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In this regard, the fossil record demonstrates two important facts. Firstly, that although relative rates have varied greatly, over geological time as a whole there has been a net excess of species originations over species extinctions i. Secondly, that virtually all the species that have ever existed are now extinct, and the extinction of every species is a natural and expected event.

Self-evidently there must always have been species at risk of extinction, i. It is difficult for many reasons to keep track of species extinction in recent time. The species involved may be unknown; it may be unclear whether some population represents a separate species or not; the individuals may be too small to be noticed without special sampling procedures; and the entire process of decline and extinction may extend over much more than an average human lifespan.

Positive evidence of extinction i. In other words, unless circumstances are exceptional, monitoring of recent extinction events has a resolution limit measured in decades, and it is thus impossible to state with precision how many species have gone extinct in any given period or to predict exactly how many species are going to become extinct by some point in the future.

From the imperfect evidence that is available, it appears that around vertebrates and nearly invertebrates have become extinct during the past years see Table 1. The number of plant extinctions is thought to be in the hundreds, although some believed to be extinct in the wild have survived in botanic gardens and seed collections. Because mammals and birds tend to be relatively well recorded, and leave recognizable skeletal remains, it is principally among these groups that known extinctions may be reasonably representative of actual extinctions.

In these two groups the known rate of extinction over the past years averages around species per hundred years. Vertebrates extint since AD The key question then is: Extinction rates have varied greatly, and species that are rare or otherwise prone to extinction must be poorly represented by fossils and so bias the record, but the average lifespan of a fossil species appears to be about four million years.

Given this average, if 10 million species existed at any one time, the extinction rate would have been about 2. Applying this factor to recent birds and mammals numbering about 10, and living species respectively the expected background extinction rate would be around one species every four hundred years and eight hundred years, respectively.

The known recent extinction rate appears to be some or times higher than background. Bias inherent in the fossil record makes it difficult to achieve greater precision in such estimates, but the general direction of the trend is well supported. Most predictions of the contemporary extinction rate are based on combining estimates of species richness in tropical forest with estimates of rate of loss of these forests; species extinction is then predicted on the basis of the general species-area relationship, under which species richness will decline as area declines.

Projections of this sort suggest the contemporary extinction rate is very high.