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The theory of the Cambrian Explosion holds that, beginning some 545 million years ago, an explosion of diversity led to the appearance over a relatively short period of 5 million to 10 million years of a huge number of complex, multi-celled organisms. Moreover, this burst of animal forms led to most of the major animal groups we know today, that is, every extant Phylum. It is also postulated that many forms that would rightfully deserve the rank of Phylum both appeared in the Cambrian only to rapidly disappear. Natural selection in many cases favored larger size, for example, hard skeletons to provide structural support - hence, the Cambrian gave rise to the first shelly animals and animals with exoskeletons (e.g., the trilobites), and the size of many animals also "exploded". By the start of the Cambrian, a large supercontinent comprising all land on Earth was breaking up into smaller land masses. This increased the area of continental shelf produced shallow seas, and an expanded diversity of environments in which animals could specialize and speciate. The debate persists today about whether the evolutionary "explosion" of the Cambrian was as sudden and spontaneous as it appears in the fossil record. The discovery of new pre-Cambrian and Cambrian fossils help, as these transitional forms support the hypothesis that diversification was well underway before the Cambrian began. More recently, the sequencing of the genomes of thousands of life forms is revealing just how many and what genes and the proteins they encode have been conserved from the Precambrian. The explosion of external form in the fossil record is what we see, but more gradual adaptation was taking place at the molecular level. Wang et. al. (1999) for example, recently conducted phylogenetic studies divergences among animal phyla, plants, animals and fungi. These researchers estimated Arthropods diverged from more primitive chordates more than 900 million years ago, and Nematodes from that lineage almost 1200 million years ago. They furthermore estimated that the plant, animal and fungi Kingdoms might have split almost 1600 million years ago. Finally, they conjecture that the basal animal phyla (Porifera, Cnidaria, Ctenophora) diverged between about 1200 and 1500 million years ago. If their research is valid, at least six major metazoan phyla appeared deep in the Precambrian, hundreds of millions of years before the oldest fossils in the fossil record. A facinating aspect of the Cambrian explosion is its speed over some 10 million years. From this it is reasonable to infer that expanded genomic complexity occured much earlier, perhaps over a billion years, prior to the morphological (phenotype) diversity that appeared in the Lower Cambrian. In recent years, research has shown that genomic complexity "happens" in many ways, including duplication and deletion of genes, cascades of genes, and, in complex organisms, entire chromosomes can be affected. Interesting also, is that such geneomic scrambling is an important mechanism in the etiology of cancers in animals. Genomic diversity is, of course, the stuff on which natural selection operates. The more numerous and complex enviroments and ecosystems provided the varying selective pressures to amplify benefitial mutation (genotypes) within populations, prune detrimental mutations, and otherwise fine-tuning genomes to enhance survival. Such fine-tuning would be different in different ecological niches. Among the famous Lagerstatten from Cambrian time, the Burgess Shale of Canada and Chengjiang, in Yunnan Province, China are the best known, having a great diversity of benthic or burrowing creatures, many of which are soft-bodied, lacking an exoskeleton. Less well known is that the American state of Utah where similar Cambrian creatures are found. If fact, some researchers believe a larger number of species are to be found in the Wheeler and Marjum Formations of Utah than in the Burgess Shale, though the fossils in Utah are far less abundant.
It is important to remember that geological history contains numerous periods of slow evolution punctuated by periods of rapid evolution, which Steven J. Gould called Punctuated Equilibrium. The rates of evolution generally depend on rates of selection, which in turn depend on rates of environmental change. It also depends upon the existing genomic diversity on which selection acts. Mutation rates tend to be slow and steady, and in the absence of environmental change, slowly accumulate in a population. It is selective pressure that weeds out the mutations that are detrimental or neutral to survival, and retains and multiplies the mutations that are beneficial within a population. For a population isolated in a new environment, rapid selection can lead to speciation, and in the Lower Cambrian, to radically new forms that we now group in the Phyla of modern times. The years ahead should see furtherance of knowledge of how and the timeline along which the Tree of Life branched, especially when proteomes of its many branches are unraveled. Still, major mysteries are likely to persist, given the amazing ability of nature to splice, dice, reassemble, swap, amplify, and silence or re-use nucleid acid sequences within the genome of living organisms.
Unlike all other jawed vertebrates, placoderms never had teeth, and did not descend from toothed ancestors. Instead, bony plates associated with the jaws performed the function of teeth, sometimes forming razor-like, literally self-sharpening edges (as can be seen in Dunkleosteus on the left). Additional peculiarities of the skull, such as nasal capsules that were not fused to the rest of the braincase, distinguish placoderms from all other jawed vertebrates. Furthermore, in 1997, a placoderm fossil from Antarctica was found to contain preserved pigment cells: iridescent silver on the ventral side (belly) and red on the dorsal side (back). Placoderms are the oldest vertebrates for which we know something about their color in life. This further implies that placoderms may have had color vision. The evolutionary history of placoderms has been compared to a brilliant light bulb that soon burns out. They were a highly successful and diverse taxon, but they lasted only about fifty million years. Contrast this with the history of sharks, which appeared at about the same time as placoderms -- but which have survived for over 400 million years! In a sense, placoderms represent an "early experiment" in the evolution of jawed fish; they radiated into a number of body shapes and ecological niches, which were occupied by other fish lineages after the extinction of the placoderms. The oldest placoderms known, found in China, appeared in the later part of the Early Silurian. However, placoderms reached their greatest diversity in the Devonian, the so-called "Age of Fishes". A number of Devonian placoderms have been found in freshwater habitats: placoderms included some of the first vertebrates to colonize fresh water. They also included the earliest vertebrates to colonize the open ocean. The Devonian saw the greatest diversity of a large number of fish taxa, including not only placoderms, but armored jawless fishes, early Chondrichthyes, and the first ray-finned and lobe-finned fishes. Many of these taxa died out around the end of the Devonian Period for reasons that are still not well understood. Placoderms survived until the very end of the Devonian, and their extinction appears to have been quite sudden, but its causes are still unknown.
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