Changes in the physical environment are major drivers of evolutionary change, either through direct effects on the distribution and abundance of species or more subtle shifts in the outcome of biological interactions. To investigate this phenomenon, we built a fossil data set of drilling gastropod predation on bivalve prey for the last 11 Myr to determine how the regional collapse in Caribbean upwelling and planktonic productivity affected predator–prey interactions. Contrary to theoretical expectations, predation increased nearly twofold after productivity declined, while the ratio of drilling predators to prey remained unchanged. This increase reflects a gradual, several-fold increase in the extent of shallow-water coral reefs and seagrass meadows in response to the drop in productivity that extended over several million years. Drilling predation is uniformly higher in biogenic habitats than in soft sediments. Thus, changes in predation intensity were driven by a shift in dominant habitats rather than a direct effect of decreased productivity. Most previous analyses of predation through time have not accounted for variations in environmental conditions, raising questions about the patterns observed. More fundamentally, however, the consequences of large-scale environmental perturbations may not be instantaneous, especially when changes in habitat and other aspects of local environmental conditions cause cascading series of effects.

The repeated return of tetrapods to aquatic life provides some of the best-known examples of convergent evolution. One comparison that has received relatively little focus is that of mosasaurids (a group of Late Cretaceous squamates) and archaic cetaceans (the ancestors of modern whales and dolphins), both of which show high levels of craniodental disparity, similar initial trends in locomotory evolution, and global distributions. Here we investigate convergence in skull ecomorphology during the initial aquatic radiations of these groups. A series of functionally informative ratios were calculated from 38 species, with ordination techniques used to reconstruct patterns of functional ecomorphospace occupation. The earliest fully aquatic members of each clade occupied different regions of ecomorphospace, with basilosaurids and early russellosaurines exhibiting marked differences in cranial functional morphology. Subsequent ecomorphological trajectories notably diverge: mosasaurids radiated across ecomorphospace with no clear pattern and numerous reversals, whereas cetaceans notably evolved toward shallower, more elongated snouts, perhaps as an adaptation for capturing smaller prey. Incomplete convergence between the two groups is present among megapredatory and longirostrine forms, suggesting stronger selection on cranial function in these two ecomorphologies. Our study highlights both the similarities and divergences in craniodental evolutionary trajectories between archaic cetaceans and mosasaurids, with convergences transcending their deeply divergent phylogenetic affinities.

An enigma of deep-sea biodiversity research is that the abyss with its low productivity and densities appears to have a biodiversity similar to that of shallower depths. This conceptualization of similarity is based mainly on per-sample estimates (point diversity, within-habitat, or α-diversity). Here, we use a measure of between-sample within-community diversity (β1H) to examine benthic foraminiferal diversity between 333 stations within 49 communties from New Zealand, the South Atlantic, the Gulf of Mexico, the Norwegian Sea, and the Arctic. The communities are grouped into two depth categories: 200–1500 m and >1500 m. β1H diversity exhibits no evidence of regional differences. Instead, higher values at shallower depths are observed worldwide. At depths of >1500 m the average β1H is zero, indicating stasis or no biodiversity gradient. The difference in β1H-diversity explains why, despite species richness often being greater per sample at deeper depths, the total number of species is greater at shallower depths. The greater number of communities and higher rate of evolution resulting in shorter species durations at shallower depths is also consistent with higher β1H values.

Taxonomic membership frequencies exhibit distributions in which groups with few numbers of subtaxa are much more common in a clade than those with more subtaxa. Here, a “broken plate” model is developed to describe such taxonomic memberships; some higher taxonomic group (the plate) is randomly subdivided into intermediate taxonomic units (plate fragments), whose sizes are dependent on the number of taxonomic subunits that they each contain. Theoretical distributions of membership frequencies produced by this model yield a superior fit to data from both modern and fossil groups, as illustrated by classifications for primarily fossil brachiopods and entirely modern mammals. The nature of these distributions is consistent with the contention that Linnaean membership frequencies result from the random partitioning of taxonomic/morphologic space. Moreover, numbers of taxa contained within hierarchically equivalent groups are unrelated, as are membership numbers at taxonomically higher and lower levels of consideration. Agreement between observed taxonomic memberships and those anticipated from the random partitioning of diversity as described by the “broken plate” model bears directly on a number of fundamental questions including the significance of extreme polytypy and inferred causes of adaptive radiation within many taxonomic groups.

