The tailbend, a downward flexure of the vertebral column, is a prominent feature of post-Triassic ichthyosaurs. However, due to the vagaries of preservation and of preparation, its presence cannot be verified in all skeletons. Computed tomography offers a solution to the problem by identifying the wedge-shaped centra that contribute to the tailbend. Leptopterygius tenuirostris appears to have a tailbend, albeit slightly downturned. There is also evidence that its close relative Eurhinosaurus longirostris had a tailbend, too.

Geographic distribution, species duration, and population size of 41 bivalve species which coexisted during the uppermost biozone (Sciponoceras gracile zone) of the Cenomanian were measured. Average species duration and correlation between geographic distribution, species duration, and population size were computed for five groups: total, infaunal, epifaunal, cemented epifaunal, and uncemented epifaunal bivalves.

Average species duration for all bivalves is less than 2 Myr. Infaunal bivalves averaged slightly higher, epifaunal slightly lower. Positive correlation was measured between species duration and geographic distribution for all five groups and statistical significance of the relationships was demonstrated for all but infaunal forms. In general no correlation was found between species duration and population size, and between population size and geographic distribution with the following exceptions: species duration and population size were significantly correlated for infaunal bivalves and uncemented epifaunal bivalves, and population size and geographic distribution were correlated for uncemented bivalves. Because these 41 fossil species coexisted at the same time and were distributed in the same space, differences in the relationships of species duration, geographic distribution and population among the various life habits indicate differences in ecological strategy and the evolutionary consequences of that difference.

Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Laurasian Late Cretaceous ecosystems, have an exceptional fossil record consisting of many species known from partial ontogenetic series, making them an ideal clade with which to conduct life-history studies. Previous research considered the Dinosaur Park Formation (DPF) of Alberta as an attritional, or time-averaged, sample and interpreted size–frequency distribution of long bones collected from the DPF with three size classes to suggest that hadrosaurids from the DPF attained near-asymptotic body size in under 3 years. This conflicted with previously published osteohistological estimates of 6+ years for penecontemporaneous hadrosaurids from the Two Medicine Formation (TMF) of Montana, suggesting either extreme variation in hadrosaurid growth rates or that size–frequency distributions and/or osteohistology and growth modeling inaccurately estimate ontogenetic age.

We tested the validity of the previously proposed size–age relationship of hadrosaurids from the DPF by significantly increasing sample size and combining data from size–frequency distributions and osteohistology across multiple long-bone elements. The newly constructed size–frequency distributions typically reveal four relatively distinct size–frequency peaks that, when integrated with the osteohistological data, aligned with growth marks. The yearling size class was heavily underrepresented in the size–frequency distribution. If not due to preservation, this suggests that either juvenile (<2 years of age) hadrosaurids from the DPF had increased survivorship following an initially high nestling mortality rate or that yearlings were segregated from adults. A growth-curve analysis revealed asymptotic body size was attained in approximately 7 years, which is consistent with hadrosaurids from the TMF. The data suggest size–frequency distributions of attritional samples underestimate age and overestimate growth rates, but when paired with osteohistology can provide unique life-history insights.

More than 500 undistorted, unfragmented pedicle valves of Ambocoelia umbonata (Conrad) (Spiriferida, Brachiopoda) were recovered from each of four levels within a Middle Devonian fossil cluster. The fossil cluster, an ellipsoidal shell accumulation measuring one meter in diameter and 2 cm in thickness, was exhumed from an exposure of the Ludlowville Shale (Hamilton Group) of western New York. Size frequency histograms indicate that the brachiopod experienced very high levels of juvenile mortality, due, probably, to the effects of high bottom turbidity. Sedimentological and paleontological evidence indicates that the cluster represents a sequence of in situ benthic associations.

Size-independent variation was estimated by calculating the eigenvalue of the minor axis on a log-transformed plot of pedicle valve lengths and widths. The eigenvalue technique eliminates the effect of allometrically induced shape changes and is applicable to multicharacter analyses of morphological variability.

Size-independent variability among the larger individuals of Ambocoelia decreases in successively younger cluster populations. The decrease is not correlated with any observed or inferred change in substratum, diversity, equitability or turbidity.

