Steve C. Wang

Origination, Extinction, And Mass Depletions Of Marine Diversity

Abstract In post-Cambrian time, five events—the end-Ordovician, end-Frasnian in the Late Devonian, end-Permian, end-Triassic, and end-Cretaceous—are commonly grouped as the “big five” global intervals of mass extinction. Plotted by magnitude, extinction intensities for all Phanerozoic substages show a continuous distribution, with the five traditionally recognized mass extinctions located in the upper tail. Plotted by time, however, proportional extinctions clearly divide the Phanerozoic Eon into six stratigraphically coherent intervals of alternating high and low extinction intensity. These stratigraphic neighborhoods provide a temporal context for evaluating the intensity of extinction during the “big five” events. Compared with other stages and substages in the same neighborhood, only the end-Ordovician, end-Permian, and end-Cretaceous extinction intensities appear as outliers. Moreover, when origination and extinction are considered together, only these three of the “big five” events appear to have been generated exclusively by elevated extinction. Low origination contributed more than high extinction to the marked loss of diversity in the late Frasnian and at the end of the Triassic. Therefore, whereas the “big five” events are clearly times when diversity suffered mass depletion, only those at the end of the Ordovician, Permian, and Cretaceous periods unequivocally qualify as globally distinct mass extinctions. Each of the three has a unique pattern of extinction, and the diversity dynamics of these events differ, as well, from the other two major diversity depletions. As mass depletions of diversity have no common effect, common causation seems unlikely.

Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity

The maximum size of organisms has increased enormously since the initial appearance of life >3.5 billion years ago (Gya), but the pattern and timing of this size increase is poorly known. Consequently, controls underlying the size spectrum of the global biota have been difficult to evaluate. Our period-level compilation of the largest known fossil organisms demonstrates that maximum size increased by 16 orders of magnitude since life first appeared in the fossil record. The great majority of the increase is accounted for by 2 discrete steps of approximately equal magnitude: the first in the middle of the Paleoproterozoic Era (≈1.9 Gya) and the second during the late Neoproterozoic and early Paleozoic eras (0.6–0.45 Gya). Each size step required a major innovation in organismal complexity—first the eukaryotic cell and later eukaryotic multicellularity. These size steps coincide with, or slightly postdate, increases in the concentration of atmospheric oxygen, suggesting latent evolutionary potential was realized soon after environmental limitations were removed.

Trophic network models explain instability of Early Triassic terrestrial communities

Studies of the end-Permian mass extinction have emphasized potential abiotic causes and their direct biotic effects. Less attention has been devoted to secondary extinctions resulting from ecological crises and the effect of community structure on such extinctions. Here we use a trophic network model that combines topological and dynamic approaches to simulate disruptions of primary productivity in palaeocommunities. We apply the model to Permian and Triassic communities of the Karoo Basin, South Africa, and show that while Permian communities bear no evidence of being especially susceptible to extinction, Early Triassic communities appear to have been inherently less stable. Much of the instability results from the faster post-extinction diversification of amphibian guilds relative to amniotes. The resulting communities differed fundamentally in structure from their Permian predecessors. Additionally, our results imply that changing community structures over time may explain long-term trends like declining rates of Phanerozoic background extinction

Estimating the diversity of dinosaurs

Despite current interest in estimating the diversity of fossil and extant groups, little effort has been devoted to estimating the diversity of dinosaurs. Here we estimate the diversity of nonavian dinosaurs at ≈1,850 genera, including those that remain to be discovered. With 527 genera currently described, at least 71% of dinosaur genera thus remain unknown. Although known diversity declined in the last stage of the Cretaceous, estimated diversity was steady, suggesting that dinosaurs as a whole were not in decline in the 10 million years before their ultimate extinction. We also show that known diversity is biased by the availability of fossiliferous rock outcrop. Finally, by using a logistic model, we predict that 75% of discoverable genera will be known within 60–100 years and 90% within 100–140 years. Because of nonrandom factors affecting the process of fossil discovery (which preclude the possibility of computing realistic confidence bounds), our estimate of diversity is likely to be a lower bound.