Crinoids were relatively unaffected by the end-Devonian Hangenberg mass extinction event. Major clades of Devonian durophagous fishes suffered significant extinctions, however, and the dominant surviving clades were biting or nipping predators. In part as a response to the Hangenberg event, early Mississippian crinoids underwent an adaptive radiation, while fish clades with a shell-crushing durophagous strategy diversified. Durophagous predators are inferred to have been more effective predators on camerate crinoids; and it is hypothesized, following the predictions of escalation, that through the early Mississippian, camerate crinoids evolved more effective anti-predatory strategies in response. We test this hypothesis of escalation by examining the changes in spinosity and plate convexity among camerate crinoids throughout this interval. A new method was formulated to test for an increase in convexity of the tegmen plates. Traits in Agaricocrinus, Aorocrinus, and Dorycrinus (Family Coelocrinidae) were tested for congruence to the escalation hypothesis, and results were mixed. Convexity of tegmen plates in Agaricocrinus, spine length/calyx diameter in Aorocrinus, calyx size in Aorocrinus, central spine length in Dorycrinus, and spine width in Dorycrinus did not have size increase trends supporting escalation. Rather than an increase in convexity, the variance of convexity in Agaricocrinus tegmen plates narrowed, which could reflect an optimum. Alternatively, morphological change consistent with the escalation hypothesis occurred in calyx size of Agaricocrinus and in lateral spine length and calyx size in Dorycrinus. Furthermore, central and lateral spine length, parameters of the spine width, and size trends support escalation when Aorocrinus and Dorycrinus are treated as a lineage. Thus, inferred escalation acted on traits differently within a single lineage and was relevant for both speciation and the diversification of a new genus.

Gaps in the fossil record can be identified in compilations of stratigraphic ranges of lower taxa classified within higher taxa. Statistical properties of such gaps suggest the following conclusions. First, the random sampling model most commonly used to correct for incompleteness in the record is inconsistent with the data. Second, the decline in taxonomic extinction rate during the Phanerozoic is not just an artifact of an increase in the completeness of the record.

Historians have debated whether pathways and events from the past to the present are influenced largely by contingency, the dependence of outcomes on particular prior conditions, or whether there is long-term emergent directional change. Previous arguments for predictability in evolutionary history relied on the high frequency of convergence, but the repeated evolution of widely favored adaptations need not imply long-term directionality. Using evidence from the fossil record and arguments concerning the metabolic evolution of organisms, I show here that power (total energy taken up and expended per unit time) has increased stepwise over time at ecosystem-level and global scales thanks to the ratchet-like, cumulative effects of competition and cooperation and to the disproportionate influence of powerful top competitors and opportunistic species on emergent ecosystem properties and processes. The history of life therefore exhibits emergent directionality at large ecosystem-wide scales toward greater power.

Stratigraphic range data are used to derive time scales on which taxonomic survivorship curves for genera and families are as nearly as possible independent of their times of origin. These time scales correct for temporal variations in overall extinction rates caused by major extinctions and the pull of the Recent. Survivorship curves for genera and families on their respective time scales are well fit by Pareto distributions differing only in their scale parameter.

