The field of sclerochronology has long been known to paleobiologists. Yet, despite the central role of growth rate, age, and body size in questions related to macroevolution and evolutionary ecology, these types of studies and the data they produce have received only episodic attention from paleobiologists since the field's inception in the 1960s. It is time to reconsider their potential. Not only can sclerochronological data help to address long-standing questions in paleobiology, but they can also bring to light new questions that would otherwise have been impossible to address. For example, growth rate and life-span data, the very data afforded by chronological growth increments, are essential to answer questions related not only to heterochrony and hence evolutionary mechanisms, but also to body size and organism energetics across the Phanerozoic. While numerous fossil organisms have accretionary skeletons, bivalves offer perhaps one of the most tangible and intriguing pathways forward, because they exhibit clear, typically annual, growth increments and they include some of the longest-lived, non-colonial animals on the planet. In addition to their longevity, modern bivalves also show a latitudinal gradient of increasing life span and decreasing growth rate with latitude that might be related to the latitudinal diversity gradient. Is this a recently developed phenomenon or has it characterized much of the group's history? When and how did extreme longevity evolve in the Bivalvia? What insights can the growth increments of fossil bivalves provide about hypotheses for energetics through time? In spite of the relative ease with which the tools of sclerochronology can be applied to these questions, paleobiologists have been slow to adopt sclerochronological approaches. Here, we lay out an argument and the methods for a path forward in paleobiology that uses sclerochronology to answer some of our most pressing questions.

Organismal metabolic rates reflect the interaction of environmental and physiological factors. Thus, calcifying organisms that record growth history can provide insight into both the ancient environments in which they lived and their own physiology and life history. However, interpreting them requires understanding which environmental factors have the greatest influence on growth rate and the extent to which evolutionary history constrains growth rates across lineages. We integrated satellite measurements of sea-surface temperature and chlorophyll-a concentration with a database of growth coefficients, body sizes, and life spans for 692 populations of living marine bivalves in 195 species, set within the context of a new maximum-likelihood phylogeny of bivalves. We find that environmental predictors overall explain only a small proportion of variation in growth coefficient across all species; temperature is a better predictor of growth coefficient than food supply, and growth coefficient is somewhat more variable at higher summer temperatures. Growth coefficients exhibit moderate phylogenetic signal, and taxonomic membership is a stronger predictor of growth coefficient than any environmental predictor, but phylogenetic inertia cannot fully explain the disjunction between our findings and the extensive body of work demonstrating strong environmental control on growth rates within taxa. Accounting for evolutionary history is critical when considering shells as historical archives. The weak relationship between variation in food supply and variation in growth coefficient in our data set is inconsistent with the hypothesis that the increase in mean body size through the Phanerozoic was driven by increasing productivity enabling faster growth rates.

One of the longest-lived, noncolonial animals on the planet today is a bivalve that attains life spans in excess of 500 years and lives in a cold, seasonally food-limited setting. Separating the influence of temperature and food availability on life span in modern settings is difficult, as these two conditions covary. The life spans of fossil animals can provide insights into the role of environment in the evolution of extreme longevity that are not available from studies of modern taxa. We examine bivalves from the unique, nonanalogue, warm and high-latitude setting of Seymour Island, Antarctica, during the greenhouse intervals of the Late Cretaceous and Paleogene. Despite significant sampling limitations, we find that all 11 species examined are both slow growing and long-lived, especially when compared with modern bivalves living in similar temperature settings. While cool temperatures have long been thought to be a key factor in promoting longevity, our findings suggest an important role for caloric restriction brought about by the low and seasonal light regime of the high latitudes. Our life-history data, spanning three different families, emphasize that longevity is in part governed by environmental rather than solely phylogenetic or ecologic factors. Such findings have implications for both modern and ancient latitudinal diversity gradients, as a common correlate of slow growth and long life is delayed reproduction, which limits the potential for evolutionary change. While life spans of modern bivalves are well studied, data on life spans of fossil bivalves are sparse and largely anecdotal. Life histories of organisms from deep time can not only elucidate the controls on life span but also add a new dimension to our understanding of macroevolutionary patterns.

