Paleontological research on predation has been expanding rapidly in scope, methods, and goals. The growing assortment of research strategies and goals has led to increasing differences in sampling strategies, types of data collected, definition of variables, and even reporting style. This methodological overview serves as a starting point for erecting some general methodological guidelines for studying the fossil record of predation. I focus here on trace fossils left by predators in the skeleton of their prey, arguably one of the most powerful sources of direct data on predator-prey interactions available in the fossil record. A critical survey of sampling protocols (data collecting strategy, sieve size, and sample size) and analytical approaches (predation intensity metrics, strategies for evaluating behavioral selectivity of predators, and taphonomic tests) reveals that various approaches can be fruitful depending on logistic circumstances and scientific goals of paleoecological projects. Despite numerous caveats and uncertainties, trace fossils left by predators on skeletons of their prey remain one of the most promising directions of research in paleoecology and evolutionary paleobiology.

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 importance of ecological interactions in driving the evolution of animals has been the focus of intense debate among paleontologists, evolutionary biologists, and macroecologists. To test whether the intensity of such interactions covaries with the secular evolutionary trend in global biodiversity, we compiled a species-level database of predation intensity, as measured by the frequency of common predation traces (drillings and repair scars ranging in age from Ediacaran to Holocene). The results indicate that the frequency of predation traces increased notably by the Ordovician, and not in the midPaleozoic as suggested by multiple previous studies. Importantly, these estimates of predation intensity and global diversity of marine metazoans correlate throughout the Phanerozoic fossil record regardless of corrections and methods applied. This concordance may represent (i) an ecological signal: long-term coupling of diversity and predation; (ii) a diversity-driven diffusion of predatory behaviors: an increased probability of more complex predatory strategies to appear at higher diversity levels; or (iii) a spurious concordance in signal capture: an artifact where rare species and less-frequent (e.g., traceproducing) predatory behaviors are both more detectable at times when sampling improves. The coupling of predation and diversity records suggests that macroevolutionary and macroecological patterns share common causative mechanisms that may reflect either historical processes or sampling artifacts.macroecology ͉ macroevolution C ongruencies and discordances between biologically relevant patterns extracted from the geological record offer key historical data about the processes that have governed the history of life. For example, correlations between secular changes in rock volume, sea level, and biodiversity (1, 2), ocean chemistry and types of biomineral skeletons (3), and local and global diversity levels (4) all opened major research avenues with broad historical relevance. More recently, key predictions of the important, and controversial, hypothesis of escalation (biotic systems have become more dangerous through the Phanerozoic, and organisms respond evolutionarily to their enemies) have been tested by using time series spanning the Phanerozoic (5). Madin et al. (5) suggest that, when the data are properly analyzed, there are no significant correlations between the long term patterns in the proportion of carnivorous marine invertebrates and the proportions of infaunal or mobile prey and between the proportion of bioturbators and the proportion of immobile epifauna. The results of Madin et al. and others did not support escalation as an important shaper in the history of Phanerozoic marine life, but the validity of their approach was subsequently debated (6-8). Here, by comparing secular changes in biodiversity (2, 4) with estimates of the intensity of predator-prey interactions, we offer a more direct test of this fundamental question shared by paleontologists, evolutionary biologists, and macroecologis...

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The Ediacara biota include macroscopic, morphologically complex soft-bodied organisms that appear globally in the late Ediacaran Period (575-542 Ma). The physiology, feeding strategies, and functional morphology of the modular Ediacara organisms (rangeomorphs and erniettomorphs) remain debated but are critical for understanding their ecology and phylogeny. Their modular construction triggered numerous hypotheses concerning their likely feeding strategies, ranging from micro-to-macrophagus feeding to photoautotrophy to osmotrophy. Macrophagus feeding in rangeomorphs and erniettomorphs is inconsistent with their lack of oral openings, and photoautotrophy in rangeomorphs is contradicted by their habitats below the photic zone. Here, we combine theoretical models and empirical data to evaluate the feasibility of osmotrophy, which requires high surface area to volume (SA/V) ratios, as a primary feeding strategy of rangeomorphs and erniettomorphs. Although exclusively osmotrophic feeding in modern ecosystems is restricted to microscopic bacteria, this study suggests that (i) fractal branching of rangeomorph modules resulted in SA/V ratios comparable to those observed in modern osmotrophic bacteria, and (ii) rangeomorphs, and particularly erniettomorphs, could have achieved osmotrophic SA/V ratios similar to bacteria, provided their bodies included metabolically inert material. Thus, specific morphological adaptations observed in rangeomorphs and erniettomorphs may have represented strategies for overcoming physiological constraints that typically make osmotrophy prohibitive for macroscopic life forms. These results support the viability of osmotrophic feeding in rangeomorphs and erniettomorphs, help explain their taphonomic peculiarities, and point to the possible importance of earliest macroorganisms for cycling dissolved organic carbon that may have been present in abundance during Ediacaran times.erniettomorphs ͉ rangeomorphs ͉ Fractofusus ͉ Pteridinium

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Ediacara fossils [575 to 542 million years ago (Ma)] represent Earth's oldest known complex macroscopic life forms, but their morphological history is poorly understood. A comprehensive quantitative analysis of these fossils indicates that the oldest Ediacara assemblage-the Avalon assemblage (575 to 565 Ma)-already encompassed the full range of Ediacara morphospace. A comparable morphospace range was occupied by the subsequent White Sea (560 to 550 Ma) and Nama (550 to 542 Ma) assemblages, although it was populated differently. In contrast, taxonomic richness increased in the White Sea assemblage and declined in the Nama assemblage. These diversity changes, occurring while morphospace range remained relatively constant, led to inverse shifts in morphological variance. The Avalon morphospace expansion mirrors the Cambrian explosion, and both events may reflect similar underlying mechanisms.