Northern Appalachian Basin deposits and associated fossils have served as exemplars for ecological-evolutionary investigations, and as the reference interval for the concept of coordinated stasis. Here, we examine faunal and environmental changes within the uppermost Hamilton and lowermost Genesee Groups of the late Middle Devonian succession of New York State. Dramatic diversity loss, faunal migrations, and ecological restructuring recognized in these strata have been used previously to define the end of the Hamilton ecological-evolutionary subunit, and, furthermore, these strata and corresponding faunal changes represent the type region for the global Taghanic Biocrisis. We present and analyze a new, high-resolution data set of post-Taghanic Genesee fossil assemblages, in which we recognize 11 biofacies corresponding to an onshore-offshore (depth) gradient. The Genesee Fauna shows an unexpectedly high taxonomic similarity to nearshore biofacies of the pre-Taghanic Hamilton Fauna, related to the persistence of siliciclastic-dominated nearshore settings through the Taghanic Biocrisis, whereas the onset of anoxic/dysoxic conditions typified offshore portions of the environmental gradient. The “Nearshore Refugium Model” of Erwin offers a possible explanation for the persistence of taxa through the biocrisis in nearshore settings. This constriction was followed by subsequent expansion of these residual taxa to offshore environments in relatively similar associations, as increased Acadian orogenic activity and resultant delta progradation increased habitable space offshore by decreasing the extent of deeper-water, oxygen-poor settings. Although taxonomic similarity was high between the Hamilton and Genesee Faunas, biofacies structure differed primarily because of tectonically driven physical transformations to the basin and associated biotic turnover. Nevertheless, the combination of high taxonomic persistence of Hamilton nearshore taxa and the introduction of relatively few new taxa in the Genesee Fauna resulted in a taxonomic holdover that was much higher than observed in the original formulation of coordinated stasis.

Recent progresses in our knowledge of mouse odontogenesis have enhanced rodent tooth morphology as a model for Evo-Devo studies. Deciphering the connection between macroevolution and microevolution, however, especially in the case of mammalian teeth, requires examples to illustrate how morphological differences among species, or higher taxa, can stem from population-level processes. In this paper we use paleontological material to study intraspecific variation of tooth morphology in the late Miocene species Progonomys clauzoni, over a short span of geological time in a restricted area. Progonomys is of particular interest as a stem genus of all murine rodents (Old World rats and mice). We use morphometrical and statistical methods to illustrate how change in the amplitude in variation at the population level through geological time is associated with the emergence of new characters. Some of these new characters, including functional ones, become fixed in parallel in distinct murine lineages. Nine million years ago, Progonomys clauzoni displayed variational properties of the developmental system shared by the Murinae, which can also explain some singular tooth characteristics that now are scattered among the diverse lineages. Further morphometric studies, however, are necessary to explain how the variety of cusp patterns observed in Progonomys clauzoni can be explained by developmental properties.

The trophic role assumed by an animal taxon in a community is best understood when the structure of the community is viewed in historical perspective (Whittaker 1977). In many environments it is not physical and behavioral limitations which guide a group's evolutionary potential, but the opportunity to exploit available adaptive zones, which, because of the nature of the fauna, were open (Hecht 1975, p. 248). [The term adaptive zone is used as defined by Van Valen (1971, p. 421) … “the niche of any taxon, especially a supraspecific one.” Van Valen distinguished adaptive zones from the “ways of life” of taxa. Adaptive zones are part of the environment and exist independently of taxa that exploit them.] It is often assumed or implied that a systematic group can, in its adaptive radiation, fill most of the available niches within a particular adaptive zone (Hecht 1975, p. 247). This type of “taxocene” analysis (Whittaker 1972, p. 218; Levandowsky and White 1977, p. 133), which promotes these views, has obvious shortcomings. It is now evident that mammals, for example, were unable to equally exploit and partition all available adaptive zones on every continent. The failure of carnivorous mammals to do this to a complete degree in Australia (see Hecht 1975, p. 247) and in South America (see Marshall 1977, p. 709) has some historical basis in the development of the stratification of the vertebrate fauna.