A preliminary survey of benthic invertebrates off central west Florida provides documentation of modern epifaunal communities on a low-gradient carbonate slope. Three large-scale biofacies occur in soft-sediment carbonate environments between 200 and 2,000 m: an Echinoderm biofacies (200–550 m) dominated by a diverse assemblage of echinoderms, gastropods, and decapod crustaceans; a Penaeid shrimp–conical mound biofacies (550–1,200 m) characterized by large bioturbation structures; and a Microbial mat biofacies (1,200–2,000 m) with only rare epifaunal invertebrates. A fourth, hard-substrate biofacies reflects the presence of localized Miocene and Pleistocene hardgrounds in water depths of 200–600 m. This illustrates that hard-substrate biofacies may be laterally correlative with soft-sediment biofacies in a slope setting, thus producing a mosaic of contrasting faunal associations. All four biofacies have low population densities, presumably as a consequence of relatively low surface productivity. All four biofacies also show biogeographic affinity with other faunas at intermediate depths in the Caribbean region. Depth-related faunal transitions on the west Florida slope correlate with substrate and current velocity. Decreasing species diversity and abundance and a biofacies transition from suspension-feeding to deposit-feeding assemblages correlate with increasing depth, a decrease in mean grain size, and an increase in organic content of the sediment. This biofacies model may provide a modern analogue for faunas of ancient low-gradient slopes such as those of Cretaceous “shelf-sea” chalks of northwestern Europe.

Since David Raup's seminal 1976 work, paleontologists have been aware of the relationship between outcrop area and diversity. By incorporating lithologic data derived from Alexander Ronov and coworkers into our own data on areas of mapped outcrops from worldwide geologic maps, we are able to establish a quantitative connection between area of sedimentary rock exposure and diversity from the same general depositional environments. Significant power-law relations are observed at both global and continental scales of consideration. The addition of data on areas of habitable area estimated from paleogeographic maps does not substantially affect these correlations between outcrop area and diversity, indicating that the relation between outcrop area and diversity is primarily a function of sampling and not a common cause such as sea level. We observe a significant diversity-area effect, first noted by Jack Sepkoski in the marine realm. Unlike Sepkoski's, however, our diversity-area effect appears to play a substantial role in influencing diversity through time; a true global diversity signal appears to be contained in the rock record despite the impacts of variable sampling.

Greater outcrop area can serve to increase estimated diversity by increasing both the sample size and the range of habitats and biogeographic provinces sampled. After standardizing for pure sampling intensity by rarefying the number of taxon occurrences, outcrop area continues to explain a substantial portion of global marine diversity. This indicates that coverage, or sampling from multiple habitats and biogeographic provinces, is even more important than sampling intensity. If we remove the effect of outcrop area from our estimate of global biodiversity, we do not observe a net increase in diversity toward the present, lending support to other studies that have not supported a substantial, long-term global increase in biodiversity during the Phanerozoic.

The late Mesozoic through early Cenozoic is an interval of significant biologic turnover and ecologic reorganization within marine assemblages, but the timing and causes of these changes remain poorly understood. Here, we quantify the pattern and timing of shifts in the diversity (richness and evenness) and ecology of local (i.e., sample level) mollusk-dominated assemblages during this critical interval using field-collected and published data sets from the U.S. Gulf Coastal Plain. We test whether the biologic and ecologic patterns observed primarily at the global level during this time are also expressed at the local level, and whether the end-Cretaceous (K/Pg) mass extinction and recovery moderated these trends. To explore whether environment had any effect on these patterns, we examine data from shallow subtidal and offshore settings.

