Effects of sampling standardization on estimates of Phanerozoic marine diversification.

Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.

Designer snails.

The Algorithmic Beauty of Sea Shells (2nd edn) by Hans Meinhardt Springer-Verlag, 1998. DM89.00/$54.95/£34.00 hbk (xi+236 pages) ISBN 3 540 63919 5.

Fossil preservation and the stratigraphic ranges of taxa.

The incompleteness of the fossil record hinders the inference of evolutionary rates and patterns. Here, we derive relationships among true taxonomic durations, preservation probability, and observed taxonomic ranges. We use these relationships to estimate original distributions of taxonomic durations, preservation probability, and completeness (proportion of taxa preserved), given only the observed ranges. No data on occurrences within the ranges of taxa are required. When preservation is random and the original distribution of durations is exponential, the inference of durations, preservability, and completeness is exact. However, reasonable approximations are possible given non-exponential duration distributions and temporal and taxonomic variation in preservability. Thus, the approaches we describe have great potential in studies of taphonomy, evolutionary rates and patterns, and genealogy. Analyses of Upper Cambrian-Lower Ordovician trilobite species, Paleozoic crinoid genera, Jurassic bivalve species, and Cenozoic mammal species yield the following results: (1) The preservation probability inferred from stratigraphic ranges alone agrees with that inferred from the analysis of stratigraphic gaps when data on the latter are available. (2) Whereas median durations based on simple tabulations of observed ranges are biased by stratigraphic resolution, our estimates of median duration, extinction rate, and completeness are not biased.(3) The shorter geologic ranges of mammalian species relative to those of bivalves cannot be attributed to a difference in preservation potential. However, we cannot rule out the contribution of taxonomic practice to this difference. (4) In the groups studied, completeness (proportion of species [trilobites, bivalves, mammals] or genera [crinoids] preserved) ranges from 60% to 90%. The higher estimates of completeness at smaller geographic scales support previous suggestions that the incompleteness of the fossil record reflects loss of fossiliferous rock more than failure of species to enter the fossil record in the first place.

Selectivity of end-Cretaceous marine bivalve extinctions.

Analyses of the end-Cretaceous or Cretaceous-Tertiary mass extinction show no selectivity of marine bivalve genera by life position (burrowing versus exposed), body size, bathymetric position on the continental shelf, or relative breadth of bathymetric range. Deposit-feeders as a group have significantly lower extinction intensities than suspension-feeders, but this pattern is due entirely to low extinction in two groups (Nuculoida and Lucinoidea), which suggests that survivorship was not simply linked to feeding mode. Geographically widespread genera have significantly lower extinction intensities than narrowly distributed genera. These results corroborate earlier work suggesting that some biotic factors that enhance survivorship during times of lesser extinction intensities are ineffectual during mass extinctions.

The role of extinction in evolution.

The extinction of species is not normally considered an important element of neodarwinian theory, in contrast to the opposite phenomenon, speciation. This is surprising in view of the special importance Darwin attached to extinction, and because the number of species extinctions in the history of life is almost the same as the number of originations; present-day biodiversity is the result of a trivial surplus of originations, cumulated over millions of years. For an evolutionary biologist to ignore extinction is probably as foolhardy as for a demographer to ignore mortality. The past decade has seen a resurgence of interest in extinction, yet research on the topic is still at a reconnaissance level, and our present understanding of its role in evolution is weak. Despite uncertainties, extinction probably contains three important elements. (i) For geographically widespread species, extinction is likely only if the killing stress is one so rare as to be beyond the experience of the species, and thus outside the reach of natural selection. (ii) The largest mass extinctions produce major restructuring of the biosphere wherein some successful groups are eliminated, allowing previously minor groups to expand and diversify. (iii) Except for a few cases, there is little evidence that extinction is selective in the positive sense argued by Darwin. It has generally been impossible to predict, before the fact, which species will be victims of an extinction event.

Geography of end-Cretaceous marine bivalve extinctions.