Approximately 50,000–11,000 years ago many species around the world became extinct or were extirpated at a continental scale. The causes of the late Pleistocene extinctions have been extensively debated and continue to be poorly understood. Several extinction models have been proposed, including two nutritionally based extinction models: the coevolutionary disequilibrium and mosaic-nutrient models. These models draw upon the individualistic response of plant species to climate change to present a plausible scenario in which nutritional stress is considered one of the primary causes for the late Pleistocene extinctions.

In this study, we tested predictions of the coevolutionary disequilibrium and mosaic-nutrient extinction models through the study of dental wear and enamel hypoplasia of Equus and Bison from various North American localities. The analysis of the dental wear (microwear and mesowear) of the samples yielded results that are consistent with predictions established for the coevolutionary disequilibrium model, but not for the mosaic-nutrient model. These ungulate species show statistically different dental wear patterns (suggesting dietary resource partitioning) during preglacial and full-glacial time intervals, but not during the postglacial in accordance with predictions of the coevolutionary disequilibrium model. In addition to changes in diet, these ungulates, specifically the equid species, show increased levels of enamel hypoplasia during the postglacial, indicating higher levels of systemic stress, a result that is consistent with the models tested and with other climate-based extinction models. The extent to which the increase in systemic stress was detrimental to equid populations remains to be further investigated, but suggests that environmental changes during the late Pleistocene significantly impacted North American equids.

Estimating biodiversity and its variations through geologic time is a notoriously difficult task, due to several taphonomic and methodological effects that make the reconstructed signal potentially distinct from the unknown, original one. Through a simulation approach, we examine the effect of a major, surprisingly still understudied, source of potential disturbance: the effect of time discretization through biochronological construction, which generates spurious coexistences of taxa within discrete time intervals (i.e., biozones), and thus potentially makes continuous- and discrete-time biodiversity curves very different. Focusing on the taxonomic-richness dimension of biodiversity (including estimates of origination and extinction rates), our approach relies on generation of random continuous-time richness curves, which are then time-discretized to estimate the noise generated by this manipulation. A broad spectrum of data-set parameters (including average taxon longevity and biozone duration, total number of taxa, and simulated time interval) is evaluated through sensitivity analysis. We show that the deteriorating effect of time discretization on the richness signal depends highly on such parameters, most particularly on average biozone duration and taxonomic longevity because of their direct relationship with the number of false coexistences generated by time discretization. With several worst-case but realistic parameter combinations (e.g., when relatively short-lived taxa are analyzed in a long-ranging biozone framework), the original and time-discretized richness curves can ultimately show a very weak to zero correlation, making these two time series independent. Based on these simulation results, we propose a simple algorithm allowing the back-transformation of a discrete-time taxonomic-richness data set, as customarily constructed by paleontologists, into a continuous-time data set. We show that the reconstructed richness curve obtained this way fits the original signal much more closely, even when the parameter combination of the original data set is particularly adverse to an effective time-discretized reconstruction.

Constraint is a universal feature of morphological evolution. The vertebral column of synapsids (mammals and their close relatives) is a classic example of this phenotypic restriction, with greatly reduced variation in the number of vertebrae compared with the sauropsid lineage. Synapsids generally possess only three sacral vertebrae, which articulate with the ilium and play a key role in locomotion. Dicynodont anomodonts are the exception to this rule, possessing seven or more sacral vertebrae while reaching a range of body sizes rivaled among synapsids only by therian mammals. Here we explore the evolution of this unusual sacral morphology in dicynodonts by (1) hypothesizing homologies of the additional sacral vertebrae, (2) using ancestral state reconstruction and phylogenetic regressions (e.g., logistic regression, Poisson regression) to track the coevolution of sacral count and body size, and (3) proposing mechanisms by which additional sacral vertebrae were incorporated during dicynodont evolution. We find that sacral vertebral morphology covaries with sacral count in consistent ways across dicynodonts, implying that sacra with a given number of vertebrae are composed of homologous elements. There is a correlation between increased sacral count and larger body size, especially at the shift from four to five sacrals near the origin of Bidentalia. Based on position, morphology, and the consistent number of presacral vertebrae among dicynodonts, we hypothesize that the additional sacrals anterior to the plesiomorphic three are duplications of the first sacral, and that a single caudosacral was incorporated by a shift in the identity of the anteriormost caudal vertebra. Although changes in sacral count appear to be correlated with shifts in body size in dicynodonts, the evolution of general morphological conservativism in the synapsid sacrum remains to be further explored.

