In North America, hipparion horses consisted of four distinct genera, Hipparion, Neohipparion, Nannippus, and Cormohipparion, known from the medial Miocene to the late Pliocene, ca. 15.5–2 ma ago. The speciation mode is determined for 10 of the 16 valid hipparion species; 50% (5) seem to have resulted from anagenesis and 50% (5) from cladogenesis. Roughly contemporaneous species of the four hipparion genera seem to proceed through similar evolutionary stages of cranial and dental characters. The mean hipparion species longevity is 3.3 ma (N = 16); the mean generic longevity is 8.4 ma (N = 4). Only two species become extinct in less than 2 ma; extinction occurs at a rate of ca. 470 μmacarthurs for species with durations between 2 and 5 ma. For selected dental characters, mean evolutionary rates (for the 16 species) were between .08 darwins (d) and .03 d with crown height evolving most rapidly. Despite the commonly held notion that fossil horses evolved rapidly, these data suggest normal or average (horotelic) evolution for hipparions, which is not surprising because they were already established in the grazing adaptive zone during their history.

A late Miocene population of Neohipparion cf. leptode from the Love Bone Bed of Florida was found to have 13 discrete annual age classes based on eruption sequences and tooth wear. The large sample size of individuals (minimum number = 229) allowed the estimation of standard population parameters, including age-specific mortality rates. The average age of death for a newborn was 3.5 years, but those individuals that survived the 64% juvenile mortality occurring during the first two years lived an average of 8.0 years. The potential longevity of this species of Neohipparion had not increased beyond that of known samples of Merychippus, despite increased crown height. The paleoenvironment of the collecting site was a wooded grassland savanna with annual wet and dry seasons. The Neohipparion population migrated into the area during the dry season but left in the wet season to give birth away from the collecting site.

The foraging paradigm of trace fossil theory has historically accorded random behavior to non-food-limited deposit-feeders and non-random behavior to food-limited feeders. A series of randomness measures derived from empirical modeling, simulation modeling, stochastic modeling and probability theory applied to foraging patterns observed in deep-sea bottom photographs from the Arctic and Antarctic yielded a behavioral continuum of increasing non-randomness. A linear regression of trace positions along the continuum to bathymetric data did not substantiate the optimal foraging efficiency-depth dependence model of trace fossil theory, except that all traces exhibited a greater optimization than that of simulated random foraging. It is hypothesized that optimization as evidenced by non-random foraging strategies represents maximization of the cost/benefit ratio of resource exploitation to risk of predation and that individual foraging patterns reflect an exploration response to the morphometry of a patchily distributed food resource. Differential predation and competition may account for the co-occurrence of random and non-random strategies within the same bathymetric zone.

A rate of extinction must specify both the amount of taxonomic change and the duration of that change. A consistent means of temporal measurement must be used to establish the duration. Terms used to describe the extinction, like catastrophic and gradual, should be quantitatively defined in relation to this framework.

Resolving rates of extinction in the stratigraphic record is limited by (1) the precision of chronostratigraphic correlations between individual sections and (2) the temporal completeness of litho- and biostratigraphic records in those sections. In this study, temporal completeness is estimated in eight of the best-known fluvial and pelagic sections spanning the K-T boundary. Temporal standardization is provided by correlations to the geomagnetic polarity time scale. A review of the literature documents lithologic criteria that might be used to infer the presence of a hiatus of some length at the biostratigraphically recognized K-T boundary. Rather than rejecting the presence of such gaps, this study argues that a hiatus of some length does exist and tries to estimate both a maximum limit for its duration and the probability that it represents a given duration below that limit.

Pelagic sections are probably more complete than fluvial ones. The section at Caravaca, Spain, can be expected to be the most complete, probably preserving sediment during each 10,000-yr interval of Chron 29R. The section in the San Juan Basin, New Mexico, is expected to be the most complete fluvial section, probably preserving sediment during each 100,000-yr interval of 29R.

