Proper identification of behavioral patterns is an important prerequisite for the identification of any trace fossil and even more so for its interpretation. For the last 70 yr, the continually advancing state of ichnological knowledge has led to a gradual recognition of recurrent patterns of organismal behavior documented in the fossil record, which in turn gave rise to the ethological categories. “Mortichnia” was proposed for traces created during a death struggle of the tracemaker but has been reported only in a few cases. Fish mortichnia so far have only been reported in one specimen recovered from the Upper Jurassic Plattenkalk of Nusplingen (SW Germany). The category mortichnia is refined herein, but remains ambiguous. Eight newly discovered unique specimens of mortichnia from Upper Cretaceous marine sediments in central Lebanon (Haqil, En Nammoura) are preserved together with their tracemakers and described herein. In addition, 14 further incomplete specimens were collected where no tracemakers are present. However, morphology and close provenance allow them to be assigned to the same ichnotaxon.

The Lebanese mortichnia originate from fish that were subjected to significant environmental or individual stress leading to their deaths. During death convulsions, their bodies created sedimentary structures with a specific recurring morphology. The ichnogenus Pinnichnus n. igen. with ichnospecies P. haqilensis and P. emmae n. ispp. is proposed for these specimens.

The Ediacaran/Cambrian transition (ECT; ~575–500 Ma) captures the early diversification of animals, including the oldest crown-group taxa of most major animal phyla alive today. Key to understanding the drivers underneath the ECT macroevolutionary patterns are the interactions of animals with one another and their environment, and how these interactions scale up to global diversity patterns. Understanding the ecology of ECT organisms is enabled by the abundance of Lagerstätten over this time period, with a relatively large proportion of soft-bodied organisms preserved, often within the communities in which they lived. Here, we review our understanding of organismal, community, and macroecology of the ECT, and how these different scales of ecological analyses relate to the macroevolutionary diversification patterns we see over this 75 Myr time period. Across all ecological scales, we find clear trends, starting with stochastic ecosystem dynamics dominated by generalist taxa in the first Ediacaran communities, to more structured, niche-driven specialist dynamics by Cambrian Epoch 2. These trends are reflected in organism functional morphology, the complexity and strength of organisms’ interactions within their communities, and large-scale metacommunity, biogeographic, and biodiversity patterns. Yet there is often a time delay between the origination of a new type of ecological interaction and when it is observed to impact the ecosystem as a whole. As such, while many modern ecological innovations were in place by the end of the Cambrian, the knock-on effects and complexity of these interactions continued to build up throughout the Phanerozoic, leading to the complex biosphere we have today.

Phacopid trilobites are well documented during the Paleozoic. Nevertheless, while 2D quantitative analyses have advanced our understanding of the morphological relationships among trilobites, the quantification of their morphological traits in 3D remains rarely documented. Based on two sets of morphological data (head and tail), 2D versus 3D shape quantification approaches were used to explore shape allometries as well as to explore how the shape variations can be explained by the phylogenetic relationships among phacopid trilobite species for the first time. We demonstrate that (1) there are similar patterns of morphological variability across taxa in 3D and 2D; (2) there are rather congruent results between 3D and 2D to discriminate taxa; (3) 2D and 3D landmarks capture different levels of detail, and the third dimension in 3D is very important for making taxonomic distinctions at the genus level; (4) there is congruity between 2D and 3D datasets for allometric patterns with results showing similar allometric slopes among species exhibiting a glabellar length decrease during growth leading to wider cephala; (5) the phylomorphospaces show tree branches that do not intersect, suggesting possible phylogenetic constraints on morphospace occupation for each species and supporting the idea that the Austerops and Morocops groups are sister clades that experienced different modes of morphological evolution; and (6) the morphological descriptors in morphometric analyses in 2D and 3D throughout phacopid evolution are effective.

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.