Models for the areal distribution of marine animals may be formulated in such a way that their validity can be tested statistically. Five such models are described here: in each case the distributions of the individual taxa are made functions of depth of water, oceanicity, and substrate type.

The simplest model proposes that benthic species are substrate-dependent: it can be verified by a series of χ2 tests. Benthic species can also be seriated along an environmental gradient: this second model can be tested by using a Monte Carlo simulation procedure. The third model proposes that planktic species are to some degree stratified in the water column, and hence that their remains are distributed additively in the underlying sediments: tests on modern and sub-fossil data sets indicate that realistic versions of this model are probably valueless for most geological work. The final models are extensions of the additive model: they are used here to demonstrate the importance of selecting an appropriate model and, in particular, of knowing the mode of life of each species before the model is selected.

Intraspecific aggression, or agonism, is a widespread intrasexual selective behavior important to understanding animal behavioral ecology and reproductive systems. Such behavior can be studied either by direct observation or inferred from wound/scar frequency in extant species but is difficult to document in extinct taxa, limiting understanding of its evolution. Among extant archosaurs, crocodylians display extensive intrasexual aggression, whereas birds show extreme visual/vocal intersexual display. The evolutionary origin of this behavioral divergence, and pattern in non-avian dinosaurs, is unknown. Here we document the morphology, frequency, and ontogeny of intraspecific facial bite lesions (324 lesions) in a large sample of tyrannosaurids (202 specimens, 528 elements) to infer patterns of intraspecific aggression in non-avian theropods. Facial scars are consistent in position and orientation across tyrannosaurid species, suggesting bites were inflicted due to repeated/postured behavior. Facial scars are absent in young tyrannosaurids, first appear in immature animals (~50% adult skull length), are present in ~60% of the adult-sized specimens, and show aggressor:victim size isometry. The ontogenetic distribution of bite scars suggests agonistic behavior is associated with the onset of sexual maturity, and scar presence in approximately half the specimens may relate to a sexual pattern. Considered in a phylogenetic context, intraspecific bite marks are consistent and widely distributed in fossil and extant crocodyliforms and non-maniraptoriform theropods, suggesting a potential plesiomorphic behavior in archosaurs. Their absence in maniraptoriform theropods, including birds, may reflect a transition from boney cranial ornamentation and crocodylian-like intrasexual aggression to avian-like intersexual display with the evolution of pennaceous feathers.

Death assemblages that occupy the upper tens of centimeters of sediment in shallow-marine settings are often subject to extensive mixing, thereby limiting their usefulness in assessing environmentally mediated compositional changes through time in the local biota. Here, we provide evidence that dense, Thalassia-rich seagrass beds preserve a stratigraphic record of biotic variation because their dense root–rhizome mats inhibit mixing. We sampled benthic mollusk assemblages at seven localities in Thalassia-rich beds around St. Croix, USVI, collecting three separate sediment intervals of ~13 cm each to a total depth of ~40 cm below the sediment–water interface, and found evidence that sedimentary intervals preserved compositional stratigraphy. Further, some localities displayed systematic, directional changes down-core. An examination of interval-to-interval changes in composition revealed that compositional variation was unique from locality to locality rather than reflecting coordinated, island-wide transitions. In general, however, relative abundances of epifaunal gastropods and small lucinid bivalves tended to decrease with depth below the sediment–water interface. Quantitative comparisons of life-to-death assemblages from each successive sedimentary interval demonstrated that the shallowest death assemblages were typically more similar to the life assemblages than were deeper assemblages, suggesting that deeper intervals provide records of earlier community states.