Results indicate that our best-known marine and terrestrial sections spanning the K-T boundary should not be expected to document biologically catastrophic rates of extinction. A sufficiently precise means of global correlation that is independent of biostratigraphy and causal hypotheses is still unavailable, and it is very unlikely that the sections examined are complete at the 100-yr or biologically catastrophic level of precision. So, although catastrophic amounts of extinction might have occurred during the K-T transition, it seems unlikely that we can distinguish episodes of extinction lasting 100 yr or less from episodes lasting as long as 100,000 yr. Consequently, acceptance of catastrophic hypotheses based on these stratigraphic records seems improbably optimistic at this time.

Estimating speciation and extinction rates is essential for understanding past and present biodiversity, but is challenging given the incompleteness of the rock and fossil records. Interest in this topic has led to a divergent suite of independent methods—paleontological estimates based on sampled stratigraphic ranges and phylogenetic estimates based on the observed branching times in a given phylogeny of living species. The fossilized birth–death (FBD) process is a model that explicitly recognizes that the branching events in a phylogenetic tree and sampled fossils were generated by the same underlying diversification process. A crucial advantage of this model is that it incorporates the possibility that some species may never be sampled. Here, we present an FBD model that estimates tree-wide diversification rates from stratigraphic range data when the underlying phylogeny of the fossil taxa may be unknown. The model can be applied when only occurrence data for taxonomically identified fossils are available, but still accounts for the incomplete phylogenetic structure of the data. We tested this new model using simulations and focused on how inferences are impacted by incomplete fossil recovery. We compared our approach with a phylogenetic model that does not incorporate incomplete species sampling and to three fossil-based alternatives for estimating diversification rates, including the widely implemented boundary-crosser and three-timer methods. The results of our simulations demonstrate that estimates under the FBD model are robust and more accurate than the alternative methods, particularly when fossil data are sparse, as the FBD model incorporates incomplete species sampling explicitly.

Sauropodomorpha included the largest known terrestrial vertebrates and was the first dinosaur clade to achieve a global distribution. This success is associated with their early adoption of herbivory, and sauropod gigantism has been hypothesized to be a specialization for bulk feeding and obligate high-fiber herbivory. Here, we apply a combination of biomechanical character analysis and comparative phylogenetic methods with the aim of quantifying the evolutionary mechanics of the sauropodomorph feeding apparatus. We test for the role of convergence to common feeding function and divergence toward functional optima across sauropodomorph evolution, quantify the rate of evolution for functional characters, and test for coincident evolutionary rate shifts in craniodental functional characters and body mass. Results identify a functional shift toward increased cranial robustness, increased bite force, and the onset of static occlusion at the base of the Sauropoda, consistent with a shift toward bulk feeding. Trends toward similarity in functional characters are observed in Diplodocoidea and Titanosauriformes. However, diplodocids and titanosaurs retain significant craniodental functional differences, and evidence for convergent adoption of a common “adaptive zone” between them is weak. Modeling of craniodental character and body-mass evolution demonstrates that these functional shifts were not correlated with evolutionary rate shifts. Instead, a significant correlation between body mass and characters related to bite force and cranial robustness suggests a correlated-progression evolutionary mode, with positive-feedback loops between body mass and dietary specializations fueling sauropod gigantism.

A range of specialized burrowing sculptures evolved convergently in at least one lingulid and in several obolid and lingulasmatid genera. The sculptures in Lingula punctata (Ordovician) indicate a burrowing process with the pedicle trailing behind the shell, similar to that of Recent Lingulidae. In the Obolidae and Lingulasmatidae, the orientation of the sculptures indicates a burrowing process with the pedicle oriented downward in the sediment. A burrowing mechanism in which the valves alternated in functioning as anchors, without active participation of the pedicle, is proposed for these forms. Infaunal life habits for several obolid genera are further indicated by the shell morphology and by the distribution of repaired shell damage.

