientist to win biology's "triple crown," capturing the International Balzan Foundation prize in 1983, the Royal Swedish Academy of Sciences' Crafoord Prize in 1999 and the Japan Society for the Promotion of Science's International Prize for Biology in 1994. He also won the U.S. National Medal of Science in 1970. He was named one of the 100 most influential scientists of all time in a 1996 book.

He wrote 25 books, five after he turned 90, and one that attempted to explain, in laymen's terms, "What Evolution Is" (2001). It had an appendix with arguments for encounters with believers in creationism.

"I'm an old-time fighter for Darwinism," he told the Harvard University Gazette in 1991. "I say, 'Please tell me what is wrong with Darwinism. I don't see anything wrong with Darwinism.' "

The Washington Post Book World said of his "The Growth of Biological Thought" (1982), "It seems safe to say that this magisterial study -- all 974 pages of it -- is one of the greatest works ever on the history of science."

He was born in Kempten, Germany, and received a medical degree from the University of Greifswald in 1925. Despite coming from a long line of physicians, he turned to zoology and received a doctorate in the field 16 months later.

Offered the chance to travel to New Guinea to collect birds of paradise, he went off to the South Seas for several years, traveling through hostile territories, battling dengue fever, malaria and dysentery while collecting specimens and developing his scientific theories.

Upon his return, he worked at the American Museum of Natural History in New York, and in 1953 moved to Harvard, where he stayed, retiring in 1975 but keeping an office. He was seen on campus as recently as last week.

On his 100th birthday, in an interview with Scientific American, he addressed the breadth of his career.

"When people ask me what is really your field, 50 years or 60 years ago, without hesitation, I would have said I'm an ornithologist. Forty years ago I would have said I'm an evolutionist. And a little later I would still say I'm an evolutionist, but I would also say I'm an historian of biology. And the last 20 years, I love to answer, I'm a philosopher of biology."

He also was a curator, running Harvard's Museum of Comparative Zoology from 1961 to 1970. He edited the journal Evolution. In 1990, he joined members of the nation's most prestigious science organization and 49 Nobel Prize winners in appealing to President George H.W. Bush to take the threat of global warming seriously and to begin immediately to curb the man-made gases that contribute to the greenhouse effect.

In 1997, The Post wrote that Dr. Mayr had become the world's loudest opponent of the search for radio signals from intelligent civilizations in other worlds.

"They're not out there," he said. "Life is almost a certainty, but it would be something like bacteria." He spoke with a mixture of conviction and outrage. "How many species have existed on the Earth since the origin of life, which was 3,850,000,000 years ago? A good answer is 1 billion. How many of those species have acquired advanced intelligence? You know the answer.

"High intelligence is just absolutely a fluke of history."

His wife of 55 years, Margarete Mayr, died in 1990.

Survivors include two daughters, five grandchildren and 10 great-grandchildren.

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