Sampling bias, gradual extinction patterns and catastrophes in the fossil record

Abstract
Catastrophic hypotheses for mass extinctions are commonly criticized because many taxa gradually disappear from the fossil record prior to the extinction. Presumably, a geologically instantaneous catastrophe would not cause a reduction in diversity or a series of minor extinctions before the actual mass extinction. Two types of sampling effects, however, could cause taxa to appear to decline before their actual biotic extinction. The first of these is reduced sample size provided in the sedimentary record and the second, which we examine in greater detail, is artificial range truncation. The fossil record is discontinuous in time and the recorded ranges of species or of higher taxa can only extend to their last known occurrence in the fossil record. If the distribution of last occurrences is random with respect to actual biotic extinction, then apparent extinctions will begin well before a mass extinction and will gradually increase in frequency until the mass extinction event, thus giving the appearance of a gradual extinction. Other factors, such as regressions, can exacerbate the bias toward gradual disappearance of taxa from the fossil record. Hence, gradual extinction patterns prior to a mass extinction do not necessarily eliminate catastrophic extinction hypotheses. The recorded ranges of fossils, especially of uncommon taxa or taxa in habitats not represented by a continuous record, may be inadequate to test either gradual or catastrophic hypotheses.