The burrowing Obolidae are likely to have been adapted to relatively high-energy environments, which required frequent reburrowing in addition to escape (upward) burrowing. The increase in bioturbation in these environments in the Middle Paleozoic may have contributed to their extinction. The Lingulidae, instead, being mostly adapted to extreme environmental conditions, could survive in marginal niches which were not intensively exploited by actively burrowing organisms.

The mammal fauna of the Willwood Formation, central Bighorn Basin, Wyoming, is ideal for paleoecological analysis because it is extensive, well studied, and continuously distributed over sediments representing the first 3 Myr of the early Eocene. The geology of the Bighorn Basin is also well known, providing a precise temporal framework and climatic context for the Willwood mammals. Previous analysis identified three “biohorizons,” based on simple counts of the first and last appearances of species. This study uses species diversity and appearance rates calculated from more extensive collections to approximate the ecological dynamic of the ancient fauna and assess whether the biohorizons were significant turnover events related to recently described climatic variation. Diversity and appearance data collected for this project are extensively corrected for uneven sampling, which varies by two orders of magnitude. Observed, standardized appearance and diversity estimates are subsequently compared with predicted background frequencies to identify significant variation. Important coincident shifts in the biotic parameters demonstrate that ecological change was concentrated in two discrete intervals ≤300 Kyr each that correspond with two of the original biohorizons. The intervals coincide with the onset and reversal of an episode of climate cooling identified directly from Bighorn Basin floras and sediments. Ecological changes inferred from the diversity and turnover patterns at and following the two biohorizons suggest short- and long-term faunal response to shifts in mean annual temperature on the order of 5–8°C.

In Baculites, a straight shelled ammonite, the constructional limits on shell shape resulting from the limited strength of nacre in tension are circumvented by a system of vaults in the phragmocone. Vaults bridge between regions of the phragmocone supported by the complex ammonite septal suture, and maintain the shell wall in compression when hydrostatic load induces bending moments. To determine how these vaults interact in the phragmocone to resist hydrostatic loading, measurements were made on a suite of Baculites specimens. In Baculites there is a statistically significant inverse relationship between circumferential curvature (radius of curvature) of the vaulted shell surface and the size of vaults spanning between sutural elements supporting the phragmocone. The inverse relationship between radius of curvature and the sizes of spans in this system of vaults results in the generation of comparable reactive forces at the ends of the vault spans where adjacent vaults interact. The equivalence of these reactive forces prevents the lateral displacement of the vault ends. Consequently, compressive stresses from adjacent vaults are superimposed on, and reduce, the tensional stress component of bending. Limiting tensile stress is of utmost importance in a lightweight shell composed of a brittle material such as nacre, which is strong in compression but weak in tension.

Baculites is particularly appropriate for this study because its straight shell is curved only in the circumferential direction, thus simplifying the problem. However, sutural patterns in coiled ammonites appear to be similarly constrained to produce vaults in the phragmocone which vary inversely in curvature and span size.

Major shifts in ecological dominance are one of the most conspicuous but poorly understood features of the fossil record. Here we examine one of the most prominent such shifts, the Ordovician shift from trilobite to brachiopod dominance of benthic ecosystems. Using an integrated database of high-resolution paleoecological samples and body size data, we show that while the average local richness and relative abundance of trilobites declined significantly through the Ordovician, the estimated standing biomass of trilobites, and by implication the amount of energy that they used, remained relatively invariant. This is attributable to an increase in the average body size of trilobite species in our data set, and especially to the widespread occurrence of the exceptionally large Middle-Late Ordovician trilobite genus Isotelus. Brachiopods increase in both mean body size and relative abundance throughout the Ordovician, so that estimates of brachiopod biomass and energetic use increase substantially between the Early and Late Ordovician. Although the data set includes a range of depositional environments, similar trends are observed in both shallow subtidal and deep subtidal settings. These results suggest that diversification of the Paleozoic Fauna did not come at the energetic expense of the Cambrian Fauna. The declining relative abundance of trilobites may reflect a combination of numerical dilution and the necessary energetic trade-offs between body size and abundance.

