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.

Despite growing attention on the influence of functional diversity changes on ecosystem functioning, a palaeoecological perspective on the long-term dynamic of functional diversity, including mass extinction crises, is still lacking. Here, using a novel multidimensional functional framework and comprehensive null-models, we compare the functional structure of Cambrian, Silurian and modern benthic marine biotas. We demonstrate that, after controlling for increases in taxonomic diversity, functional richness increased incrementally between each time interval with benthic taxa filling progressively more functional space, combined with a significant functional dissimilarity between periods. The modern benthic biota functionally overlaps with fossil biotas but some modern taxa, especially large predators, have new trait combinations that may allow more functions to be performed. From a methodological perspective, these results illustrate the benefits of using multidimensional instead of lower dimensional functional frameworks when studying changes in functional diversity over space and time.

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 (Ϸ1.9 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.body size ͉ Cambrian ͉ oxygen ͉ Precambrian ͉ trend D espite widespread scientific and popular fascination with the largest and smallest organisms and numerous studies of body size evolution within individual taxonomic groups (1-9), the first-order pattern of body size evolution through the history of life has not been quantified rigorously. Because size influences (and may be limited by) a broad spectrum of physiological, ecological, and evolutionary processes (10-16), detailed documentation of size trends may shed light on the constraints and innovations that have shaped life's size spectrum over evolutionary time as well as the role of the body size spectrum in structuring global ecosystems. Bonner (17) presented a figure portraying a gradual, monotonic increase in the overall maximum size of living organisms over the past 3.5 billion years. The pattern appears consistent with a simple, continuous underlying process such as diffusion (18), but could also reflect a more complex process. Bonner, for example, proposed that lineages evolve toward larger sizes to exploit unoccupied ecological niches. For decades, Bonner's has been the only attempt to quantify body size evolution over the entire history of life on Earth, but the data he presented were not tied to particular fossil specimens and were plotted without consistent controls on taxonomic scale against a nonlinear timescale. Hence, we have lacked sufficient data on the tempo and mode of maximum size change to evaluate potential first-order biotic and abiotic controls on organism size through the history of life.Here, we document the evolutionary history of body size on Earth, focusing on the upper limit to size. Use of maximum size allows us to assess constraints on the evolution of large body size and avoids the more substantial empirical difficulties in determining mean, median, or minimum size for all life or even for many individual taxa. For each era within the Archean Eon (4,000-2,500 Mya) and ...

The process of evolution hinders our ability to make large-scale ecological comparisons—such as those encompassing marine biotas spanning the Phanerozoic—because the compared entities are taxonomically and morphologically dissimilar. One solution is to focus instead on life habits, which are repeatedly discovered by taxa because of convergence. Such an approach is applied to a comparison of the ecological diversity of Paleozoic (Cambrian–Devonian) and modern marine biotas from deep-subtidal, soft-substrate habitats. Ecological diversity (richness and disparity) is operationalized by using a standardized ecospace framework that can be applied equally to extant and extinct organisms and is logically independent of taxonomy. Because individual states in the framework are chosen a priori and not customized for particular taxa, the framework fulfills the requirements of a universal theoretical ecospace. Unique ecological life habits can be recognized as each discrete,n-dimensional combination of character states in the framework. Although the basic unit of analysis remains the organism, the framework can be applied to other entities—species, clades, or multispecies assemblages—for the study of comparative paleoecology and ecology. Because the framework is quantifiable, it is amenable to analytical techniques used for morphological disparity. Using these methods, I demonstrate that the composite Paleozoic biota is approximately as rich in life habits as the sampled modern biota, but that the life habits in the modern biota are significantly more disparate than those in the Paleozoic; these results are robust to taphonomic standardization. Despite broadly similar distributions of life habits revealed by multivariate ordination, the modern biota is composed of life habits that are significantly enriched, among others, in mobility, infaunality, carnivory, and exploitation of other organisms (or structures) for occupation of microhabitats.

The taxonomic and ecologic composition of Earth's biota has shifted dramatically through geologic time, with some clades going extinct while others diversified. Here, we derive a metric that quantifies the change in biotic composition due to extinction or origination and show that it equals the product of extinction/origination magnitude and selectivity (variation in magnitude among groups). We also define metrics that describe the extent to which a recovery (1) reinforced or reversed the effects of extinction on biotic composition and (2) changed composition in ways uncorrelated with the extinction. To demonstrate the approach, we analyzed an updated compilation of stratigraphic ranges of marine animal genera. We show that mass extinctions were not more selective than background intervals at the phylum level; rather, they tended to drive greater taxonomic change due to their higher magnitudes. Mass extinctions did not represent a separate class of events with respect to either strength of selectivity or effect. Similar observations apply to origination during recoveries from mass extinctions, and on average, extinction and origination were similarly selective and drove similar amounts of biotic change. Elevated origination during recoveries drove bursts of compositional change that varied considerably in effect. In some cases, origination partially reversed the effects of extinction, returning the biota toward the pre-extinction composition; in others, it reinforced the effects of the extinction, magnifying biotic change. Recoveries were as important as extinction events in shaping the marine biota, and their selectivity deserves systematic study alongside that of extinction.

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The high concentration of molecular oxygen in Earth's atmosphere is arguably the most conspicuous and geologically important signature of life. Earth's early atmosphere lacked oxygen; accumulation began after the evolution of oxygenic photosynthesis in cyanobacteria around 3.0-2.5 billion years ago (Gya). Concentrations of oxygen have since varied, first reaching near-modern values ~600 million years ago (Mya). These fluctuations have been hypothesized to constrain many biological patterns, among them the evolution of body size. Here, we review the state of knowledge relating oxygen availability to body size. Laboratory studies increasingly illuminate the mechanisms by which organisms can adapt physiologically to the variation in oxygen availability, but the extent to which these findings can be extrapolated to evolutionary timescales remains poorly understood. Experiments confirm that animal size is limited by experimental hypoxia, but show that plant vegetative growth is enhanced due to

The average body size of brachiopods from a single habitat type increased gradually by more than two orders of magnitude during their initial Cambrian-Devonian radiation. This increase occurred nearly in parallel across all major brachiopod clades (classes and orders) and is consistent with Cope's rule: the tendency for size to increase over geological time. The increase is not observed within small, constituent clades (represented here by families), which underwent random, unbiased size changes. This scale-dependence is caused by the preferential origination of new families possessing initially larger body sizes. However, this increased family body size does not confer advantages in terms of greater geological duration or genus richness over families possessing smaller body sizes. We suggest that the combination of size-biased origination of families and parallel size increases among major, more inclusive brachiopod clades from a single habitat type is best explained by long-term, secular environmental changes during the Paleozoic that provided opportunities for body size increases associated with major morphological evolution.body volume ͉ origin of clades ͉ macroevolutionary trend ͉ species selection ͉ maximum likelihood