Analysis of the end-Cretaceous mass extinction, based on 3514 occurrences of 340 genera of marine bivalves (Mollusca), suggests that extinction intensities were uniformly global; no latitudinal gradients or other geographic patterns are detected. Elevated extinction intensities in some tropical areas are entirely a result of the distribution of one extinct group of highly specialized bivalves, the rudists. When rudists are omitted, intensities at those localities are statistically indistinguishable from those of both the rudist-free tropics and extratropical localities.

Extinction from a paleontological perspective.

Extinction of widespread species is common in evolutionary time (millions of years) but rare in ecological time (hundreds or thousands of years). In the fossil record, there appears to be a smooth continuum between background and mass extinction; and the clustering of extinctions at mass extinctions cannot be explained by the chance coincidence of independent events. Although some extinction is selective, much is apparently random in that survivors have no recognizable superiority over victims. Extinction certainly plays an important role in evolution, but whether it is constructive or destructive has not yet been determined.

Nonconscious intelligence in the universe.

Animals lacking humanoid intelligence have evolved systems indistinguishable in function, if not in structure, from systems built by humans. Although radio communication has never been verified in animals, it is completely feasible biologically. If such systems are present in non-intelligent organisms on other planets, then our chances of detecting life in the universe by current SETI methods are greatly enhanced.

Large-body impact and extinction in the Phanerozoic.

The kill curve for Phanerozoic marine species is used to investigate large-body impact as a cause of species extinction. Current estimates of Phanerozoic impact rates are combined with the kill curve to produce an impact-kill curve, which predicts extinction levels from crater diameter, on the working assumption that impacts are responsible for all "pulsed" extinctions. By definition, pulsed extinction includes the approximately 60% of Phanerozoic extinctions that occurred in short-lived events having extinction rates greater than 5%. The resulting impact-kill curve is credible, thus justifying more thorough testing of the impact-extinction hypothesis. Such testing is possible but requires an exhaustive analysis of radiometric dating of Phanerozoic impact events.

A kill curve for Phanerozoic marine species.

A kill curve for Phanerozoic species is developed from an analysis of the stratigraphic ranges of 17,621 genera, as compiled by Sepkoski. The kill curve shows that a typical species' risk of extinction varies greatly, with most time intervals being characterized by very low risk. The mean extinction rate of 0.25/m.y. is thus a mixture of long periods of negligible extinction and occasional pulses of much higher rate. Because the kill curve is merely a description of the fossil record, it does not speak directly to the causes of extinction. The kill curve may be useful, however, to li¿mit choices of extinction mechanisms.

Crater taphonomy and bombardment rates in the Phanerozoic.

Impact structures can be catalogued according to the age of the rocks in which they are now found ("country rocks"). The observed frequency distribution of craters by age of country rock is shown to be statistically indistinguishable from the predictions of a simple model in which it is assumed that the survival time for craters is the same as that for their target rocks. Other models are considered, but do not match the data. A lower limit of the rate of bombardment through the Phanerozoic, based only on documented craters, is 0.13 +/- 0.09 events/ma/10(8) km2 for craters with diameters > or = 10 km and 0.09 +/- 0.08 events/ma/10(8) km2 for craters with diameters > or = 20 km. The data allow, but do not demand, an increase in meteorite flux over the Phanerozoic but do not allow any significant decrease. We estimate that only about 6% of the existing terrestrial impact structures of diameter greater than 10 km have been discovered to date, and only 16% of those with diameter greater than 20 km.

The case for extraterrestrial causes of extinction.

The dramatic increase in our knowledge of large-body impacts that have occurred in Earth's history has led to strong arguments for the plausibility of meteorite impact as a cause of extinction. Proof of causation is often hampered, however, by our inability to demonstrate the synchronism of specific impacts and extinctions. A central problem is range truncation: the last reported occurrences of fossil taxa generally underestimate the true times of extinction. Range truncation, because of gaps in sedimentation, lack of preservation, or lack of discovery, can make sudden extinctions appear gradual and gradual extinctions appear sudden. Also, stepwise extinction may appear as an artefact of range truncation. These effects are demonstrated by experiments performed on data from field collections of Cretaceous ammonities from Zumaya (Spain). The challenge for future research is to develop a new calculus for treating biostratigraphic data so that fossils can provide more accurate assessments of the timing of extinctions.