Immunologically reactive protein substances were extracted from bone samples of an American mastodon (Mammut americanum), 10,200 yr old by radiocarbon dating, and from muscle samples of three woolly mammoths (Mammuthus primigenius), 10,000, 40,000 and 53,000 yr old, respectively. The mastodon samples contained significant quantities of the amino acids hydroxylysine and hydroxyproline, both of which are usually found in collagens and not in albumins. Using these products and other comparable extracts, as well as sera and purified collagens from modern elephants and other living mammals, as test antigens, immunological comparisons were carried out with the following antisera: rabbit anti-mastodon bone; chicken anti-mammoth muscle; chicken anti-elephant muscle; rabbit anti-elephant albumin and rabbit anti-elephant collagen, as well as with rabbit antisera to purified albumins and collagens of other mammals. For the first time, mastodon bone was found to have elephant-like proteins, which elicited antibodies that reacted strongly with collagen and serum proteins of extant elephants. Mammoth muscle strongly reacted with anti-elephant collagen and anti-elephant albumin, but the concentrations of the recoverable mammoth collagen and albumin decreased with increasing chronological age of the mammoth specimens. Nevertheless, in the immunological comparisons, the mammoth was closer to Asian (Elephas maximus) and African (Loxodonta africana) elephants than to the mastodon; in turn, the mastodon was closer to these elephantid species than to mammals outside the order Proboscidea.

Analysis of dental wear and damage is becoming an increasingly important tool in unraveling the trophic ecology of a wide range of vertebrates, and when applied to fossils it can provide evidence of both diet and feeding kinematics that is independent of morphological analysis. Conodonts have the best fossil record among vertebrates and their skeletal elements are known to exhibit surface wear and damage generated in vivo as a consequence of their function as teeth. We report the results of the first systematic survey and analysis of the frequency and extent of this wear and damage in conodonts (based on P1 elements from a range of Carboniferous genera). This has revealed that wear and damage are remarkably common, present in all conodont elements sampled. Multivariate analysis reveals that patterns of wear and damage differ significantly among different conodont taxa, and exploratory ANOVA and linear discriminant analyses show that wear and damage differ according to the position of taxa in an onshore-offshore gradient, and whether they are likely to have had a benthic or pelagic mode of life. The incidence of denticle tip spalling in particular is higher in more-offshore environments and in taxa likely to have had a pelagic mode of life. Aspects of the data also reflect the occlusal kinematics of the elements, providing a means of testing hypotheses of element function. Our results have wide-ranging implications for unlocking the fossil record of conodonts, by, for example, furnishing direct evidence of the diet-mediated processes that may have driven observed patterns of evolutionary change, and reducing the confounding effects of depth segregation when using conodonts in isotope-based paleotemperature studies.

Many studies have examined temporal changes in insect feeding on angiosperm leaves, but none have considered variability within a single stratigraphic level. If spatial variability within a level is high, a single sample will not adequately represent the level and may either mask true temporal changes or create spurious ones. In order to measure the spatial variability in fossil insect feeding damage, I collected 12 replicate samples from two laterally extensive carbonaceous shale beds (55.2 and 52.6 Ma) from the early Eocene of the Bighorn Basin, Wyoming. Over 2800 fossil angiosperm leaves were scored for presence or absence of 50 insect damage morphotypes. Damage frequency, diversity, and composition were computed for both the bulk flora and individual plant species in each sample, and variation within a bed was compared with differences between the two beds. Differences in diversity and composition between beds were significantly greater than variations within a bed, and intra-bed variation was primarily due to differing floral composition. Damage frequency within a bed, however, was more variable than diversity. Damage diversity and composition reflect the number of insect species present, whereas damage frequency also depends on the number of insects present, which may be much more variable over small distances.

