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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.

NASA Program Exobiology↗

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.

Non-NASA Center↗

Ten years in the library: new data confirm paleontological patterns

A comparison is made between compilations of times of origination and extinction of fossil marine animal families published in 1982 and 1992. As a result of ten years of library research, half of the information in the compendia has changed: families have been added and deleted, low-resolution stratigraphic data been improved, and intervals of origination and extinction have been altered. Despite these changes, apparent macroevolutionary patterns for the entire marine fauna have remained constant. Diversity curves compiled from the two data bases are very similar, with a goodness-of-fit of 99%; the principal difference is that the 1992 curve averages 13% higher than the older curve. Both numbers and percentages of origination and extinction also match well, with fits ranging from 83% to 95%. All major events of radiation and extinction are identical. Therefore, errors in large paleontological data bases and arbitrariness of included taxa are not necessarily impediments to the analysis of pattern in the fossil record, so long as the data are sufficiently numerous.

NASA Discipline Number 52-40↗

A Cretaceous-Tertiary mass extinction? Were most of Earth's species killed off?

For the past decade, the scientific and popular press have carried frequent articles about a catastrophic mass extinction that supposedly destroyed the majority of the earth's species, including the dinosaurs, approximately 65 million years ago. Since 1980, more than 2000 papers and books have dealt with some aspect of a mass extinction at the Cretaceous-Tertiary (K/T) boundary. One authoritative estimate of the severity of the extinctions is that 60-80% of all the living species became extinct at this boundary (Raup 1988). There appears to be a general acceptance of the fact that such a great catastrophe did occur. Most of the argument among scientists now is devoted to the determination of the cause. In this article, I argue that the species changes at the K/T boundary were neither sudden nor catastrophic. They were most likely caused by a regression of sea level that led to a decrease in primary production.

NASA Discipline Exobiology↗

Tektite-bearing, deep-water clastic unit at the Cretaceous-Tertiary boundary in northeastern Mexico

The hypothesis of Cretaceous-Tertiary (K-T) boundary impact on Yucatan, Mexico, predicts that nearby sites should show evidence of proximal impact ejecta and disturbance by giant waves. An outcrop along the Arroyo el Mimbral in northeastern Mexico contains a layered clastic unit up to 3 m thick that interrupts a biostratigraphically complete pelagic-marl sequence deposited at more than 400 m water depth. The marls were found to be unsuitable for determining magnetostratigraphy, but foraminiferal biostratigraphy places the clastic unit precisely at the K-T boundary. We interpret this clastic unit as the deposit of a megawave or tsunami produced by an extraterrestrial impact. The clastic unit comprises three main subunits. (1) The basal "spherule bed" contains glass in the form of tektites and microtektites, glass spherules replaced by chlorite-smectite and calcite, and quartz grains showing probable shock features. This bed is interpreted as a channelized deposit of proximal ejecta. (2) A set of lenticular, massive, graded "laminated beds" contains intraclasts and abundant plant debris, and may be the result of megawave backwash that carried coarse debris from shallow parts of the continental margin into deeper water. (3) At the top, several thin "ripple beds" composed of fine sand are separated by clay drapes; they are interpreted as deposits of oscillating currents, perhaps a seiche. An iridium anomaly (921 +/- 23 pg/g) is observed at the top of the ripple beds. Our observations at the Mimbral locality support the hypothesis of a K-T impact on nearby Yucatan.

NASA Discipline Number 52-40↗

A model of onshore-offshore change in faunal diversity

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.

NASA Discipline Number 52-40↗

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 inverted question markmit choices of extinction mechanisms.

NASA Discipline Number 52-40↗

Periodicity in extinction and the problem of catastrophism in the history of life

The hypothesis that extinction events have recurred periodically over the last quarter billion years is greatly strengthened by new data on the stratigraphic ranges of marine animal genera. In the interval from the Permian to Recent, these data encompass some 13,000 generic extinctions, providing a more sensitive indicator of species-level extinctions than previously used familial data. Extinction time series computed from the generic data display nine strong peaks that are nearly uniformly spaced at 26 Ma intervals over the last 270 Ma. Most of these peaks correspond to extinction events recognized in more detailed, if limited, biostratigraphic studies. These new data weaken or negate most arguments against periodicity, which have involved criticisms of the taxonomic data base, sampling intervals, chronometric time scales, and statistical methods used in previous analyses. The criticisms are reviewed in some detail and various new calculations and simulations, including one assessing the effects of paraphyletic taxa, are presented. Although the new data strengthen the case for periodicity, they offer little new insight into the deriving mechanism behind the pattern. However, they do suggest that many of the periodic events may not have been catastrophic, occurring instead over several stratigraphic stages or substages.