IDENTIFYING HETEROGENEITY IN RATES OF MORPHOLOGICAL EVOLUTION: DISCRETE CHARACTER CHANGE IN THE EVOLUTION OF LUNGFISH (SARCOPTERYGII; DIPNOI)

Quantifying rates of morphological evolution is important in many macroevolutionary studies, and critical when assessing possible adaptive radiations and episodes of punctuated equilibrium in the fossil record. However, studies of morphological rates of change have lagged behind those on taxonomic diversification, and most authors have focused on continuous characters and quantifying patterns of morphological rates over time. Here, we provide a phylogenetic approach, using discrete characters and three statistical tests to determine points on a cladogram (branches or entire clades) that are characterized by significantly high or low rates of change. These methods include a randomization approach that identifies branches with significantly high rates and likelihood ratio tests that pinpoint either branches or clades that have significantly higher or lower rates than the pooled rate of the remainder of the tree. As a test case for these methods, we analyze a discrete character dataset of lungfish, which have long been regarded as “living fossils” due to an apparent slowdown in rates since the Devonian. We find that morphological rates are highly heterogeneous across the phylogeny and recover a general pattern of decreasing rates along the phylogenetic backbone toward living taxa, from the Devonian until the present. Compared with previous work, we are able to report a more nuanced picture of lungfish evolution using these new methods.

Macroevolutionary Patterns In The Evolutionary Radiation Of Archosaurs (Tetrapoda: Diapsida)

The rise of archosaurs during the Triassic and Early Jurassic has been treated as a classic example of an evolutionary radiation in the fossil record. This paper reviews published studies and provides new data on archosaur lineage origination, diversity and lineage evolution, morpho- logical disparity, rates of morphological character change, and faunal abundance during the Triassic-Early Jurassic. The fundamental archosaur lineages originated early in the Triassic, in concert with the highest rates of character change. Disparity and diversity peaked later, during the Norian, but the most significant increase in disparity occurred before maximum diversity. Archo- saurs were rare components of Early-Middle Triassic faunas, but were more abundant in the Late Triassic and pre-eminent globally by the Early Jurassic. The archosaur radiation was a drawn-out event and major components such as diversity and abundance were discordant from each other. Crurotarsans (crocodile-line archosaurs) were more disparate, diverse, and abundant than avemeta- tarsalians (bird-line archosaurs, including dinosaurs) during the Late Triassic, but these roles were reversed in the Early Jurassic. There is no strong evidence that dinosaurs outcompeted or gradually eclipsed crurotarsans during the Late Triassic. Instead, crurotarsan diversity decreased precipitously by the end-Triassic extinction, which helped usher in the age of dinosaurian dominance.

Cope’s rule in the evolution of marine animals

Getting bigger all the time In todays world, many animal species are large, with even larger species only recently extinct, but the first animals to evolve were tiny. Was this increase in size due to active selection or to some more random process? Heim et al. test the classic hypothesis known as Copes rule, which posits that there is selection for increasing body size. They analyzed a data set that spans over 500 million years and includes more than 17,000 marine animal species. In support of Copes rule, body volumes have increased by over five orders of magnitude since the first animals evolved. Furthermore, modeling suggests that such a massive increase could not have emerged from a random process. Science, this issue p. 867 Diversification produced a 150-fold increase in the mean size of marine animals over the past 542 million years. Cope’s rule proposes that animal lineages evolve toward larger body size over time. To test this hypothesis across all marine animals, we compiled a data set of body sizes for 17,208 genera of marine animals spanning the past 542 million years. Mean biovolume across genera has increased by a factor of 150 since the Cambrian, whereas minimum biovolume has decreased by less than a factor of 10, and maximum biovolume has increased by more than a factor of 100,000. Neutral drift from a small initial value cannot explain this pattern. Instead, most of the size increase reflects differential diversification across classes, indicating that the pattern does not reflect a simple scaling-up of widespread and persistent selection for larger size within populations.

