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CHAPTER 9APALEOBIOLOGYPART I. PRECAMBRIAN LIFE
1. INTRODUCTION1.1
Life on Earth began early. The oldest undoubted fossils are about 3500million years old, back in the early Archean—and, as you will see in a latersection of this chapter, the earliest fossil organisms are very similar to organismsthat are abundant and successful today. These organisms, although primitivecompared to advanced metazoans (like us), are, in an absolute sense, ratheradvanced in their physiology. It’s generally agreed that the earliest life, earlierthan that represented by the oldest fossils, must have been much simpler and lesssophisticated biochemically The clear implication is that
life must have evolved much earlier than 3.5 Ga
—although it is doubtful if fossils much older than thatwill ever be found. Why not? Perhaps largely because the sedimentary rocks inwhich they would have been preserved are no longer around, in pristine,unmetamorphosed condition, for us to scrutinize.
1.2
This section purports to tell you something about the nature andevolution of life in the Precambrian (more specifically, up to the latter part of theProterozoic; for the rise of eukaryotic organisms in general, and multicellulareukaryotes in particular, late in the Proterozoic, see the following section). For thesake of full disclosure here, I should point out that I am a novice in biology! I feelmyself to be especially inadequate when it comes to the biochemistry of metabolicprocess and the molecular biology of replication and inheritance. With thatdisclaimer, however, I have attempted here to present to you the basics of thefossil record of the Precambrian and some of its implications for the earlyevolution of life. If you have a special interest in paleontology, and in particularthe earliest life you might consider going into the literature. I have provided afairly long list of materials, largely review papers by specialists in the field.
1.3
Just to set the stage at this early point in the chapter, Figure 9-1 is adiagram that shows, in a very generalized way, the known distribution of stromatolites and microbial microfossils through geologic time. The clearmessage from Figure 9-1 is that
the Archean fossil record is scanty but real, and the Proterozoic fossil record is far more abundant 
. The terms used in Figure 9-1,as well as some of the place names, will be elaborated later in this section.182
 
 
Figure by MIT OCW.
Figure 9-1: Diagram showing the known distribution of stromatolites and microbialrnicrofossils through geologic time
2. THE KINGDOM OF LIFE2.1
Back when I was a child, the standard idea was that life was divided intothe plant kingdom and the animal kingdom. (In fairness to biologists, I shouldpoint out that the experts knew then that the situation was more complicated:what to do about fungi, for example?). Progress on deciphering the “tree of life”(that is, the evolutionary development of the various life forms known today,through geologic time, from a presumably common original ancestor) has beentruly spectacular in recent years. With the development of techniques for183
   P   R   E   C   A   M   B   R   I   A   N   P   R   O   T   E   R   O   Z   O   I   C   A   R   C   H   E   A   N
01234Age
(Ga)
   W   i   d  e  s  p  r  e  a   d ,   A   b  u  n   d  a  n   t   M   i  c  r  o   f  o  s  s   i   l  s   W   i   d  e  s  p  r  e  a   d ,   A   b  u  n   d  a  n   t   S   t  r  o  m  a   t  o   l   i   t  e  s
   M   i  c  r  o   b   i  a   l   S   t  r  o  m  a   t  o   l   i   t  e  s   M   i  c  r  o   b   i  a   l   F  o  s  s   i   l  s
????Hamersley GroupFortescue GroupInsuzi GroupOnverwacht GroupWarrnawoonaFig Tree GroupIsua SupracrustalsFormation of Earth
(Oldest Known SedimentaryRocks)
   P   H   A   N   E   R   O  -   Z   O   I   C
 
examining the genetic material carried by RNA and DNA, it is now possible totrace evolutionary lineages back in time. (I would have included a “backgroundsection” on that, but I think that it would have taken me weeks, not just days, toproduce an adequate one, given the very steep learning curve I would have faced.)Biologists now have a fairly generally accepted picture of the “tree of life”.Before considering that, however, we need to deal with some rather intricateterminology for organisms and their metabolic processes.
2.2
The basic distinction among organisms is between
unicellular organisms
and
multicellular organisms
. Among the unicellular organisms, thereis another fundamental distinction: between
 prokaryotes
and
eukaryotes
.Prokaryotes are relatively primitive unicellular organisms in which the protoplasmand the genetic material are encased within a cell wall of some kind but thegenetic material is not located within a nucleus. Eukaryotes are relativelyadvanced unicellular organisms in which the genetic material is enclosed in aspecial nucleus within the cell. Prokaryotes reproduce asexually, whereaseukaryotes reproduce sexually. (The implication of that is that the pace of evolution was very slow early on, but quickened with the advent of eukaryotes,because sexual reproduction allows the ever-arising mutations to spread rapidlythrough an interbreeding population, leading to Darwinian evolution throughnatural selection processes.) The earliest fossils are believed to be prokaryotic;eukaryotic organisms evolved in the course of the Proterozoic.
2.3
All organisms can be classified as either
heterotrophic
or
autotrophic
.A heterotrophic organism obtains its raw-material energy resources (its “food”)from preexisting organic matter (or abiotic organic molecules). An autotrophicorganism synthesizes its own organic materials by conversion of simple inorganiccompounds, making use of a variety of external energy sources.
2.4
There seems to be a general consensus that
the very earliest organismsmust have been heterotrophs
, because heterotrophs can be simpler in theirmetabolic processes than autotrophs. In that view, the more advanced autotrophiclife forms evolved from early heterotrophs. The problem with widespreadheterotrophy is that eventually the organisms use up whatever stock of “food” isavailable, and without autotrophy, no more “food” is produced—except perhapsby the continuing inorganic synthesis of abiotic organic molecules, which wouldnot have been produced in any great abundance.
2.5
Depending on the form of energy utilized, autotrophs can be classifiedas either
chemoautotrophic
or
 photoautotrophic
. Chemoautotrophic organismsuse chemical energy to fuel their processes of biochemical synthesis. Suchorganisms are known to exist today, perhaps most spectacularly in the deep ocean,in the vicinity of hydrothermal vents (called, picturesquely, “black smokers”)associated with mid-ocean spreading ridges. Photoautotrophic organisms usesolar energy for their processes of synthesis.184
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