Extinction has always been a controversial subject. Even after
              decades of intense study, researchers are still arguing over the
              cause of the best-known mass extinction, the death of the
              dinosaurs. And theres a topical slant in the idea that
              - even today - species are regarded as "endangered" if they are
              relatively susceptible to extinction. Then too, what thinking person
              could have avoided wondering whether our own species is slated for
              extinction and, if so, how long we have before we meet our fate?
            
 The disappearing Mammoth
            
 Two centuries ago extinction held centre-stage in an even more
              fundamental debate: whether the Bible should be interpreted literally.
              The bones of a very large fossil elephant (a Mammoth)
              had been collected in the then new world of North America. Discussing
              the Mammoth at the National Institute of Sciences and Arts in Paris
              in 1796, the French anatomist Baron Georges Cuvier argued that the
              bones belonged to a unique species of which no living examples were
              known. It was extremely unlikely that such a large animal could
              have gone undetected if its descendants had survived to the present
              day, so Cuvier pronounced the species extinct.
            
 This conclusion contradicted religious dogma of the time. A benevolent
              creator, it was assumed, would not allow any of his creation to
              disappear from the Earth. In this way, Cuvier fired off one of the
              first salvos in a debate between theologians and natural historians
              that culminated 60 years later with the writings of Charles Darwin
              - and still continues in some quarters of society to this day.
            
 Observing extinctions
            
 Since Cuvier's time, extinctions have studied primarily by palaeontologists
              - scientists who investigate the remains of ancient life. For all
              that is said in popular books, articles and television programmes,
              the study of extinction is a surprisingly problematic and young
              field of inquiry. Extinction is one of the most common observations
              in palaeontology, but it remains very difficult to understand precisely
              why a particular ancient plant or animal species went extinct.
            
 Much of this difficulty stems from the way that scientists observe
              an extinction. Stated rather badly, we study extinction by observing
              what isnt there. When we collect a fossil from a rock deposit,
              we assume that the species represented by this fossil lived at the
              place and time the rock was formed. If we sample a younger deposit
              and find that the species is not present, and if it never reappears
              in still younger deposits, we assume the species has become extinct,
              and that the time of extinction corresponds to the date of the youngest
              observed fossils.
            
 But there may be other explanations for the absence of the species
              in younger deposits. For example, the species may have become extinct
              in this particular region, but survived to later times in other
              areas. Or the younger deposits may record an environment in which
              the species never occurred in the first place. A third possibility
              is that natural selection changed the appearance of the species
              to such an extent that palaeontologists mistakenly assign younger
              members to a different species.
            
 A chart of extinctions
            
 These problems mean that we are only just beginning to compile
              reliable data on how extinctions have varied over time. And such
              data is essential if we want to test different hypotheses on what
              has caused extinctions.
            
            
              
                | Click the
                  image to see a larger version 
  Figure 1. Extinctions over the past 600 million years - mass
                  extinctions show up as peaks superimposed on a general decline
                  in extinctions (diagonal line). The mass extinction marked as
                  'Maastrichtian' was the death of the dinosaurs.
 | 
              
                | Reference: Sepkoski, JJ, Jr. 1994. Extinction and the fossil record. Geotimes
                  March 15 - 17
 
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             Figure 1 shows one of the latest summaries of extinction data.
              It concentrates on invertebrates - such as snails, clams and coral
              species - that lived in ancient oceans. The graph shows the number
              of extinctions that occurred within successive small subdivisions
              of geological time - called stages - which are typically a few million
              years in duration.
            
 There are two particularly noteworthy features. First, there is
              a general decline in the number of extinctions over the last 600
              million years. Secondly, there are several major peaks
              of extinction, which are roughly evenly spaced. Both of these features
              are spurring a vigorous debate among palaeontologists.
            
 Extinctions on the decline
            
 The overall trend in Figure 1 shows the number of extinctions
              decreasing with time. At face value, it means that a species existing
              in the past was more likely to be wiped out than a species existing
              now. Is this really the case, and - if so - why?
            
 There are currently three explanations.
            