We analyzed the oxygen isotope composition of biogenic apatite phosphate (δ18Op) in fossil tooth enameloid to investigate the paleoecology of Late Cretaceous sharks in the Gulf Coastal Plain of Alabama, USA. We analyzed six different shark taxa from both the Mooreville Chalk and the Blufftown Formation. We compared shark δ18Op with the δ18Op of a co-occurring poikilothermic bony fish Enchodus petrosus as a reference for ambient conditions. Enchodus petrosus tooth enamel δ18Op values are similar between formations (21.3‰ and 21.4‰ Vienna Standard Mean Ocean Water [VSMOW], respectively), suggesting minimal differences in water δ18O between formations. Most shark taxa in this study are characterized by δ18Op values that overlap with E. petrosus values, indicating they likely lived in similar habitats and were also poikilothermic. Ptychodus mortoni and Cretoxyrhina mantelli exhibit significantly lower δ18Op values than co-occurring E. petrosus (P. mortoni δ18Op is 19.1‰ VSMOW in the Mooreville Chalk; C mantelli δ18Op is 20.2‰ VSMOW in the Mooreville Chalk and 18.1‰ VSMOW in the Blufftown Formation). Excursions into brackish or freshwater habitats and thermal water-depth gradients are unlikely explanations for the lower P. mortoni and C. mantelli δ18Op values. The low P. mortoni δ18Op value is best explained by higher body temperature relative to surrounding temperatures due to active heating (e.g., mesothermy) or passive heating due to its large body size (e.g., gigantothermy). The low C. mantelli δ18Op values are best explained by a combination of mesothermy (e.g., active heating) and migration (e.g., from the Western Interior Seaway, low-latitude warmer waters, or the paleo–Gulf Stream), supporting the hypothesis that mesothermy evolved in lamniform shark taxa during the Late Cretaceous. If the anomalous P. mortoni δ18Op values are also driven by active thermoregulation, this suggests that mesothermy evolved independently in multiple families of Late Cretaceous sharks.

The Cretaceous/Paleogene (K/Pg) mass extinction was a pivotal event in Earth history, the latest among five mass extinctions that devastated marine and terrestrial life. Whereas much research has focused on the global demise of dominant vertebrate groups, less is known about changes among plant communities during the K/Pg mass extinction. This study investigates a suite of 11 floral assemblages leading up to and across the K/Pg boundary in northeastern Montana constrained within a well-resolved chronostratigraphic framework. We evaluate the impact of the mass extinction on local plant communities as well as the timing of post-K/Pg recovery. Our results indicate that taxonomic composition changed significantly from the Late Cretaceous to Paleocene; we estimate that 63% of latest-Cretaceous plant taxa disappeared across the K/Pg boundary, on par with other records from North America. Overall, taxonomic richness dropped by ~23–33% from the Late Cretaceous to the Paleocene, a moderate decline compared with other plant records from this time. However, richness returned to Late Cretaceous levels within 900 kyr after the K/Pg boundary, significantly faster than observed elsewhere. We find no evidence that these results are due to preservational bias (i.e., differences in depositional environment) and instead interpret a dramatic effect of the K/Pg mass extinction on plant diversity and ecology. Overall, plant communities experienced major restructuring, that is, changes in relative abundance and unseating of dominant groups during the K/Pg mass extinction, even though no major (e.g., family-level) plant groups went extinct and communities in Montana quickly recovered in terms of taxonomic diversity. These results have direct bearing on our understanding of vegetation change during diversity crises, the differing responses of plant groups (e.g., angiosperms vs. gymnosperms), and spatial variation in extinction and recovery timing.

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.

Preserved records of tooth–bone interactions, known as tooth marks, can yield a wealth of information regarding organismal behavior and ecology. For this reason, workers in a wide range of disciplines, but particularly paleontology, have inspected and interpreted these features for decades. Although previous studies have gleaned invaluable insights, they have also described tooth marks using terminological frameworks that have been incompletely defined, have incorporated behavioral hypotheses in definitions, and/or have been inconsistently applied. To address these problems, we introduce the category-modifier (CM) system, the first system to both sort tooth marks into clearly defined main categories and use descriptive modifiers to characterize their appearance more precisely. The CM system is designed to apply to a wide range of vertebrates, to enable comparisons across disciplines and studies, and to help researchers keep their investigations into behavioral hypotheses free of circular reasoning.

