Paleobiology was founded 50 years ago to provide an outlet for biological paleontology, with an emphasis on investigating evolutionary patterns and processes that could apply generally across the history of life. While the intellectual and financial prospects for Paleobiology were uncertain in the beginning (Sepkoski 2012; Valentine 2009), this 50th anniversary issue testifies to its overwhelming success. Fifty years of anything well done deserves a celebration. These moments are a time for reflection and a time for imagining future directions. With this introduction, we outline briefly the start of the journal and two landmark anniversary issues, the 10th and the 25th. No special issue can adequately survey all research themes in a field as intellectually rich as paleobiology. However, these anniversary issues offer a snapshot of research directions, and they can trace the shift and expansion of established fields and mark the emergence of new ones. We end by outlining the contributions to the 50th anniversary issue that summarize current themes and future directions for the field.

Prospective and early-career paleontologists deserve an accurate assessment of employment opportunities in their chosen field of study. Drawing on a wide range of sources, we have produced an admittedly incomplete analysis of the current status and recent trends of permanent academic employment in the discipline. Obtaining more complete longitudinal data on employment trends is a major difficulty; this is a challenge that needs to be addressed. The number of job seekers is far in excess of available positions. There has been a clear erosion in the number of academic paleontologists in the United States, a trend exacerbated in recent years. The decline, in constant dollars, of federal funding for paleontological research has potential strong negative impacts on future hiring. The loss of paleontology positions has also had a deleterious effect on our professional societies, which have seen a loss of regular (professional) membership, although student membership remains strong. These trends also potentially negatively impact efforts to diversify the field. Professional societies need to better coordinate their efforts to address these serious issues. Individual paleontologists also must become more effective advocates for the importance and relevance of our science.

The spatial distribution of individuals within ecological assemblages and their associated traits and behaviors are key determinants of ecosystem structure and function. Consequently, determining the spatial distribution of species, and how distributions influence patterns of species richness across ecosystems today and in the past, helps us understand what factors act as fundamental controls on biodiversity. Here, we explore how ecological niche modeling has contributed to understanding the spatiotemporal distribution of past biodiversity and past ecological and evolutionary processes. We first perform a semiquantitative literature review to capture studies that applied ecological niche models (ENMs) to the past, identifying 668 studies. We coded each study according to focal taxonomic group, whether and how the study used fossil evidence, whether it relied on evidence or methods in addition to ENMs, spatial scale of the study, and temporal intervals included in the ENMs. We used trends in publication patterns across categories to anchor discussion of recent technical advances in niche modeling, focusing on paleobiogeographic ENM applications. We then explored contributions of ENMs to paleobiogeography, with a particular focus on examining patterns and associated drivers of range dynamics; phylogeography and within-lineage dynamics; macroevolutionary patterns and processes, including niche change, speciation, and extinction; drivers of community assembly; and conservation paleobiogeography. Overall, ENMs are powerful tools for elucidating paleobiogeographic patterns. ENMs are most commonly used to understand Quaternary dynamics, but an increasing number of studies use ENMs to gain important insight into both ecological and evolutionary processes in pre-Quaternary times. Deeper integration with traits and phylogenies may further extend those insights.

Life-history traits such as dispersal affect population attributes like gene flow, which can have consequences for speciation and extinction rates over macroevolutionary timescales. Here we use the Cheilostomatida, a monophyletic order of marine bryozoans, to test whether a life-history trait, larval brooding, affected the origination and extinction rates of genera throughout their fossil record. Cheilostome lineages that brood their larvae have shorter larval dispersal distances than non-brooding lineages, which has led to the hypothesis that the evolution of larval brooding decreased gene flow, increased origination, and drove their Cretaceous diversification. Brooding cheilostomes are far more diverse than non-brooding cheilostomes today, but it remains to be shown that brooding lineages have a higher origination rate than non-brooders. We fit time-varying Pradel seniority capture–mark–recapture models to look at the effect of brooding on origination and extinction rates during the Cretaceous cheilostome diversification, the Cretaceous/Paleogene mass extinction and recovery, and through the Cenozoic. Our results support the hypothesis that brooding affects origination rate, but only in the Cenomanian to Campanian. Extinction rates do not differ between brooding and non-brooding genera, and there is no regime shift specific to the Cretaceous/Paleogene mass extinction. Our work illustrates the importance of using fossil occurrences and time-varying models, which can detect interval-specific diversification differentials.

