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436  INTRODUCTION TO PALEOBIOLOGY AND THE FOSSIL RECORD



                      Placoderms were fearsome predators, some of       Other Devonian fishes were more modern
                      them, like Dunkleosteus from the Late Devo-     in appearance. The fi rst shark-like chondrich-

                      nian of North America, reaching the impres-     thyans, or cartilaginous fishes, came on the
                      sive length of 10 m. This was the largest       scene during the Early Devonian. Acanthodi-

                      animal that had lived until then, and its size   ans were small fishes, mostly in the range
                      and fearsome jaws may explain why so many       50–200 mm in length, and they bore numer-

                      Devonian fi shes were armored.                   ous spines at the front of each fin and in






                               Box 16.4  Genome duplications and vertebrate evolution

                        Vertebrates have larger genomes than other animal groups. The genome is the entire sequence of
                        genes contained on all the chromosomes within the nuclei of cells. Various worms and insects have

                        around 15,000 genes in their genomes, while the figure is 31,000 for humans, 30,000 for the mouse

                        and 38,000 for the pufferfish. However, vertebrates do not just have more genes than invertebrates,
                        they have two, four or even eight copies of many individual invertebrate genes. At one time, molecu-
                        lar biologists thought that humans had as many as 100,000 genes, but the reduced fi gure  was
                        established in 2004 after the intense gene sequencing efforts of the Human Genome Project. What
                        does genome size mean?
                           Some have suggested that genome size maps on to the complexity of an organism. Surely, a single-
                        celled bacterium does not need many genes because it does not do much, and vertebrates, as much
                        more complex organisms, would need more genes. Humans ought to have the largest genomes since
                        we are somehow very complex and important. In fact, genome size is only loosely related to bodily
                        complexity: the largest genome reported so far comes from a lungfish! Much of the genome is so-

                        called junk DNA, or at least duplicate genes and non-coding sections, so the functional genome size
                        might be a better correlate of function or bodily complexity.
                           Whether functional or not, molecular biologists have proposed that there were at least
                        three  genome duplication events (GDEs) in the history of vertebrates – times when evolutionary
                        change was dramatic and large sectors of the genome duplicated. GDEs are identified at the

                        origin of vertebrates, the origin of gnathostomes and the origin of teleosts, the hugely diverse modern

                        bony fishes (Furlong & Holland 2004). Could the evolutionary jump have caused the GDE, or

                        perhaps the GDE stimulated rapid and fundamental reorganization of the fishes at these three
                        points?
                           Donoghue and Purnell (2005) suggest that molecular biologists have been misled. By omitting
                        fossils, they see artificial morphological jumps in their cladograms, and then link this to the

                        postulated GDE. In fact, when fossils are inserted, the “jumps” seem less clear. For the origin of
                        gnathostomes, biologists have compared lampreys with sharks, and there is a wide gulf
                        between these two groups, so suggesting quite a leap in terms of anatomic change and in terms of
                        genome duplication. However, when fossils are inserted (Fig. 16.7), seven major ostracoderm and
                        placoderm clades fall between the living groups, and the evolutionary transition is stretched. Some
                        of the fossil groups (especially pteraspidimorphs, conodonts and placoderms) were diverse, and it is
                        not clear that the GDE drove, or permitted, a single dramatic burst of speciation, as had been
                        proposed. Further, it is not clear that there was a single reorganization of anatomy associated with
                        the origin of jaws and the GDE: the fossils show step-by-step character changes over a long
                        interval.
                           This is a developing field of study. The claim that genome duplication can drive major bursts

                        of evolution is dramatic, and perhaps overstated. Paleontologists can make profound contribu-
                        tions in new areas of science by working hand-in-hand with molecular and developmental
                        biologists.
                           Read more through http://www.blackwellpublishing.com/paleobiology/.
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