The case for extraterrestrial causes of extinction.

The dramatic increase in our knowledge of large-body impacts that have occurred in Earth's history has led to strong arguments for the plausibility of meteorite impact as a cause of extinction. Proof of causation is often hampered, however, by our inability to demonstrate the synchronism of specific impacts and extinctions. A central problem is range truncation: the last reported occurrences of fossil taxa generally underestimate the true times of extinction. Range truncation, because of gaps in sedimentation, lack of preservation, or lack of discovery, can make sudden extinctions appear gradual and gradual extinctions appear sudden. Also, stepwise extinction may appear as an artefact of range truncation. These effects are demonstrated by experiments performed on data from field collections of Cretaceous ammonities from Zumaya (Spain). The challenge for future research is to develop a new calculus for treating biostratigraphic data so that fossils can provide more accurate assessments of the timing of extinctions.

The role of extraterrestrial phenomena in extinction.

In the several years since the Alvarez report of anomalously high iridium concentrations at the Cretaceous-Tertiary boundary, evidence for the involvement of meteorite impacts in biological extinction has increased dramatically. Much more research will be needed, however, before meteorite impact is established as a general causal factor in extinction. Of ever greater long-term interest is the possibility that other extraterrestrial forces have had important influences on the evolution of life. To recognize the effects of such forces, it will be necessary to coordinate the research of astronomy and paleontology so that testable predictions can be formulated. It is possible that known, systematic changes in the Solar System or Galaxy have had effects on global biology and that these effects have been preserved in the paleontological record.

Patterns of generic extinction in the fossil record.

Analysis of the stratigraphic records of 19,897 fossil genera indicates that most classes and orders show largely congruent rises and falls in extinction intensity throughout the Phanerozoic. Even an ecologically homogeneous sample of reef genera shows the same basic extinction profile. The most likely explanation for the congruence is that extinction is physically rather than biologically driven and that it is dominated by the effects of geographically widespread environmental perturbations influencing most habitats. Significant departures from the congruence are uncommon but important because they indicate physiological or habitat selectivity. The similarity of the extinction records of reef organisms and the marine biota as a whole confirms that reefs and other faunas are responding to the same history of environmental stress.

Mass extinction: a commentary.

Four neocatastrophist claims about mass extinction are currently being debated; they are that: 1, the late Cretaceous mass extinction was caused by large body impact; 2, as many as five other major extinctions were caused by impact; 3, the timing of extinction events since the Permian is uniformly periodic; and 4, the ages of impact craters on Earth are also periodic and in phase with the extinctions. Although strongly interconnected the four claims are independent in the sense that none depends on the others. Evidence for a link between impact and extinction is strong but still needs more confirmation through bed-by-bed and laboratory studies. An important area for future research is the question of whether extinction is a continuous process, with the rate increasing at times of mass extinctions, or whether it is episodic at all scales. If the latter is shown to be generally true, then species are at risk of extinction only rarely during their existence and catastrophism, in the sense of isolated events of extreme stress, is indicated. This is line of reasoning can only be considered an hypothesis for testing. In a larger context, paleontologists may benefit from a research strategy that looks to known Solar System and Galactic phenomena for predictions about environmental effects on earth. The recent success in the recognition of Milankovitch Cycles in the late Pleistocene record is an example of the potential of this research area.

Numerical simulations and the problem of periodicity in the cratering record.

Ages of craters in the record of impacts on earth may be uniformly period, totally random, or a mixture of the two. These alternatives are studied through numerical simulation wherein time-series analysis is performed on real and simulated sequences to which random noise has been added to represent age-dating uncertainty. We conclude that the real record is most likely to have been generated by a mixture of random and periodic impacts, with the random events constituting the majority.