Geometric morphometrics facilitates the quantification and visualization of variation in shape and proportion through the comparison of homologous features. Eublastoidea, a Paleozoic echinoderm clade with a conservative body plan, is an ideal group for morphometric analysis, because their plate junctions are homologous and identifiable on all species. Eublastoids have previously been grouped taxonomically by generalized shape types (e.g., globose). These shapes are often used in taxonomic descriptions and as characters in phylogenetic analyses. The underlying homology of these broad shape types has never been explored. Herein we apply the first comprehensive use of three-dimensional geometric morphometrics (3D GMM) on fossil echinoderms to investigate taxonomic assignments, temporal distribution, and whether the varying proportions of skeletal elements that produce the gross thecal morphology are distinguishable. Taxonomic assignments specifically at the ordinal and family levels show varying amounts of overlap in morphospace, suggesting that many assignments may not be reevaluated. Our results suggest that none of the generalized shape types are distinct in morphospace and, therefore, likely do not capture the homologous changes in taxa. The plate circlet ratios showed trends specifically relating to the deltoid plate circlet, which has the most variability. We reanalyzed previous work and subsetted our data to be more comparable and found that there are key differences between methodologies and landmarks that will require future evaluation. Applying modern technological methods to previously explored questions allows for an updated understanding of this important fossil clade and provides a framework for others to assess fossil clades in a similar manner.

The trace fossil record provides important insights into the evolution of early animals during the Ediacaran/Cambrian transition, with changes in ichnodiversity through time and between environments informing on the diversification of major body plans, behaviors, and niches. To quantify variation in the diversity of trace fossils across this critical interval, we propose a measure of trace fossil dissimilarity (ichnodissimilarity) based on vector calculation. Furthermore, by comparing discrepancies between the angular bisector and mean vector of two sets of vectorized fossil data, we are able to weigh the relative contribution of increases and decreases in the variation of occurrences of taxa. We used this metric to analyze an expansive dataset of Ediacaran/Cambrian trace fossils. The results allowed us to quantify the diversification of traces across this transition, informing on the timing of first appearance of different behaviors (e.g., foraging, grazing, and resting) and functional groups. By interpreting the results in the context of environmental changes and advancements in motility and sensory capabilities, we were able to pinpoint the onset and sequence of the Fortunian diversification event, Cambrian information revolution, and agronomic revolution, shedding light on the evolution of organismal body plans, behaviors, and locomotion during the Ediacaran/Cambrian transition. We identified two phases of origination and expansion during the divergence of early animal traces. Furthermore, by analyzing shallow- and deep-marine trace fossils, we were able to uncover evidence for a more rapid diversification of traces in shallow-marine environments, with progressive niche partitioning through the Ediacaran to Cambrian.

The last mass extinction event some 66 million years ago at the Late Cretaceous/Paleogene boundary caused the extinction of many clades, including the non-avian dinosaurs. Turtles, as well as several other vertebrate clades, survived. However, the debate about whether the diversity of turtles was affected during this event is still ongoing. Here, I calculate a global turtle diversity curve at the species level that shows that the diversity of turtle species was already in decline since the Campanian, before the extinction event, and was further reduced during the Danian. The sample coverage of turtle occurrences at the stage level is also calculated and discussed.

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.

It is a pleasure to be invited to contribute to the celebration of the publication of “punctuated equilibria” (“punk eek”) 50 years ago—the canonical version I did with Steve Gould (Eldredge and Gould 1972) at the behest of Tom Schopf for his visionary project to inject more thought, more interpretation and theory, into the working lives of paleontologists.

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.

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.