Recovery of trilobite faunas following a mass extinction during the Upper Cambrian Sunwaptan Stage (equal to the “Ptychaspid Biomere”) involved increases in both within-habitat (alpha) diversity and in levels of biofacies differentiation. This pattern parallels, at a lower hierarchical level, the global increase in Phanerozoic marine invertebrate species richness (Sepkoski et al. 1981), which was influenced by changes in provinciality and in alpha diversity. Diversification slowed markedly in subtidal shelf carbonate habitats in the upper half of the stage, but continued in carbonate shelf margin environments, possibly due to higher rates of immigration from off-shelf sites. Trilobite clades appearing early in the stage have low centers of gravity, reflecting high immigration rates during and shortly after an interval of mass extinction. Expansion of shallow-water carbonate deposition in the upper half of the stage was accompanied by the spread of carbonate bank biofacies. Families occurring in these biofacies gradually increased in species richness and tended to have high centers of gravity.

Eurypterids are generally considered to comprise a mixture of active nektonic to nektobenthic predators and benthic scavenger-predators exhibiting a mode of life similar to modern horseshoe crabs. However, two groups of benthic stylonurine eurypterids, the Stylonuroidea and Mycteropoidea, independently evolved modifications to the armature of their anterior appendages that have been considered adaptations toward a sweep-feeding life habit, and it has been suggested the evolution toward sweep-feeding may have permitted stylonurines to capture smaller prey species and may have been critical for the survival of mycteropoids during the Late Devonian mass extinction. There is a linear correlation between the average spacing of feeding structures and prey sizes among extant suspension feeders. Here, we extrapolate this relationship to sweep-feeding eurypterids in order to estimate the range of prey sizes that they could capture and examine prey size in a phylogenetic context to determine what role prey size played in determining survivorship during the Late Devonian. The mycteropoid Cyrtoctenus was the most specialized sweep-feeder, with comblike appendage armature capable of capturing mesoplankton out of suspension, while the majority of stylonurines possess armature corresponding to a prey size range of 1.6–52 mm, suggesting they were suited for capturing small benthic macroinvertebrates such as crustaceans, mollusks, and wormlike organisms. There is no clear phylogenetic signal to prey size distribution and no evolutionary trend toward decreasing prey sizes among Stylonurina. Rather than prey size, species survivorship during the Late Devonian was likely mediated by geographic distribution and ability to capitalize on the expanding freshwater benthos.

The fossil record of mammal dentition provides crucial insight into key ecological and functional transitions throughout mammalian evolutionary history. For cetaceans, both extant clades differ markedly from their stem ancestors; neither retains the differentiated dentition or the tribosphenic molars characteristic of Mammalia. We used quantitative measures of dental complexity across fossil and living cetaceans to identify a trend toward dental simplicity through the Neogene. Both extant cetacean clades depart from the ancestral mammalian condition and concurrently converge upon a reduced and simplified dentition; modern mysticetes all have become entirely edentulous (at birth), and living odontocetes possess teeth as single-rooted, conical pegs. These two parallel trends accompany major shifts in feeding strategy (i.e., filter feeding in mysticetes and echolocation in odontocetes), suggesting that these evolutionary innovations for prey acquisition are enabling factors for the loss of prey processing and subsequent convergence on dental simplification.

With a single complete mandible and 56 mandibular symphyseal fragments of various sizes, the Late Cretaceous Hungarian azhdarchid material has been considered one of the most extensive monospecific pterosaur assemblages in the world. Representing a broad size range, these elements have been thought to demonstrate a developmental series of Bakonydraco galaczi. As such, they were ideal to test whether absolute size and/or morphology reliably indicate relative ontogenetic stages in this pterosaur. Forty-five specimens were selected for multivariate morphometrics and classified into four size classes. After acquiring the morphometric data set, we thin-sectioned eight symphyses representing all size groups and classified them into relative ontogenetic stages based on qualitative microstructural inspection prior to quantitative histological analyses. Microstructural characters suggestive of developmental state were then quantified for intra- and interindividual uni- and multivariate analyses to test the correspondence among the results of qualitative and quantitative analyses. In contrast to our expectations, histological features identified the smallest specimen as an adult and not an early juvenile. The substantial size difference between this specimen and other adults, along with its distinct microanatomical and histological features, implies the presence of at least two pterosaur taxa in this symphysis assemblage. This hypothesis is further supported by multivariate morphometrics, which separate the smallest symphyses from all other specimens that form one continuous group. Although the latter group also shows considerable size variability in corresponding ontogenetic stages, this suggests developmental plasticity rather than the presence of even more taxa, and indicates that symphysis size and morphology are poor indicators of skeletal maturity in these animals. Hence, bone histology is an important independent test of the assessment of ontogenetic stage using size and morphology.

