Marine Animal Genera Research Articles

Selection by differential survival among marine animals in the Phanerozoic

The Gaia hypothesis posits that the Earth and its biosphere function as a single self-stabilizing system, but a key challenge is explaining how this could have arisen through Darwinian evolution. One theory is that of “selection by differential survival,” in which a clade’s extinction probability decreases with age as it accumulates adaptations resisting environmental disturbances. While this is hard to assess during early Earth history, we can assess whether this process operated among marine animal genera throughout the Phanerozoic. To that end, we analyzed time ranges of 36,117 extinct animal genera using fossil occurrence data from the Paleobiology Database in order to calculate marine metazoan extinction age selectivity, extinction rates, and speciation rates over the Phanerozoic. We identify four signatures of selection by differential survival: lower extinction rates among older lineages, heritability and taxonomically nested propagation of extinction resistance, reduced age selectivity during rare environmental perturbations, and differential extinction rather than speciation as the primary driver of the phenomenon. Evidence for this process at lower taxonomic levels also implies its possibility for life as a whole – indeed, the possibility of Gaia.

Environmental controls on geographic range size in marine animal genera

Here I test the hypothesis that temporal variation in geographic range size within genera is affected by the expansion and contraction of their preferred environments. Using occurrence data from the Paleobiology Database, I identify genera that have a significant affinity for carbonate or terrigenous clastic depositional environments that transcends the Database's representation of these environments during the stratigraphic range of each genus. These affinity assignments are not a matter of arbitrarily subdividing a continuum in preference; rather, genera form distinct, nonrandom subsets with respect to environmental preference. I tabulate the stage-by-stage transitions in range size within individual genera and the stage-by-stage changes in the extent of each environment. Comparing the two shows that genera with a preference for a given environment are more likely to increase in geographic range, and to show a larger average increase in range, when that environment increases in areal extent, and likewise for decreases in geographic range and environmental area. Similar results obtain for genera with preferences for reefal and non-reef settings. Simulations and subsampling experiments suggest that these results are not artifacts of methodology or sampling bias. Nor are they confined to particular higher taxa. Genera with roughly equal preference for carbonates and clastics do not have substantially broader geographic ranges than those with a distinct affinity, suggesting that, at this scale of analysis, spatial extent of preferred environment outweighs breadth of environmental preference in governing geographic range. These results pertain to changes over actual geologic time within individual genera, not overall average ranges. Recent work has documented a regular expansion and contraction when absolute time is ignored and genera are superimposed to form a composite average. Environmental preference may contribute to this pattern, but its role appears to be minor, limited mainly to the initial expansion and final contraction of relatively short-lived genera.

Determinants of early survival in marine animal genera

Genera by their very nature are expected to be monotypic and geographically and environmentally restricted at their origin, and most genera do not endure past their stage of first appearance. At the same time, those genera that do endure have a capacity to expand greatly in geographic range, environmental breadth, and species richness. Here we ask what it is that allows some genera and not others to survive past their inception. Using occurrence data from the Paleobiology Database, we find that initial geographic range has the strongest effect on survival, followed by environmental breadth, with the effect of species richness weaker on average. The effect of geographic range is strongest if measured as the distances spanned by the occurrences of a genus rather than the number of distinct areas in which a genus lives. We document substantial secular variation in selectivity of early survival. The most striking aspect of this variation is that survival is only weakly selective among genera that first appear during the Mesozoic. By following genera beyond their stage of first appearance, we find that selectivity with respect to all factors becomes systematically stronger as cohorts age and genera become more differentiated in range, breadth, and richness. This may help account for a previously identified statistical effect of genus age on the chances of survival.

FossilPlot, an Excel-based Computer Application for Teaching Stratigraphic Paleontology Using the Sepkoski Compendium of Fossil Marine Genera

FossilPlot is a new Excel-based spreadsheet application of the Sepkoski Compendium designed for educational use in paleontology and historical geology courses. This free software allows students to compile any combination of marine animal genera to generate graphs depicting diversity curves and stratigraphic ranges for the Phanerozoic. Sample exercises are provided to show how this software can be integrated into lecture, lab and field studies in university-level paleontology courses. The large volume and full accessibility of the dataset gives instructors a versatile tool to enhance active learning of the dynamic history of life.