Over the last 50 years, paleobiology has made great strides in illuminating organisms and ecosystems in deep time through study of the often-curious nature of the fossil record itself. Among fossil deposits, none are as enigmatic or as important to our understanding of the history of life as Konservat-Lagerstätten, deposits that preserve soft-bodied fossils and thereby retain disproportionately large amounts of paleobiological information. While Konservat-Lagerstätten are often viewed as curiosities of the fossil record, decades of study have led to a better understanding of the environments and circumstances of exceptional fossilization.Whereas most types of exceptional preservation require very specific sets of conditions, which are rare but can occur at any time, Seilacher noted the problem of “anactualistic” modes of exceptional preservation, defined as modes of fossilization that are restricted in time and that no longer occur. Here, we focus on anactualistic preservation and the widely recognized overrepresentation of Konservat-Lagerstätten in the Ediacaran and early Paleozoic. While exceptional fossil deposits of Ediacaran, Cambrian, and Early Ordovician age encompass a number of modes of fossilization, the signal of exceptional preservation is driven by only two modes, Ediacara-type and Burgess Shale–type preservation. Both are “extinct” modes of fossilization that are no longer present in marine environments. We consider the controls that promoted widespread anactualistic preservation in the Ediacaran and early Paleozoic and their implications for the environmental conditions in which complex life first proliferated in the oceans.

Body mass is an important facet of reconstructing the paleobiology of fossil species and has, historically, been estimated from individual skeletal measurements. This paper demonstrates the potential advantages of estimating body mass using 3D geometric morphometrics on limb bones, which allows size to be explicitly contextualized within the functional morphology of the animal. Geometric morphometrics of the humerus and femur is used to estimate body mass in domestic dogs and wild canids, and the resulting estimates are compared with estimates made using limb bone dimensions and centroid size. In both groups, 3D methods produced more accurate estimates of body mass than linear dimensions. Additionally, centroid size was a poor predictor of body mass and should not be preferred over linear measurements. The use of 3D methods also reveals specific aspects of shape that are associated with different sizes. In general, relatively heavier individuals were associated with more robust bones and wider articulation sites, as well as larger attachment sites for muscles related to flexion and extension of the shoulder and hip joints. The body-mass equations constructed based on dogs were further evaluated on wild canids, as a test of their potential efficacy on fossil canids. With some adjustments, the body-mass estimation equations made for domestic dogs were able to reliably predict the mass of wild canids. These equations were then used to estimate body mass for a selection of fossil canids: Canis latrans, 16 kg; Aenocyon dirus, 67 kg; Phlaocyon multicuspus, 8 kg; and Hesperocyon gregarius, 2.5 kg.

The evolution of mammalian innovations like elevated growth rates, endothermy, and live birth has been the subject of paleobiological work for decades. Bone histology provides one of the best lines of evidence for assessing growth rates and life-history traits in the fossil record. However, little ontogenetic information is available for nonmammalian cynodonts, the stock lineage that eventually gave rise to mammals. Here, I report the bone histology of two traversodontid cynodonts from the Triassic Manda Formation of Tanzania. Using two femoral size series, I correlate bone tissue composition and limb size in Scalenodon angustifrons and Luangwa drysdalli. Fifteen individuals were analyzed from seven penecontemporaneous localities to assess intraspecific histovariation within traversodontid ontogenetic development for the first time. My results show that Scalenodon and Luangwa have disparate life histories despite being similarly sized contemporaries. Luangwa is characterized by parallel-fibered bone that transitions to woven-parallel bone early in ontogeny, interpreted as a growth spurt. This increase in growth rate is seen in small- and middle-sized individuals but is resorbed and remodeled in the largest, skeletally mature individual. By contrast, Scalenodon is characterized by woven-parallel tissue in early ontogeny. However, femur size is not correlated with changes in bone tissue composition, as multiple individuals show peripheral slower-growing tissue regardless of size, interpreted as highly developmentally plastic growth. Together, these results demonstrate that coeval members of Traversodontidae show disparate life histories. The underlying mechanisms to explain different life histories in these taxa are likely due to (1) intrinsic differences in growth rates and (2) varying degrees of developmentally flexible growth. The implication of this work is that intraspecific variation in growth dynamics may be more widespread than currently understood in cynodonts and that size is not a good indicator of maturity for some species.