Stable carbon and oxygen isotope ratios were measured for carbonate in samples of hadrosaurid tooth enamel and dentine, and gar scale ganoine and dentine from five geologically “contemporaneous“ (two-million-year resolution) and geographically distant late Campanian formations (Two Medicine, Dinosaur Park, Judith River, Kaiparowits, and Fruitland) in the Western Interior Basin. In all cases, isotopic offsets were observed between enamel and dentine from the same teeth, with dentine being characterized by higher and more variable carbon and oxygen isotope ratios. Isotopic offsets were also observed between gar ganoine and hadrosaur enamel in all sites analyzed. Both of these observations indicate that diagenetic overprinting of enamel isotope ratios did not entirely obfuscate primary signals. Decreases in carbon and oxygen isotope ratios were observed in hadrosaur enamel from east to west, and overlap in isotope ratios occurred only between two of the sampled sites (Dinosaur Park and Judith River Formations).

The lack of isotopic overlap for enamel among localities could be due to diagenetic resetting of isotope ratios such that they reflect local groundwater effects rather than primary biogenic inputs. However, the large range in carbon isotope ratios, the consistent taxonomic offsets for enamel/ganoine data, and comparisons of enamel-dentine data from the same teeth all suggest that diagenesis is not the lone driver of the signal. In the absence of major alteration, the mostly likely explanation for the isotopic patterns observed is that hadrosaurids from the targeted formations were eating plants and drinking waters with distinct isotopic ratios. One implication of this reconstruction is that hadrosaurids in the Late Cretaceous of the Western Interior did not migrate to an extent that would obscure local isotopic signatures.

We show that the decline in the extinction rate during the Phanerozoic can be accurately described by a logarithmic fit to the cumulative total extinction. This implies that extinction intensity is falling off approximately as the reciprocal of time. We demonstrate that this observation alone is sufficient to explain the existence of the proposed power-law forms in the distribution of the sizes of extinction events and in the power spectrum of Phanerozoic extinction, results that previously have been explained by appealing to self-organized critical theories of evolutionary dynamics.

The temporal dimension of fossil sequences provides a critical component to the study of intraspecific dynamics and species formation. Here I report on the branching and subsequent morphological evolution of two gastropods (Melanopsis fossilis and M. vindobonensis) from the Late Miocene of the Pannonian basin in eastern and central Europe. Although morphological divergence between species is rapid, intermediates between the two species co-occur with typical individuals for approximately 1 m.y. and then disappear. The long-term persistence of intermediates followed by their ultimate disappearance is a pattern that, to my knowledge, has not been previously observed.

Distinguishing genetic from ecophenotypic influences on shell form in freshwater prosobranchs is difficult. Nevertheless, consideration of the temporal, geographic, lithologic, and paleoecologic patterns of this sequence suggests that the morphologic differences between M. fossilis and M. vindobonensis had some genetic basis. Whether these forms were initially morphs of a single species or two species with some hybridization between them is impossible to determine. In either case, the morphological changes that resulted in M. vindobonensis were rapid, but the attainment of complete isolation between M. fossilis and M. vindobonensis apparently did not occur until approximately 1 m.y. later.

