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MACROEVOLUTION AND THE TREE OF LIFE  117



                                         a 14 q 14  p 14  b 14  f 14  o 14  e 14  m 14  F 14  n 14  r 14  w 14  y 14  v 14  x XIV
                                                                                                     14
                                                                                                      XIII
                                                                                                      XII
                                                                                                     XI
                                           a 10          f 10      m 10 E 10  F 10      w 10      z 10  X
                                            a 9          f  9       m 9                w 9       z 9  IX
                                             a 8        f  8  k 8  l 8  m 8         u 8  w 8     z 8  VIII
                                              a 7      f  7  k 7  l 7  m 7         u 7   w 7    z 7  VII
                                               a 6   f 6    k 6   m 6               u 6       z 6    VI
                                                a 5     d 5  k 5  m 5                 u 5     z 5    V
                                                  a 4   d 4  l 4  m 4                       z 4      IV
                                                   a 3      l 3  m 3                 t 3   z 3       III
                                                     a 2   s 3  m 2                  t 2  z 2        II
                                                      a 1   m 1                         z 1          I
                                                          A  B  C  D  E    F    G H   I  K L





             (a)                        (b)
             Figure 5.1  (a) Charles Darwin. (b) Branching diagram of phylogeny, the only illustration in On the
             Origin of Species (1859). It shows how two species, A and I, branch and radiate through time. The
             units I–XIV are time intervals of variable length, and the lower case letters (a, b, c) represent new
             species.


             Darwin laid the framework for evolutionary      often represented as a branching tree diagram.
             biology 150 years ago. Despite millions of      Darwin’s idea was that life had diversifi ed to
             essays, books and web sites discussing evolu-   millions of species by the continued splitting
             tion, no one has yet falsifi ed Darwin’s theory   of species from a common stem (Fig. 5.1).
             of evolution by natural selection, and so it    Indeed, he proposed that all of life, modern
             stands as the core of modern biology and        and ancient, could be followed back down the
             paleontology, just as Isaac Newton’s laws       phylogenetic tree to a single point of origin:
             stand at the heart of much of modern physics.   modern evidence confi rms  this  remarkable
             And yet, surprisingly, Darwin is quoted, and    insight.
             misquoted, by many special-interest groups        Darwin’s branching diagram also explained
             who want to use him in support of, or against,   for the first time the meaning of the natural

             their views of politics, sociology and religion.   hierarchy of life that Linnaeus had discovered
             So it is important to understand what Darwin    100 years earlier (Box 5.1). This natural inclu-
             said, how his insights affect science today, and   sive branching hierarchy is the basis of modern
             how paleontology relies on modern evolution     approaches to discovering the tree of life, the
             as its basis.                                   single great evolutionary tree that links all
               Charles Darwin’s On the Origin of Species     species living and extinct, from the modern
             (1859) is usually remembered as the book        biodiversity of over 10 million species, back
             that made the case for natural selection as the   to a single hypothetical species 3500 million
             mechanism of evolution, sometimes called        years ago in the Precambrian. Paleontological

             “survival of the fittest” (see pp. 118–19).      aspects of evolution, such as the tree of life
             Since the time of Darwin, evolution has been    and studies of processes over thousands and
             seen in action in the laboratory and in the     millions of years, are sometimes called mac-
             field, and paleontologists use the principles of   roevolution (“big evolution”) to distinguish

             evolution to understand how species origi-      them from  microevolution (“small evolu-
             nate. The origin of species is also core to a   tion”), all the smaller-scale and shorter-term
             second theme in Darwin’s writings, namely       processes studied by biologists and geneticists
             phylogeny, or the pattern of evolution that is   in the laboratory or in the fi eld.
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