Memoir 4: An Analysis of the History of Marine Animal Diversity

According to when they attained high diversity, major taxa of marine animals have been clustered into three groups, the Cambrian, Paleozoic, and Modern Faunas. Because the Cambrian Fauna was a relatively minor component of the total fauna after mid-Ordovician time, the Phanerozoic history of marine animal diversity is largely a matter of the fates of the Paleozoic and Modern Faunas. The fact that most late Cenozoic genera belong to taxa that have been radiating for tens of millions of years indicates that the post-Paleozoic increase in diversity indicated by fossil data is real, rather than an artifact of improvement of the fossil record toward the present.Assuming that ecological crowding produced the so-called Paleozoic plateau for family diversity, various workers have used the logistic equation of ecology to model marine animal diversification as damped exponential increase. Several lines of evidence indicate that this procedure is inappropriate. A plot of the diversity of marine animal genera through time provides better resolution than the plot for families and has a more jagged appearance. Generic diversity generally increased rapidly during the Paleozoic, except when set back by pulses of mass extinction. In fact, an analysis of the history of the Paleozoic Fauna during the Paleozoic Era reveals no general correlation between rate of increase for this fauna and total marine animal diversity. Furthermore, realistically scaled logistic simulations do not mimic the empirical pattern. In addition, it is difficult to imagine how some fixed limit for diversity could have persisted throughout the Paleozoic Era, when the ecological structure of the marine ecosystem was constantly changing. More fundamentally, the basic idea that competition can set a limit for marine animal diversity is incompatible with basic tenets of marine ecology: predation, disturbance, and vagaries of recruitment determine local population sizes for most marine species. Sparseness of predators probably played a larger role than weak competition in elevating rates of diversification during the initial (Ordovician) radiation of marine animals and during recoveries from mass extinctions. A plot of diversification against total diversity for these intervals yields a band of points above the one representing background intervals, and yet this band also displays no significant trend (if the two earliest intervals of the initial Ordovician are excluded as times of exceptional evolutionary innovation). Thus, a distinctive structure characterized the marine ecosystem during intervals of evolutionary radiation—one in which rates of diversification were exceptionally high and yet increases in diversity did not depress rates of diversification.Particular marine taxa exhibit background rates of origination and extinction that rank similarly when compared with those of other taxa. Rates are correlated in this way because certain heritable traits influence probability of speciation and probability of extinction in similar ways. Background rates of origination and extinction were depressed during the late Paleozoic ice age for all major marine invertebrate taxa, but remained correlated. Also, taxa with relatively high background rates of extinction experienced exceptionally heavy losses during biotic crises because background rates of extinction were intensified in a multiplicative manner; decimation of a large group of taxa of this kind in the two Permian mass extinctions established their collective identity as the Paleozoic Fauna.Characteristic rates of origination and extinction for major taxa persisted from Paleozoic into post-Paleozoic time. Because of the causal linkage between rates of origination and extinction, pulses of extinction tended to drag down overall rates of origination as well as overall rates of extinction by preferentially eliminating higher taxa having relatively high background rates of extinction. This extinction/origination ratchet depressed turnover rates for the residual Paleozoic Fauna during the Mesozoic Era. A decline of this fauna's extinction rate to approximately that of the Modern Fauna accounts for the nearly equal fractional losses experienced by the two faunas in the terminal Cretaceous mass extinction.Viewed arithmetically, the fossil record indicates slow diversification for the Modern Fauna during Paleozoic time, followed by much more rapid expansion during Mesozoic and Cenozoic time. When viewed more appropriately as depicting geometric—or exponential—increase, however, the empirical pattern exhibits no fundamental secular change: the background rate of increase for the Modern Fauna—the fauna that dominated post-Paleozoic marine diversity—simply persisted, reflecting the intrinsic origination and extinction rates of constituent taxa. Persistence of this overall background rate supports other evidence that the empirical record of diversification for marine animal life since Paleozoic time represents actual exponential increase. This enduring rate makes it unnecessary to invoke environmental change to explain the post-Paleozoic increase of marine diversity.Because of the resilience of intrinsic rates, an empirically based simulation that entails intervals of exponential increase for the Paleozoic and Modern Faunas, punctuated by mass extinctions, yields a pattern that is remarkably similar to the empirical pattern. It follows that marine animal genera and species will continue to diversify exponentially long into the future, barring disruption of the marine ecosystem by human-induced or natural environmental changes.