Thalattosuchians are crocodylomorphs mainly known from marine strata of Early Jurassic to Early Cretaceous age. They represent the earliest crocodylomorph radiation to an aquatic habitat and their evolutionary history offers very few records from freshwater settings. Here, we report several exquisitely preserved thalattosuchian skulls attributed to a derived teleosaurid from a pedogenic horizon located at the base of a fluvial series of alternating silts and sandstones of the Phu Kradung Formation (Upper Jurassic) of northeastern Thailand. Using laser ablation multicollector inductively coupled mass spectrometry (MC-ICP-MS) on tooth enamel and dentine, we measured isotopic ratios of strontium (87Sr/86Sr) to test the habitat of these teleosaurids. In addition, Sr concentrations of the dental tissues were estimated from the calibrated signal intensities of the Sr isotope measurements. The dataset includes bioapatite (teeth or scales) of eight terrestrial and five aquatic vertebrates. Theropods exhibit lower Sr concentrations both in enamel and dentine compared to others groups, a pattern in accordance with the calcium biopurification process, which predicts that Sr concentrations in the body of vertebrates decrease up the trophic chain. It also excludes the possibility that diagenesis has completely overprinted the Sr isotope compositions of the fossil assemblage, which exhibits a homogeneous 87Sr/86Sr signature above the Late Jurassic seawater value. Values for teleosaurid teeth are in the range of other values for vertebrates in the continental assemblage and imply that these crocodylomorphs did not migrate between freshwater and marine habitats at least in the time constraint of the mineralizing tooth. This result represents the first demonstration that a population of teleosaurids was established for a prolonged time in a freshwater environment. Whether the ability of teleosaurids to inhabit freshwater habitats is a secondary adaptation or whether it is plesiomorphic and inherited from freshwater ancestors is discussed.

Despite more than a century of interest, body-mass estimation in the fossil record remains contentious, particularly when estimating the body mass of taxa outside the size scope of living animals. One estimation approach uses humeral and femoral (stylopodial) circumferences collected from extant (living) terrestrial vertebrates to infer the body masses of extinct tetrapods through scaling models. When applied to very large extinct taxa, extant-based scaling approaches incur obvious methodological extrapolations leading some to suggest that they may overestimate the body masses of large terrestrial vertebrates. Here, I test the implicit assumption of such assertions: that a quadratic model provides a better fit to the combined humeral and femoral circumferences-to-body mass relationship. I then examine the extrapolation potential of these models through a series of subsetting exercises in which lower body-mass sets are used to estimate larger sets. Model fitting recovered greater support for the original linear model, and a nonsignificant second-degree term indicates that the quadratic relationship is statistically linear. Nevertheless, some statistical support was obtained for the quadratic model, and application of the quadratic model to a series of dinosaurs provides lower mass estimates at larger sizes that are more consistent with recent estimates using a minimum convex-hull (MCH) approach. Given this consistency, a quadratic model may be preferred at this time. Still, caution is advised; extrapolations of quadratic functions are unpredictable compared with linear functions. Further research testing the MCH approach (e.g., the use of a universal upscaling factor) may shed light on the linear versus quadratic nature of the relationship between the combined femoral and humeral circumferences and body mass.

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 Cretaceous–Paleogene (K/Pg) extinction appears to have been geographically heterogeneous for some organismal groups. Southern Hemisphere K/Pg palynological records have shown lower extinction and faster recovery than in the Northern Hemisphere, but no comparable, well-constrained Southern Hemisphere macrofloras spanning this interval had been available. Here, macrofloral turnover patterns are addressed for the first time in the Southern Hemisphere, using more than 3500 dicot leaves from the latest Cretaceous (Maastrichtian) and the earliest Paleocene (Danian) of Argentine Patagonia. A maximum ca. 90% macrofloral extinction and ca. 45% drop in rarefied species richness is estimated across the K/Pg, consistent with substantial species-level extinction and previously observed extirpation of host-specialized leaf mines. However, prior palynological and taxonomic studies indicate low turnover of higher taxa and persistence of general floral composition in the same sections. High species extinction, decreased species richness, and homogeneous Danian macrofloras across time and facies resemble patterns often observed in North America, but there are several notable differences. When compared with boundary-spanning macrofloras at similar absolute paleolatitudes (ca. 50°S or 50°N) from the Williston Basin (WB) in the Dakotas, both Maastrichtian and Danian Patagonian species richnesses are higher, extending a history of elevated South American diversity into the Maastrichtian. Despite high species turnover, our analyses also reveal continuity and expansion of leaf morphospace, including an increase in lobed and toothed species unlike the Danian WB. Thus, both Patagonian and WB K/Pg macrofloras support a significant extinction event, but they may also reflect geographically heterogeneous diversity, extinction, and recovery patterns warranting future study.