Occupancy statistics in ecology and paleontology are biased upward by the fact that we generally do not have solid data on species that exist but are not found. The magnitude of this bias increases as the average occupancy probability decreases and as the number of sites sampled decreases. A maximum-likelihood method is developed to estimate the underlying distribution of occupancy probabilities of all species based only on the sample of observed species with nonzero occupancy. The method is based on determining the probability that the number of occupied sites will take on any specific value for a given occupancy probability, integrated over the entire distribution of occupancy probabilities. If the shape of the underlying distribution is well modeled, the resulting occupancy estimates circumvent the bias inherent in failing to observe some species and the fact that this bias depends on the number of sites. For occupancy data on marine animal genera drawn from the Paleobiology Database, the underlying distribution is reasonably approximated as a right-truncated log-normal, but the methods developed can be extended to any distribution. Examples are presented to illustrate some observations that are robust and others that need to be revised in light of this bias correction. The method is compared to a recently developed, distribution-free approach to the same problem.

Sexual dimorphism is common in many extant animals, but it is difficult to demonstrate in fossil species. Working with material from the Late Cretaceous of the U.S. Coastal Plain, we herein analyze sexual dimorphism in ostracodes from the superfamily Cytheroidea, a group whose extant members have males that are relatively more elongate than females. We digitized outlines of more than 6000 individual ostracode valves or carapaces, extracted size (area) and shape (length-to-height ratio) information, and used finite mixture models to assess hypotheses of sexual dimorphism. Male and female clusters can be discerned in nearly all populations with sufficient data, resulting in estimates of size and shape dimorphism for 142 populations across 106 species; an additional nine samples are interpreted to consist only of females. Dimorphism patterns varied across taxa, especially for body size: males range from 30% larger to 20% smaller than females. Magnitudes of sexual dimorphism are generally stable within species across time and space; we can demonstrate substantial evolutionary changes in dimorphism in only one species, Haplocytheridea renfroensis. Several lines of evidence indicate that patterns of sexual dimorphism in these ostracodes reflect male investment in reproduction, suggesting that this study system has the potential to capture variation in sexual selection through the fossil record.

The first lower molar length, width, and posterior loop width were analyzed for ten Cosomys primus populations from horizontal deposits ranging in elevation from 3,000 to 3,295 feet. Sedimentation rates based on radiometric dates for two ashes and rates derived in previous studies suggest that these sediments span an interval of time that ranges from 45,000 to 164,000 years ago. Nonmetric data included the presence of enamel pits, prism folds, and crenulations; the number of triangles; and relative dentine tract heights. Coefficients of variation for the metrics were low, 4 to 7, and a one-way ANOVA revealed no significant differences between any of the populations. Changes in the mean phenotype were less than two phenotypic standard deviations and were not correlated with elevation. The presence of enamel pits, prism folds, and crenulations correlated only with less occlusal wear.

Thus, these microtine rodent populations appear to exhibit stasis over a time period of at least 45,000 and possibly as much as 164,000 years, unless the deposits accumulated more rapidly than previously estimated.

Diachronic variation in insectivore remains from a deeply stratified cave deposit in southeastern North America is described. The paleobotanical record for the region is congruent with variation in the soricid (shrew) and talpid (mole) faunal assemblages. Both faunal and floral records reflect considerable climatic change since the full Wisconsinan in this unglaciated portion of eastern North America. Variations in the insectivore assemblages indicate the presence of a more equable climate during the late Pleistocene. Continental climates ushered in at the end of the Pleistocene resulted in a marked decrease in insectivore diversity. Relative frequency changes in taxa throughout the Holocene reflect the mid-Holocene Climatic Optimum.

Where do species that become important players in ecosystems evolve? This simple yet crucial question must be answered if we want to understand how the biosphere is rejuvenated following a crisis. We cannot simply assume that the environments in which we find fossil remains of a given species, or living populations of a species, are the environments in which that species evolved. Take the most obvious example: Fossil human skeletons have been unearthed by the hundreds in North America, but all available evidence points to a human origin in Africa. We can often identify the general geographic origins of species and clades thanks to fossil occurrences and the application of phylogenetic techniques; but can we do likewise for more ecological aspects of the environment? Advances in population biology and in paleobiology now permit us to outline a hypothesis of the circumstances most favorable to the evolution of abundant, widespread, or ecologically powerful species, those with adaptations that are selectively advantageous across many environments, and large short-term and long-term effects in ecosystems.