 The first suggests that the effect isnt real. It occurs
              because palaeontologists are more familiar with modern marine organisms
              than with species from the remote past. When dealing with recent
              fossils, palaeontologists can be fairly confident that they are
              classifying them correctly into different species and groups of
              species (genera). When they look back to more ancient and unfamiliar
              fossils, they are more likely to be fooled into separating them
              out into many more genera. So theres a higher chance that
              one of these - artificially small - ancient genera would become
              extinct.
            
 The second invokes marine organisms moving from the general ocean
              into marginal environments, for example around the edges
              of continents. Here, they might be sheltered from environmental
              changes, and so suffer less extinctions. On the other hand, the
              marginal environment might play the opposite role. It could be so
              extreme that the species had to evolve into a hardier form, which
              was then immune to all but the most profound episodes of global
              environmental change.
            
 The third (and most recent) explanation argues that conditions
              in the world's oceans have become progressively more favourable
              for marine life. In 1994, Prof. Ron Martin of The University of
              Delaware summerized geochemical evidence that the oceans themselves
              have changed over geological time, by studying isotopes of elements
              such as sulphur-34, strontium-87 and carbon-13. These trends match
              the declining extinction curve.
            
            
              
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                | photo:
                  Norman Macleod 
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             Ocean changes
            
 In Martin's model, the oceans of 600 million years ago were very
              different from those of today. Their circulation was sluggish, with
              large areas of the ocean floor covered by waters with little or
              no dissolved oxygen. They received large amounts of nutrient washed
              off from the barren continents (plants and soils had not yet evolved).
              And ocean plankton was relatively inefficient at producing organic
              materials through photosynthesis.
            
 As time went by, the oceans became more dynamic as plate tectonics
              pushed the continents around. Life on land trapped the nutrients
              that once ran off into the seas. As a result, the oceans became
              host to a greater diversity of plankton. This, in turn, meant they
              could support a diverse array of species, increasing the stability
              of marine ecosystems - and decreasing the probability of extinction.
            
 Mass extinctions: the three culprits
            
 The second important pattern we can see in Figure 1 are peaks,
              where large numbers of species (up to 90%) become extinct within
              a short period of time. These are termed mass extinctions. The most
              famous - though not the most severe - involved the death of
              the dinosaurs about 65 million years ago.
            
 Scientists have proposed many different mechanisms to account
              for the pattern of mass extinctions over the last 600 million years.
              Currently, the most popular are: a fall in sea level; vast eruptions
              of basalt; and the impact of an asteroid or comet.
            
 Interestingly, its not the direct effects that cause the
              damage. In all three cases, researchers explain the extinctions
              through subsequent drastic changes in the environment. For example,
              they may cool the Earth by promoting the formation of clouds, or
              lead to global warming as greenhouse gases are released. Nitrogen
              and sulphur compounds thrown into the atmosphere may produce acid
              rain, and also destroy the ozone layer.
            
 Patterns of extinction
            
 Generally, its difficult link any particular catastrophic
              event to a subsequent mass extinction. Instead palaeontologists
              comparing the pattern of mass extinctions as a whole with the patterns
              of each of the three possible mechanisms. Figure 2 compares the
              record of mass extinctions with geological records of sea-level
              falls, flood-basalt eruptions, and impacts of bolides (asteroids
              or comets).
            
            
              
                | Click the
                  image to see a larger version 
  Figure 2. The record of extinctions (bottom), compared with
                  - from top - major sea-level changes, giant eruptions of basalt,
                  and impact craters.
 | 
              
                | References: Courtillot, V, Jaeger, J-J, Yang, Z, Féraud, G, Hofmann,
                  C. 1996. The influence of continental flood basalts on mass
                  extinctions: where do we stand? In Ryder, G, Fastovsky, D, and
                  Gartner, S, eds. The Cretaceous-Tertiary event and other catastrophes
                  in earth history: The Geological Society of America, Special
                  Paper 307, 513-525. Grieve, R, Rupert, J, Smith, J, Therriault,
                  A. 1996. The record of terrestrial impact cratering. GSA Today
                  5: 193-195
 Hallam, A. 1992. Phanerozoic sea-level changes. New York; Columbia
                  University Press.
 