Stratigraphic paleobiology uses a modern understanding of the construction of the stratigraphic record—from beds to depositional sequences to sedimentary basins—to interpret patterns and guide sampling strategies in the fossil record. Over the past 25 years, its principles have been established primarily through forward numerical modeling, originally in shallow-marine systems and more recently in nonmarine systems. Predictions of these models have been tested through outcrop-scale and basin-scale field studies, which have also revealed new insights. At multi-basin and global scales, understanding the joint development of the biotic and sedimentary records has come largely from macrostratigraphy, the analysis of gap-bound packages of sedimentary rock. Here, we present recent advances in six major areas of stratigraphic paleobiology, including critical tests in the Po Plain of Italy, mass extinctions and recoveries, contrasts of shallow-marine and nonmarine systems, the interrelationships of habitats and stratigraphic architecture, large-scale stratigraphic architecture, and the assembly of regional ecosystems. We highlight the potential for future research that applies stratigraphic paleobiological concepts to studies of climate change, geochemistry, phylogenetics, and the large-scale structure of the fossil record. We conclude with the need for more stratigraphic thinking in paleobiology.

A central goal in ecology is investigating the impact of major perturbations, such as invasion, on the structure of biological communities. One promising line of inquiry is using co-occurrence analyses to examine how species’ traits mediate coexistence and how major ecological, climatic, and environmental disturbances can affect this relationship and underlying mechanisms. However, present communities are heavily influenced by anthropogenic behaviors and may exhibit greater or lesser resistance to invasion than communities that existed before human arrival. Therefore, to disentangle the impact of individual disturbances on mammalian communities, it is important to examine community dynamics before humans. Here, we use the North American fossil record to evaluate the co-occurrence structure of mammals across the Great American Biotic Interchange. We compiled 126 paleocommunities from the late Pliocene (4–2.5 Ma) and early Pleistocene (2.5–1 Ma). Genus-level co-occurrence was calculated to identify significantly aggregated (co-occur more than expected) and segregated (co-occur less than expected) genus pairs. A functional diversity analysis was used to calculate functional distance between genus pairs to evaluate the relationship between pair association strength and functional role. We found that the strength distribution of aggregating and segregating genus pairs does not significantly change from the late Pliocene to the early Pleistocene, even with different mammals forming the pairs, including immigrant mammals from South America. However, we did find that significant pairs, both aggregations and segregations, became more similar in their functional roles following the Plio-Pleistocene transition. Due to different mammals and ecological roles forming significant associations and the stability of co-occurrence structure across this interval, our study suggests that mammals have fundamental ways of assembling that may have been altered by humans in the present.

The fitness of groups is often considered to be the average fitness among constituent members. This assumption has been useful for developing models of multilevel selection, but its uncritical adoption has held back our understanding of how multilevel selection actually works in nature. If group fitness is only equal to mean member fitness, then it is a simple task to erode the importance of group-level selection in all multilevel scenarios—species selection could then be reduced to organismal selection as easily as group selection can. Because selection from different levels can act on a single trait, body size, for example, there must be a way to translate one level of fitness to another. This allows the calculation of the contributions of selection at each level. If high-level fitness is not a simple function of member fitness, then how do they interlace? Here we reintroduce Leigh Van Valen’s argument for the inclusion of expansion as a component of fitness. We show that expansion is an integral part of fitness even if one does not subscribe to the energetic view of fitness from which Van Valen originally derived it. From a hierarchical perspective, expansion is the projection of demographic fitness from one level to the next level up; differential births and deaths at one level produce differential expansion one level above. Including expansion in our conceptual tool kit helps allay concerns about our ability to identify the level of selection using a number of methods as well as allowing for the various forms of multilevel selection to be seen as manifestations of the same basic process.