Assemblages from both settings recovered to pre-extinction diversity levels rapidly, in less than 7 million years. Following initial recovery, diversity remained unchanged in both settings. The trajectory of ecological restructuring was distinct for each setting in the wake of the K/Pg extinction. In offshore assemblages, the abundance and number of predatory carnivorous taxa dramatically increased, and surficial sessile suspension feeders were replaced by more active suspension feeders. In contrast, shallow subtidal assemblages did not experience ecological reorganization following the K/Pg extinction. The distinct ecological patterns displayed in each environment follow onshore-offshore patterns of innovation, whereby evolutionary novelties first appear in onshore settings relative to offshore habitats. Increased predation pressure may explain the significant ecological restructuring of offshore assemblages, whereby the explosive radiation of predators drove changes in their prey. Habitat-specific ecological restructuring, and its occurrence solely during the recovery interval, implies that disturbance and incumbency were also key in mediating these ecological changes.

The concept of coordinated stasis, manifest as a pattern of long intervals of concurrent taxonomic and ecologic persistence separated by comparatively abrupt periods of biotic change, has been challenged in recent studies that claim a lack of prolonged persistence of taxa and associations. A key problem has been the difficulty of distinguishing faunal change owing to localized, short-term environmental fluctuation or patchiness from that indicating regionally pervasive, long-term evolutionary or ecological change. Here, we use an extensive database from the Middle Devonian Hamilton Group of the Appalachian Basin to test for taxonomic and ecologic persistence within this ecological-evolutionary subunit, a succession of purported relative stability. Replicate samples collected from many localities and stratigraphic horizons over a wide geographic area allow us to address the effects of small-scale environmental variation and localized faunal patchiness while exploring basin-scale variation in faunal composition within and between the formations of the Hamilton Group.

Observed stratigraphic distributions of fossils are consistent with a scenario in which all taxa are present from bottom to top of the Hamilton Group, and absences result only from sampling failure. Although small-scale variation in faunal composition indeed does occur, there is no more variation among formations than occurs within them. Assemblages from different formations, whether they are defined by taxonomic or ecologic composition, are statistically indistinguishable according to several independent metrics, including ANOSIM and a maximum likelihood estimation that evaluates stratigraphic turnover using Bayesian “Information Criterion.” Simulated data sets indicate that test results are most consistent with species-level extinction of 2.6% per Myr within the Hamilton Group, far lower than the Givetian rate of 11.5% per Myr generic extinction derived from a global database. Such faunal persistence over the ~5.5 Myr encompassed by this unit is consistent with the pattern of coordinated stasis. Earlier studies showing greater amounts of temporal turnover in Hamilton Group faunas are likely influenced by their smaller geographic scale of analysis, suggesting that regional studies done elsewhere may yield similar results.

Seasonal temperature variation at any location is most strongly regulated by the annual cycle of solar insolation, which is controlled by latitude and orbital parameters; other factors are continentality, albedo feedbacks, stratification and advection, and water depth. Seasonal minima, maxima, and range vary with change in mean annual temperature (MAT) across latitudes; seasonal variation in ocean-water temperature is highest in the mid-latitudes. Seasonal temperature range at depth is likely to be less than that at the sea surface. The chemistry of accretionary biogenic carbonate can preserve a record of seasonal temperature variation experienced by an organism over some portion of its ontogeny. Sampling early parts of ontogenies will yield a more complete temperature record because growth is faster (so temporal resolution is higher) and more likely to be continuous throughout the year. Biases due to preferential season of growth can be recognized by characteristic patterns of isotope data when plotted versus distance along the ontogenetic trajectory. Multiple years are needed to reliably characterize seasonal temperature range and inter-annual variability for a stratigraphic horizon in a given region; single-shell studies are less useful in this regard unless they contain very long records. Data from multiple time intervals or regions using the same taxon, sampled the same way, and from the same facies can establish meaningful trends in average temperature and seasonal range. Deep-time studies exploring seasonal temperature extremes and ranges can be coupled with proxies for mean annual temperature to offer a more complete understanding of changing climate conditions through time.