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             Looking at Figure 2, it is clear that impacts show the worst record
              of association with mass extinction events. Sizeable impacts are
              as likely to have occurred during low-extinction stages as during
              high-extinction stages. In fact, theres only one compelling
              example of an association between a large impact and a major extinction
              event. The Chicxulub event of 65 million years ago was the largest
              impact of the last 600 million years, and third largest in the known
              history of our planet. It blasted out a large crater in Mexico.
              At the same time, we find the Maastrichtian mass extinction event,
              also known as the K/T mass extinction. This was the disaster in
              which the dinosaurs perished, and it is ranked as the third to fifth
              largest extinction event of the last 600 million years.
            
 Moving on to the second possible mechanism, abrupt falls in sea
              level show a rather better level of association with extinctions.
              Each of the three largest mass extinctions during the last 250 million
              years (the time interval for which we have the most accurate time
              resolution) corresponds to a major sea-level change. This supports
              the idea that rapid falls in sea level have a detrimental effect
              on the diversity of marine invertebrates.
            
 Giant eruptions
            
 But the strongest association - by far - comes with the third
              prospective extinction mechanism, as seen in the record of continental
              flood-basalt eruptions. These are vast outpourings of basaltic lava,
              similar to the eruptions responsible for the Hawaiian Island chain,
              but taking place entirely on land.
            
 Perhaps the best known is the series of eruptions that took place
              on the Indian subcontinent beginning at the very end of the Maastrichtian
              stage, 66.5-64.5 million years ago. They poured out over one million
              cubic kilometres of lava in just over one million years, forming
              the vast Deccan Traps. The eruption did not take place continuously
              over the entire million years, but episodically in massive lava
              flows that have no counterpart in human history.
            
 Since we have no direct experience of such cataclysmic eruptions,
              it is difficult to imagine (much less model) the climatic effects.
              Regardless, the close association between the flood-basalt record
              and the pattern of extinction events is very difficult to explain
              away as mere coincidence. In addition, its worth noting that
              the three largest extinctions of the last 250 million years took
              place during times of combined sea-level fall and flood-basalt eruption.
              Of course, the Maastrichtian event is also associated with the Chicxulub
              impact. However, this event is by no means the largest mass extinction
              to have occurred and, according to available data on the timing
              and magnitudes of impacts, several extinction events of equal and
              greater magnitude are not associated with large asteroid impacts.
            
 Weighing up the evidence
            
 As far as any single extinction event is concerned, in all probability
              we will never know all the factors involved. Nevertheless, the existing
              data provide an insight into the factors that are repeatedly associated
              with extinction events in the geological record.
            
 A simple statistical analysis of the data in Figure 2 reveals
              the relative strength of the association between each type of catastrophe
              and mass extinctions as follows:
            
              -  sea-level change 7 out of 14
              
-  continental flood basalt eruptions (over the last 250 million
                years) 10 out of 10
              
-  asteroid impact 1 out of 17.
            
Based on current data, tectonic factors - giving rise to flood-basalt
            volcanism and sea-level fall - appear to exhibit the greatest level
            of association with large-scale extinction events over the course
            of the last 600 million years. Extraterrestrial impacts - current darlings of the media coverage
              of extinctions - have certainly played an important role in Earth
              history and may have enhanced the death of the dinosaurs.
              However, asteroid impacts do not appear to be the primary agents
              responsible for the overall patterns in the geological extinction
              record.
            
 
            Dr Norman MacLeod is an international expert in the
              study of extinctions. After a BSc in geology from the University
              of Missouri, he received an MSc from the Southern Methodist University
              and a PhD from the University of Texas, both in palaeontology. Norm
              MacLeod has researched at the University of Michigan and Princeton
              University, and is now a member of the Palaeontology Department
              at The Natural History Museum, London. As well as extinctions, his
              research interests include the quantitative analysis of morphology
              and the analysis of evolutionary patterns. 
            
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