Ostracoderms, Paleozoic jawless stem-gnathostomes, are characterized by distinctive bony shields covering the front of their bodies. These headshields exhibit significant variations in morphology across species, boasting frontal, lateral, and dorsal processes. Ostracoderms represent pivotal intermediaries between modern jawless and jawed vertebrates, so understanding their biology and ecology is crucial for unraveling the selective pressures that shaped the early evolution and diversification of jawed vertebrates, which now dominate vertebrate diversity. This study employs virtual paleontology techniques and phylogenetic comparative methods to explore the hydrodynamic and ecological implications of these processes, focusing on pteraspidomorphs, the most diverse ostracoderm group. The analysis reveals widespread convergence in the arrangement and development of headshield processes. Lateral processes enhance hydrodynamic efficiency and generate lift, while combined lateral and dorsal processes provide stability in rolling, yawing, and pitching. Frontal processes reduce drag in many cases. These findings illuminate the enigmatic roles of ostracoderm headshields, showing how the dimensions and arrangement of their processes are biomechanically linked to a range of functions and ecological roles. Collectively, this highlights the intricate evolutionary pathways of lifestyles and ecologies within stem-gnathostomes, challenging the idea of a unidirectional trend toward more active lifestyles in vertebrate evolution and suggesting diverse ecological roles for ostracoderms.

Every organism interacts with a host of other organisms of the same and different species throughout its life. These biotic interactions have varying influences on the reproduction and dispersal of the organism, and hence also the population and species lineage to which the organism belongs. By extension, biotic interactions must contribute to the macroevolutionary patterns that we observe in the fossil record, but exactly how, when, and why are research questions we have been asking before the start of the journal Paleobiology. In this contribution for Paleobiology’s 50th anniversary, we present a brief overview of how paleobiologists have studied biotic interactions and their macroevolutionary consequences, recognizing paleontology’s unique position to contribute data and insights to the topic of interspecies interactions. We then explore, in a semi-free-form manner, what promising avenues might be open to those of us who use the fossil record to understand biotic interactions. In general, we emphasize the need for increased effort surrounding the understanding of ecological details, integration of different types of information, and model-based approaches.

Mass extinctions are natural experiments on the short- and long-term consequences of pushing biotas past breaking points, often with lasting effects on the structure and function of biodiversity. General properties of mass extinctions—exceptionally severe, taxonomically broad, global losses of taxa—are starting to come into focus through comparisons among dimensions of biodiversity, including morphological, functional, and phylogenetic diversity. Notably, functional diversity tends to persist despite severe losses of taxonomic diversity, whereas taxic and morphological losses may or may not be coupled. One of the biggest challenges in synthesizing and extracting general consequences of these events has been that they are often driven by multiple, interacting pressures, and the taxa and their traits vary among events, making it difficult to link single stressors to specific traits. Ongoing improvements in the taxonomic and stratigraphic resolution of these events for multiple clades will sharpen tests for selectivity and help to isolate hitchhiking effects, whereby organismal traits are carried by differential survival or extinction of taxa owing to other organismal or higher-level attributes, such as geographic-range size. Direct comparative analyses across multiple extinction events will also clarify the impacts of particular drivers on taxa, functional traits, and morphologies. It is not just the extinction filter that deserves attention, as the longer-term impact of extinctions derives in part from their ensuing rebounds. More work is needed to uncover the biotic and abiotic circumstances that spur some clades into re-diversification while relegating others to marginal shares of biodiversity. Combined insights from mass extinction filters and their rebounds bring a macroevolutionary view to approaching the biodiversity crisis in the Anthropocene, helping to pinpoint the clades, functional groups, and morphologies most vulnerable to extinction and failed rebounds.