NASA Discipline Number 52-40↗

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.

NASA Program Exobiology↗

Continental-pelagic carbonate partitioning and the global carbonate-silicate cycle

A carbonate-silicate geochemical cycle model is developed and used to explore dynamic and climatic consequences of constraints on shallow-water carbonate burial and possible carbon loss to the mantle associated with sea-floor subduction. The model partitions carbonate deposition between shallow-water and deep-water environments and includes carbon fluxes between the mantle and lithosphere. When total lithospheric carbonate mass is constant, there are two stable steady states, one in which the carbonate burial flux is mostly continental and another in which it is mostly pelagic. The continental steady state is characterized by a low metamorphic CO2 flux to the atmosphere and predominantly shallow-water carbonate burial. The pelagic steady state is characterized by a high metamorphic CO2 flux and predominantly deep-water carbonate burial. For reasonable parameter values, when total lithospheric carbonate mass is allowed to vary, the model oscillates between predominantly continental and predominantly pelagic modes. Model results suggest that carbonate deposition patterns established during the Cenozoic may be pushing the Earth system from the continental to the pelagic mode on a time scale of 10(8) yr, with a possible consequent order-of-magnitude increase in the metamorphic CO2 flux to the atmosphere.

NASA Program Exobiology↗

Evolutionary dynamics of plants and animals: a comparative approach

Patterns of longevity and rate of appearance of taxa in the fossil record indicate a different evolutionary dynamic between land plants and marine invertebrates. Among marine invertebrates, rates of taxonomic turnover declined through the Phanerozoic, with increasingly extinction-resistant, long-lived, clades coming to dominate. Among terrestrial vascular plants, rates of turnover increased through the Phanerozoic, with short-lived, extinction-prone clades coming to dominate from the Devonian to the present. Terrestrial vertebrates appear to approximate the marine invertebrate pattern more closely than the plant record. We identify two features which individually or jointly may have influenced this distinction. First, land plants continuously invaded stressful environments during their evolution, while marine invertebrates and terrestrial vertebrates did not. Second, the relative structural simplicity and indeterminate mode of plant growth vs. the relative structural complexity and determinate mode of animal growth may have influenced the timing of major clade origin in the two groups.

Review↗

Proximal impact deposits at the Cretaceous-Tertiary boundary in the Gulf of Mexico: a restudy of DSDP Leg 77 Sites 536 and 540

Restudy of Deep Sea Drilling Project Sites 536 and 540 in the southeast Gulf of Mexico gives evidence for a giant wave at Cretaceous-Tertiary boundary time. Five units are recognized: (1) Cenomanian limestone underlies a hiatus in which the five highest Cretaceous stages are missing, possibly because of catastrophic K-T erosion. (2) Pebbly mudstone, 45 m thick, represents a submarine landslide possibly of K-T age. (3) Current-bedded sandstone, more than 2.5 m thick, contains anomalous iridium, tektite glass, and shocked quartz; it is interpreted as ejecta from a nearby impact crater, reworked on the deep-sea floor by the resulting tsunami. (4) A 50-cm interval of calcareous mudstone containing small Cretaceous planktic foraminifera and the Ir peak is interpreted as the silt-size fraction of the Cretaceous material suspended by the impact-generated wave. (5) Calcareous mudstone with basal Tertiary forams and the uppermost tail of the Ir anomaly overlies the disturbed interval, dating the impact and wave event as K-T boundary age. Like Beloc in Haiti and Mimbral in Mexico, Sites 536 and 540 are consistent with a large K-T age impact at the nearby Chicxulub crater.

NASA Discipline Exobiology↗

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.

NASA Discipline Exobiology↗

Insect diversity in the fossil record

Insects possess a surprisingly extensive fossil record. Compilation of the geochronologic ranges of insect families demonstrates that their diversity exceeds that of preserved vertebrate tetrapods through 91 percent of their evolutionary history. The great diversity of insects was achieved not by high origination rates but rather by low extinction rates comparable to the low rates of slowly evolving marine invertebrate groups. The great radiation of modern insects began 245 million years ago and was not accelerated by the expansion of angiosperms during the Cretaceous period. The basic trophic machinery of insects was in place nearly 100 million years before angiosperms appeared in the fossil record.

NASA Discipline Exobiology↗