The Red Queen revisited: reevaluating the age selectivity of Phanerozoic marine genus extinctions

Abstract Extinction risk is inversely related to genus age (time since first appearance) in most intervals of the Phanerozoic marine fossil record, in apparent contradiction to the macroevolutionary Red Queens Hypothesis, which posits that extinction risk is independent of taxon age. Age-dependent increases in the mean species richness and geographic range of genera have been invoked to reconcile this genus-level observation with the presumed prevalence of Red Queen dynamics at the species level. Here we test these explanations with data from the Paleobiology Database. Multiple logistic regression demonstrates that the association of extinction risk with genus age is not adequately explained by species richness or geographic range: there is a residual association between age and extinction risk even when range and richness effects are accounted for. Throughout most of the Phanerozoic the age selectivity gradient is highest among the youngest age cohorts, whereas there is no association between age and extinction risk among older age cohorts. Some of the apparent age selectivity of extinction in the global fauna is attributable to differences in extinction rate among taxonomic groups, but extinction risk declines with genus age even within most taxonomic orders. Notable exceptions to this pattern include the Cambrian–Ordovician, latest Permian, Triassic, and Paleocene intervals. The association of age with extinction risk could reflect sampling heterogeneity or taxonomic practice more than biological reality, but at present it is difficult to evaluate or correct for such biases. Alternatively, the pattern may reflect consistent extinction selectivity on some as-yet unidentified covariate of genus age. Although this latter explanation is not compatible with a Red Queen model if most genus extinctions have resulted from biological interactions, it may be applicable if most genus extinctions have instead been caused by recurrent physical disturbances that repeatedly impose similar selective pressures.

Improved Confidence Intervals For Estimating The Position Of A Mass Extinction Boundary

Abstract Marshall (1995) used the distribution of the endpoints of 50% range extensions added to the stratigraphic ranges of individual taxa to bracket the position of an extinction boundary. Here we describe two improvements to Marshalls method. First, we show that more precise estimates of the position of such a boundary may be obtained using range extensions with confidence levels of less than 50% (e.g., 20%). Second, we introduce a new method of calculating confidence intervals that explicitly takes into account the position of the highest fossil find. Incorporating these improvements leads to confidence intervals for simulated data sets that are approximately four times more precise than those obtained by using Marshalls (1995) original method and approximately twice as precise as those using other published methods. We provide a look-up table that shows for different numbers of taxa the confidence level that should be used to maximize the precision of the estimated position of the extinction boundary, while ensuring that the boundary still lies within the stratigraphic interval bounded by at least one range extension. Unlike some other methods, our method is nonparametric and does not make the restrictive assumption of uniform preservation and recovery potential. We apply the method to Macellaris (1986) ammonite data from the late Cretaceous of Seymour Island, Antarctica.

Within- And Among-Genus Components Of Size Evolution During Mass Extinction, Recovery, And Background Intervals: A Case Study Of Late Permian Through Late Triassic Foraminifera

Abstract One of the best-recognized patterns in the evolution of organismal size is the tendency for mean and maximum size within a clade to decrease following a major extinction event and to increase during the subsequent recovery interval. Because larger organisms are typically thought to be at higher extinction risk than their smaller relatives, it has commonly been assumed that size reduction mostly reflects the selective extinction of larger species. However, to our knowledge the relative importance of within- and among-lineage processes in driving overall trends in body size has never been compared quantitatively. In this study, we use a global, specimen-level database of foraminifera to study size evolution from the Late Permian through Late Triassic. We explicitly decompose size evolution into within- and among-genus components. We find that size reduction following the end-Permian mass extinction was driven more by size reduction within surviving species and genera than by the selective extinction of larger taxa. Similarly, we find that increase in mean size across taxa during Early Triassic biotic recovery was a product primarily of size increase within survivors and the extinction of unusually small taxa, rather than the origination of new, larger taxa. During background intervals we find no strong or consistent tendency for extinction, origination, or within-lineage change to move the overall size distribution toward larger or smaller sizes. Thus, size stasis during background intervals appears to result from small and inconsistent effects of within- and among-lineage processes rather than from large but offsetting effects of within- and among-taxon components. These observations are compatible with existing data for other taxa and extinction events, implying that mass extinctions do not influence size evolution by simply selecting against larger organisms. Instead, they appear to create conditions favorable to smaller organisms.