Ichthyosauria, Plesiosauria, and Metriorhynchidae were apex predators in Mesozoic oceanic trophic networks. Previous stable oxygen isotope studies suggested that several taxa belonging to these groups were endothermic and that some of them were homeothermic organisms. However, these conclusions remain contentious owing to the associated uncertainties regarding the δ18O value and oxygen isotope fractionation relative to environmental seawater. Here, we present new bioapatite phosphate δ18O values (δ18Op) of Ichthyosauria, Plesiosauria, and Metriorhynchidae (Middle Jurassic to Early Cretaceous) recovered from mid- to high paleolatitudes to better constrain their thermophysiology and investigate the presence of regional heterothermies. The intraskeletal δ18Op variability failed to reveal distinct heterothermic patterns within any of the specimens, indicating either intrabody temperature homogeneity or an overriding diagenetic overprint of the original biological δ18Op bone record. Body temperature estimates have been reassessed from new and published δ18Op values of well-preserved isolated teeth, recently revised Mesozoic latitudinal δ18O oceanic gradients, and 18O-enrichment factors of fully aquatic air-breathing vertebrates. Our results confirm that Ichthyosauria were homeothermic endotherms (31°C to 41°C), while Plesiosauria were likely poikilothermic endotherms (27°C to 34°C). The new body temperature estimates of the Metriorhynchidae (25°C to 32°C) closely follow ambient temperatures and point to poikilothermic strategy with no or little endothermic ability. These results improve our understanding of Mesozoic marine reptile thermoregulation and indicate that due to their limited body temperature variations, the δ18Op values from Ichthyosauria fossil remains could be used as valuable archives of Mesozoic oceans δ18Osw values that may help improve paleoenvironmental and paleoclimatic reconstructions.

Ancient changes in the biosphere, from organismic traits to wholesale ecosystem changes, can be aligned with climate forcing across the Phanerozoic. Clear examples of abrupt climate warming causing biodiversity crises are primarily found between the Permian and Paleogene periods. During these times, catastrophic events occurred, resembling the extreme climate scenarios projected for the near future. The paleobiologic literature around these events generally supports the hypothesis that abrupt climate change was a dominant trigger of extinction and/or ecological crisis. When climate change and climate history are considered, virtually all post-Paleozoic global biotic events can be confidently attributed to climatic change, with abrupt warming (hyperthermal events) leaving the most consistent fingerprint. The combined stress of deoxygenation and warming are sufficient to explain marine extinction patterns across most hyperthermal events. Although ocean acidification may have contributed, the direct role of pH on the extinction toll of organisms is not consistently demonstrated. Future research can enhance the correspondence between the magnitudes of climatic changes and their biological impacts, even though observed rates of change cannot currently be compared across different timescales. Mimicking multi-scale approaches in modern ecology, paleontological approaches to climate impact research will benefit from specifically targeting scaling relationships.

Toothed whales (odontocetes) make use of high-frequency sounds to echolocate, differing significantly from their sister group baleen whales (mysticetes), which make use of low-frequency sound for long-distance communication. This divergence in auditory ability has led to considerable speculation as to how hearing functioned in the ancestral archaeocetes, and when the specializations of modern species arose. Numerous studies have attempted to infer auditory capabilities from morphological correlates valid in modern species. Here, we build upon these previous methods with a focus on cochlear structures that have well-understood links to function. We combine this with information on the sound conduction apparatus to chart the evolutionary trajectory of cetacean hearing. Our results suggest an initial move toward low-frequency specialization in early Eocene cetaceans, which coincides with the appearance of new sound conduction pathways. This paved the way for the later movement toward higher-frequency hearing in protocetids; however, the ultra-high- and low-frequency hearing specializations of both modern cetacean clades evolved after their divergence. We use these data to test the hypotheses that evolutionary brain size increases in cetaceans were related to the origin of high-frequency echolocation. We show that no shift in relative brain size coincides with any changes toward high-frequency perception. However, this does not rule out a role for other changes in hearing ability such as some simple forms of echolocation, similar to that suggested for hippopotamuses or bowhead whales, which may have been present in even the earliest cetaceans.

Humans have dramatically transformed ecosystems over the previous millennia and are potentially causing a mass extinction event comparable to the others that shaped the history of life. However, only a fraction of these impacts has been directly recorded, limiting conservation actions. Conservation paleobiology leverages geohistorical records to offer a long-term perspective on biodiversity change in the face of anthropogenic stressors. Nevertheless, the field's on-the-ground contributions to conservation outcomes are still developing. Here, we present an overview of directions in which paleobiological research could progress to aid conservation in the coming decades using elasmobranchs (sharks, rays, and skates)—a highly threatened group with a rich fossil record—as a model. These research directions are guided by areas of overlap between an expert-led list of current elasmobranch conservation priorities and available fossil and historical records. Four research topics emerged for which paleobiological research could address open questions in elasmobranch science and conservation: (1) baselines, (2) ecological roles, (3) threats, and (4) conservation priorities. Increasingly rich datasets and novel analytical frameworks present exciting opportunities to apply the elasmobranch fossil record to conservation practice. A similar approach could be extended to other clades. Given the synthetic nature of these research topics, we encourage collaboration across timescales and with conservation practitioners to safeguard the future of our planet's rapidly disappearing species.