Many marine benthic metazoans must stabilize themselves upon the seafloor for survival, and as a result their morphologies are controlled in part by local substrate conditions. The Agronomic Revolution (AR), spurred by increasing vertical bioturbation during the Ediacaran–Cambrian transition, permanently altered the nature of shallow marine substrate conditions and led to a major shift in adaptive strategies among benthic metazoans. These ecological and evolutionary changes, known as the Cambrian Substrate Revolution (CSR), are generally understood from observations of benthic metazoan fossils across the Ediacaran/Cambrian boundary, but the timing and geographic extent of this transition are less well known. This analysis attempts to constrain the temporal and spatial pattern of the AR and CSR by performing a global-scale paleoecological analysis of the adaptive strategies of benthic fauna living during the Cambrian. This analysis focused on Burgess Shale-type (BST) faunas because of their exceptional preservation, and was conducted through direct observation of fossil specimens, analysis of data compiled from the Paleobiology Database, and literature review. From these analyses, faunal groups are assigned a metric, the Substrate Adaptability Index (SAI), that relates the overall affinity the fauna demonstrates toward either Proterozoic-style (SAI=0) or Phanerozoic-style (SAI=1) substrate conditions. The results of this analysis demonstrate that most early and middle Cambrian faunas were mixtures of Phanerozoic- and Proterozoic-style adaptive strategists, suggesting that Proterozoic-style substrates were still influential in controlling adaptive strategies in marine environments until at least that time. This is further supported by ichnofabric analysis of many of these localities, where overall bioturbation levels are exceedingly low, indicating a lack of mixed-layer development and the prevalence of firm Proterozoic-style substrates well into the Cambrian.

Large herbivores can act as keystone species that strongly influence their communities. During the Pliocene and Pleistocene, Australia was dominated by a number of large to gigantic marsupial herbivore taxa. Many of these have been understudied quantitatively with regard to their ecology; and identifying the diet of these species will improve our understanding of not only their ecologies, but also of past environments. Recent research has found that cranial morphology among kangaroos and wallabies corresponds with foraging behaviors and mechanical properties of preferred plant tissues. Here we apply shape analysis and computational biomechanics to test the hypothesis: that feeding ecology is associated with craniofacial morphology across a taxonomically broad sample of diprotodont herbivores. Based on our results we predict the diet of an extinct short-faced kangaroo, Simosthenurus occidentalis. We find that biting behaviors are reflected in craniofacial morphology, but that these are more a reflection of the hardest bites required for their lifestyle, rather than diet composition alone. A combination of a very short face, robust musculoskeletal features, and dental arrangements predict that S. occidentalis was a browser, capable of consuming particularly resistant, bulky plant matter. These features were largely conserved among other short-faced kangaroos and may have offset the unpredictable availability of quality forage during the climatically variable Pleistocene epoch, contributing to their prolific diversification during this time.

The taxonomic and ecologic composition of Earth's biota has shifted dramatically through geologic time, with some clades going extinct while others diversified. Here, we derive a metric that quantifies the change in biotic composition due to extinction or origination and show that it equals the product of extinction/origination magnitude and selectivity (variation in magnitude among groups). We also define metrics that describe the extent to which a recovery (1) reinforced or reversed the effects of extinction on biotic composition and (2) changed composition in ways uncorrelated with the extinction. To demonstrate the approach, we analyzed an updated compilation of stratigraphic ranges of marine animal genera. We show that mass extinctions were not more selective than background intervals at the phylum level; rather, they tended to drive greater taxonomic change due to their higher magnitudes. Mass extinctions did not represent a separate class of events with respect to either strength of selectivity or effect. Similar observations apply to origination during recoveries from mass extinctions, and on average, extinction and origination were similarly selective and drove similar amounts of biotic change. Elevated origination during recoveries drove bursts of compositional change that varied considerably in effect. In some cases, origination partially reversed the effects of extinction, returning the biota toward the pre-extinction composition; in others, it reinforced the effects of the extinction, magnifying biotic change. Recoveries were as important as extinction events in shaping the marine biota, and their selectivity deserves systematic study alongside that of extinction.

