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                                                                                 Origin of Life
The history of life tends to move in quick and quirky episodes, rather than by gradual improvement.


DAWN OF OCEAN LIFE

Haikouella


Seen here with different fossils on both the obverse and reverse, it is a very unique specimen from the Chengjiang Biota. One one side is a mass mortality of Haikouella lanceolata, thought by its describers to be the earliest craniate-like chordate. This fish-like animal has many similarities to the contemporaneous Yunnanozoon lividum, but differs in several aspects: it has a discernible heart, dorsal and ventral aorta, gill filaments, and a neural chord. For all these reasons, Chen, Huang, and Li identified it in the seminal Nature paper as a chordate. The debate rages on, but whatever the outcome, this creature was much like the ancestor of all the vertebrates. It derives its generic name from its resemblance to the modern day lancet Amphioxus. Such assemblages have been mentioned in the literature, and must represent some catastrophe that overcame them in mass.


Xidazoon


The reverse holds an unusual fossil that is a rare member of the Vetulicolians, an enigmatic group which some scientists place in their own phylum (Phylum Vetulicolia). They are thought to have been swimmers that either were filter feeders or detritivores. One researcher places them with the Urochordates, giving them strong affinity with the Phylum Chordata. At present, there is no agreement as to their systematic placement. This one is quite unusual with a circular mouth which was thought to have served it in filter-feeding.

Most major animal groups appear for the first time in the fossil record some 545 million years ago on the geological time scale in a relatively short period of time known as the Cambrian explosion. While some scientists believe there was indeed an explosion of diversity, others believe that such rapid acceleration of evolution is not possible; they posit that there was an extended period of evolutionary progression of all the animal groups, the evidence for which is lost in a sparsely populated fossil record.

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.


SLAB CONTAINING SPECIMENS of Pteridinium from Namibia shows a prominent organism from the earth's first multicellular fauna, called Ediacaran, which appeared some 600 million years ago. The Ediacaran animals died out before the Cambrian explosion of modern life. These thin, quilted, sheetlike organisms may be ancestral to some modern forms but may also represent a separate and ultimately failed experiment in multicellular life. The history of life tends to move in quick and quirky episodes, rather than by gradual improvement. To understand the events and generalities of life's pathway, we must go beyond principles of evolutionary theory to a paleontological examination of the contingent pattern of life's history on our planet - the single actualized version among millions of plausible alternatives that happened not to occur. Such a view of life's history is highly contrary both to conventional deterministic models of Western science and to the deepest social traditions and psychological hopes of Western culture for a history culminating in humans as life's highest expression and intended planetary steward.

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.

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Early Paleozoic Rise of Fish

Dunkleosteus

Dunkleosteus, a marine predator from the Devonian of Ohio that is a member of the most diverse subgroup of the placoderms, the Arthrodira. Dunkleosteus reached estimated sizes of up to twenty feet (six meters) in length -- which makes Dunkleosteus and related species among the largest animals, of any kind, in the Devonian. Most arthrodires, however, were smaller than Dunkleosteus and may have fed on molluscs, crustaceans, and other invertebrates.

Bothriolepis

Bothriolepis from Escuminac Bay, Québec, representing the placoderm subgroup Antiarcha. The armored head of Bothriolepis was only about four inches (ten cenitmeters) across. The unusual "paddles" on either side of the boxy armored body are the pectoral fins, which were armored in Bothriolepis and related species; the hole in the top of the head housed both the nostrils and the eye sockets, set close together. Other placoderm taxa included flattened, stingray-like forms (rhenanids), forms with long spines (petalichthyids), and slender, streamlined forms with crushing tooth plates (ptyctodontids).

The extinct armored fishes known as placoderms make up what is considered to be the earliest branch of the gnathostome family tree -- the earliest branch of the jawed fishes. Placoderms bore heavy bony armor on the head and neck, often with an unusual joint in the dorsal armor between the head and neck regions; this joint apparently allowed the head to move upwards as the jaw dropped downwards, creating a larger gape. However, most of the body was either naked or, less commonly, covered with small scales. Both of the placoderms shown above had long bodies with heterocercal (asymmetrical, shark-like) tails extending past the head armor.

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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THE HISTORY OF LIFE

Late Proterozoic

Just before the Cambrian, in the Late Proterozoic, large, multi-cellular life that had soft bodies (jellyfish, etc.) are found in the fossil record.



Cambrian

The base of the Cambrian is defined as the occurrence of the first-hard shelled fossils. This does not mean that complex life did not exist before the Cambrian, just that the fossil record is much more unlikely to record soft-bodied life. One of the most common fossils in the Cambrian is the trilobite (Wisconsin state fossil), which are shown here.

Artist drawing of the bottom of the Cambrian shallow sea floor, showing trilobites (imagine these crawling around on the Cambrian sea floor at Devil's Lake state park 550 m.y. ago!).



Ordovician

Ordovician life continued to increase in complexity from that of the Cambrian, and appearance of extensive coral reefs occurred. This was accompanied by extensive limestone formation, such as is well shown by the Ordovician limestone rocks that can be found surrounding Madison (look at the road outcrops next time!).

Ordovician life included a larger diversity of predators, including straight-shelled "nautiloid-like" or "squid-like" creatures shown here.

Another example of the diverse Ordovician life in the shallow inland seas of the continents. Note the complex flora (plants) and fauna (animals).



Silurian

Marine life in the Silurian continued much like the Ordovician, with abundant corals, bivalves (clam-like animals), trilobites, and nautilus-like predators.

A major change, however, in Silurian life was the appearance of fish.



Devonian

Devonian life in the shallow inland seas continued much as it had in the Silurian, with extensive coral reefs and ocean plants.

Another artist's rendering of Devonian marine life.


The major change in life in the Devonian occurred not in the oceans but on land. Fr most of Earth's history, including the early part of the Paleozoic, the land was devoid of plant life - it must have looked quite desolate. However, land plants became common on land in the Devonian, paving the way for life to move out of the oceans and on to the land. The first amphibians are found in the Devonian.



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