Analysis of length-frequency data using the ELEFAN (Electronic Length-Frequency Analysis) approach and software is widely used to quantify the growth, mortality, longevity, and related parameters of Recent marine animals. Here we analyze a sample (n = 211) of the Ediacaran metazoan Parvancorina minchami Glaessner, 1958, from the Vendian siliciclastic marine deposits of the southeastern White Sea region, Russia. The results fit a von Bertalanffy equation with the parameters L∞ = 2 cm, K = yr−1 (with t0 not estimated) and an instantaneous rate of mortality (M) of 1.44 yr−1, implying M/K ≈ 2, as commonly occurs in Recent small invertebrates. These parameter values also imply a longevity for P. minchami of about 4 yr. The concepts and approach used here, previously applied to an Ordovician trilobite and a Cambrian radiodont, suggest that inferences on growth, mortality, longevity, and related parameters can be obtained from suitable size-frequency samples of long-extinct metazoans, opening new vistas on their growth dynamics and functional roles in ancient ecosystems.

The Neotropics host the highest diversity of plants on the Earth today and have done since at least the late Paleogene (~58 Ma). Several mechanisms have been proposed to explain this elevated diversity, but the empirical patterns of Neotropical plant diversification that would test key aspects of those mechanisms are still unclear. We use an extensive palynological database from northern South America to characterize patterns of extinction, origination, and diversity and their possible drivers since the Paleogene. The foreland Llanos basin of Colombia preserves the evolutionary history of Neotropical vegetation as well as the geological evolution of northern South America, offering a unique opportunity to study the relationship between the geological and fossil records. The palynological record of the Llanos basin has been intensely studied mainly for oil exploration, and we use this information to infer the evolutionary history of Neotropical vegetation in Colombia during the Cenozoic. There is no straightforward relationship between global temperature and Neotropical plant diversity. Nevertheless, environmental change had an important influence on the dynamics of diversification, especially during volatile climate intervals such as the Paleocene–Eocene and the Pleistocene. Pulses of regional extinction were driven by large-scale temperature excursions, including both warming and cooling phases. Time-lagged origination pulses results in rapid floral replacement on a timescale of 1 Myr. Origination and extinction are essentially balanced on long timescales, leading to a near-zero long-term net diversification rate. Regional geological events, like the uplift of the Andean Cordillera, and changes in paleogeography also played an important role in Neotropical plant diversification.

The first compilations of Proterozoic eukaryote diversity, published in the 1980s showed a dramatic peak in the Tonian Period (1000–720 Ma), interpreted as the initial radiation of eukaryotes in the marine realm. Over the decades, new discoveries filled in the older part of the record and the peak diminished, but the idea of a Tonian radiation of eukaryotes has remained strong, and is now widely accepted as fact. We present a new diversity compilation based on 181 species and 713 species occurrences from 145 formations ranging in age from 1890 Ma to 720 Ma and find a significant increase in diversity in the Tonian. However, we also find that the number of eukaryotic species through time is highly correlated with the number of formations in our dataset (i.e. eukaryote-bearing formations) through time. This correlation is robust to interpretations of eukaryote affinity, bin size, and bin boundaries. We also find that within-assemblage diversity—a measure thought to circumvent sampling bias—is related to the number of eukaryote-bearing formations through time. Biomarkers show a similar pattern to body fossils, where the rise of eukaryotic biosignatures correlates with increased sampling. We find no evidence that the proportion of eukaryote-bearing versus all fossiliferous formations changed through the Proterozoic, as might be expected if the correlation reflected an increase in eukaryote diversity driving an increase in the number of eukaryote-bearing formations. Although the correlation could reflect a common cause such as changes in sea level driving both diversification and an increase in sedimentary rock volume, we favor the explanation that the pattern of early eukaryote diversity is driven by variations in paleontological sampling.