Previous studies of cranial shape have established a consistent interspecific allometric pattern relating the relative lengths of the face and braincase regions of the skull within multiple families of mammals. In this interspecific allometry, the facial region of the skull is proportionally longer than the braincase in larger species. The regularity and broad taxonomic occurrence of this allometric pattern suggests that it may have an origin near the base of crown Mammalia, or even deeper in the synapsid or amniote forerunners of mammals. To investigate the possible origins of this allometric pattern, we used geometric morphometric techniques to analyze cranial shape in 194 species of nonmammalian synapsids, which constitute a set of successive outgroups to Mammalia. We recovered a much greater diversity of allometric patterns within nonmammalian synapsids than has been observed in mammals, including several instances similar to the mammalian pattern. However, we found no evidence of the mammalian pattern within Therocephalia and nonmammalian Cynodontia, the synapsids most closely related to mammals. This suggests that the mammalian allometric pattern arose somewhere within Mammaliaformes, rather than within nonmammalian synapsids. Further investigation using an ontogenetic series of the anomodont Diictodon feliceps shows that the pattern of interspecific allometry within anomodonts parallels the ontogenetic trajectory of Diictodon. This indicates that in at least some synapsids, allometric patterns associated with ontogeny may provide a “path of least resistance” for interspecific variation, a mechanism that we suggest produces the interspecific allometric pattern observed in mammals.

Lungfishes are known for, and indeed take their name from, their bimodal respiratory abilities. All three extant genera can use their lungs to extract oxygen from the atmosphere, although their reliance upon this capability differs among taxa. Lungs are considered primitive for the Osteichthyes, however the distinctive buccal pump mode of air gulping exhibited by extant lungfishes appears to be a specialization. It is associated with a number of derived skeletal characters (cranial ribs, long parasphenoid stalk, midline gap between palatal tooth plates) that first appeared during the Devonian. These have been described individually, but in no Devonian lungfish has their three-dimensional (3D) spatial relationship been reconstructed and analyzed. Here we present the 3D morphology of Rhinodipterus, a Mid-Late Devonian lungfish from Australia and Europe, based on synchrotron tomography and conventional microtomography scans.

Unlike less crownward contemporaneous lungfishes such as Griphognathus and Chirodipterus, Rhinodipterus has a full set of skeletal buccal pump components that can be directly compared to those of extant lungfishes, suggesting that it made more extensive use of air breathing than other Gogo or Bergisch Gladbach genera. This is interesting in relation to the environmental context as Gogo and Bergisch Gladbach are both marine, contrasting with the frequently hypoxic tropical to subtropical fresh water environments inhabited by modern lungfishes. The evolution of buccal pump-supported lung ventilation was evidently not necessarily associated with a transition to non-marine habitats.

When asked how he might sum up his long life, an aged Benjamin Franklin replied that he had, at least, been useful to his fellows. Paleobiology is still young, as our journal has just completed 20 years and now begins this season of its majority, while the vibrancy of our subject certainly proclaims a vigorous incipiency. But academic generations are short, a mere 5 years or so, and Paleobiology has therefore enjoyed adequate time to demand judgment by accomplished results, not simple promise. When, fifteen years ago (6, 1, 80), I wrote a similar piece to celebrate the completion of this journal's first five volumes, I spoke in my title about “the promise of paleobiology”; now I must refer to “the task of paleobiology” and delineate what has been accomplished, and how much we have yet to do.

Numerous methods have been developed to estimate the position of a mass extinction boundary while accounting for the incompleteness of the fossil record. Here we describe the point estimator and confidence interval for the extinction that are optimal under the assumption of uniform preservation and recovery potential, and independence among taxa. First, one should pool the data from all taxa into one combined “supersample.” Next, one can then apply methods proposed by Strauss and Sadler (1989) for a single taxon. This gives the optimal point estimator in the sense that it has the smallest variance among all possible unbiased estimators. The corresponding confidence interval is optimal in the sense that it has the shortest average width among all possible intervals that are invariant to measurement scale. These optimality properties hold even among methods that have not yet been discovered. Using simulations, we show that the optimal estimators substantially improve upon the performance of other existing methods. Because the assumptions of uniform recovery and independence among taxa are strong ones, it is important to assess to what extent they are satisfied by the data. We demonstrate the use of probability plots for this purpose. Finally, we use simulations to explore the sensitivity of the optimal point estimator and confidence interval to nonuniformity and lack of independence, and we compare their performance under these conditions with existing methods. We find that nonuniformity strongly biases the point estimators for all methods studied, inflates their standard errors, and degrades the coverage probabilities of confidence intervals. Lack of independence has less effect on the accuracy of point estimates as long as recovery potential is uniform, but it, too, inflates the standard errors and degrades confidence interval coverage probabilities.